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ACN IN ASL)S

OF THE

SOUTH AFRICAN MUSEUM

VOL CME Oe I

a al oder lite eign) :

7 i) MR nf

fm v oh Te Le %

Au * D Re ay he

ve yi \ % nT v

ANNALS

OF THE

BoOUTH AFRICAN MUSEUM

VOLUME. XLII

OMe Ms a yu an

eee ate ‘o

ail E il

RTE

} cS

Qi>

his 4

e. Ul ee

tat

PRINTED FOR THE ;

TRUSTEES OF THE SOUTH AFRICAN MUSEUM

: AND FOR THE

GEOLOGICAL SURVEY OF SOUTH AFRICA

"1953 — 1956

BY THE NATIONAL COMMERCIAL PRINTERS LIMITED,

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TRUSTEES OF THE SOUTH AFRICAN MUSEUM.

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Dr. M. R. Drennan, M.B., Ch.B., F.R.S.A.Afr.-

CouNCILLOR H. E. GEARING.

Cale SIBBEeTr,, ].P:

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SENATOR D. H. van ZyYL.

SCIENTIFIC STAFF OF THE SOUTH AFRICAN MUSEUM.

KEPPEL HARCOURT BARNARD, M.A., D.Sc., F.L.S., Director.

ALBERT JOHN HeEsSsE, B.Sc., Ph.D., Assistant in charge of the Entomological

Department. .

Lizuwe DrrK Boonstra, D.Sc., Assistant in charge of the Palaeontological:

| Department.

Miss E. MARGARET SHAW, B.A., Assistant in charge of the Ethnological and.

- Numismatic Departments.

Miss G. Joyce Lewis, Ph.D., Assistant in charge of the Botanical Depart-—

ment. |

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

LIST OF CONTRIBUTORS.

L. D. BOoOoNSTRA.

HH.

Note on some Rhynchosaurian remains from Tanganyika Territory

Report on a collection of Fossil Reptilian bones from Tanganyika

Territory

Suggested clarification or the axehonne satis of the South

African Titanosuchians : 7

The Gorgonopsians: Aelurognathus don and ‘Hipnesaiee

boonstrai reconstructed ;

The cranial morphology and taxonomy a the Tapinocemlakel

genus Struthiocephalus 2 I at OES Sa

The Lower Jaw Articulatory region in some Pristerognathid

Therocephalians 2 ie

The Pristerognathid ne Occohalians: fro the Tapinoce naan

zone in the South African Museum wae

The cranial structure of the Titanosuchian Anieccnuee

The smallest Titanosuchid yet recovered from the Karroo

Paranteosaurus gen. nov.: a Titanosuchian Reptile

Struthiocephalellus: a new Deinocephalian

The Girdles and Limbs of the South African Debceplaln

B. S. COOKE.

Some Fossil Mammals in the South African Museum collections

. BE, EWER & R. SINGER,

Fossil Carnivora from MHopefield

. H. HAUGHTON.

Phosphatic-Glauconitic Deposits off the West Coast of South Africa

. N. KEEN 6 R. SINGER.

Further Fossil Suidae from Hopefield

. K. MILLER @ W. M,. FURNISH.

Tertiary Nautiloids dredged near Cape of Good Hope

. SINGER 5 E, N. KEEN.

Fossil Suiformes from Hopefield

PAGE

108

149

157

180

185

Lor

335

329

350

327

169

NEW GENERIC NAMES PROPOSED IN THIS VOLUME.

meosatsanius (iherocephalidace) Boonstra «900.0; . 0. 2k ke 62

Micranteosaurus (Anteosauridae) Boonstra . . . . . . Cy 150

Neomegacyclops nom.nov. for Megacyclops Broom 1931 preocc., Boonstra 7

Pearanteosaurus (Anteosatiridae) Boonstra 50. a ee 57

Parascapanodon (Titanosuchidae) Boonstra .. SEN AONE We full tecvedt as ge

Pristerognathoides (Therocephalidae) Boonstra . . . . . . 94

Pristerosaurus (Therocephalidae) Boonstra . : : ; BN SO

Reeuemestes| (imerocephatdae)) Boonstra Woo ee 8

Struthiocephalellus (Struthiocephalidae) Boonstra . . . . . . 184

Semupazonsen) (Lherapsidac) (Boonstra: ake 9

Therioides (Therocephalidae) Boonstra PIN a Pu atic AR Te NR oat! A CR

DATE, OF JSSUE,.OF THE: PARTS.

Part 1, April, 1953.

Part 2, January, 1954.

Part 3, January, 1955.

Part 4, March, 1956.

PLATE,

I-V.

Vii

VEE VEL

TX.

X-XIV.

DY RV.

XVII.

XVIII.

XIX.

XX-XXIV.

XXV.

XXXVI.

XXVIT-XXIX.

XXX, XXXII.

AXXITI.

XXXII.

XXXIV.

XXXV.

LIST. OF , PLATES.

Anteosaurus abeli.

Anteosaurus minor.

Anteosaurus vorsterl.

Anteosaurus abeli.

Aelurognathus microdon.

Hipposaurus boonstrai.

Struthiocephalus.

Micranteosaurus parvus. |

Homoioceras bainii.

Mesochoerus lategani.

Struthiocephalus whaitsi.

Aturia lotzi,

Crocuta spelaea.

Lycaon pictus magnus, and Crocuta spelaea.

Mellivora capensis.

Mesochoerus lategani, and paiceae.

Mesochoerus, juvenile.

Tapinochoerus meadowsi.

Aelurognathus

Aepyceros

Agnosaurus

Alcelaphus

Alopecognathus .

Anteosaurus

Anthodon

Antidorcas

Aonyx :

Archaeosuchus

Arctocephalus

Aturia 5 oe

Aulacephalodon

Avenantia

23,

Balaena

Balaenoptera

Bubalus

Cadulus

Canis

Carcharias

Carcharodon

Cephalophus

Chlamys

Connochaetes

Conus

Crocuta

Cuspidaria

Cynariognathus

Damaliscus

Dentalium

Diceros

Dicynodon

Dinartamus .

Dinocynodon

Dinopolus

Dinosphageus

Dinosuchus

Eccasaurus

Enobius

Equus

Fusus

LIsh OF

PAGE.

165

BANS

26, 108,

327, 329,

766.

24,

334

GENERA.

Gazella

Giraffa

Glanosuchus

Herpestes

Hippopotamus

Hipposaurus

Hippotragus

Histiophorus

Homoioceras

Hyaena

Hylochoerus

Isurus

Jonkeria

Kannemeyeria

Keratocephalus

Lamiasaurus

Leptailurus

Loxodonta

Lycaenops

Lycaon

Lystrosaurus

Maraisaurus

Megacyclops

Megaptera

Mellivora

Mesochoerus

Mesoplodon

Metridiochoerus

Micranteosaurus

Mormosaurus

Moschognathus

Moschoides

Moschops

Moschosaurus

Natica

Notochoerus

Neomegacyclops

Notohipparion

23;

164,

Cars

29,

62,

170,

353)

156,

177,

PAGE.

164

166

335

170

166

333

161

336

353

333

285

212

340

165

Omochoerus

Orca

Ostrea

Papio vee:

Paranteosaurus

Parapapio

Parascapanodon

Pecten

Pedetes

Pelea

Peloroceras

Pelosuchus

Phacochoerus

Phocosaurus

Pleurotoma

Pnigalion

Potamochoerus .

Pristerognathoides .

Pristerognathus

Pristerosaurus

Procavia

Propalaeonyx

Ptomalestes

Pusionella

Pycnodonta

157;

164,

Raphicerus

Redunca :

Rhachiocephalus

Scapanodon

PAGE.

269

Scaphonyx

Scullya

Scymnosaurus

Staganosuchus

Stenaulorhynchus

Strepsiceros bl hae

Struthiocephalellus

Struthiocephaloides

Struthiocephalus

Struthionops

Suricata

54>

Sus

Sylvicapra

Syncerus .

Tangagorgon

Tapinocephalus

Tapinochoerus

Taurocephalus

Taurotragus

Terebratula

Terebratulina

Thecodontosaurus

Therioides

AOS Higa

Titanognathus

Titanosuchus

Tragelaphus

Trirachodon

Xenophora

Ziphius

PAGE.

334

333

ANNALS

OF THE

SOUTH AFRICAN MUSEUM

VOLUME XLII

Descriptions of the Palaeontological Material collected by the South African

Museum and the Geological Survey of South Africa.

PART I, containing :—

A Note on some Rhynchosaurian Remains from Tanganyika Territory.

By L. D. Boonstra, D. Sc. (With two text-figures.)

: A Report on a collection of Fossil Reptilian Bones from Tanganyika

Territory. By L. D. Boonsrra, D.Sc. (With seven text-figures.)

A suggested Clarification of the Taxonomic Status of the South African

Titanosuchians. By L. D. Boonstra, D.Sc. (With Plates I-IX.)

The Gorgonopsians, Aelurognathus microdon and Hipposaurus boonstrat,

reconstructed. By L. D. Boonsrra, D.Sc. (With Plates X-XVI.)

The Cranial Morphology and Taxonomy of the Tapinocephalid genus

Struthiocephalus. By L. D. Boonstra, D.Sc. (With Plate XVII and

_ ‘six text-figures.) _

The Lower Jaw Articulatory Region in some Pristerognathid Thero-

cephalians. By L. D. Boonstra, D.Sc. (With four text-figures.)

ISSUED APRIL 1953 PRICE 12s. 6d.

PRINTED FOR THE

TRUSTEES OF THE SOUTH AFRICAN MUSEUM

| AND THE ,

GEOLOGICAL SURVEY OF SOUTH AFRICA

BY THE RUSTICA PRESS LIMITED, COURT ROAD, WYNBERG, CAPE

PONINIACES

OF THE

SOUTH AFRICAN MUSEUM

VOLUME XLII

1. A Note on some Rhynchosaurian Remains from Tanganyika Territory. By LizEUWE

D. BoonstraA, D.Sc.

(With 2 text-figures.)

From Mr. G. M. Stockley I have received three tins containing fossil remains

from near Msamara, in the Tunduru district of the Southern Province of

Tanganyika Territory, for determination. This material ($7662) consists of

weathered fragments of skull, limb-bones and vertebrae. The largest skull

fragment consists of part of the left upper jaw. The limb-bones include a

proximal third of a right femur, a weathered distal end of a femur, a bit of

humerus, scapula and tibia. The vertebral material consists of the weathered

centra of dorsal and caudal vertebrae showing few distinctive features.

SKULL (fig. 1)

This weathered and rolled fragment consists of part of the left jugal, maxilla

and palatine. On the lateral surface two nutritive foramina pierce the maxilla.

The dentigerous border of the maxilla is convex antero-posteriorly. ‘The

ventral surface of the maxilla is studded with a large number of worn peg-like

teeth varying in diameter from 2 to 4 mm. These teeth are somewhat indefi-

nitely arranged, but there does appear to be some indication of an alignment

in four rows. The teeth on the inner or lingual margin of the maxilla are

aligned in a fairly definite row. On the outer or labial margin and anteriorly

the teeth are larger than those situated medio-posteriorly. The width of the

dentigerous surface is wider posteriorly than anteriorly. Medial to the maxillary

teeth there lies a groove filled with matrix, and the maxillo-palatine suture

would lie along this groove. Lingual to this groove there lies a row of teeth on

the labial border of the palatine, with at least three more indefinite rows of

teeth lingually. As is the case with the maxilla the width of the dentigerous

surface is posteriorly wider than anteriorly. As preserved, the length of the

maxillary dentigerous surface is 100 mm., the width anteriorly 7 mm. and

posteriorly 20 mm.

The nature of the dentigerous plate of the maxillo-palatine indicates that we

have here a fragment of a Rhynchosaurian skull showing particular affinities

Won. XLII. PART I.

SUL 23 1952

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

to Scaphonyx australis described by Von Huene from South America, Hyperoda-

pedon gordom described by Huxley from Scotland, and Hyperodapedon huxleyi

described by Lydekker from India. The length of the dentigerous surface in

this specimen from Tanganyika is, as preserved, greater than in these three

known forms.

Fic. 1.—Maxillo-palatine complex of Scaphonyx

stockleyi. a, lateral view (x 4). 6, ventral view (x 4).

S.A.M. 11704.

In H. gordoni the teeth are placed in rows much more definitely than in the

Tanganyika specimen; in the former there are two rows on the maxilla whereas

in the latter there are possibly four rows; and the width of the dentigerous

surface of the maxilla is also smaller in H. gordoni.

In H. huxleyi the teeth are also in much more definite rows, and the number

of rows on the maxilla exceed those on the palatine. In the Indian beast the

crowns form triangular pyramids whereas in the Tanganyika beast the crowns

are bluntly conical.

In Scaphonyx australis the teeth are of very similar shape and have very nearly

the same arrangement. But, whereas the anterior maxillary teeth are larger

than the posterior ones in the Tanganyika specimen, the reverse is the case in the

South American specimen.

NOTE ON SOME RHYNCHOSAURIAN REMAINS 3

From the available evidence, admittedly scanty, it would thus appear that

we have here a type distinct from all the hitherto known Rhynchosaurians, but

apparently approaching fairly closely the skull fragment described by Von Huene

from South America and by him referred to as Scaphonyx australis. I propose

that the Tanganyika beast be included in Smith Woodward’s genus Scaphonyx,

but to be distinguished from the hitherto known species and to be known under

the name Scaphonyx stockleyi n.sp.

Type.—Skull fragment in the South African Museum, 5.A.M. 11704.

Femur (fig. 2)

The proximal third of a weathered right femur is preserved. The post-axial

corner with the external trochanter is missing. Noteworthy features of this

fragment are: the caput femoris is bent sharply dorsally; the internal tro-

Fig. 2.—Proximal end of femur of Scaphonyx africanus. a, ventral view (xX #).

b, dorsal view (x4). c¢, pre-axial view (x4). d, articular surface (x }). S.A.M

11705.

chanter forms a prominent bulbous structure and is abruptly demarcated from

the caput, and is situated some distance distally from the proximal articulatory

surface; the inter-trochanteric fossa is comparatively shallow, but, because of

the curvature of the bone and the presence of a low ridge, is clearly separated

from the more distal ventral surface of the bone; the proximal articulatory

surface is wide dorso-ventrally; the external trochanter was probably not

strongly developed and the ilio-femoralis flange not extensive.

This femur is about half the size of that (S344) described by Haughton under

the name Stenaulorhynchus stockleyi but is obviously of a related type. Apart from

the difference in size it differs in the shape of the caput; the internal trochanter

is situated further distally; is more bulbous and is abruptly separated from the

caput without a connecting neck. This femur, although also only half the size,

shows closer affinities to that of Scaphonyx fischeri and Scaphonyx australis as

described and figured by Von Huene. But here also the nature of the internal

trochanter is strikingly different, and the femur cannot be considered as being

that of either of these two South American species.

4. ANNALS OF THE SOUTH AFRICAN MUSEUM

If Von Huene is correct in assigning his skull fragment and femoral fragment

to the same species (Scaphonyx australis), we have a dentigerous plate of 80 mm.

associated with a femur proximally 80 mm. wide. In the Tanganyika material

we have a dentigerous plate of at least 100 mm., and we would thus expect

that the associated femur would have a width proximally of about 100 mm.

Actually the femur under consideration cannot proximally be of a greater width

than 50 mm. The femur can thus only be of a younger beast or of a smaller

species. ‘The ossification of the femoral fragment does not appear to be that of

an immature half-grown animal, and is thus in all probability that of a species

of half the size of Scaphonyx australis. For this species I propose the name

Scaphonyx africanus.

Type.—Proximal third of a right femur, S.A.M. 11705.

The discovery by Mr. Stockley of these definite Rhynchosaurian remains in

Tanganyika, together with the Rhynchosaurian limb-bones of Stenaulorynchus

stockleyi previously described by Haughton from Nyjalila, will enable the strati-

graphists to attempt a closer correlation of these beds with those that have

yielded the species of Hyperodapedon, Scaphonyx, and the other closely related

genera. The discovery of better-preserved material may make it necessary to

remove these species from the genus Scaphonyx to a new genus. Until such time

I do not think it necessary to create a new genus for these African forms.

2. A Report on a collection of Fossil Reptilian Bones from Tanganyika Territory. By

LizEuWE D. Boonstra, D.Sc.

(With 7 text-figures)

INTRODUCTION

Recently I received from Mr. G. M. Stockley eight cases of fossil reptilian

bones collected from the Ruhuhu Coalfield region. This collection was obtained

from three distinct localities. ‘Those labelled $346 come from the Matomondo

area, $559 from the Ngaka-Kingori Hill area, and $340 from the Nijalila-

Mkongeleko area. The geological horizon of the first two localities is given as

the Lower “Bone Bed’ of the Songea Series, and the third as the Upper ‘Bone

Bed’ of the same series.

Stockley states that the bones ‘were picked up from the surface and thus are

much weathered’. It is clear that all the fragments collected from a locality

were given the same number, and each number undoubtedly includes fragments

derived from a number of different skeletons. I have no means, other than the

state of preservation, of determining the degree of association of the various

fragments. For the greater part the fragments thus have to be considered

individually. No evidence is given by the collector that the fragments bearing

the same number were obtained or derived from the same horizon within the

bed. On the contrary, it is evident on morphological grounds that from the

Upper ‘Bone Bed’ there are at least four groups of specimens and that these

were derived from four different levels within the bed. It should be stressed

that in future collecting attention should be paid to the relative levels within

the bed from which each specimen is collected.

A. MATERIAL FROM THE LOWER ‘BONE BED’

1. Anomodont Skull Fragments

Under the label $346 there are a small number of weathered pieces which

include fragments identifiable as parts of a fairly large anomodont skull with

a narrow parietal crest. ‘This feature apparently excludes reference to the

Aulacephalodon group of genera, and these fragments must thus represent a large

species of the genus Dicynodon.

Under the label $559 there are a large number of weathered pieces—cranial

as well as post-cranial. Of the recognizable skull fragments the following

identifications have been possible, mainly on pieces of the intertemporal region

bearing the pineal foramen and the structures immediately surrounding it.

(a) Inone piece of the intertemporal region of a fairly large dicynodont there

is a medium-sized oval foramen situated at the posterior end of a depression

formed by the preparietal. The parietal crest is of medium width and height.

The distinguishing feature in this fragment is the nature of the preparietal.

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

From the anterior border of the foramen the sides of the preparietal diverge in

anterior direction. The preparietal thus meets the frontal in a long transverse

suture. The whole of the bone lies in a depression. Direct comparison proved

the identity of this specimen with a form from the Luangwa Valley of Northern

Rhodesia which I described under the name Dicynodon roberti. This specimen

now bears the South African Museum Cat. No. 11706. _

(5) In another frag-

ment (fig. 1) the pineal

foramen pierces a large

oval boss which is almost

entirely formed by the

preparietal. The pos-

terior third of the boss is

formed by the parietal,

but this bone only

stretches two-thirds up

the posterior surface of the

boss and is thus excluded

from the border of the

pineal foramen. The

: preparietal completely

a surrounds the foramen,

which is of medium size

and nearly circular. The

foramen is directed

anteriorly, and the plane

in which it lies makes an

angle of 45° with the

plane of the dorsal sur-

face of the skull. Pos-

teriorly, the surface of

the boss, here formed

mainly by the parietal,

a, dorsal view of pineal seston (4). , lateral view (lef of OP V2°8i Maia

pineal boss (x 4). The parietal, which has

only a small exposure, is

then covered by the two post-orbitals which meet on the median line. Anteriorly,

the boss is abruptly marked off from the surface of the frontals by a semi-

circular groove. A near approach to this condition of the pineal region, in the

more fully known Karoo species, is shown by some of the species of the genus

Platycyclops, where the foramen is also completely surrounded by the preparietal

raised in the form of a boss. A nearer approach still is shown in the species of

the genus Megacyclops, where the parietal participates in and forms the posterior

part of the boss. In the present specimen the shape of the boss differs remarkably

REPORT ON COLLECTION OF FOSSIL REPTILIAN BONES 7

from that of M. whaitsi of the South African Karoo and of M. rugosus described

by Haughton from Tanganyika, but agrees in other features, particularly in

being so definitely anteriorly directed and in the manner in which the parietal

participates in its formation. It differs from M. usilzensis of Von Huene in the

slope of the boss and in the fact that the parietal is excluded from the border of

the foramen. It is thus evident that we have here a new species of Megacyclops*

and this fragment can be designated as the type specimen of Neomegacyclops

cyclops n. sp. This specimen now bears the South African Museum Cat. No.

i 70'7-

nature of the pineal region is strongly reminiscent of that known in the genus

Rachiocephalus of the Endothiodon zone of the South African Karoo and may be

provisionally referred to that genus. This specimen now bears the South African

Museum Cat. No. 11708.

(d) Another identifiable fragment is of the central part of a massive occiput.

The basisphenoidal tubera are exceptionally massive and are, furthermore,

peculiar in that they lie practically horizontally instead of approaching the

vertical, so that the foramen ovale is directed nearly completely posteriorly,

whereas the more usual direction is nearly completely ventrally. This condition

is not known to exist in the genera Megacyclops and Rachiocephalus. This fragment

is thus not derived from either of the large skulls that yielded the pineal regions

mentioned above. In none of the larger Anomodonts examined by me nor in

the published descriptions have I seen basisphenoidal tubera of a similar

nature. This fragment thus indicates the existence in Tanganyika of a new type

of the larger Anomodonts, of which one hopes that a more complete skull may

soon be found and be designated as the type of this new species. This specimen

now bears the South African Museum Cat. No. 11709.

2. Anomodont Limb-bones

Included in the collection are a number of weathered distal and proximal

ends of some large humeri, femora and ulnae. ‘The former two are of about the

same size and nature as the humerus and femur described by Haughton and

referred to as Eocyclops? and Megacyclops respectively. In addition to these there

are some proximal and distal ends of humeri and femora of Anomodonts of

smaller size which could be associated with a skull of the size of a form like

Dicynodon huenei—a species peculiar to Tanganyika. These specimens now bear

the South African Museum Cat. Nos. 11710 and 11742.

3. Pareiasaurian Vertebrae

Two medium-sized vertebrae are preserved in a weathered condition. These

are typically Pareiasaurian, and are smaller than those of the larger genera

which predominate in the Tapinocephalus zone of the Union. They are also

* I find that the name Megacyclops which Broom (1931) proposed for this anomodont genus is

preoccupied for a crustacean (Kiefer, 1927). I propose that this anomodont genus be known

under the new name Neomegacyclops nom. nov.

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

smaller than, and do not show the peculiarly narrow elongated dorsal spine so

characteristic of, the vertebrae described by Haughton from locality Big, west

of Kingori. These vertebrae prove the presence in Tanganyika of Pareiasaurs

probably closely related to the medium-sized types of the South African Karoo.

These now bear the South African Museum Cat. No. 11743.

4. Therapsid Cranial Material

Among the mass of weathered fragments from the Ngaka-Kingori Hill area

there are two incomplete snouts much weathered, and one fairly complete but

Fic. 2.—Tangagorgon tenuirostris n. g. et n. sp. S.A.M. Cat. No. 11744.

Lateral view of skull (x 3).

somewhat distorted and weathered skull. The more complete skull is of a

Gorgonopsian and the two snouts appear to be Therocephalian.

(a) The more complete specimen (figs. 2 and 3) has the outer surface con-

siderably eroded, so that it is difficult to determine the sutures. The skull is

of medium size. The chief measurements are:

Maximum@length ora i ee

Basioccipital—Premaxilla 72 te

Premaxilla—-Pineal’foramen’ 2...) ie

Premaxalla““Orbit <0" en es

Widthacross'squamosals’)) "525 1

Imterorbital width" oe ee a ee AOS; Se

Intertemporal“width® (o's ea ee

Widthacross' canines) iS ee ce el a se

Height from Mx. edge—median suture . . . . 50 ,,

Height of mentum ene nner iO.

Length of molar, series: Left,(4 teeth) )\...4..9 ee

Right (@ teeth); 4.5) 29) eee

Lengthyotuncisomseresien 5.) sane

Dental formula: Right, i5, co+1, m3.

Left, 15,.cr--1, m4.

25 »

REPORT ON COLLECTION OF FOSSIL REPTILIAN BONES e)

The snout is long, high and narrow. Anterior to the orbits the cross-section

is squarish. The orbits are situated in the posterior half of the skull and are

laterally directed. The temporal openings are of medium size, oval, and directed

as much dorsally as laterally. The pineal foramen is small and posteriorly

situated. The frontals apparently form

only a small part of the orbital border.

The snout is higher than wide. ‘The

maxilla is deep. The interparietal is only

slightly inclined downwards from the

general dorsal surface and is more a bone

of the dorsal surface than of the occiput.

From the posterior edge of the inter-

parietal the occiput descends practically

vertically. The nature of the rest of the

occiput is obscured by the presence of the

proatlantal arches, atlas and axis. The

teeth are poorly preserved. On the left

there are preserved five medium-sized

incisors, a fairly weak canine with the tip

of a replacing? canine just emerging, and

the roots of four small molars. On the

right side the last of the five incisors is

small (a replacing tooth ?), the canine has

been lost and a replacing? one is emer-

ging; the crowns of three small molars

are preserved, number 3 having fallen out.

The characters enumerated above do

not occur together in any known Gorgo-

nopsian, and I therefore designate this

specimen as the type of a new genus and

species— T angagorgon tenuirostris. ‘This type

specimen now bears the South African Fic. 3.—Tangagorgon tenuirostris n. g. et

‘

’

eof

Museum Cat. iy : n. sp. S.A.M. Cat. No. 11744. Dorsal

Sutth (Cee NG: 744 . view of skull with proatlas, atlas and

(6) The smaller snout fragment is axis. (x 4)

nearly completely stripped of bone but

shows the roots of five strong and large incisors. This specimen, which. is

probably Therocephalian, now bears the South African Museum Cat. No. 11745.

medium-sized skull. The tip of the snout is weathered away and no upper

incisors are preserved. The lower edge of the dentary is also weathered away.

A moderately sized canine is represented by a root, and four irregularly spaced

and directed molars are present. The molars are implanted on a flange of bone

lying in a plane medial to the general lateral maxillary surface. The lacrymal is

of small antero-posterior extent and its intra-orbital surface is pierced by two

IO ANNALS OF THE SOUTH AFRICAN MUSEUM

foramina. The accompanying sketch gives the general shape and proportions.

The affinities of this Therocephalian are uncertain, but the nature of the lacry-

mal indicates that it is not a very primitive form, and that its affinities would lie

with the Therocephalian forms younger than those from the Tapinocephalus zone.

This specimen now bears the South African Museum Cat. No. 11746.

B. MATERIAL FROM THE UPPER ‘BONE BED’

Under the field number $340 there were six cases of weathered fragments.

The majority of these, mostly small pieces, are so worn as to be indeterminable.

In the workshop few contacts were found and only a small number could be

fitted together. The determinable pieces consist of cranial fragments, a large

number of vertebrae, parts of the pelvic and shoulder girdles, three complete

Fic. 4.—Therocephalian sp. S.A.M. Cat. No. 11746. Lateral

view of incomplete and weathered snout. (x $)

limb-bones and a large number of ends of various limb-bones. The following

reptile groups are represented: Pareiasaurians, Anomodonts, Rhynchosau-

rians, Pseudosuchians, Cynodonts, and a Theropodous Dinosaur.

1. Pareiasaurians

Of this group there is preserved a single weathered distal end of a small right

humerus. It is much smaller than the specimen from Tanganyika figured by

Haughton and described under the new specific name Anthodon minusculus. In

Haughton’s specimen the greatest width across the condyles is 87 mm., whereas

in this specimen it is only 63 mm. In all other determinable features the two

specimens agree very closely. It would thus appear that we have here a humeral

end of a smaller species or more probably of a juvenile specimen of Anthodon

minusculus.

In the material from Tanganyika described by Haughton some errors of

labelling are mentioned by him. The type specimen bears the number $342

REPORT ON COLLECTION OF FOSSIL REPTILIAN BONES Ii

and a second specimen the number $350; the former number being given to

specimens from a locality (Njalila) in the Upper ‘Bone Bed’ and the latter

number to specimens from a locality (below and west of Kingori) in the Lower

‘Bone Bed’. Haughton assumed the number $342 to be an error in the labelling.

Now, the present specimen bears the number $340, which refers to a locality

(Njalila-Mkongeleko) in the Upper “Bone Bed’. The weight of evidence in

regard to the labelling is thus that all these specimens are derived from the Upper

‘Bone Bed’. Palaeontologically, however, the Lower ‘Bone Bed’ is the horizon

indicated, unless we consider it probable that this species survived practically

unchanged into the Upper “Bone Bed’. Until this uncertainty is removed these

specimens must be cited with caution in drawing stratigraphical or faunistic

conclusions.

2. Anomodonts

Fragments of skulls, lower jaws and various limb-bones prove the presence

in these beds of species of medium sized to large Anomodonts.

(a) A well-preserved right humerus of medium size (fig. 5) can, despite the

paucity of our knowledge of the post-cranial skeleton of the numerous species of

Anomodonts, be referred to the genus Lystrosaurus. ‘The distal condyles are

undeveloped, the antero-ventral line is only weakly indicated, the lateral median

line undeveloped, and no definite scar for the medial humeral head of the

triceps is present. These characters exclude the terrestrial genera of the Anomo-

donts, and agree with the condition manifested in the species of the genus

Lystrosaurus. In the absence of a skull no specific determination is possible, and

it is advisable, until further data are available, to refer to this specimen as

Lystrosaurus sp. ‘The chief measurements are:

Greatest lencthen \ 4a ee ee, 4m

Greatest;proximal width= 377 © ;,

Greatest aistal.width =~.) 2 2/2 61...

Shattw cs Gh oe bves 5 ram:

This specimen now bears the South Atican Museum Cat. No. 11748.

(b) Under the number $340 are included about a dozen ends of limb-bones.

Among these are seven distal and three proximal ends of humeri, three distal

and two proximal ends of femora which, with slight individual differences,

agree well with the corresponding bones of Kannemeyeria. ‘The widths across the

distal ends of the humeri are: 180, 160, 158, 154, 148, 139 and 125? mm.

These humeral ends are manifestly different from the humerus figured by

Haughton under the name Focyclops(?) sp. These specimens now bear the

South African Museum Cat. No. 11749. Some fragments of the pelvic and

pectoral girdle may also very well belong to this genus. In addition to these

limb-bones there are also preserved the ends of much smaller Anomodont humeri

and femora. So that, contrary to our experience in the South African Karoo,

small Anomodonts survived comparatively late in Tanganyika. These bones

now bear the South African Museum Cat. No. 11750.

ANNALS OF THE SOUTH AFRICAN MUSEUM

"(¢ x) smoata Jeunxoid pure so1ojue “[esrop ‘TeVUSA UT snJoUINET ys “gPL11 ‘oN VED ‘Fry's ‘ds sninvsousdI—G oy

e hy) Jniid

Wie

REPORT ON COLLECTION OF FOSSIL REPTILIAN BONES 13

a quadrate, and some pieces of occiput. One occiput, with its tripartite condyle

and its characteristically shaped and ventrally directed basisphenoidal tubera

nearly surrounding the foramen ovale, is strongly reminiscent of the same struc-

tures in the skull of Kannemeyerta and may with confidence be referred to that

genus. Some pieces of maxilla with the canine roots preserved may also be

included in this determination. These fragments now bear the South African

Museum Cat. No. 11751.

(d) Three other occipital fragments are more massive. Here the condyle is

rounded with no grooves tripartitioning it. ‘The basioccipital tubera, though

massive, do not descend so far ventrally and are situated some distance apart.

A very similar condition is shown by a number of the larger Aulacephalodon-like

forms, but as in the large number of described forms this area is neither figured

nor described, closer comparison is not possible, and for more specific deter-

minations it is necessary that we have better descriptions of known forms or

better-preserved skulls from this area. These specimens now bear the South

African Museum Cat. No. 11752.

3. Cynodonts

One very badly eroded fragment consists of the anterior two-thirds of a fairly

small Cynodont skull. The outer surface is so badly weathered that the structure

is indeterminable, and on the palatal surface little more than the teeth sockets

are preserved. These indicate that we have here a specimen of the genus

Trirachodon. This specimen now bears the South African Museum Cat. No.

11755.

4. Rhynchosaurians

The collection includes the following bones, or parts of bones, determinable

as Rhynchosaurian: a complete humerus, two proximal and seven distal ends

of humeri, six proximal and four distal femoral ends, parts of the pelvic and

pectoral girdles, a large number of disarticulated vertebrae, a jaw fragment and

two incomplete occiputs.

(a) The Humerus (fig. 6).—Direct comparison of these humeral elements to

the type humeral end of Stenaulorhynchus stockleyt proves them to be specifically

identical. In size and in the proportions there are considerable differences in

the hitherto described material and that at present under consideration, as will

be apparent from the following table:

Type Para- Huene’s Spec. Spec. Spec. Specs.

a b

type Spec. c geen fe

via, length. 22) se) E00 : ? 145 166 G 2 a vats fiat

Max. width across prox.end 10! 77 92 110 120 ? OE niet nan

Piatt ae (eee - ee AL X< 30) OTOL On KOO)" 39> 241 ? ? iii a Vi

Proc. lat.—prox.end . . 83 57 60? 81 94. ? abidots sal ig

Proc. med.—prox.end . 85 63 88? IOI 110 ? be sO

Thickness at med. corner . 37 26 33 31 40 Y Ee ia aol

Max. width across dist. end 2 ? 80 100 ? OSE SOr a7! AO,

ANNALS OF THE SOUTH AFRICAN MUSEUM

suimeiq *(# xX) smorA [ewrxoid puv solazue ‘Jessop “fen

‘assaH] ‘[ vy aq Aq

oA Ul snuouinyy 1ysry “SSL 11 ‘ON VeD “Py'sS. “2py205 snysudsojnouajs—'g “O14

REPORT ON COLLECTION OF FOSSIL REPTILIAN BONES I5

In the light of our present limited knowledge it is advisable to consider these

differences of size and proportions as age and/or sex variations and not as

denoting any specific distinctness.

(b) The Femur.—In the collection there are six proximal and four distal ends

of femora. Direct comparison with the type material proves these bones to be

specifically identical to Haughton’s Stenaulorhynchus stockley. As in the type

material there is considerable variation in the various dimensions:

, a b c d e ie

Max. width over trochanter . 93 73 72 71 65 64

Max. thickness of prox.end . 58 y 44 53 46 52

Trochanter—prox.end. . . 37 2 Be: 15 22 BT

There is, moreover, considerable variation in point of size, shape and position

of the trochanter:

In a the slightly bulbous trochanter is connected with the proximal surface

by a somewhat constricted neck.

In 6 the trochanter is less bulbous and the neck less constricted.

In ¢ the trochanter is hardly thickened, and instead of a neck a sharp ridge

connects it to the proximal surface.

In d the trochanter lies nearly in the same plane as the proximal surface, and

in proximal view appears as a tongue-like extension of the proximal surface,

without any neck.

In e the bulbous trochanter connects with a short neck to the side of the

proximal end to form a distinct step.

In f a thickened ridge attaches the trochanter to the proximal end.

In the specimen recently described by me under the name Scaphonyx africanus

the bulbous trochanter is separated from the proximal surface by a much greater

step than in e. The amount of variation shown by the above specimens impels

one to reconsider the position of Scaphonyx africanus. ‘This must now be considered

as an extreme variant of a femur of Stenaulorhynchus stockley1, and the name given

by me becomes a synonym of Stenaulorhynchus stockley: Haughton.

The distal femoral ends of our material agree very well with that of the type

material and the shaft of one specimen with that figured by Von Huene.

essentials with the corresponding bones figured by Von Huene in his paper on

Stenaulorhynchus. , |

(d) The Occiput.—tIncluded in the material there are two imperfect and

weathered occipital fragments that appear to be identical to the occiput

described by Von Huene and figured in fig. 4 in his paper on Stenaulorhynchus

and in fig. 6 in his “Die Verwantschaftsgeschichte der Rhynchosauriden des

Siidamerikanischen Gondwanalandes’. A very eroded fragment in a different

type of matrix shows a part of the maxilla and dentary which also appears to be

Rhynchosaurian. All these Rhynchosaurian bones now bear the South African

Museum Cat. No. 11753.

ANNALS OF THE SOUTH AFRICAN MUSEUM

‘(| x ) smora yeuutxoid pue so1a}ue “fesiop ‘fenuaA UIs

‘assay [vy ‘aq Aq suimeiqgg

nooumy wysry “PSL11 on VO “y's ‘ds ‘u sosuavyruvsuny snyonsouosnjg—'L, *o1.J

REPORT ON COLLECTION OF FOSSIL REPTILIAN BONES a7)

5. Pseudosuchians (fig. 7)

Also bearing the field number $340 is a well-preserved right humerus. This

bone shows undoubted and close relationship to the corresponding bone in the

material collected by Dr. Nowack in the Njalila area, and included in the type

material described by Von Huene under the name Stagonosuchus nyassicus, and

representing a new genus and species of Stagonolepid Pseudosuchians. Although

closely resembling Von Huene’s form, the following comparative table of

measurements in mm. will show that some noteworthy differences in size and

proportions exist:

St. nyassicus St. tanganytkaensis

Masamanriencth, 2. 2 320 200

Max. width across prox.end . . 180 12

Max. width across dist.end. . . 130 93

Maxewidth ofshaft . 9. 2... 50 38

roe lat—prox.end.° . . . . 70 52

Ectepicondylar flange—dist. end. 30 23

These differences appear to be of a specific nature, and I propose that this

new species be known under the name Stagonosuchus tanganyikaensis. The main

points of difference between nyassicus and tanganyikaensis may be enumerated:

in nyassicus the processus lateralis lies in the same plane as the median corner of

the proximal surface, whereas in tanganyikaensis the medial corner is not deflected

and thus not situated so far distally; in tanganyikaensis the bicipital fossa is

deeper and circumscribed much more definitely; the angle between the planes

in which the proximal and distal ends lie is less in tanganyikaensis than in nyassicus ;

proportionally the distal end is wider, the shaft has a greater maximum width

and is relatively shorter, the processus lateralis extends further distally in

tanganyikaensis than in nyassicus. ‘This humerus now bears the South African

Museum Cat. No. 11754. :

6. Dinosaurs

Among the large number of Rhynchosaurian vertebral elements in this

collection I found a fairly small caudal vertebra lacking the upper part of the

neural spine, of a Theropodous Dinosaur. The chief measurements are:

Greatest lensth ofcentrum ~~. . .. 40 mm.

Grcatestaheicht of centrum) isc js o 34 5,

Greatesu width, of centrum >.2;.., ..-. 20

I have compared this vertebra with the Thecodontosaurus caudal vertebrae in

our Museum and find a fairly close agreement, especially with the anterior

caudals of a specimen from the Red Beds of the Stormberg Series of the Union.

This specimen now bears the South African Museum Cat. No. 11793.

CONCLUSIONS

From the above account it is thus evident that in this collection there are

from the Lower “Bone Bed’ no forms showing any close relationship to the fauna

of the TYapinocephalus zone. ‘The Anomodonts, Pareiasaurs and Therapsids it

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

contains are all manifestly akin to species from the Endothiodon and Cistecephalus

zones of the Karoo of the Union. The beds represented in the Matomondo and

Ngaka-Kingori Hill areas are thus homotaxial to the upper two zones of the

Lower Beaufort of the Union.

The assemblage from the Upper “Bone Bed’ of the Njalila-Mkongeleko area,

containing as it does small to medium-sized Anomodonts, Anomodonts of the

Aulacephalodon group, a Lystrosaur, a Kannemeyeria, a Gynodont, a Rhyncho-

saurian, a Pseudosuchian and a Theropodous Dinosaur, is related to the fauna

known from the beds of the Karoo ranging from the top of the Cistecephalus zone

of the Lower Beaufort right up to the Red Beds of the Stormberg. A more

detailed recording of the relative levels in which the various fossils occur will

enable the stratigrapher to subdivide the Upper “Bone Bed’ of this area into

beds respectively homotaxial to the Middle and the Upper Beaufort, the Mol-

teno and the Red Beds. It is probable that in this area some of the Anomodonts

were actually derived from the top of the Lower Beaufort.

ACKNOWLEDGMENTS

My thanks are due to Mr. G. M. Stockley and his collectors for the oppor-

tunity of examining this interesting collection, and to Dr. A. J. Hesse for two

of the illustrations.

REFERENCES

Boonstra, L. D. ‘A Report on some Karroo Reptiles from the Luangwa Valley, Northern

Rhodesia’, Q.7.G.S., xciv. 1938.

wate L. D. ‘A Note on Some Rhynchosaurian Remains from Tanganyika Territory’,

Ann. S. Afr. Mus., xli, p. 1. 1952.

Brot, F., and ScHRODER, J. ‘Beobachtungen an Wirbeltieren der Karrooformation, xxiv,

and ‘Uber Theriodontier-Reste aus der Karrooformation Ostafrikas’, Sitzb. Bay. Akad. Wiss.

1936.

Lceron S. H. ‘On Karroo Vertebrates from Nyasaland’, Trans. Geol. Soc. S. Afr., xxix. 1926.

Haucuton, S. H. ‘On a Collection of Karroo Vertebrates from Tanganyika Territory’, Q.7.G.S.,

Ixxxviil. 1932.

Huene, F. von. ‘Uber Rhynchosaurier und andere Reptilien aus den Gondwana Ablagerungen

Stidamerikas’, Geol. Pal. Abh., 17/1/1929.

‘Kurzer Uberblick iiber die terrestrischen Wirbeltierfaunen der jungeren Gondwana-

zeit’, Centr. f. Min. usw., Abt. B, No. 6. 1933.

‘Stenaulorhynchus, ein Rhynchisauride der Ostafrikanischen Obertrias’, Nova Acta Leo-

poldina, vi, 36. 1938.

‘Ein grosser Stagonolepide aus der jungeren Trias Ostafrikas’, N. Jahr. f. Min. usw.,

Bd. 80, Abt. B. 1938.

‘Die Alterbeziehungen der siidamerikanischen Gondwana Fauna’, Physis., xiv. 1939.

‘Die Lebensweise der Rhynchosauriden’, Pal. Zeit., xxi, 3. 1939.

‘Die Karroofauna im ostafrikanischen Ruhuhu Gebiet’, Zenérbl. f. Min. usw., Abt. B

NOs 2a

‘Ein bese Pseudosuchier und ein Saurischier aus den ostafrikanischen Mandaschichten’,

N. Jahr. f. Min. usw., Bd. 81, Abt. B. 1939.

‘Die Verwantschaftsgeschichte der Rhynchosauriden des sitidamerikanischen Gondwana-

landes’, Physis., xiv. 1939.

‘Die Saurier der Karroo-, Gondwana- und verwandten Ablagerungen in faunistischer,

biologischer und phylogenetischer Hinsicht’, NV. Jahr. f. Min. usw., Bd. 83, Abt. B. 1940.

‘Die Anomodontier des Ruhuhu-Gebietes in der Tubinger Sammlung’, Aeon braphica,

XClV. 1942.

eee aus dem Ruhuhu-Gebiet’, Pal. Zeit., xxiii, 3/4. 1944.

PARRINGTON, F. R. “On the ‘Tooth-Replacement in ’Theriodont Reptiles’, Phil. Trans. Roy. Soc.,

B, 226, 532. 1936.

‘On the Cranial Anatomy of Cynodonts’, P..Z.S., cxvi, 2. 1946.

Youne, C. C. ‘The Triassic Vertebrate Remains of China’, Am. Mus. Nov., No. 1324. 1946.

3. A suggested clarification of the Taxonomic Status of the South African Titanosuchians.

By Lizuwe D. Boonstra, D.Sc.

(With Plates I-I1X)

INTRODUCTION

Being engaged on a morphological study of the Deinocephalians the need has

arisen to establish some order in the systematics of the group. In an assessment

of the importance of the various morphological features the existence of a

number of generic and specific names, not associated with at least some of the

main morphological characters, is merely confusing and an encumbrance.

In the literature there are about half a dozen generic and a somewhat greater

number of specific names that signify next to nothing. One is led to wonder why

they were ever created at all. One realizes that due to the nature of the material

dealt with in palaeontology, it is not always possible to describe only relatively

well-preserved specimens. ‘There is some justification, although one doubts the

wisdom, if a newly discovered fragment establishes the existence of a group

hitherto unknown to science (as, for instance, Owen’s creation of Titanosuchus

Jerox), but there is no justification for subsequent authors to create additional

names for specimens that do little more than prove that there are more indivi-

duals of the group besides the original individual. It is, of course, fully realized

that the vertebrate palaeontologist generally deals with individuals and that

his species mostly refer to single individuals and are hardly ever the norm of a

good series.

In the Titanosuchians sixteen generic names have been created up to date.

Of these only five (Jonkerta, Dinosphageus, Anteosaurus, Dinophoneus, and Dino-

suchus) are based on reasonably fully preserved skulls, one (Phoneosuchus) on a

good lower jaw, and the other ten (Tztanosuchus, Archaeosuchus, Dinartamus,

Scapanodon, Lamiasaurus, Dinocynodon, Enobius, Scullya, Dinopolus, and Titanogna-

thus) are all based on either skull fragments or parts of dentaries.

In two former papers, in 1935 and 1936, I have shown that two of the above

names must fall away—Dzinophoneus and Phoneosuchus being both synonyms of

Jonkeria. In the sequel a further reduction of generic names will be proposed.

At this stage I am only considering the cranial material on which, in any case,

the authors have mainly created their new genera and species. I have critically

examined most of the described specimens and for the rest have extracted the

pertinent points from the descriptions of the authors concerned. In addition

there is at the South African Museum a large collection, mainly collected by

myself in recent years, of undescribed material which includes a dozen or so

really good skulls. I now wish to present tentatively, for criticism by colleagues,

a solution of the taxonomic maze formed by the plethora of names. With due

Hg)

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

regard to the fetish of the law of priority I have attempted to include as many

of the fragmentary specimens as is possible instead of simply regarding them as

generically or specifically indeterminate.

SEX AND AGE IN THE TITANOSUCHIANS

In addition to the fragmentary nature of many of the types there are other

difficulties in arriving at a reasonable classification. With the small number of

individuals known we have little to indicate sex in the Titanosuchians. It must,

however, be kept in mind that a smaller size, lighter build and a lesser degree

of pachyostosis, particularly with regard to bosses, may indicate the female of

the species.

Without a reasonably long series of specimens it is difficult to determine the

effects of age in the Titanosuchian skull. In the Tapinocephalian, Moschops,

Gregory found an increasing pachyostosis, especially of the postorbital bar, a

character indicating increasing age. In some of the Titanosuchians, especially

of the Anteosaurus group, there is a slight variation in the thickening of the post-

orbital bar, but whether this is due to age or sex is uncertain. In the Jonkeria

group there are marked differences in size unaccompanied by other differences

of generic value. Here size is not considered a character of generic value and at

most may be regarded as specific, but in some cases may very well be a character

of full maturity. Thus Dinosphageus is considered to be a large species of Jonkeria,

but may very well prove to be a fully grown specimen of one of the previously

described species of Jonkerta. ‘The strength and size of the dentary, particularly

of the mentum, have been used as a character of systematic value, but is it not

probable that a moderately strong dentary with a sloping mentum may become

massive with a fairly upright mentum with increasing age? This was one

reason for regarding the name Enodius as unnecessary.

THE DENTITION

The dentition of the Deinocephalians presents some very interesting condi-

tions. A detailed study of the modifications and their implications should be

undertaken. Here I can only touch on some aspects. A study of the teeth is

made very difficult because of the poor preservation in most specimens. In the

majority of cases the crowns are not preserved and we are forced to study stumps

and cross-sections at varying levels which are not directly comparable. The

presence of a lingual crushing surface internal to a talon in many teeth very

materially affects the nature of the cross-section at different levels. Where

crowns are preserved the intractable nature of the matrix of the Tapinocephalus

zone makes it difficult to free them from the matrix satisfactorily.

The Titanosuchians have a heterodont dentition with incisors, canine and

postcanines. In some forms the incisors are fairly short with a labial talon and

a lingual crushing surface, in others the incisors are long to very long with a

talon long to very long and the lingual crushing surface is poorly developed or

represented only by a cingulum, or wholly absent. ‘The canines are strong to

TAXONOMIC STATUS OF SOUTH AFRICAN TITANOSUCHIANS DON

very strong, simple, pointed teeth of a typical carnivorous nature with, in some

cases, a possibly serrated posterior cutting edge. What little is known of the

postcanines indicates a considerable variation in the nature of the crowns—

some have flattened crowns with serrations, others short or longish conical teeth.

Now, is the heterodont Titanosuchian dentition to be morphologically

derived from the homodont dentition of the Tapinocephalians? The Tapino-

cephalian teeth all have a talon, pointed or somewhat flattened, and a lingual

crushing surface varying in strength, from which the different Titanosuchian

types of teeth could be morphologically derived. The medium-sized incisors of

a form like Jonkeria, with its short talon and definite lingual crushing surface,

would then represent a more primitive stage than the long incisors of some of

the Anteosaurians which are all talon with no crushing surface. The stages in

this transition could then be used for taxonomic purposes. But, in Dinartamus,

Broom found a mixture of these two types of incisors and the question arises

whether this may not represent a difference in the consecutive sets of incisors.

We do not know how many times the Titanosuchians replaced their incisors.

Hitherto only two sets have been described. On this subject the external nature

of the teeth gives little information and we have to rely on fortuitous weathering

and fracturing or sectioning. The material at my disposal affords no evidence

as to whether the Jonkerian type of incisor is the deciduous type and the Anteo-

Saurian type typical of a later set of teeth. Until we have evidence supporting

Broom’s observations on Dinartamus I propose to consider the presence or

absence of the lingual crushing surface in the incisors to be a specific character

present in the consecutive sets of teeth, and not dependent on age.

Besides having these two types of incisors the Titanosuchians also possess a

variable number of incisors—as described, from 0 to 5. In those specimens

where the premaxilla is edentulous the condition can be interpreted in a

number of ways: the incisors may have been shed postmortemly, or being

juvenile they have not yet erupted, or being gerontic they have been lost. The

nature of the material being what it is, it is difficult to decide which explanation

best fits each case. Sectioning through the alveolar region is not always possible

in order to base a count on the nature of the alveoli. The usual number of

incisors is 5 but in some forms it is 4 or even 3. Can the number be considered

a character of systematic value? If the juvenile and mature incisors are of the

same size then obviously the premaxilla of a young animal could only house a

smaller number than that of the mature animal. In using the number of

incisor teeth as a taxonomic character the age of the animal must thus be taken

into account.

The number of postcanine teeth in the Titanosuchians varies greatly, from 1

to 19, and is of doubtful systematic value. In some, these teeth are small; in

others, strong; in some specimens they are regularly spaced and in others quite

irregular. The two sides are often dissimilar, e.g. in Enobius there are, according

to Broom, in the left dentary ‘. .. portions of three molars. There certainly

have been four, and very probably there have been five’, and on the right side

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

Broom found only one tooth behind the canine. In general it can be stated that

in the Jonkerias the postcanines are regularly spaced and form a long series,

whereas in the Anteosaurians they are irregular and the series is short.

With the material at my disposal I can state that the dentary usually has one

incisor less than the corresponding premaxilla. It would also appear that there

are fewer postcanines in the dentary than in the opposing maxilla.

Teeth on the palatines occur in the Anteosaurians and in the fragment called

Scullya, but have as yet not been recorded in the Jonkerias although I found

some indication in Jonkeria ingens (A.M.N.H. No. 5608).* These teeth are

mostly seen only in section but in $S.A.M. No. 11592 there is preserved a small

pointed tooth slightly recurved. In the Anteosaurians the palatine teeth are

situated on a prominent elevation, semilunar or reniform in shape. Their

function seems clear—the incisors and canines tear out a lump of flesh from the

victim and this is then held by the raised palatine teeth as an intermediate

stage in the swallowing process. The palatine teeth situated on this characteris-

tic boss constitute a diagnostic character of value.

KEY FOR THE GENERA

Bearing the above remarks in mind we may now proceed with our attempt

to arrange the Titanosuchians in some reasonable order. As a preliminary step,

and in practice of considerable value, to facilitate the process of a rough-and-

ready sorting out of the 14 genera, I propose making use of the following

key:

A. Forms without bosses on postorbital bar and angular, incisors not long

and with step: 7

1. With many regular postcanines, no palatine boss—fonkeria, Dinospha-

geus, Dinopolus.

2. With few postcanines—Dzinariamus.

B. Forms with bosses on postorbital bar and angular, incisors long and with-

out step:

3. With variable irregular postcanines, prominent palatine boss—Antéeo-

saurus, Dinosuchus.

4. With variable irregular postcanines, probably with palatine boss—

Titanosuchus, Scapanodon, Archeosuchus, Lamiasaurus (snout), Dinocynodon,

Scullya, Titanognathus, Enobuus.

REDEFINITION OF GENERA

With our increased knowledge of the Titanosuchians it has become advisable

to redefine the valid genera:

* For the institutions housing the specimens here referred to the following abbreviations are used:

A.M.N.H., American Museum of Natural History, New York. S.A.M., South African

Museum, Cape Town. B.M., British Museum (Natural History), London. T.M., Trans-

vaal Museum, Pretoria. A.K., Alte Akademie, Munich. K.M., Kimberley Museum,

Kimberley.

TAXONOMIC STATUS OF SOUTH AFRICAN TITANOSUCHIANS 23

I. Jonkeria van Hoepen 1916.

The genotype 7. truculenta is based upon an excellent skull and lower jaw.

Io.

It.

T3.

Skull size—this is medium to large, not massive.

Bosses—there are no prominent bosses.

Additional pachyostosis—the parietals form a prominent ridge

thickened round the pineal foramen but do not form a well-demar-

cated boss.

Palatine—there is no prominent boss, but the palatine is possibly

dentigerous.

Premaxilla—the dentigerous border does not curve upwards and

there is also no corresponding upward sweep of the dentary.

Incisors—are of medium length with labial talon pointed and with

well-developed lingual crushing surface.

Pineal foramen—penetrates anterior part of the parietals and is thus

some distance from the occipital border.

Snout—is relatively long, broader than high, the frontals are only

slightly swollen, the premaxilla is fairly flat.

Squamosal—does not extend far ventrally and does not sweep far

posteriorly, posterior to the interparietal. There is also little lateral

sweep of the squamosal.

Temporal fossa—this is fairly large, with the antero-posterior and

dorso-ventral diameters moderate and approximately equal, not

extending much laterally, as the squamosal does not sweep much

outwards.

Intertemporal width—this is small to moderate; the parietals are

laterally pinched in to form a fairly high and narrow parietal crest.

Dentary—this is strong but not massive, with sloping mentum.

- 4—5 1 14—19

Memeaormiula ie, C7.) P-€ aaaqe

Synonym. As thus defined Dinosphageus and Dinopolus become congeneric

with Jonkeria.

Anteosaurus Watson 1921.

The genotype, A. magnificus, is based upon the major portion of a skull

somewhat weathered.

Tee

Die

Skull size—this is small to very large, slightly to very massive.

Bosses—a medium to very prominent boss is present on the dorsal

part of the postorbital bar, and there is a prominent oval boss on the

angular, the boss on the jugal is absent or low and moundlike to very

prominent.

Additional pachyostosis—the parietals are much thickened but do

not form a crest, greatly thickened around the pineal foramen to

form a mound or well-demarcated circular boss; the frontals are

sreatly thickened to produce a medium to very prominent swelling.

HII.

ANNALS OF THE SOUTH AFRICAN MUSEUM

4. Palatine—there is a prominent semilunar ridge-like or reniform boss-

like eminence carrying irregular small pointed teeth.

5. Premaxilla—the dentigerous border curves antero-dorsally to form

an obtuse angle with the maxillary border and there is little corre-

sponding upward sweep of the anterior part of the dentary.

6. Incisors—are long to very long, the labial talon forming most or all

of the tooth, with the lingual crushing surface greatly reduced to form

little more than a cingulum, or are altogether absent.

7. Pineal foramen—penetrates the posterior part of the parietals and is

thus near the occipital border.

8. Snout—is of medium length, higher than broad, the frontals are

greatly swollen, the premaxilla is dorsally swollen and demarcated

from the maxilla by a groove.

9g. Squamosal—extends moderately to far ventrally and sweeps far

posteriorly, i.e. much posterior to the occipital surface of the inter-

parietal, and sweeps far to very far outwards.

10. Temporal fossa—this is large, with the dorso-ventral diameter

ereater than the antero-posterior, extending much laterally due to

the outward sweep of the squamosal.

11. Intertemporal width—this is moderate to large, the parietals are

laterally somewhat pinched in, but do not form a high and narrow

crest.

12. Dentary—this is strong and fairly to very massive, with fairly upright

mentum.

13. Dental formula—i.3=; ct pee

Synonyms. As thus defined Dinosuchus becomes congeneric with

Anteosaurus.

Dinartamus Broom 1923.

The genotype, D. vanderbyli, is based upon portions of a skull, probably

associated.

Skull size—this is probably large, not massive.

Bosses—no evidence is preserved.

Additional pachyostosis—no evidence of this is preserved.

Palatine—no evidence of a palatine boss is preserved.

Premaxilla—the border does not curve upwards.

Incisors—according to Broom, ‘the first is of Deinocephalian type,

but the other three incisors may have had pointed crowns. There is

some little indication that this may have been so.’

Pineal foramen—this area is not preserved.

Snout—this is weathered.

Squamosal—this is not preserved.

Temporal fossa—this region is not preserved.

Intertemporal width—this region is not preserved.

Soe ee eae ee

ee Ce pes

= a

rv.

TAXONOMIC STATUS OF SOUTH AFRICAN TITANOSUCHIANS 25

12. Dentary—in the associated specimen the dentary is apparently fairly

massive with a fairly upright mentum.

Ay a | 5

13. Dental formula—according to Broom, 1.3, c.;, p.c.3.

Titanosuchus Owen. 1879.

The genotype, 7. ferox, is based upon upper and lower jaw fragments

showing sections of the teeth roots. The only diagnostic features that can

be determined in the genotype are: dentary strong and massive with

mentum apparently fairly upright, and dental formula: i3 : C4 s pc.

Other forms considered here are:

Titanosuchus cloete: (Broom 1903) is based on a piece of massive dentary,

with the incisors lacking a lingual crushing surface and with the dental

fonmaulas i. 4. °C. 1,0 p.c. 4+.

Scapanodon (Broom 1904). ‘The type species is based upon two imperfect,

badly preserved jaws showing a series of teeth: 1.2+,c. 1, p.c. 11+.

Archaeosuchus (Broom 1905). ‘The type of the type species is a partial

maxilla with some teeth; according to Broom c. 1, p.c. 8 (in 1932 Broom

gives 7 postcanines). |

Lamiasaurus (Watson 1914). Only the snout is considered here. The

premaxillary border is not dorsally directed and the dental formula is

MMPOrn. (C1, p.c. 3-4.

Dinocynodon (Broom 1929). For the type dentary Haughton (1915)

states that the symphysis is massive and square, and the dental formula is

1.4, C. I, p.c. 11+, and Broom gives as the only generic character ‘the

extreme flattening of the large canine’. Now, in a specimen of Anteo-

saurus, there is a canine on the point of being shed which is also flattened,

and this character can thus hardly be considered of generic value.

Scullya (Broom 1929). The type species is based upon ‘a very badly

crushed snout’ with the dental formula i. 5, c. 1, p.c. 12, and the only

other characters are the possible presence of teeth on the palatine and the

dentary massive.

Titanognathus (Broili and Schroder 1935). The type of the type species

consists of skull fragments showing, “Schadel mit schmaler und steil vom

pramaxillaren Kieferrand aufsteigender Schnauze, sehr gross. Prae-

maxillarer Kieferrand gegeniiber dem maxillarer stark in die Hohe

gezogen, Symphysenregion des Unterkiefers entsprechend erhoht gegenii-

ber dem riickwartigen Abschnitt des Dentale. Zahnformel: 1. Ho Guts

p-c: aoe

Enobius (Broom 1923). The type of the type species consists of two

dentaries; mentum massive and squarish, with the dental formula c. 1,

DCI Be pcre

From the above it is quite evident that this series of fragmentary ‘types’

affords no very trustworthy bases on which they can be distinguished from

ANNALS OF THE SOUTH AFRICAN MUSEUM

each other generically. I propose lumping all these forms together and to

redefine the genus Tztanosuchus compositely as follows:

Cary Bodo

Skull size—this is very probably large and massive.

Bosses—are probably present.

Additional pachyostosis—of this no evidence is preserved.

Palatine—this is not preserved.

Premaxilla—the dentigerous border curves antero-dorsally with an

associated step-up of the anterior part of the alveolar border of the

dentary.

Incisors—these are apparently long, with no lingual crushing surface.

Pineal foramen—no evidence is preserved.

Snout—is probably as in Anteosaurus.

Squamosal—is not preserved.

Temporal fossa—this area is unknown.

Intertemporal width—this area is not preserved.

. Dentary—this is strong and massive with a squarish and upright

mentum.

ii el eS 1 (il =154:

Dentaliormulas We Gs Ps eee

Synonyms: Scapanodon, Archaeosuchus, Lamiasaurus (snout), Dinocynodon,

Scullya, Enobius, and Titanognathus.

THE SpPEcIFIC NAMES OF THE ABOVE FORMS

Until a detailed study is completed it is impossible to make any statement on

the validity of the specific names given by authors. Meanwhile a list is appended

of the names as they stand at the present moment in the literature, but under

the genera—Anteosaurus, Dinartamus, Jonkerta and Titanosuchus—as defined above.

Genus Anteosaurus :

A. abeli Boonstra 1952. Plates I-V and IX.

Type. A good skull and lower jaw. S.A.M. No. 11296.

A. magnificus Watson 1921.

Type. Major part of skull. B.M. No. 3595.

A. minor (Broom) 1929.

Type. Fragment of skull. B.M. No. 5742.

Referred specimens:

S.A.M. No. 11492. A somewhat weathered skull.

S.A.M. No. 11694. A good skull. (Plate V1, figs. 1, 2.)

Additional diagnostic features revealed by these two skulls are: In

S.A.M. No. 11492, the left premaxilla has no incisors preserved, but on

the right side there are three incisors, fairly long, with no indication of

a lingual step; in S.A.M. No. 11694 no incisors are present, the denti-

gerous boss on the palatine is prominent and semilunar in outline; there

is no indication of a jugal boss. The dental formula is i. 0-3, c. I,

Picai7=e:

TAXONOMIC STATUS OF SOUTH AFRICAN TITANOSUCHIANS 27

A. vorstert (Broom) 1936.

Type. A good skull. T.M. 265, Broom’s Dinosuchus vorstert.

Referred specimen: a good skull. S.A.M. No. 11577. (Plates VII and

VIII.)

Genus Dinartamus :

D. vanderbyli Broom 1923.

Type. Skull fragments. Coll. Broom.

Genus Jonkeria:

J. angusticeps (Broom) 1929.

Type. Good lower jaw. A.M.N.H. No. 5633. Broom’s Phoneosuchus

angusticeps.

F. haughton (Broom) 1939. 3

‘Type. Fairly good skull. S.A.M. No. 4343. Broom’s Dinosphageus haugh-

tont.

J. ingens (Broom) 1923.

Type. Fair skull. A.M.N.H. No. 5634. Broom’s Dinophoneus ingens.

Synonym. 7. pugnax (Broom) 1929. Fairly good skull and lower jaw.

A.M.N.H. No. 5608.

7. truculenta van Hoepen 1916.

Type. Good skull and lower jaw. T.M. 212.

f. vanderbylt (Broom) 1929.

Type. Good skull. A.M.N.H. No. 5620.

J. spp. The lack of identifiable cranial elements makes the specific status of

J. crassus (Broom) 1929, Type A.M.N.H. No. 5577, uncertain. The skull

referred to Scapanodon duplessist by Broom 1923 is undoubtedly a Jonkeria

species. Further study may prove synonomy with one of the named species

and until then it is best left unnamed. Dzénopolus atrox Broom 1923, Type

T.M. 274, consists of a snout. The incisors are short, the anterior three

with ledge, the outer without ledge, according to Broom. The lower

canine peculiar, but otherwise it falls under Jonkeria as here defined. ‘These

characters may indicate that it is a distinct species—onkeria atrox Broom,

bridging the gap between Dinartamus and Jonkeria.

Genus Titanosuchus :

T. cloetei (Broom) 1903.

Type. Piece of dentary. S.A.M. No. 731.

IT. dubius (Haughton) 1915.

Type. Major part of dentaries. S.A.M. No. 2759. Broom’s Dinocynodon

dubius.

T. ferox Owen 1879.

Type. Upper and lower jaw fragments. B.M. No. 49370.

T. gigas (Broom) 19209.

Type. A very badly crushed snout. Coll. Broom. Broom’s Scullya gigas.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

T. lotzi (Broili and Schréder) 1936.

Type. Skull fragments. A.K. (no number given). Broili and Schréder’s

Titanognathus lotzi.

T. strubent (Broom) 1923.

Type. Partial dentaries. K.M. (no number given). Broom’s Enobius stru-

bent.

Incertae sedis. ‘The following forms do not merit separate generic rank and may

best be regarded as species of Yitanosuchus, whose specific validity is

questionable.

T. duplessist (Broom) 1904.

Type. Imperfect badly preserved jaws. S.A.M. No. 769. Broom’s Scapano-

don duplessist.

T. cairncrosst (Broom) 1905.

Type. Part of maxilla. S.A.M. No. 916. Broom’s Archaeosuchus cairncrosst.

T. newtont (Watson) 1914.

Type. Snout. B.M. No. 49385. Watson’s Lamiasaurus newtoni.

REFERENCES

Boonstra, L. D. ‘A note on the synonomy of the two deinocephalians, Dinophoneus ingens Broom

and Jonkeria pugnax (Broom).’ S. Afr. J. Sci., vol. 32, pp. 329-31. 1935.

“The cranial morphology of some titanosuchid deinocephalians.’ Bull. Amer. Mus. Nat.

Hist., vol. 72, pp. 99-116, pls. 9-17. 1936.

Miljoene Jare Gelede in die Karoo. Johannesburg, Voortrekkerpers, 125 pp. 1948.

°’n Nuwe Titanosuchiérsoort (Anteosaurus abeli).’ Tydsk. Wet. en Kuns, xu, bls. 142-9.

1952.

Broom, R. ‘On evidence of a new species of Titanosuchus (T. cloetei).’ Ann. S. Afr. Mus., vol. 4,

pp. 142-3. 1903.

‘Notice of a new fossil reptile (Scapanodon Duplessisi) from the lower Karroo beds of Prince

Albert, Cape Colony.’ Rec. Albany Mus., vol. 1, pp. 182-3. 1904.

‘Notice of some new fossil reptiles from the Karroo beds of South Africa.’ ibid., vol. 1,

[2]0e ee eh Soe

comparison of the Permian reptiles of North America with those of South Africa.’

Bull. Amer. Mus. Nat. Hist., vol. 28, pp. 197-234. 1910.

‘On the structure of the skull in the carnivorous deinocephalian reptiles.’ Proc. Zool. Soc.

London, pp. 661-84. 1923.

‘On the carnivorous mammal-like reptiles of the family Titanosuchidae.’ Ann. Transvaal

Mus., vol. 13, pp. 9-36, pl. 4. 1929.

The mammal-like reptiles of South Africa and the origin of mammals. London, H. G. and G.

Witherby, xvi. pp. 376. 1932.

‘On some new genera and species of Karroo fossil reptiles, with notes on some others.’

Ann. Transvaal Mus., vol. 18, pp. 349-86. 1936a.

‘On the structure of the skull in a new type of dinocephalian reptile.’ Proc. Zool. Soc.

London, pp. 733-42. 19360.

Bro, F., and ScHRODER, J. ‘Ein Dinocephalen-Rest aus den unteren Beaufort-Schichten.’

Sitzber. Bayrischen Akad. Wiss., pp. 93-114. 1935.

Grecory, W. K. ‘The skeleton of Moschops capensis Broom, a dinocephalian reptile from the

Permian of South Africa.’ Bull. Amer. Mus. Nat. Hist., vol. 56, pp. 179-251, pls. 1-21. 1926.

HavucutTon, S. H. ‘On two new therocephalians from the Gouph.’ Ann. S. Afr. Mus., vol. 12,

PP- 55-7: 1915.

Hoepen, E. C. N. van. ‘A new Karroo reptile.’ Ann. Transvaal Mus., vol. 5, suppl. 3, no. 3, I p.

1916.

Owen, R. ‘Description of fragmentary indications of a huge kind of theriodont reptile ( Titano-

suchus ferox Ow.) from Beaufort West, Gough Tract, Cape of Good Hope.’ Quart. Four. Geol.

Soc., vol. 35, pp. 189-99, pl. 11. 1879.

Watson, D. M. S. ‘The Deinocephalia, an order of mammal-like reptiles.’ Proc. Zool. Soc.

London, pp: 749-86, pls. 4, 5. 1914.

‘The bases of classification of the Theriodontia.’ ibid., pp. 35-98. 1921.

-azis Teanjeu + ynoqe ‘MoIA [ese “Q6SI1 “ON “W'V'S ‘uouttoods odAyT, “vaysuoog 1aqv sninvsoajup

snp POW YD) 2 80104 |

Plate

mn. 5. Afr. Mus., Vol. XLII

Peer Cas)

Plate u

Ann. S. Afr. Mus., Vol. XLII

*pasojsaa MUL JAMO] JO oLSuUy

Plate yr

fr. Mus., Vol. XLII

Ama. S.

“OZIS jeanjyeu

Node ‘MOIA [R191e'T ‘obSP CoN CIW'V'S ‘9dA1-o1Fr ‘Bsuo0g 773qD

SNANDSOIJU’

Plate iv

Ann. S. Afr. Mus., Vol. XLII

‘posojsod JNOUS Jo Jed JOIQJUW ‘9ZIS [Vanjeu F ynoqe ‘Mola vszjVT “GSOS1I “ON “WYy'S ‘edAl-o1-) ‘vajsuoog ijaqp snunpsoajuyy

Ann. S. Afr. Mus., Vol. XLII Plate v

Anteosaurus abeli Boonstra. Co-type. S.A.M. No. 11293. Dorsal view, about } natural size. Anterior

part of snout restored.

Ann. S. Afr. Mus., Vol. XLII Plate

}

Anteosaurus minor (Broom). Referred specimen. S.A.M. No. 11694. About } natural size. Fig. 1, lateral view.

Fig. 2, ventral view.

Plate vu

Ann. S. Afr. Mus., Vol. XLII

‘OZIS [VANVU fF INOGV SMOTIA [V19]U'T

uguil

)

ads

PotTIJo YY ‘(wool ) 14948400 SNANDSOAU

Ann. S. Afr. Mus., Vol. XLII | aie som |

Anteosaurus vorsteri (Broom). Referred specimen. S.A.M. No. 11577. Dorsal view, about

4 natural size.

Plate 1x

Ann. S. Afr. Mus., Vol. XLII

(-ojoyd siag ayeuoiseNy) ‘sayeyd SuroSei0j ay} ul poansy sed4j-o9 pure odA} 9y} JO sT[Nys 9y} UO

paseq ‘peoy ay} Jo Joyseyd ut uoNoNssuOdaI ozIs-oyt] JO Ydersojoyd “esuoOg 179q) siinDsoajUp

4. The Gorgonopsians, Aelurognathus microdon and Hipposaurus boonstrai,

reconstructed. By L. D. Boonstra, D.Sc.

(With Plates X—-XVI)

In 1934 I described a new species of Gorgonopsian under the name Aelurogna-

thus microdon. ‘The specimen consisted of a skull, much of the vertebral column,

an excellent pectoral girdle, a good fore-limb with most of the carpus, a good

pelvis and most of the hind-limb. (Plate X.)

The fragile nature of the preserved bones has made it impossible to attempt

a free mount of the skeleton as preserved. During 1937 Mr. J. Drury, then

modeller to the South African Museum, modelled in plaster of paris all the

elements preserved, two-thirds natural size. Utilizing our knowledge derived

from other gorgonopsians species (see op. cit.) the missing bones were restored.

The resulting mount could be considered fairly accurate and a photograph

of it was published in the ‘Report of the South African Museum for 1937’ and

republished in a popular booklet Miljoene Fare Gelede in die Karoo. Apparently

both these have passed unnoticed by overseas colleagues, and Colbert in his

study of Lycaenops ornatus does not mention either of these publications.

An augmented set of new photographs of the modelled skeleton (Plates XI-

XIII) and of the animal reconstructed ‘in the flesh’ (Plate XIV), also by Drury,

is here presented for comparison with the excellent set of photographs published

by Colbert of the remarkable free-mount of Lycaenops ornatus as mounted by

Charles Lamb after preparation by Jeremiah Walsh.

Aelurognathus microdon has been reconstructed with 29 presacral vertebrae

(including the proatlas) measuring 750 mm.; 3 sacrals (90 mm.); 29 caudals

(585 mm.). All these measurements are projections. The total projected length

of the skeleton is 1,530 mm. and measured over the curvature of the back is

1,650 mm. The shoulder height is 570 mm. and at the hips the height is 368 mm.

THe STANCE OF AELUROGNATHUS

The following remarks should be read in conjunction with the section in

Colbert’s paper, ‘The Skeleton as a Whole’, as, after his masterly account,

written with the help derived from Schaeffer’s movie-film of the alligator, I can

here be brief and confine myself to the conditions in Aelurognathus without

repetition of comparisons with other Therapsids.

Drury’s model is mounted showing Aelurognathus in the standing position

based on views held in 1937, some of which have since been modified necessita-

ting some alterations to the original mount. (Plate XIV.)

The curvature of the spine, as mounted, is probably over-accentuated with

the apex of the curve too far forward and I now think that in life the back

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

would have been much straighter with the curve nearer the sacrum. In this I

would be in agreement with Colbert.

The ribs form a complete presacral set. On this point I had in 1937 corrected

the erroneous view expressed in 1934 as to the probable absence of lumbar ribs

in Lycaenops, thereby anticipating Colbert’s criticism of 1948 and thus adeeane

its validity in advance.

The skull hangs downward, dog-fashion, and, in harmony with this, I would

now prefer the neck to show a more pronounced downward curve.

The pectoral girdle was preserved in approximately natural articulation and

‘this position is retained with little correction in the model. As mounted

Aelurognathus has a deep chest—much deeper than Lycaenops as shown in

Colbert’s figure 22. In lateral view the scapula has a slight backward tilt when

the animal is in the standing position. When executing a stride, the forward side

would show the scapula tilting further backwards with its posterior edge everted,

i.e. there would be some rotation of the scapula on its long axis. In anterior

view the scapular girdle is V-shaped, but, with the scapular blade somewhat

curved, the top edge with the cartilaginous suprascapula would not stand

excessively away from the ribs. This I believe to be the natural position of the

pectoral girdle, for in this position the glenoid articulation would appear to

function properly. When executing a stride the left humerus would, in the

forward position, be directed somewhat laterally, and the right humerus be

directed backwards close in to the body. To keep both the humeri in articula-

tion the direction in which the glenoids face must be in keeping. This would be

achieved by a lateral sigmoidal curvature of the spine accompanied by a slight

movement of the scapular girdle as a whole, and also of its two halves indivi-

dually, in relation to the clavicular girdle. Thus the left half of the scapular

girdle would move forwards, sag slightly and rotate on its long axis so that the

glenoid is directed more outwards and, at the same time, the right half would

concomitantly rotate so that the right glenoid is directed slightly inwards from

the backwardly facing position.

In the mount, with the animal in the standing position, i.e. nearly half-way

through the stride, the humerus is directed obliquely outwards with the distal

end appreciably lower than the proximal end, and the elbow is thus everted

but less anteriorly and downwardly directed than would be the case at the

commencement of the stride.

The femur, in the 1937 mount, is directed obliquely forward and outward with

the distal end much lower than the acetabulum. I now think that the femur was

too much everted and that in life the knee would be closer in to the body.

This correction has now been made to the mount as shown in the accompanying

new photographs—on the right side the femur is brought in closer to the body

than the left side. In the stride of the back limbs the spine would be flexed in

the lumbar region to complete the sigmoid curve which the movement of the

fore-limbs initiated.

Plate x

Ne See Air. Muss Vol; SIT

6 ‘ON (IW V'S ‘921s Teanyeu

- jNOGqV

od nyis Ut YY ul yno prey UOPOLILUL snyJDUsOINJap

U0}9]

IAS

‘azis Jeanjyeu $ ynoqe ‘MoIA [eJOJV YT “UoposINU SNYDUTOMNJayy JO UOJI[BYS PeT[Ppoyy

—_=

—_

Afr.

Ann.

Plate xu

‘ozis Jeanyeu # ynoqe ‘MOIA [BS

1 LOCL “UOposdiut SNYIDUDOANIIYT jo UO)JI[IAS PYTPP POA

AW is, ‘ .

& ‘: : .

— as as ei ee ad a ae

~AA A eS s

Ol, DULIL!

Ann. S. Afr. Mus., V

onbiqQ “UOPOLITU SNY TOL a0 njap jo UOJo[I zs P2T[2PPN

Plate xin

—

Plate x1v

ol. XLII

Ann. S. Afr. Mus., \

‘(uunasnyy Uvolapy YING) Aanaqy *[ Aq Ysop otf} Ur, UoposnU

SNYJVUTOANJAP JO |PPOW

Plate xv

Ann. S. Afr. Mus., Vol. XLII

‘ozIs [eanyeu ? ynoqe ojoyg “sJoyNe oY} Aq stred-jo-19}se]d ur ozIs [enyeuU poy[apour wysu00g sninvsodquyy

Plate xvt

Wolk OOIUL

cf)

Ann. S. Afr. Mus

uno f, ado’ ‘slag apouoispyy aq sYydvadojoyd I]t

uoog §

‘UOTONASUOIIT JIAO IU} JO MolA onbyqo

WwW.

imosodquyy

————_—

AELUROGNATHUS MICRODON AND HIPPOSAURUS BOONSTRAI 31

In the reconstruction the feet are probably mounted slightly too much in the

semi-digitigrade position. They should, in the standing position, be more

plantigrade. At the commencement of the stride they would be wholly planti-

grade, and, at the completion of the stride, the digitigrade position would be

obtained just before the foot is lifted from the ground. The fore-feet are

correctly shown as forwardly directed, and, with the knees drawn in to the body

in the corrected mount, the hind-feet also assume a more natural forward

position than was the case in the 1937 mount.

For comparison I have included here some hitherto unpublished photographs

of a life-sized reconstruction of Hipposaurus boonstrat which I have myself

modelled in plaster, with due acknowledgment to Colbert for ideas derived

from his Lycaenops paper. (Plates XV and XVI.)

Eipposaurus is here represented in the walking position and the greater

straightness of the back is noteworthy. The right fore-foot, at the commence-

ment of the stride, is flat on the ground; the left, nearly at the completion of

the move, is semi-digitigrade. The left hind-foot, just after the commencement

of the stride, is semi-plantigrade with the knee probably everted a little too far;

the right hind-foot is just about to be lifted and swung forwards after having

completed the stride.

REFERENCES

BoonstrA, L. D. ‘A Contribution to the Morphology of the Gorgonopsia.’ Ann. S. Afr. Mus. 31,

P- 137- 1934.

Report of the South African Museum for 1937, p. 9 and plate facing p. 18. 1938.

Muljoene Fare Gelede in die Karoo. Voortrekkerpers, Johannesburg. p. 51. 1948.

‘Bewoners van die Oer-Karoo’, Die Huisgenoot, Cape Town, 6 Aug., p. 23. 1948.

“‘Wereld van 250 Miljoen Jaar Gelede.’ Die Burger, Cape Town, 11 May, p. 3. 1950.

Bystrow, A. P. ‘Rekonstuktionversuche einiger Vertreter der Nord-Dwina Fauna.’ Akad.

Nauck, U.S.S.R., 4, p. 289. 1935.

Co.Bert, E. H. “The Mammal-like Reptile, Lycaenops.’ Bull. Am. Mus. Nat. Hist., 89, 6, p. 357.

1948.

5. The cranial morphology and taxonomy of the Tapinocephalid genus Struthiocephalus.

By L. D. Boonstra, D.Sc.

(With Plate XVII and 6 text-figures)

The one important genus of the Tapinocephalid Deinocephalians! of which

the cranial structure has hitherto not been adequately described is Struthio-

cephalus.

Up to date five species of Struthiocephalus have been created. Few details

of the cranial structure were published before I re-examined the type skull?

and described the two new species, duplessisi® (since transferred to the new genus

Struthtocephaloides*) and akraalensis.4 Of Broom’s rheedert® and Olson and Broom’s

milleri® little more is known than can be gathered from Broom’s sketches of the

outlines of the skull and the main skull openings—temporal fossae, orbits,

nostrils and the pineal foramen.

In the collection of the South African Museum there is, besides the type skulls

of whatis: (S.A.M. 2678), and akraalensis (S.A.M. 3719), an excellently preserved

skull of the species whaitst (S.A.M. 11591) showing nearly all the features of the

dorsal, lateral, occipital and palatal surfaces and the right side of the brain-case,

and a juvenile skull (S.A.M. 11493) of the same species. The skull of the

mounted specimen which has associated with it much of the postcranial skeleton

(S.A.M. 3012) is crushed and shows little of the detailed structure.’

The structural details determined in these five specimens are here pooled to

give an account of the cranial morphology of the genus but the figures are all

of the one specimen of whatisi (S.A.M. 11591).

THE GENERAL SHAPE AND FORM

The generic name very aptly describes the shape of the skull as being very

similar to the shape of the head of an ostrich. Anterior to the orbital border the

snout is anteriorly directed, long, shallow but fairly broad and, in relation to

the part of the skull posterior to the orbits, appears weak. With the lower jaw

occluded the muzzle appears much less weak as the mentum is quite massive.

The skull is low—the width over the quadratojugals being much greater than

the height in this plane. The bones of the posterior part of the skull are all

strongly pachyostosed, whereas those of the snout are only relatively moderately

thickened with the outer bone surface smooth. The transition from the posterior

part of the dorsal surface to the dorsal and lateral surface of the snout is thus

abrupt but less abrupt than in Mormosaurus. As the strong pachyostosis does not

include the lacrimal and only affects the posterior part of the prefrontal the

transition from the smooth snout to the rough posterior part lies further back

than in Taurocephalus and Mormosaurus. The pachyostosis in Struthiocephalus is

not general over the posterior part of the outer surface—the ‘cheek’ being still

CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 33

fairly light and its surface smooth, and on the dorsal surface the strong thicken-

ing is still localized, emanating from distinct centres. The pachyostosis forms

a strong rugose, dorsally somewhat bulbous, postorbital bar; a prominent

rugose boss surrounding the pineal foramen and a peculiar naso-frontal boss.

(The first is reminiscent of the condition in the Titanosuchid Anteosaurus,

_where it is very strongly developed, and the last a feature which it has in

common with the Tapinocephalid Keratocephalus.)

The orbits, situated in the posterior half of the skull, are large, round and

directed forwards and outwards with the thickened postero-dorsal half of the

orbital border strongly overhanging.

The nostrils are large and elongated, situated on the dorsal surface mainly

dorsally, and only slightly laterally directed and are well back from the anterior

edge of the snout, close to each other and separated by a strong internasal bar.

The temporal fossa is fairly large, higher than long, and the pachyostosis of

the postorbital bar and the posttemporal arch has not reduced its size much;

its anterio-posterior diameter is still greater than in Mormosaurus and much

greater than in the slitlike fossa of Tapznocephalus. Dorsally, it extends medially

to form a bay encroaching into the parietal region (in akraalensis the fossa

approaches the condition in Mormosaurus).

The interparietal width is moderate, due to a pinching-in laterally of the

parietals to form the dorso-median bay of the temporal fossa.

Due to the forward position of the quadrate, which lies anterior to the plane

of the orbit, the lower jaw is short and the maximum gape of the jaws is com-

paratively small. :

The anterior teeth of the upper jaw are directed much anteriorly but the

intermeshing teeth of the dentary are directed dorsally.

THE BONES OF THE DoRSAL AND LATERAL SURFACES (figs. I and 2)

The matrix of the Tapinocephalus-zone being notoriously intractable, the

determination of sutures in most specimens from this horizon is extremely

difficult. ‘Thus Haughton in his description of S. whaitst and Broom in the case

of S. rheedert and Olson and Broom in 'S. millert have, together, only figured parts

of two sutures. It has only been through a laborious process of ‘artificial

weathering’ by dilute hydrochloric acid that I have been able to determine

most of the sutures, but some still remain indeterminable and others uncertain.

The premaxillaries (P.Mx.) together form a large part of the snout. From

the anterior border they stretch posteriorly to past the middle of the skull.

From its anterior border each premaxilla narrows, where it forms the inner

border of the nostril, then it stretches as a long tapering bone posteriorly, where

it lies in a groove of the nasals.

The nasals (N.) are long, narrow bones, which in their postero-median part

are grooved to house the posterior tongue of the premaxillaries; posterior to

the limits of the premaxillaries they meet on the dorsal surface in the median

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

line and are here thickened to form the anterior and major part of the naso-

frontal boss.

The septomaxillaries (S.Mx.) appear to be small splint-like bones forming

the outer border of the nostrils, but their limits are uncertain in most specimens.

The maxillaries (Mx.) are the largest bones of the snout, being long but

shallow. Posteriorly a dorsal prong just meets the prefrontal, but in some speci-

mens the lacrimal is intercalated, and a ventral prong extends far posteriorly

with its upper edge applied to the lower border of the jugal. In between these

two prongs lie the anterior ends of the lacrimal and jugal.

Fic. 1. Struthiocephalus whatisi. Lateral view of the skull (S.A.M.

11591) (x4). All the figures are not perspective drawings but pro-

jections drawn with the aid of a pantograph. The lateral view of

the skull and that of the braincase are projected on to the median

(sagittal) plane, the dorsal and ventral views are projected on to

the plane of the alveolar border, and the occipital view at right

angles to the plane of the alveolar border.

The lacrimal (L.) ventrally meets the upper edge of the jugal in a straight

suture and these two bones together extend anteriorly into the posterior fork

of the maxilla. The lacrimal is in its anterior extent as in Mormosaurus and thus

stretches much further anteriorly than it does in Taurocephalus. It forms only a

small part of the relatively unthickened anterior orbital border. Dorsally the

prefrontal does not, in S.A.M. 11591, altogether exclude the lacrimal from

contact with the nasal; in the other specimens a tongue of the prefrontal is

intercalated between the nasal and lacrimal. (In Taurocephalus the lacrimal

does not meet the nasal, but in Mormosaurus it does.)

The jugal (J.) is a strong bone; like the bones of the snout it is not greatly

thickened and its surface is not rugose but smooth. It forms the antero-ventral

comparatively unthickened border of the orbit. It extends far ventrally as a

tapering element to be separated (in S.A.M. 11591) from the quadratojugal

by a narrow incisure. Anteriorly it stretches far as a prolongation, with the

lacrimal, into the posterior fork of the maxilla. The posterior border forms a

shallow curve and is thus not deeply indented by an anterior wedge of the

squamosal as in Mormosaurus and Taurocephalus.

CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 35

The prefrontal (Pr.F.) is much thickened along its lateral edge to form the

antero-dorsal thick, rugose and rounded portion of the orbital border. In all the

specimens, except S.A.M. 11591, it has an anterior tongue which separates the

lacrimal from the nasal. It does not extend much posteriorly, but is thickest

here. Medially it is thinner and there is thus a hollow between the thickened

orbital border and the naso-frontal boss.

The frontal (F.) is a large element of the dorsal skull roof. Its median part is

roughly rectangular and from here three tongues extend: one, anteriorly inter-

Fic. 2.—Struthiocephalus whaitsi. Dorsal

view of the skull (S.A.M. 11591) (x4).

calated between the nasal and prefrontal; the second, entering the thickened

rugose upper orbital border and, the third, is a wedge between the postfrontal

and the parietal. Anteriorly, the frontals meet the nasals and here form the

posterior minor part of the naso-frontal boss. For the rest the frontals are not

greatly pachyostosed and in whaitsi form a shallow saddle between the naso-

frontal and parietal bosses, whereas in akraalensis the frontals are thickened in

the median line with laterally a deep depression which is further laterally

bounded by a strong ridge formed by the postfrontal and postorbital. Pos-

teriorly, the frontals meet the parietals in a nearly straight frontal suture just

anterior to the parietal boss surrounding the pineal foramen.

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

The postfrontal (Po.F.) forms the dorso-posterior corner of the orbital border

which is here greatly thickened and very rugose. Its suture with the postorbital

is in Most specimens uncertain but in whazts: (S.A.M. 11591) it is a large bone

forming the dorsal swollen part of the postorbital bar, and in akraalensis it forms

part of the ridge lying lateral to the depression in the surface of the frontal. This

identification of the large size of the postfrontal in Struthiocephalus leads one to

suspect that this element is a much larger bone than it has hitherto been thought

to be in most Tapinocephalia. The pachyostosis of the postfrontal has resulted

in the postorbital being practically excluded from the dorsal surface of the skull

and thus forms only the lower part of the postorbital bar. Another result of

the thickening of the postfrontal has been that together with the enlarged and

thickened prefrontal the frontal tends to become excluded from the orbital

border.

The parietals (P.) together form the greater part of the cranial roof. In their

antero-median portion a large elevated and rugose boss is developed and is

pierced by a large round pineal foramen. In their posterior half the parietals

form a narrowed dorsal surface as they are here laterally pinched in. This

pinching-in is less evident in akraalensis. Here a sharp edge separates the dorsal

surface from a lateral surface, which forms the dorsal part of the median or

inner face of the temporal fossa. ‘This lateral parietal surface, extending on to

the posttemporal arch to meet the squamosal, effectively prevents the post-

orbital from meeting the squamosal at this level. The medio-dorsal bay of the

temporal fossa thus formed by the pinching-in of the parietals laterally is clearly

shown in whaitsi, but in akraalensis the temporal fossa is more a continuous slit

with this bay not clearly demarcated. Posteriorly, the parietals are buttressed

by the interparietal in their median part and, more laterally, by the strong

tabulars.

The postorbital (P.O.) is a massive element forming the lower part of the

thickened postorbital bar which in its upper postfrontal part is bulbous on a

scale just less than in the Titanosuchid Anteosaurus. ‘The postorbital bar 1s

strong, wide and fairly rugose in whaitsi, and very strong, very wide and strongly

rugose in akraalensis. Posteriorly, the postero-lateral flange of the parietal lies

between the posterior process of the postorbital and the squamosal. In a

juvenile specimen of whaits1, S.A.M. 11493, the posterior process of the post-

orbital still stretches far posteriorly, but even here does not meet the squamosal.

Ventrally, the postorbital forms an overlapping suture with the squamosal.

The squamosal (Sq.) is the main constituent bone of the ‘cheek’. It is a

strong thickened element but, as in the jugal, its outer surface is not rugose but

smooth in whaztsi, but with pits and rugae in akraalensis. Anteriorly, it meets the

jugal in a long curved suture with no anterior wedge-shaped process as in

Mormosaurus and Taurocephalus. Its postero-ventral corner overlaps on the outer

surface of the quadratojugal. Dorsally, it is overlapped by the ventral edge of

the postorbital, and, further, posteriorly, it forms the thickened lower border

of the temporal fossa. From here it sweeps upwards to form most of the anterior

CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 37

face of the posttemporal arch and here its dorsal end overlaps the postero-

lateral flange of the parietal, where this bone forms the inner upper margin of

the temporal fossa. Postero-ventrally, the squamosal forms the thickened

rounded postero-ventral edge of the skull. This rounded border forms the lateral

wall of the wide and deep auditory groove, which groove lies mostly in the

squamosal. Medially to this groove, the squamosal forms a strong and prominent

ridge, medio-ventrally buttressed by the paroccipital and further dorsally

wholly formed by the tabular. This ridge, thus composed of squamosal, tabular

and paroccipital, forms the median wall of the auditory groove and from it

originated the strong depressor muscle of the mandible.

The tabular (Tab.) in dorsal view is seen to form the posterior half of the

dorsal part of the posttemporal arch, supporting the anterior half formed by the

flange of the parietal and the up-sweeping flange of the squamosal. In lateral

view, the tabular is seen to form the lateral part of the posterior edge of the

skull.

The interparietal (I.P.) in dorsal view, shows its upper edge where it forms

the posterior buttress to the parietal in the median part of the posterior margin

of the skull.

The quadratojugal (Q.J.) in dorsal and lateral views is seen to form the

antero-ventral corner of the ‘cheek’. Along its posterior border it is clasped by

the squamosal and its inner surface supports the quadrate. Its dorsal margin

does not abut against the jugal but is separated from it by a narrow incisure.

THE OccrpuT (fig. 3)

The occiput in Struthiocephalus forms a large surface, much broader than high

and nearly semicircular in outline. It is shallowly concave from side to side.

In the median line it is nearly ver-

tical with its dorsal edge slightly

further posteriorly and here it lies in

a plane nearly at right angles to the

plane of the maxillary alveolar bor-

der. In the median line there is a

ridge, which runs from the foramen

magnum to the upper edge of the

occipital surface. In whats: this

ridge is wedge-shaped, broad dor-

sally and tapering to the upper

edge of the foramen magnum,

a ee an eM eo whereas in akraalensis this ridge is

straight and narrow with a sharp

edge forming the median line with a deep depression lateral to it. In

this species the occiput has its ventral part situated much anteriorly so that it is

no longer nearly at right angles to the alveolar border but forms an obtuse

angle with the alveolar plane and an acute angle with the dorsal surface. This

may be due to post-mortem dorso-ventral pressure.

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

The condyle is directed postero-ventrally so that the skull would normally

hang somewhat downwards. It forms a stout rounded knob, dorsally excavated

by a groove leading into the foramen magnum. ‘The foramen magnum is large and

oval. The posttemporal fossae are small slits, bounded dorsally by the supra-

occipital and ventrally by the paroccipital. The lateral outer border of the

occipital plate is formed by the squamosal and median to this lies the deep

auditory groove, whose inner wall is formed by a strong and prominent ridge

to whose formation the tabular, squamosal and paroccipital contribute. Ven-

trally the condyles of the quadrates lie far anteriorly to the plane of the occiput.

In occipital view the basioccipital condyle forms the median part of the ventral

edge, and laterally the quadrate rami of the pterygoids together form a third

of the ventral edge of the skull.

In only one specimen (S.A.M. 11591) is the occiput well preserved, and even

here the sutures between the basioccipital and exoccipital and between the

supraoccipital and interparietal cannot be determined. :

The basioccipital (B.Oc.) apparently forms the whole of the condyle with no

participation by the exoccipital, which appears to be a small element lying

dorso-laterally in a plane anterior to that of the condyle. A groove on the

dorsal surface of the basioccipital leads into the foramen magnum. A notochordal

pit lies in the centre of a shallow concavity in the postero-ventral surface of the

condyle.

The supraoccipital (S.Oc.) appears to form the major part of the median

portion of the occipital plate. Laterally it stretches to the inner base of the

prominent ridge composed of the tabular, squamosal and paroccipital and

forming the inner rampart of the auditory groove, and here it meets the tabular

in a long vertical suture. In its ventro-lateral part the supraoccipital forms the

upper border of the slit-like posttemporal fossa. Medially and laterally to the

posttemporal fossa it meets the paroccipital and enters the upper edge of the

jugular foramen. Dorsally its junction with the interparietal cannot be deter-

mined, but with this bone it forms the median occipital ridge to which it

contributes the lower, narrower part.

The interparietal (I.P.) forms the dorso-median part of the occipit but it is

uncertain how far it stretches ventrally before meeting the supraoccipital. It

forms the dorsal part of the median ridge, which, in whaitst, is broad in its inter-

parietal part but in akraalensis is sharp and narrow throughout.

The tabular (Tab.) as I have determined its limits, forms only a small part

of the occipital surface as it appears not to stretch far medially. It forms the

greater part of the prominent lateral ridge lying medially to the auditory groove.

Dorsally the tabular flanks the postero-lateral parietal flange and the upsweeping

dorsal squamosal flange and thus forms the postero-dorsal part of the temporal

arch. This part of the tabular is especially strong in akraalensis, and here its

dorsal edge is wide, strong and rugose.

The paroccipital (P.Oc.) in occipital view is seen to be a strong bar medially

abutting against the basioccipital and stretching laterally to the lateral ridge

CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 39

where its dorso-lateral corner forms the most prominent part of this ridge. Its

ventro-lateral edge overlaps and supports the quadrate and dorsally it meets the

supraoccipital and forms the lower border of the small slit-like posttemporal

fossa and that of the small foramen jugale. Medially to the jugular foramen it

meets the ventral edge of the exoccipital.

The quadrate (Q) in occipital view shows a squarish posterior surface with,

ventrally, paired strong rounded knobs separated by a broad groove together

forming the ginglymoid articulatory surface for the articular. Laterally the

quadrate is overlapped by the squamosal and flanked by the quadratojugal.

Dorsally the posterior face of the quadrate is overlapped by the paroccipital.

Medially the long quadrate ramus of the pterygoid is applied to its inner surface

ventral to where the expanded distal end of the stapes abuts against the quadrate.

The quadratojugal (Q.J.) in posterior view is seen to form the latero-ventral

corner of the skull. Dorsally its posterior surface is overlapped by the squamosal.

The stapes (St.) is only partly exposed in occipital view. It is a stout rod with

its proximal end obscured by the paroccipital and its distal expanded end is

seen to be applied to the inner face of the quadrate.

The pterygoid (Pt.) only shows its long quadrate ramus in occipital view.

This is seen to extend very far posteriorly with its dorsal edge overlapping the

distal end of the stapes and its postero-lateral end applied to the inner face of

the quadrate. |

If my interpretation of the relations of the interparietal, supraoccipital,

tabular and paroccipital is correct the structure of the occiput in Struthiocephalus

differs greatly from the condition in Mormosaurus, Taurocephalus, Tapinocephalus

and Moschops. In these forms the tabular has a much greater occipital surface

and the supraoccipital is a much smaller bone. In Taurocephalus and Moschops

the tabular forms the whole of the dorsal border of the posttemporal fossa, but

in Tapinocephalus the tabular, as in Struthiocephalus, is excluded from the post-

temporal fossa.

THE VENTRAL SURFACE OF THE SKULL (fig. 4)

The palate and the basis crani lie in the same plane, with the strong lateral

pterygoidal rami extending ventral to this plane and the suspensorium lying

still further ventrally. Striking is the very anterior position of the articulatory

surfaces of the quadrates, which lie anterior to the posterior third of the skull.

The suborbital fossae are small, the choanae large and oval and the interpterygoid

vacuity is a narrow slit not extending between the prevomers.

The basioccipital (B.Oc.), in ventral view, is seen to carry a strong condyle

pear-shaped in outline. Postero-ventrally the condyle is circularly excavated

round the notochordal pit. Anteriorly to the condyle the basioccipital forms a

squarish plate of bone directed antero-ventrally to meet the surface of the

basisphenoid at an obtuse angle in a not very secure ankylosis. This surface

carries a low median ridge flanked by shallow oval depressions and the anterior

and lateral edges are rounded. Laterally the basioccipital is flanked by the

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

small exoccipital, whose limits are uncertain. Anterior to the jugular foramen

the basioccipital is strongly supported by the paroccipital which it meets in a

firm curved suture.

The basisphenoid (B.Sph.) posteriorly meets the basioccipital at an angle so

that the two ventral surfaces subtend an obtuse angle. The postero-lateral

corner of the basisphenoid forms the anterior border of the foramen ovale.

Anteriorly the basisphenoid extends as a blunt wedge in between the pterygoids

but the exact position of the suture is uncertain and its probable position is

eceywe

Fic. 4.—Struthiocephalus whaitsi. Ventral

view of the skull (S.A.M. 11591) (x §).

given in broken lines. In the median line the basisphenoid carries a low, sharp

keel, lateral to whose anterior end lie the carotid foramina.

The pterygoids (Pt.) form a large part of the ventral surface. In the middle

of the skull each pterygoid consists of a thin plate of bone meeting the basi-

sphenoid obliquely along a long edge whose exact position has not been deter-

mined but which I believe stretches in anterior direction to the median line

from the notch leading into the pituitary fossa. In the median line the pterygoid

meets its fellow to form a sharp median keel. Anterior to this keel lies the inter-

pterygoid slit. Lateral to the median keel the pterygoid is deeply excavated

and lateral to this wide and deep groove lies the quadrate ramus of the ptery-

goid. This is a deep sheet of bone lying obliquely in the skull nearly at right

CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 41

angles to the plane of the palate. The quadrate ramus extends far posteriorly

and here its outer surface is applied to the inner face of the quadrate, well

behind the plane of the condyle of the quadrate, and there is thus no anterior

process of the quadrate to meet the pterygoid. The distal end of the stapes thus

passes over the upper edge of the quadrate ramus of the pterygoid to reach the

medial face of the quadrate. In no other Deinocephalian is the quadrate ramus

of the pterygoid known to extend so far posterior to the quadrate condyle,

which fact emphasizes how far the quadrate has shifted in an anterior direction.

The lateral ramus of the pterygoid is only moderately strong and has no

great ventral extent. This is in strong contrast to the condition in the Titano-

suchids where the ramus is very strongly developed. In Struthiocephalus the ramus

does not extend far laterally as it does in Taurocephalus and all the ‘Titanosuchids.

The lateral edge of the ramus is supported by a descending process of the ecto-

pterygoid, which, on account of the narrowness of the lateral pterygoid ramus,

has a greater palatal face than in all other Deinocephalians. As in all Deino-

cephalians the lateral ramus is connected with the quadrate ramus by a web

of bone reducing the size of the suborbital vacuity of which it forms the antero-

median border. Anteriorly the exact limits of the pterygoid are indetermined

but are probably as indicated by broken lines in the figures.

The ectopterygoid (Ec.Pt.) has a larger palatal surface than in other Deino-

cephalians. Its anterior and median limits are not clearly shown but it appears

to descend along the lateral edge of the lateral pterygoid ramus which it

buttresses. Its posterior edge forms the antero-lateral border of the suborbital

fossa. Postero-laterally it abuts against the jugal in a sigmoid suture and

laterally against the maxilla.

The palatine (Pal.), from where the suture with the pterygoid and ecto-

pterygoid appears to lie, stretches antero-laterally as a thickened bone to form

the rounded lateral two-thirds of the choanal border, and has its lateral edge

applied to the inner maxillary surface where it flanks the alveolar border. Near

the median line the palatine ends with a short sharp ridge running parallel

to the median line. The two palatines thus do not meet each other, as median

to their inner ridged borders a posterior tongue of the prevomers intervenes.

Just lateral to the ridged inner edge there lies a rounded mound on which there

are indications of the roots of a small number of small palatine teeth.

The prevomers (P.V.) are strong elements together forming a massive inter-

choanal bar. Anteriorly they underlie the inner surface of the premaxillaries

with their anterior edges bevelled. Posteriorly they widen and overlie the

palatines postero-laterally and in the median line send a tongue posteriorly in

between the ridged inner edges of the palatines. Anteriorly the median suture

is open and the interchoanal bar is here grooved, whereas in the posterior half

of the bar a keel is developed along the median line.

The premaxillary (P.Mx.) alveolar border is very massive. Each premaxilla

carries three strong teeth which are directed antero-ventrally. Posterior to the

functional teeth there are indications of crowns which may be either replacing

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

or replaced teeth. In a juvenile specimen of whaitsi (S.A.M. 11493) where the

crowns of the teeth are just erupting the labial edge of the premaxillaries is

sharp and appears to form a cutting edge functioning as such until the teeth are

sufficiently developed. .

The maxilla (Mx.) has its alveolar border anteriorly massive and wide but

then it tapers rapidly in posterior direction, and behind the last tooth a sharp

edge is continued by the jugal, sweeping down towards the quadrato-jugal.

Stumps of teeth and infilled alveoli in most specimens show that there were ten

to eleven maxillary teeth. The anterior four are large, the fifth appreciably

smaller, the sixth abruptly smaller and the series then decreases gradually in

size in posterior direction. In the juvenile specimen of whaits: (S.A.M. 11493)

the labial edge of the maxilla, as is the case in the premaxilla, is sharp and

during immaturity apparently forms a cutting edge.

The jugal (J) in ventral view has a narrow and deep flange of bone sweeping

from the sharp outer edge of the maxilla down towards the quadratojugal.

Internally and dorsally to this sharp edge the jugal is thickened and extending

internally forms the lateral border of the suborbital fossa, and anteriorly it meets

the ectopterygoid in a sigmoid suture.

The quadrate (Q.) has its articulatory condyle very prominent in ventral view

as this forms the most ventral part of the skull. The articulatory surface has a

median trochlear surface bounded internally and externally by longitudinally

oval condyles. ‘The rounded articulatory surfaces are sharply demarcated from

both the posterior and the anterior face of the upper part of the bone. Laterally

a ridge bounded medially and laterally by a groove lies lateral to the condyle

in the plane of the ventral edge of the quadratojugal. Dorsally the posterior

surface of the quadrate is seen wedged in between the paroccipital internally

and the quadratojugal and squamosal externally. Internally the long quadrate

ramus of the pterygoid is applied to the medio-posterior face of the quadrate.

The extremity of the ramus extends far posteriorly of the plane of the quadrate

condyle and there is no anterior process of the quadrate to meet the quadrate

ramus of the pterygoid. No foramen quadrati could be located with certainty but

it may be small and may lie just medio-dorsally of the ridge on the quadrate

lying laterally to the condyle.

The stapes (St.) is in position in S.A.M. 11591 on both sides. It is a stout rod

with expanded distal and proximal ends. The distal end applied to the inner

face of the quadrate is underlain by the long quadrate ramus of the

pterygoid and has its postero-distal corner much expanded. No foramen can be

determined.

The quadratojugal (Q.J.) in ventral view is seen to form the angle of the

‘cheek’. Its ventral edge lies in a plane dorsal to the condyle of the quadrate

from which it is separated by a ridge bounded on both sides by a groove. The

posterior surface of the quadratojugal is seen to be overlapped by the descending

posterior squamosal process.

CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 43

The squamosal (Sq.) in ventral view forms the outer edge of the posterior

third of the skull. Median to this edge lies the auditory groove whose inner wall

is formed by a ridge formed by the squamosal, paroccipital and tabular.

Further ventrally the squamosal supports the posterior face of the quadratojugal

and quadrate.

The paroccipital (P.Oc.) shows a much greater face in ventral view than it

does in occipital view. This rotation of the paroccipital from the occipital plane

on to the ventral surface is one of the results of the forward shift of the quadrates

and is a character distinguishing all Deinocephalians from the other Therapsids.

Fic. 5.—Struthiocephalus whaitsi. The right side of the braincase in

lateral view (S.A.M. 11591) (x4). The suborbital bar, postorbital

bar and occipital plate fractured and here seen in section. The right

quadrate ramus of the pterygoid and the right stapes are also seen

in section where they have been fractured.

The paroccipital is a strong massive element acting as a very firm connecting

link between the bones of the ‘cheek’ and the suspensorium lying laterally and

the basioccipital of the cranial base. Posteriorly the thickening of the paroccipi-

tal and supraoccipital have all but obliterated the posttemporal fossa which is

only preserved as a narrow slit. The postero-lateral corner of the paroccipital

forms the most prominent part of the strong ridge from which the depressor

mandibulae originates. Antero-medially the paroccipital forms the posterior

half of the border of the foramen ovale.

In ventral view the tabulars, interparietal and supraoccipitals are seen to lie

well posterior to the basioccipital condyle with the first two forming the posterior

edge of the skull which, as is the case in most Tapinocephalids, is nearly a

straight line, whereas in the Titanosuchids this edge is concave. In akraalensis

more of the occiput is seen in ventral view than is the case in whaitst.

THE BRAINCASE IN LATERAL VIEW (fig. 5)

In a specimen of whaitsi (S.A.M. 11591) a fracture through the posttemporal

arch, postorbital and suborbital bars has enabled me to prepare the lateral

surface of the braincase on the right side. But after the removal of the intractable

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

matrix from the temporal fossa and orbit the surface of the internal bones thus

exposed is not sufficiently clear so as to determine the limits of the constituent

bones with any great degree of certainty. The accompanying figure shows what

structural details have been determined. The parasphenoid (P.S.) is the largest

element forming a large part of the fenestrated septum. A dorsal sheet of bone

supports the ventral edge of the sphenethmoid and a well-developed anterior

process is directed obliquely forwards. Of this part of the parasphenoid Efremof

says that in many cases, ‘den vordere Forsatz des Parasphenoid knorpelig

blieb’, whereas in fact the parasphenoid is an os investitiens and thus not an

element preformed in cartilage.

Of the sphenethmoid (S.E.) only the lower part can be seen where it rests on

the upper edge of the dorsal parasphenoidal process.

The prootic (P.O.) is seen wedged in the postero-dorsal corner and its rela-

tions with the sphenethmoid are uncertain.

The opening for the trigeminus and the fenestra of the fossa hypophyseos

are situated as shown in the figure.

THE Lower JAw

In S.A.M. 11693 most of the dentaries are preserved and in 8.A.M. 11493

the crushed posterior half of the right mandibulary ramus is present, but in both

only the outer surface could be prepared. What could be determined of the

structure I have included in the composite figure accompanying the description

of Struthtocephaloides duplessist.

The hinge of the lower jaw lies very far forward, in the plane of the orbit.

The dentary forms nearly two-thirds of the ramus and its mentum is massive

and fairly upright. The teeth are directed upwards and

only slightly outwards to intermesh with the labially

directed teeth of the upper jaw. The angular has a large

outer flange and the surangular has a strong rounded

dorsal border curving upwards and forwards from the

articular.

THE DeEntTiTION (fig. 6)

In all specimens stumps, imperfect crowns and empty

alveoli are all that are preserved. In fragments of the

Fic. 6.—Struthiocepha- lower jaw of S.A.M. 11591 a few crowns of the teeth

lus whaitsi. A tooth i :

of the lower jaw of the lower jaw are preserved and one is figured here.

(S.A.M. 11591) All the teeth have a labial talon or pointed cusp at

(x4). a, lingual } i :

view. 6, lateral view. whose base there lies lingually a cup-shaped base with

a serrated edge.

The three teeth in the premaxilla and the first four maxillary teeth are large

with long labial cusps. Then abruptly the fifth tooth is much smaller with a

short labial talon and then the rest of the teeth decrease still further in size

posteriorly.

CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 45

A tooth just erupting in the lower jaw (S.A.M. 11591), which is thus as yet

unabraded, shows three longitudinal ridges on the lingual surface of the talon.

With use the talon is abraded and the ridges disappear. Through use the

serrations on the labial edge of the cup are also worn down and lost.

There are indications that in the anterior teeth there can be at least three

successive sets of teeth.

TAXONOMIC DIAGNOSES

As Mormosaurids I have grouped together (1936) the genera Mormosaurus,

Struthiocephalus and Taurocephalus. The following amended diagnosis for this

group is suggested: Skull large, long and moderately wide; snout long or

fairly long and shallow; cranial bones strongly thickened with parietal, naso-

frontal and postorbital bosses undeveloped, distinctly developed or tending to

coalesce in the general pachyostosis; postorbital bar moderately wide to wide

and massive; the facial surface fairly or very abruptly demarcated from the

thickened cranial surface; temporal fossae fairly small with the dorso-ventral

diameter appreciably or not very much greater than the antero-posterior;

intertemporal region fairly narrow to moderately wide and laterally distinctly

or only slightly bayed; parietals entering supratemporal border; quadrate

ramus of the pterygoid extending only up to or well posterior to the quadratic

condyle, which is situated far forward; tabular entering or not entering the post-

temporal fossa; no differentiation of teeth into incisors, canines and postcanines.

GENERIC DIAGNOSES

A. Mormosaurus

1. Snout short, very shallow, facial surface not extending posterior to the

anterior orbital border, very abrupt transition from the facial to the cranial

surface.

2. Dorsal cranial surface very strongly pachyostosed, with the centres of

thickening coalesced and transition on to face very abrupt along a very

definite transverse line forming a transverse wall.

3. Additional pachyostosis:

a. Nasal boss laterally confluent with the thickening of the postorbital bar

and the prefrontal and thus forming a transverse wall from orbit to

orbit.

6. Parietal boss not very distinct, confluent with the general machyostasts

of the parietal and frontal.

c. Postorbital bar wide and massive but without distinct bulbous boss and

thus flowing evenly on to the general dorsally thickened surface.

d. Posttemporal arch greatly thickened and rugose.

e. Orbits not visible in dorsal view and in dorsal view the postorbitals do

not form the lateral border of the skull.

f. Antero-posterior diameter of temporal fossa small and fossa transversely

oval.

g. Dorsal parietal surface fairly wide, laterally indistinctly bayed.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Dentition feeble, uniform.

Quadrate moderately far forward with the quadrate ramus of the pterygoid

not extending much posterior to the plane of the quadratic condyle, not

underlying the distal end of the stapes. ;

The lateral ramus of the pterygoid does not form a prominent transverse bar

and the width across the transverse rami is small.

The intersquamosal width is large.

The basioccipital condyle is directed posteriorly.

The frontal appears to be small and is excluded from the orbital border, the

supraoccipital is wide and low, the tabular with a moderate occipital face

and apparently enters the posttemporal fossa, the lacrimal meets the nasal,

the squamosal with an anterior wedge into the jugal. (This diagnosis is

based on Watson’s description and figures.)

B. Struthiocephalus

Snout long, shallow, facial surface extending far back, posterior to the

anterior orbital border, fairly abrupt transition on to the cranial surface.

Dorsal cranial surface strongly pachyostosed, but centres of greatest thicken-

ing still distinct and the transition on to the face fairly abrupt but not along

a definite line and thus not forming a transverse wall.

Additional pachyostosis :

a. Naso-frontal boss not laterally confluent with the pachyostosis of the

postorbital bar and the prefrontal and thus not forming a transverse wall

from orbit to orbit.

b. Parietal boss distinct.

Postorbital bar moderately wide to wide, fairly massive to massive, with

or without distinct bulbous boss and with or without a ridge de aneotne

it from the dorsal surface.

d. Posttemporal arch moderately to shea thickened, fairly smooth to

rugose.

e. Orbits just visible in dorsal view, and in dorsal view the postorbitals do

not form the lateral border of the skull.

f. Antero-posterior diameter of temporal fossa moderate to fairly small and

fossa oval to slitlike.

g. Dorsal parietal surface narrow to moderately wide, laterally distinctly

or indistinctly bayed.

Dentition well developed with 14 teeth of which 3 are on the premaxilla;

the fifth maxillary tooth is abruptly smaller than those anterior to it and

then posteriorly the series gradually decreases in size.

Quadrate far forward with the quadrate ramus of the pterygoid extending

much posterior to the plane of the quadratic condyles, underlying the distal

end of the stapes.

The lateral ramus of the pterygoid forms a strong and prominent bar but

the width across the transverse rami is small.

{olny Ay

CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 47

. The intersquamosal width is moderate to fairly moderate.

. The basioccipital condyle is directed postero-ventrally.

. The frontal is of medium size and enters the orbital border, the supra-

occipital is wide but apparently fairly high, the tabular with a small occipi-

tal face and excluded from the posttemporal fossa, the lacrimal meets or

does not meet the nasal, the squamosal without anterior wedge into the

jugal.

C. Taurocephalus

. Snout of medium length, fairly high, facial surface not extending posterior

to the anterior orbital border, transition on to cranial surface not abrupt

but through a gentle curve.

. Dorsal cranial surface fairly strongly pachyostosed, but centres of greatest

thickening still distinct and the transition on to the face through a gentle

curve and no transverse wall is thus formed.

. Additional pachyostosis:

a. No distinct naso-frontal boss present, and from orbit to orbit the dowel

surface is evenly convex.

__b, The parietal boss forms a rounded mound, highest round the pineal

__ border but with its edges running into the general parietal surface.

c. Postorbital bar fairly wide and massive, though extending very far

laterally it is dorsally not bulbous and flows evenly on to the generally

thickened dorsal surface; the width over the postorbitals is very great

and in dorsal view they form the lateral edge of the skull.

d. Posttemporal arch moderately thick and smooth.

e. Orbits just visible in dorsal view anterior to the postorbital bar which

here forms the lateral border of the skull in dorsal view.

jf. Antero-posterior diameter of the temporal fossa fairly great and thus

broadly oval.

- g. Dorsal parietal surface wide and laterally disenctly bayed.

. -Dentition well developed with 21 teeth of which 4 are in the premaxilla;

there is an evenly graded decrease in size in posterior direction.

Quadrate not very far forward; the quadrate ramus of the pterygoid not

extending pésterior to the plane of the quadratic condyles, only underlying

the anterior corner of the distal end of the stapes; quadrate with anterior

process meeting the quadrate ramus of the pterygoid.

. The lateral ramus of the pterygoid forms a strong and prominent bar and

the width across the transverse rami is large.

. The intersquamosal width is moderate, just more than the inter-postorbital

width.

. The basioccipital condyle is directed posteriorly.

. The frontal is large and enters the orbital border, the supraoccipital is

narrow and high, the tabular has a large occipital face and enters the post-

temporal fossa, the lacrimal does not meet the nasal, the squamosal with a

sharp anterior wedge into the jugal.

ANNALS OF THE SOUTH AFRICAN MUSEUM

D. Struthiocephaloides®

Snout long, shallow, facial surface extending far back, posterior to the

anterior orbital border, transition on to the cranial surface not abrupt.

Dorsal cranial surface strongly pachyostosed, but centres of greatest thicken-

ing still fairly distinct, but the transition on to the face not abrupt, with no

indication of a transverse interorbital step.

Additional pachyostosis:

a. No naso-frontal boss (but with a swelling just anterior to the posterior

end of the premaxilla in cavifrons).

b. Parietal boss distinct in cavifrons but in duplessist undeveloped or indistinct

due to confluence with the general pachyostosis of the parietal and

frontal.

c. Postorbital bar very wide, massive, but without a bulbous boss in its

dorsal part.

d. Posttemporal arch moderately to strongly thickened, fairly smooth.

e. Orbits just to plainly visible in dorsal view, and in dorsal view the post-

orbitals do not form the lateral border of the skull.

f. Antero-posterior diameter of temporal fossa moderate to fairly large,

fossa oval or narrow (in cavifrons the fossa lies obliquely in the skull).

g. Dorsal, parietal surface wide, laterally not pinched in.

Dentition well developed with 14 teeth of which 3 are in the premaxilla,

and from the 5th the teeth gradually decrease in size.

Quadrate far forward, with the quadrate ramus of the pterygoid extending

much posterior to the plane of the quadratic condyles.

The lateral ramus of the pterygoid forms a fairly strong and prominent bar

but the width across the transverse rami is small.

The intersquamosal width is moderate to large.

The basioccipital condyle is directed much ventrally.

The frontal is fairly small, entering the orbital border in duplessist but not in

cavifrons, the tabular excluded from the temporal fossa, the lacrimal does not

meet the nasal, the squamosal without anterior wedge in to the jugal.

E. Struthionops

Snout fairly short and high, facial surface not extending posterior to anterior

orbital border, transition to cranial surface not abrupt but through a gentle

curve.

Dorsal cranial surface moderately pachyostosed, centres of greatest thickening

distinct, no interorbital step.

Additional pachyostosis:

a. Naso-frontal boss very low and not confluent with the thickening on the

prefrontals.

b. The parietal boss is a fairly prominent mound, highest round the pineal

foramen and with its outer edges running into the general parietal

surface.

Ann. S. Afr. Mus., Vol. XLII

Plate

XVII

eee

RECONSTRUCTION OF STRUTHIOCEPHALUS

1 as exhibited in the South

laster of the whole anima

ion in p

. Modelled by the author.

ife-sized reconstruct

from the |

African Museum

Photograph of the head of Struthiocephalus taken

CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 49

Postorbital bar fairly narrow and slender with dorsal swelling.

d. Posttemporal arch fairly weak and smooth.

e. Orbits large and well visible in dorsal view, postorbitals form the lateral

border of the skull as seen in dorsal view.

f. Antero-posterior diameter of temporal fossa fairly large and broadly oval.

g. Dorsal parietal surface wide, but laterally slightly, pinched in.

4. Dentition with probably 14-15 teeth evenly decreasing in size in posterior

direction.

5. Quadrate not far forward.

6. The lateral ramus of the pterygoid strong and prominent but transverse

width across the rami moderate.

The intersquamosal width is large, just more than the inter-postorbital width.

The basioccipital condyle is not preserved.

g. The frontal is large, but is just excluded from the orbital border, the lacrimal

just meets the nasal, the squamosal without a wedge into the jugal.

DIAGNOSES FOR THE SPECIES OF STRUTHIOCEPHALUS

S. whaitsi

1. Naso-frontal, parietal and bulbous boss on postorbital prominent.

2. Snout moderately long, shallow and fairly weak.

3. Moderately broad over squamosals.

4. Pineal foramen moderately far from occipital border.

5. Postorbital bar wide.

6. Temporal fossa fairly rounded.

7. Median occipital ridge wedge-shaped with rounded edges.

8. Nares well back.

9g. Quadrate well forward.

10. Interorbital width large.

11. Interparietal width small.

12, Posttemporal arch moderately strong.

S. rheederi

Naso-frontal, parietal and bulbous boss on postorbital prominent.

Snout fairly long, moderately shallow but broad.

Broad over squamosals.

Pineal foramen very near occipital border.

Postorbital bar probably wide (not evident from Broom’s account).

Temporal fossa slit-like.

Median occipital ridge not figured or described by Broom.

Nares very far back.

Quadrate not figured or described by Broom.

10. Interorbital width very large.

11. Interparietal width large.

12. Posttemporal arch moderately strong.

SOS A Pee a es

S. milleri

1. Naso-frontal and parietal bosses very prominent and bulbous boss on postorbital bar

moderately prominent.

Snout very long and very shallow.

Relatively narrow over squamosals.

Pineal foramen near occipital border.

Postorbital bar narrow.

Temporal fossa fairly rounded.

Median occipital ridge not figured or described by Olson and Broom,

Nares fairly near premaxillary edge.

Quadrate not figured or described by Olson and Broom.

5O ANNALS OF THE SOUTH AFRICAN MUSEUM

10. Interorbital width small.

11. Interparietal width fairly small.

12. Posttemporal arch robust.

‘S. akraalensis

1. Naso-frontal boss large but low, parietal boss prominent, postorbital bar dorsally

greatly thickened with the development of a longitudinal ridge raised above the

surface of the frontal which thus appears excavated.

2. Snout very long and shallow.

3. Broad over the squamosals.

4. Pineal foramen far from occipital border.

5. Postorbital bar very wide and strongly rugose.

6. ‘Temporal fossa slit-lke.

7. Median occipital ridge straight with sharp median edge.

8. Nares far back.

g. Quadrates very far forward.

10. Interorbital width very large.

11. Interparietal width very large.

12. Posttemporal arch very robust.

THe Curer MEASUREMENTS COMPARED

whaitst rheedert milleri akraalensts

Length 558 624 480 655

Width : 345 432 300 390 ©

Interorbital width . 170 180 130 175

Interparietal width. : Sete ste OO 140 105 167

Pineal foramen to occipital edge shine ti 65 48 - 44 go

Narisito Bo Mixedge uae 85 114 52 95

Keys To FActnirATE Rapip TAxonomic IDENTIFICATION

Deinocephalia. Large therapsids, with pachyostosed skulls, with large quadrates,

quadratojugals forming the corner of the ‘cheek’, premaxillaries with

long facial exposure.

Deinocephalians with differentiated carnivorous dentition, cranial bones

not greatly pachyostosed, snout long, low parietal crest, ‘quadrates not

situated far anteriorly.

Medium-sized to large Deinocephalians with undifferentiated herbi-

vorous dentition, cranial bones slightly to very greatly pachyostosed, in

all but the Moschosaurids the snout is weakened, there is no parietal

crest (except in Riebeeckosaurus and Avenantia); the quadrates are

situated moderately to very far anteriorly.

‘Titanosuchia.

Tapinocephalia.

GROUPS IN THE TAPINOCEPHALIA

Little pachyostosis

Great pachyostosis

Facial bones thickened to run evenly on to the cranial

surface

Facial bones not so thickened as to run 1 evenly on to the

cranial surface .

Snout very much shortened and ‘weakened

Snout not very much shortened and weakened

oF BP Po

GENERA OF THE MORMOSAURUS-GROUP

Skull with abrupt step from face to cranial surface

Skull without abrupt step :

Distinct naso-frontal boss present

Distinct naso-frontal boss absent ‘

Width over postorbitals not very great and postorbitals not

forming lateral edge of skull in dorsal view .

Width over parietals great and ieee forming lateral

edge of skull in dorsal view :

Width across jugals much less than across postorbitals

Width across jugals not much less than across postorbitals

D RYN so

Moschosaurus-group

Moschops-group

T apinocephalus-group

. | Mormosaurus-group

Mormosaurus

Struthiocephalus

Struthiocephaloides

T aurocephalus

Struthionops

CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS Hy

THE SPECIES OF STRUTHIOCEPHALUS

ie onout very Jong iand, very, shallow) 33%) .) G4. mallert

2. Snout not very long and very shallow . Ae aie :

3. With prominent ridge median to ee boss Ne 12) akraalensis

4. Without this ridge ' hs ey ed) 8

5. Pineal foramen very near occipital border . Bian NOs bi oy WIMECUENE

6. Pineal foramen not near occipital border . . . . . = whaitsi

DIscussIoN

It would be unwise to attempt a discussion of the relationships of the Deino-

cephalians with other orders and inter se until all the material in the South

African Museum has been studied. At this stage I am confining my remarks to

some points which the study of Struthiocephalus has brought to the fore.

Age in the Struthiocephalus skull

A specimen of whavist (S.A.M. 11493) has a skull just as long as that of the

type (S.A.M. 2678) and another specimen referred to this species (S.A.M.

11591), but in it the upper teeth are just commencing to erupt and the outer

edge of both premaxillaries and maxillaries is sharp and appears to form a

functioning cutting edge until the teeth are fully erupted. All the cranial bones

are still little affected by any pachyostosis. The postorbital bar is narrow and

lightly built with only its dorsal part slightly expanded and here the postfrontal

is large and has not yet overgrown the postorbital. ‘The posterior flange of the

postorbital forming the dorsal border of the temporal fossa is little reduced and

though not meeting the squamosal extends far posteriorly and is not encroached

on or overhung by the parietal. ‘The antero-posterior diameter of the temporal

fossa is nearly equal to the dorso-ventral diameter. The posttemporal arch is

not thickened and does not encroach into the fossa. ‘The quadrates still lie in

a plane posterior to the anterior orbital border. The whole outer surface is

still smooth and free from rugosities. Unfortunately the state of preservation

does not reveal the structure of the occiput and the ventral aspect of the skull.

What could be determined in this juvenile skull is however sufficient to show

that many of the characters peculiar to the adult Tapinocephalian skull are

mainly due to the pachyostosis which increasing age brings about and which

obscures its essential therapsid nature.

The Cranial Pachyostosis

Apart from the fact that in both Moschops and Struthiocephalus an intensifica-

tion of the thickening of the cranial bones takes place during the life of the

individual, it also appears to be a process that can be traced phyletically.

In Moschosaurus, which is undoubtedly the most primitive of the South African

Tapinocephalians, there is very little pachyostosis and in Agnosaurus it is also

slight, whereas in all the other known forms a lesser or greater degree of bone-

thickening is apparent. Less of the skull is affected in the Mormosaurus-group

than in the Moschops- and Tapinocephalus-groups. The thickening chiefly affects

the supra- and interorbital region, the intertemporal skull roof, the post-

orbital bar; the posttemporal arch and the face and ‘cheek’ are relatively little

52 ANNALS OF THE SOUTH AFRICAN MUSEUM

affected. The thickening in these regions very materially affects the size and

shape of the temporal fossa, the orbits, the braincase and the mechanism of the

lower jaw.

The Supra- and Interorbital region

In the Mormosaurus-group the moderate to fairly strong pachyostosis lies

posterior to the plane of the anterior orbital border. It is least in Struthiocephalus,

Struthiocephaloides, and Struthionops, and here the relatively unaffected snout

extends posteriorly to the plane of the anterior orbital border where the transi-

tion on to the supraorbital thickened part is not very abrupt. This transition is

not very abrupt chiefly because the pachyostosis has not affected the lacrimal,

prefrontal, nasal and frontal to any great extent, only the posterior part of the

prefrontal and nasal and the medio-anterior part of the frontal being affected.

The thickening here does not extend transversely across the skull from orbit to

orbit, but is in evidence at three separate centres, viz. the tops of the two post-

orbital bars and in the naso-frontal boss. In Taurocephalus more of the prefrontal

and the posterior part of the nasals are affected and the above-mentioned three

centres tend to coalesce and the transition from the face on to the cranium is by

a gentle curve. In Mormosaurus the process has extended further and the pachyos-

tosis extends anterior to the plane of the anterior orbital border with confluence

of the three centres to form a very distinct transverse step raised high above the

surface of the snout.

In the Tapinocephalus-group this process has progressed much further in

Tapinocephalus to include all the prefrontal, frontal and the posterior part of the

nasal to a plane well in advance of the anterior orbital border and the descent

on to the surface of the snout is very abrupt from the very high transverse

rampart. (But in Keratocephalus this region has remained essentially Stéruthio-

cephalus-like with an accentuation of the naso-frontal boss.)

In the Moschops-group the pachyostosis is continued into the bones of the snout

but is so graded that instead of forming a step there is an even curve from the tip

of the snout on to the supra- and interorbital region.

The Intertemporal Skull Roof

In Moschosaurus and Agnosaurus the intertemporal width is small with the sides

pinched in and with little thickening of the parietal. In the Mormosaurus-group

this width is increased with the stages in this process shown by the five genera

Struthiocephalus, Struthtocephaloides, Struthionops, Taurocephalus and Mormosaurus, in

this order. In Struthiocephalus this widening has advanced least but the direction

of this process is evident in the series formed by its constituent species. The loss

of the dorsal bay to the temporal fossa can be traced through the species of

Struthiocephalus to Mormosaurus.

In the Tapinocephalus-group the intertemporal width attains its greatest

dimensions and there is no sign of the bay.

In Moschops a parallel process of widening of the intertemporal region has

taken place but there is still evidence of the bay.

CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 53

The thickening of the parietal is at first mainly in evidence round the pineal

foramen. In Struthiocephalus the thickness of the parietal at the pineal foramen

varies from 40 to 80 mm. and in a Tapinocephalid species from Gunyanka’s

Kraal in Southern Rhodesia it reaches the enormous thickness of 310 mm. With

the increasing pachyostosis of the parietal the boss round the pineal foramen

tends to become engulfed by the general thickening and in the Rhodesian

specimen there is little evidence of a separate pineal boss.

The Postorbital Bar

The postorbital bar is light and slender in Moschosaurus and Agnosaurus,

moderate in Taurocephalus and Struthionops, strong to very strong in Struthio-

cephalus and Struthiocephaloides, very strong and massive in Mormosaurus, and very

wide, strong and very massive in Tapinocephalus. In Struthiocephalus the dorsal

end of the bar formed by the postfrontal has a tendency to form a bulbous boss

reminiscent of the condition in the Titanosuchid Anteosaurus, but in Tapino-

cephalus especially this is incorporated in the general extensive pachyostosis.

An increasing massiveness of the posttemporal arch parallels that of the post-

orbital bar, and together they have the effect of reducing the antero-posterior

diameter of the temporal fossa.

Adverse Effects of the Pachyostosis

The downward growth of the roof-bones, together with the reduction of the

angle between skull roof and occiput, boxes in the braincase in the dorso-

posterior corner of the skull. The encroachment of the posttemporal arch and

postorbital bar very greatly reduces the size of the temporal fossa and conse-

quently of the adductor mandibulae. The thickened bone overhanging the

orbits very materially restricts the field of vision. No doubt the increasing

pachyostosis indicates the road which led to extinction.

REFERENCES

14 Watson, D. M.S. P.Z.S., p. 749. 1914.

1b Haucuton, S. H. Ann. S. Afr. Mus., xii, 2, p. 52. 1915.

1¢ Grecory, W. K. Bull. Am. Mus. Nat. Hist., 61, p. 179. 1926.

id Broom, R. Mammal-like Reptiles of South Africa, p. 18. 1932.

1€ BoonstRA, L. D. Bull. Am. Mus. Nat. Hist., 72, p. 75. 1936.

if Erremov, J. A. Nova Acta Leopoldina, N.F. 9, 39, p. 155. 1940.

* Boonstra, L. D. Ann. Mag. Nat. Hist., xii, 5, p. 455. 1952.

3 Boonstra, L. D. ibid., p. 509. 1952.

4 Boonstra, L. D. S. Afr. Four. Sci., 48, 8, p. 247. 1952.

5 Broom, R. P.Z.S., p. 302. 1937.

§ Otson, E. C., and Broom, R. Journ. Pal., 4, 7, p. 615. 1937.

7 Havucuton, S. H. Report of the South African Museum for 1915, p. 4. 1916.

8 Boonstra, L. D. Tydskr. Wet. Kuns, xii, 2, p. 237. 1952.

® Boonstra, L. D. ibid. xii, 2, p. 246.

10 BoonstrA, L. D. Ann. Mag. Nat. Hist. xii, 5, p. 988. 1952.

11 Boonstra, L. D. Tydskr. Wet. Kuns. xii, 2, p. 242. 1952.

6. The Lower Faw Articulatory Region in some Pristerognathid Therocephalians. By

LiEuwE D. Boonstra, D.Sc.

(With 4 text-figures)

In the course of the preparation of a paper, mainly of a taxonomic nature,

for the Annals of the South African Museum on the hundred-odd specimens of

Therocephalians from the Tapinocephalus-zone preserved in the South African

Museum, I have determined certain points in the structure of the posterior part

of the lower jaw and the relations of the quadrate, quadratojugal and squamosal.

A short account of the structure in some of the better-preserved specimens is

presented here. New genera and species mentioned here will be fully described

in the forthcoming taxonomic paper referred to above.

SCYMNOSAURUS FEROX (fig. 1)

A specimen collected by me on the farm Rietkuil in the district of Beaufort

West (S.A.M. 9084) consists of a fairly good skull and parts of some limb-bones.

By direct comparison with the type in our collection I have identified it with

Scymnosaurus ferox. Natural weathering has partly exposed the mandibular

articulatory region, but also unfortunately removed the lateral surface of the

quadrate, quadratojugal and the lateral edge of the squamosal.

The dentary has a concave posterior edge overlapping the angular and its

dorso-posterior coronoid process does not extend above the subtemporal arch

but is directed more posteriorly in the direction of the squamosal.

The angular hasa fairly large lateral face with a prominent dorso-ventral ridge

and a lesser ridge roughly at right angles. The anterior half of its ventral edge

is fairly thick, but the posterior half has a double thin edge—the outer being the

reflected flange. Posteriorly the outer (reflected) flange extends to the surangular

and ventrally it underlaps the articular. Dorsally there is a small notch, anterior

to which the angular apparently overlapped the outer surangular face. This

dorsal notch, not exposing the outer surface of the inner angular sheet, but

instead the external surangular surface, thus has different relations than in other

Pristerognathids where this region has been described. A notch in the ventral

edge indicates the level to which the reflected flange usually extends posteriorly

in other Pristerognathids.

Externally the surangular shows a thickened rounded and curved dorsal and

postero-dorsal girder forming the edge of the mandible. Further ventrally the

surangular curves firmly round the lateral surface of the articular and, with the

internal prearticular, transmits the stresses arising anteriorly to the articular

bone.

The articular is a comparatively small bone of peculiar shape; externally a

tongue extending dorsally is anteriorly clasped by the surangular and posteriorly

ARTICULATORY REGION IN PRISTEROGNATHID THEROCEPHALIANS

I b.

Fic. I.—Scymnosaurus ferox. S.A.M. 9084, Rietkuil, Beaufort West. (x # nat. size.)

a. Lateral view. The lateral surface of the quadrate, quadratojugal and squamosal has

been weathered away so that these elements are seen in section.

b. Cross-section of above at A.B.

c. Oblique postero-lateral view. The lateral edge of the squamosal, quadrate and

quadratojugal weathered and seen in section.

These and subsequent figures are all ortho-projections taken with a pantograph.

An.—angular. Art.—articular. B.Oc.—basioccipital. B.Sph.—basisphenoid. Cor.—coronoid.

D.—dentary. Ep.Pt.—epipterygoid. Ex.Oc.—exoccipital. F.J.—foramen jugale. I.P.—inter-

parietal. J.—jugal. Pa.—parietal. P.O.—postorbital. P.Oc.—paroccipital. Pr.Art.—prearticular.

Pr.Ot.—prootic. P.T.F.—posttemporal fenestra. Pt.—pterygoid. Q.—quadrate. Q.J.—quadrato-

jugal. S.A.(n)—surangular. S.Oc.—supra-occipital. Sq.—squamosal. St.—stapes. Tab.—tabular.

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

forms the front face for the articulation with the quadrate condyle; it then

extends posteriorly to form the ventral part of the articulatory surface, but no

retro-articular process is developed. In posterior view it is seen that the articular

has a dorsal spur, which, if it extends anteriorly, must form a ridge dividing the

articulatory surface into a smaller lateral concavity and a larger median

concavity. The quadratic condyle would thus be bipartite with a smaller

rounded outer and an inner larger roller-shaped condyle. Medio-ventrally the

under-surface of the articular is clasped by the prearticular, which transmits the

greater part of the thrust from the anterior part of the mandible to the articular.

Only part of the quadrate is exposed in lateral and in posterior view. Its

lateral surface has been somewhat weathered away so that the drawing (fig. 14)

really shows a section face. The apparent bipartite nature of the quadratic

condyle has been noted above. In section the outer condyle has a circular

outline. From the antero-dorsal corner of this condyle the lateral edge of the

quadrate is seen extending dorsally to meet the squamosal. Further medially

a flange of the squamosal clasps the quadrate from above. From the antero-

median corner of the quadrate a process stretches antero-medially to meet the

quadrate ramus of the pterygoid. In posterior view it is seen that from the

external condyle the quadrate extends medially, forming a transverse mass

forming the major part of the articulatory surface. Dorsal to this transverse

body the quadrate extends dorsally as a strong pillar of which the posterior |

surface is overlapped by a descending sheet of the squamosal. The medial face

of the quadrate abuts against the latero-anterior corner of the strong paroccipital.

On the median surface of the internal condyle there appears to be a shallow

depression for the reception of the distal end of the stapes. The tympanic

membrane must have been situated in this region, but except for the ventral

squamosal edge no other points of attachment can be determined.

The quadratojugal is only seen in section, where it is seen as a triangular

element with its base resting on the postero-dorsal surface of the rounded

external condyle and its apical part lying posterior to the ascending column of

the quadrate and in the hollow between this and the descending flange of the

squamosal. ‘The median limits of the quadratojugal and the quadrate foramen

are in posterior view hidden by the descending flange of the squamosal.

The squamosal has its lower antero-lateral edge weathered away and a

section medial to this is shown in the figure. There is thus exposed much of the

quadratojugal, the upper part of the quadrate and the internal forwardly

directed domed sheet of the squamosal. All of which would be covered laterally

by the squamosal when fully present. In posterior view the descending sheet

of the squamosal covers the quadratojugal and most of the quadrate above the

condyles. Its ventral edge is concave and probably served as an edge of attach-

ment for the tympanic membrane. The median edge of the descending sheet

of the squamosal forms a backwardly directed ridge where it abuts against the

paroccipital. This ridge forms the median wall of the auditory groove, which is

thus situated directly dorsal of the probable location of the tympanic membrane.

ARTICULATORY REGION IN PRISTEROGNATHID THEROCEPHALIANS 57

ll b.

Fic. I1.—Therioides cyniscus Gen. et Sp. Nov. S.A.M. 11888, Vindraers-

fontein, Beaufort West (x #3). a, lateral view. 6, occipital view.

53. : ANNALS OF THE SOUTH AFRICAN MUSEUM

Anterior to the descending sheet of the squamosal the paroccipital appears to

abut against the median surface of the ascending pillar of the quadrate.

TTHERIOIDES CyNiscus Gen. et Sp. Nov. (fig. IT)

On the farm Vindraersfontein, Beaufort West, I collected a fair skull and

fore-limb which I believe to be new and for which I propose the name T heriozdes

cyniscus gen. et sp. nov. and the genotype bears the S.A.M. No. 11888. This

new genus may preliminarily be described as a Pristerognathid with dental

formula i. 6, c. 1, p.c. 6-7, postcanines slender and small, mentum fairly

strong and squarish, quadrate situated far ventrally, maximum length about

270 mm., snout as high as broad (55 mm.), orbit well in posterior half of skull.

In Therioides the posterior edge of the dentary sweeps sharply in posterior

direction and is directed more in the direction of the squamosal than towards

the dorsally situated temporal opening.

The angular has a smooth surface in its dorsal part; antero-ventrally there is

a groove and another groove is directed postero-ventrally. On its lower border

a thin sheet of bone lying medially projects below the edge of the outer surface.

The reflected lamina is strongly developed but does not extend to the surangular.

A deep notch lies dorsal to the reflected lamina and the bone surface in the

notch is formed by the angular. In lateral view the surangular shows as a

curved girder, anteriorly lying medially of the upper angular edge. Postero-

ventrally it firmly clasps the articular. The articular is small with a slight

indication of a retro-articular process. Medio-ventrally the articular is under-

lapped by the prearticular.

The quadrate in lateral view shows little more than the circular outline of the

outer condyle above which the quadratojugal lies. In posterior view the trans-

versely situated condylar part is partly exposed and the stout dorsal pillar-like

body of the bone is seen to be overlapped by the squamosal. Medio-ventrally

the rounded corner of the quadrate appears to form a surface to receive the

concave distal end of the stapes.

The quadratojugal is not clearly exposed but the visible bone immediately

above the external rounded condyle of the quadrate is apparently the lateral

part of the quadratojugal.

The postero-lateral corner of the squamosal is indentured immediately above

what I believe to be part of the quadratojugal. The ventral border of the

posterior descending flange of the squamosal is shallowly concave and this edge

probably served for the attachment of the tympanic membrane. Lateral to

where the squamosal abuts against the distal end of the paroccipital, the squa-

mosal has a high, thin sigmoid rampart forming the median wall of the auditory

groove.

The ventro-distal corner of the paroccipital has a sharp edge to which the

tympanic membrane was in part attached. The stapes is a slender rod in which

I have not been able to detect a foramen. Its distal end has its dorsal corner

prolonged and this, tipped with an extrastapedial cartilage, was probably

ARTICULATORY .REGION IN PRISTEROGNATHID THEROCEPHALIANS 59

2 ‘ =

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Ill b.

Fic. I11.—Pristerognathus roggeveldensis Sp. Nov. S.A.M. 9356a, Roggekloof, Sutherland (x #).

a, lateral view. 6, dorsal view. A horizontal fracture makes it possible to remove the left tem-

poral, post- and suborbital arches and show the underlying mandibular bones in horizontal

section and a dorsal view of the quadrate and quadratojugal.

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

attached to the tympanic membrane, and the relative positions are such that

the connection would be to the centre of the apparently circular tympanic

membrane.

PRISTEROGNATHUS ROGGEVELDENSIS Sp. Nov. (fig. III)

Among some Pareiasaurian and Deinocephalian bones collected by Mr.

A. R. E. Walker some forty years ago, I found a distorted Therocephalian skull

(S.A.M. No. 9356a) representing a new species of ‘Therocephalian, which I am

provisionally including in the genus Pristerognathus although Seeley’s genotype

is of such a nature that identification with it as norm is hazardous. Specific

characters of the new species, roggeveldensis, are: dental formula 1. 6, c. 1,

p.c. 5, maximum length of skull 255 mm., snout long, broader than high

(52: 50mm.); orbit in posterior half ofskull; dentary with truncated posteriorly

directed coronoid process. Due to the distortion the subtemporal bar is pushed

up revealing much of the quadrate complex which in life would be covered.

The dentary has its coronoid process truncated and directed posteriorly.

The outer surface of the angular has a number of radiating ridges separated

by shallow grooves. The reflected lamina does not stretch far posteriorly and

the notch is shallow. In lateral view the surangular has the same girder-like

shape as in the previous forms and the articular is also very similar.

In lateral view the quadrate shows the typical circular outline of the external

condyle clasped by the articular. From the condyle a thin sharp ridge stretches

postero-dorsally. Anterior to this ridge and lying in a more median plane the

antero-lateral face of the body of the quadrate is seen with its dorsal edge over-

lapped by the squamosal, which here forms a domed antero-medially directed

sheet of bone. Posterior to the lateral ridge on the quadrate is some bony

substance which I believe represents the small splint-like quadratojugal. In

dorsal view it is seen that the quadrate lies well laterally in the skull and is

situated in the angle formed by the subtemporal arch and the antero-medially

directed domed sheet of the squamosal lying in the posterior part of the temporal

space.

The antero-medial corner of the quadrate abuts against the slender quadrate

ramus of the pterygoid and in fig. IIIb its relations to the epipterygoid can also

be seen.

An UNIDENTIFIED PRISTEROGNATHID (fig. [Va)

A shepherd on the farm Dikbome, Laingsburg District, found some weathered

fragments of a Pristerognathid skeleton (S.A.M. 11959). One fragment is the

remains of the right articulatory region of the lower jaw. Here the quadratojugal

has been lost and the lateral aspect of the quadrate is well exposed. The con-

dylar part of the quadrate is not fully exposed but appears to be a transversely

placed roller-shaped mass posteriorly separated from the ascending pillar by a

well-defined groove. The condylar mass projects laterally as a process rounded

ARTICULATORY REGION IN PRISTEROGNATHID THEROCEPHALIANS 61

oao@®m 2 @

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IV a.

S.A.

Fic. [V.—a, unidentified Pristerognathid. S.A.M. 11959, Dikbome, Laingsburg. Lateral view

(x #). 6, Maraisaurus parvus Gen. et Sp. Nov. S.A.M. 11944, Buffelsvlei, Beaufort West. Lateral

view. Left. (x #.) c, Maraisaurus parvus. Lateral view. Right. (x %.) d, Alopecognathus angustiori-

ceps Sp. Nov. S.A.M. 9342, Kroonplaas, Beaufort West. Lateral view (x 2).

ses

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62 ANNALS OF THE SOUTH AFRICAN MUSEUM

ventrally and dorsally hollowed out. It is in this dorsal hollow that the base of

the quadratojugal must have rested. Above the external condyle the ascending

body of the quadrate forms a rounded edge dorsally meeting the internally

domed squamosal sheet. Anterior to this ridge the quadrate has a concave

lateral face. The ascending process of the quadratojugal must have passed up

along the posterior face of this ridge on the quadrate. On the antero-median

lower corner of the quadrate a sectional face indicates where the pterygoid

ramus of the quadrate emerged. The antero-dorsal face of the quadrate is

covered by the domed sheet of the squamosal.

Posteriorly the descending sheet of the squamosal extended to the level of the

upper edge of the condylar surface thus exposing little of the posterior face of

the quadrate.

The relations of the angular, surangular and articular are in essentials as in

the previously described forms.

MARAISAURUS PARVUS Gen. et Sp. Nov. (fig. [V6 and c)

Mr. Johannes Marais, who has for a number of years been actively interested

in the fossils found on his farm, has given me a small Pristerognathid skull

obtained on Buffelsvlei, Beaufort West (S.A.M. 11944). Unfortunately the

snout has been broken off obliquely in front of the orbits so that the nature of

the dentition remains unknown, but the rest of the skull shows it to be

undoubtedly a Pristerognathid. For this form I propose the name Maraisaurus

parvus in recognition of the work of its finder. This genus is characterized by

the small size of the skull (probable maximum length 162 mm..), large orbits

(34 X26 mm.), narrow snout, squarish in cross-section; dentary not extending

far posteriorly.

The posterior border of the dentary does not curve much in dorso-posterior

direction.

The angulars are not very well preserved as is evident from the differences in

the figures (a and b) of the two sides. It is, however, clear that the notch is

situated much dorsally and that the reflected flange does not extend far pos-

teriorly. ‘The relations of the surangular and articular agree in essentials with

the forms described above.

On the right side the lateral surfaces of the bones forming the jaw articulation

have been weathered and the quadratojugal lost. As preserved the quadrate

appears to be a roughly rectangular element. The condyle is a transversely

situated roller with a shallow groove demarcating two nearly equal convex

surfaces. Posteriorly above the condyle there is a transverse groove medially

receiving the descending plate of the squamosal and laterally the base of the

quadratojugal. On the left side a weathered bony edge is preserved which in

fig. IVb I have labelled the quadratojugal believing it to flank the quadrate

edge as seen in fig. [Va above the outwardly directed process of the external

condyle.

ARTICULATORY REGION IN PRISTEROGNATHID THEROCEPHALIANS 63

ALOPECOGNATHUS ANGUSTIORICEPS Sp. Nov. (fig. [Vd)

This new species is based on a well-preserved complete skull found by me on

the farm Kroonplaas, Beaufort West (S.A.M. 9342). It differs from the geno-

type in its smaller size (252 mm.) and in having a more slender skull.

On the right side the articulatory region is fairly well exposed. An indentured

descending sheet of the squamosal overlaps the ascending parts of the quadrate

and quadratojugal. ‘The quadratojugal has its base resting on the upper surface

of the external quadratic condyle, which is much the smaller of the two parts of

the condyle. In postero-ventral view the distal corner of the paroccipital, the

concave squamosal edge and the internal border of the quadratic condyle form

- two-thirds of a circle and it is in this space where the tympanic membrane must

have been situated.

The reflected flange of the angular extends to the surangular and no angular

notch is present.

With the above determinations we now know that the quadratojugal does

not form part of the articulatory condyle in a number and probably in all

Therocephalians. In addition to the above forms a similar condition has been

found by Broom in Lycedops and Hyenosaurus,and in Trochosaurus my figure of the

ocerput (Ann. S. Afr. Mus:, vol. XXXI, fig. 10, p. 229) falls in line with the

above interpretation.

The nature of this region in Therocephalians affords further evidence for

the derivation of the Cynodonts from the Therocephalians. In addition to the

cynodonts hitherto known to have the same relations of the quadrate to the

quadratojugal I have a cynograthid from Winnaarsbaken (S.A.M. 11264)

where the quadratojugal lies well laterally to the condyle. The well-defined

auditory groove of the cynodonts is undoubtedly foreshadowed in its essential

relations by the less well-defined groove seen in the Therocephalians.

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SOUTH AFRICAN MUSEUM

: VOLUME XLII

_ Descriptions of the Palaeontological Material collected by the South African

Museum and the Geological Survey of South Africa.

PART II, containing: —

‘i 7, The Pristerognathid Therocephalians from the Tapinocephalus-zone

ff in the South African Museum. By L. D. Boonstra, D.Sc. (With

ae 22 text-figures.)

: _ 8. The Cranial Structure of the Titanosuchian: Anteosaurus. By L. D.

iy Boonstra, D.Sc. (With 22 text-figures.)

sy 9g. Ihe smallest Titanosuchid yet recovered from the Karroo. By L. D.

jm Boonstra, D.Sc. (With Plate XVIII and 5 text-figures.)

yi 10. Paranteosaurus Gen. Nov.: A Titanosuchian Reptile. By L. D.

P _ Boonstra, D.Sc. (With 2 ‘text-figures.)

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7. The Pnsterognathid Therocephalians from the Tapinocephalus-zone in

the South African Museum. By L. D. Boonstra, D.Sc.

(With 22 text-figures)

In the course of my study of the fauna of the TAPINOCEPHALUS-zone the

collection of Therocephalians in the collection of the South African Museum

| has grown to number nearly a hundred specimens. The majority belong

to the family Pristerognathidae. Unfortunately, well preserved specimens

are rare and, although we have some good skulls, most of the specimens

are weathered snouts and in only a few cases have we postcranial elements

associated with the cranial material. In this paper I propose to deal only

| with the cranial material.

The Pristerognathids represent the bulk of the small to fairly large

carnivores in the fauna of the Tapinocephalus-zone. With them occur

members of some other Therocephalian families viz. Lycosuchidae and

Scylacosauridae and as a lesser element some Scaloposauridae, which were

_ probably insectivorous. The balance of the carnivorous section in the fauna

is formed by a few small Gorgonopsians and then we have the large and

massive Titanosuchid Deinocephalians.

The food source of these carnivores consisted mainly of the number of

small herbivorous Dicynodonts and the abundant large Pareiasaurs and

| Tapinocephalid Deinocephalians. Some of the Pristerognathids were

| probably carrion eaters.

Family Pristerognathidae

D1aGnosIs

Fairly small to large early carnivorous Therocephalians from the

Tapinocephalus-zone with a dental formula varying within the limits,

a, C.—, pee The incisors and canines usually well developed and

the posi-canines, with few exceptions, relatively feebly developed. Skull

long, fairly high and narrow; snout long, fairly narrow; sagittal crest high

and sharp; temporal fossa moderately long and broad; low to fairly low over

post-orbital arches; lower jaw strong, dentigerous ramus of dentary not

curved and relatively long, post-dentary part of mandible strong, strong

coronoid process. Prefrontal and _ postorbital well developed; and

postirontal well to fairly well developed; jugal arch moderate to strong;

frontal with moderate entry into orbital border or excluded from it;

septomaxilla with well developed facial surface; pineal foramen medium to

WOL. XLII. PART II.

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

fairly large; parietal narrow with sharp crest; paroccipital strong; occiput fairly

high and broad, deeply concave; epipterygoid narrow; anterior palatal openings

medium to long; suborbital vacuities large; palate flat; with only the anterior —

ramus of the pterygoid bearing teeth; basisphenoidal tubera of medium size,

parasphenoidal keel deep; suspensorium posteriorly situated high up and only

moderately laterally displaced.

Within the family two groups can be recognised which may yet have to

be considered as subfamilies:

a. Those with 6 incisors and with skulls where the length is double

or more the breadth.

b. Those with 5 incisors or with an inconstant 6th and where the skull

length is less than double the breadth.

GENERAL DESCRIPTION

Incisors, canines and post-canines have a sharp posterior cutting edge

which is serrated — the serrations vary from fine to fairly coarse. The

incisors are moderately curved teeth with a more or less flattened extremity

forming a curved cutting edge. They vary from fairly weak to very strong

seizing and tearing teeth. The first incisor with its fellow form a pair of

teeth closely set to each other, with the root rounded in cross section and

always smaller than the second incisor. The second to the 5th incisor are

subequal or decrease in size backwards. In the forms with six incisors the

6th tooth is inconstant in some genera and in the others it is mostly much

smaller and weaker than its predecessors. There is always only one canine;

except in one genus the canine is always a strong curved dog-tooth with its

extremity more pointed than is the case in the incisors; the upper canine is

situated far forward in the maxilla (except in one species) and to house its

long strong root the maxillary edge has grown downwards so that the

premaxillary edge is situated at a higher level; usually the alveolar border

curves evenly upwards anterior tc the canine, but in some the transition is by

a distinct step. The diastema between last incisor and canine varies

considerably in length — sometimes the canine follows immediately after

the last incisor. Behind the canine the diastema to the first postcanine also

varies considerably, but only in one specimen does the first postcanine lie

right up against the canine.

The number of postcanines varies considerably in the Pristerognathids —

from 2 to 9. The postcanines of the family, considered as a group, must be

described as weakly developed. They are usually small slightly curved teeth

either closely set or well spaced. In only one genus (Cynarniognathus) do

they form a set of well developed teeth. They are often irregular and of

little functional importance.

In the dentary there are apparently always 3 incisors, which in the closed

jaws are overlapped by the upper incisors. The lower canine lies lingually

THE PRISTEROGNATHID THEROCEPHALIANS 67

and anteriorly to the upper canine and its point in the closed jaws is housed

in the anterior section of the “‘choanae’’. Little is known of the postcanines

of the dentary — they lie lingually to those of the maxilla.

It is thus evident that the Pristerognathids in their feeding relied on the

anterior part of the jaws; the large canine acting as piercing instrument to

effect the kill and these together with the incisors were used to hold the

victim and then acted as the instruments for tearing away pieces of flesh.

Those with weaker incisors were probably carrion eaters. In most Pristerog-

nathids the postcanines acted chiefly as protruding points preventing flesh from

slipping out of the mouth. The transverse pterygoidal rami limited the action

of the lower jaw to movement in a vertical plane and there was thus no

chewing or cutting by the postcanines. The small set of recurved teeth on the

pterygoidal ridge served to hold the flesh in the stage preliminary to deglutition.

The postfrontal is well developed in some species, whereas in others it is

only a fairly small splintlike bone; it has only a small posterior extent, in

some forms extending up to the plane of the fronto-parietal suture, in others

a little further and at most to the level of the pineal foramen. It seldom

extends further posteriorly than the postorbital. The posterior tongue is

usually confined to the dorsal surface but sometimes lies on the lateral face

of the parietal crest.

The postorbital varies in the extent of its participation in the postorbital

bar, which is slender to fairly slender, never widened; in some forms it has

a small entry into the orbital border and the temporal border; in others the

entry into the orbital border is small, but the entry into the temporal border

is large; this depends on how high the dorsal jugal ramus extends and to

what extent the jugal is confined to the posterior half of the postorbital bar.

A postero-dorsal tongue of the postorbital sometimes just enters the

supratemporal border where it is applied to the lateral face of the parietal

or it sometimes develops quite a fair-sized sheet of bone flanking the parietal,

but seldom extends to just behind the plane of the pineal foramen.

The jugal is always well developed; it is triradiate with a dorsal, anterior

and posterior ramus. The anterior ramus forms the lower part of the orbital

border and a varying portion of the lower part of the posterior orbital border;

it mostly forms a sharp pointed wedge in between the lacrimal and maxilla,

but this is sometimes truncated. The dorsal ramus sometimes forms the whole

posterior half of the postorbital bar and thus this part of the temporal border;

in other forms its dorsal extent is much smaller and this part of the temporal

border is formed by the postorbital. The posterior ramus always extends

very far posteriorly as a long pointed wedge underlying the anterior ramus of

the squamosal, together forming the temporal arch. Usually the jugal forms

nothing or very little of the upper border of the temporal arch, but in those

forms where the anterior ramus of the squamosal is short it forms the anterior

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

portion of the upper border of the temporal arch. (This long posterior ramus

of the jugal in the Pristerognathids resembles the condition seen in Cynodonts

much more than does the jugal of the Gorgonopsians. )

The squamosal is always well developed; it has an anterior ramus extending

far anteriorly to form most of the upper border of the lower temporal arch

where it overlies the jugal; its ventral ramus extends far ventrally, laterally

largely covering the quadratojugal and posteriorly covering most of the

posterior face of the quadrate, which is housed in a recess in the antero-ventral

face of the descending ramus of the squamosal; the medially directed ramus

of the squamosal forms the posterior border of the temporal fossa and is

here applied to the outer face ot the parietal; this overlap of the squamosal

on the parietal is fairly small so that the postorbital and squamosal only flank

the lateral face of the parietal in its extreme anterior and posterior part

respectively, leaving the parietal to form nearly all of the upper border of

the temporal fossa. Where the squamosal meets the strong paroccipital the

edge of the squamosal is everted to form the auditory groove. The sweep of the

squamosal posteriorly carries the posterior border of the temporal arch far back

in the long narrow headed forms and thus forms a deeply concave occiput,

whereas in the shorter broader-headed forms there is only a moderate backward

sweep with a shallowly concave occiput. There is only a moderate lateral

sweep of the squamosal so that the temporal fossa never becomes very wide.

There is more lateral sweep in those forms where the posterior sweep is least.

In dorsal view the Pristerognathids show little more of the upper occipital

bones than their dorsal edges; the occiput is thus upright. In many forms the

condyle is clearly visible in dorsal view, but in the longheaded forms it is

either just visible or just hidden by the backward sweep of the parietals.

The quadratic condyles lie well below the level of the occipital condyle

and in a plane well below that of the alveolar border of the maxilla; they

have migrated very little in lateral direction so that the outer edge of the

quadrate ramus of the pterygoid is fairly straight; there has also been no

forward migration and they still occupy a posterior position.

The quadratojugal is much reduced and is a small element not entering the

condylar surface but resting with its base on the ledge of the quadrate above

the outer part of the condyle and with two weak dorsal processes fitting into

recesses in the overlapping squamosal.

The large prefrontal has a rounded ridge separating a dorsal from a

lateral face, but farther anteriorly the snout is rounded with no abrupt

transition from dorsal to lateral surface.

The lacrimal is small and has little anterior extension due to the upward

growth of the maxilla; it lies in the preorbital depression which is usually

fairly shallow and extending as a shallow groove in the direction of the

canines; but in some forms this depression is deep with sharp margins and

THE PRISTEROGNATHID THEROCEPHALIANS 69

anteriorly ends abruptly. The homology of the gland occupying the preorbital

depression is uncertain.

The long and strong root ot the canine is housed in a strong rounded

bulge of the maxilla, which is consequently a high bone. The outer surface

of the maxillary bulge often has an ornamentation of sub-crocodilian pitting,

but in some forms it appears to be fairly smooth. It is pierced by a number

of nutritive foramina.

The nasals are long and fairly narrow; together they are hourglass-shaped;

the constricted waist being due to the upward growth of the maxilla

overlapping the lateral edge.

In the lower jaw the symphysis is always weak and unankylosed, with

the splenial just entering the symphysis and forming a very subordinate part

and not visible ventrally. The angular has a large outer face with a notch

and a reflected lamina.

Systematic Descriptions

Genus Scymnosaurus Broom.

Broom, 1903, p. 152. Genotype, S. ferox Broom.

Large Pristerognathids with dental formula i.5, c.1I, p.c.2-4; incisors and

canines large and strong, postcanines fairly weak to medium, varying in

number — 2, 3 or 4. (In one specimen a small 6th incisor has been seen,

but only on the one side.)

Skull very large (Max. Length 375 to 475 mm.); preorbital hollow fairly

shallow, not sharply demarcated and really not more than a groove stretching

from the orbit in the direction of the canine; septomaxilla and septomaxillary

foramen well developed; frontal either excluded or possibly just entering the

supraorbital border; snout probably always slightly broader than high, slightly

narrowed between orbits and canines; orbits partly in anterior half of skull.

Mandibular symphysis, formed solely by dentaries, weak, unankylosed,

mentum sloping little or moderately.

Scymnosaurus ferox Broom. (Figs. I, Ila.)

Broom, 1903, p. 152. Holotype. S.A.M. 632. Locality unknown. Coll.

unknown. A well preserved but imperfect snout.

There are 3 to 4 postcanines; mentum moderately sloping and symphysis

weak; maximum length of skull 375 to 381 mm. (as reconstructed); frontal

possibly still entering orbital border.

S.A.M. 632 (Fig. Ia).

Broom’s holotype consists of the anterior third of the mandibular ramus,

the dentigerous border of the premaxillaries and the maxillaries up to just

behind the 3rd postcanine on the right and the 2nd postcanine on the left

side. The horizontal fracture through the snout shows the dorsal surface

of the anterior third of the palate, which is as figured by Broom

moos, Pl. 18, fig, 9).

70 ANNALS OF THE SOUTH AFRICAN MUSEUM

The first four incisors are

strong, whereas the 5th is

much shorter and this tooth

shows fine serrations on its

sharp posterior edge; the robust

canine has fine serrations on

its sharp posterior edge; only

3 postcanines are preserved

and these decrease rapidly in

size in posterior direction; the

postcanines bear coarser serra-

tions on the posterior edge.

REFERRED SPECIMENS

S.A.M. 3430 (Fig. 10).

Janwillemsfontein,

Prince Albert, ,,,,Coll.

Haughton & Whaits.

This is an imperfect snout

in which is_ preserved the

symphysis and the major part

of the premaxillaries and

maxillames’ In’) "both pre-

maxillaries 5 incisors are

preserved and here the 5th is

only slightly shorter than the

anterior 4 incisors. But on

the left side there is a stump

of what appears to be a small

6th incisor. On the right side

a stump of the canine is

preserved, as also stumps of

the 2nd and 4th postcanines

and empty alveoli of the rst

and 3rd. On the left side the

canine has dropped out and

so has the Ist postcanine, but

stumps of the 2nd and 3rd

are preserved.

The maxilla overlaps the

premaxilla right up to the

level of the anterior edge of

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Fic. 1.—a, Scymnosauvus ferox. ype:

S.A.M. 632. (x %-) ~ Right sidetverssnenc

fragment. 6b, Scymnosaurus ferox. S.A.M.

3430. (x 2.) Right side of snout fragment.

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Lettering of Figures:

A.—angular; Art.—articular; B.O..—basioc-

cipital; B.S.—basisphenoid; D.—dentary; Ep.

Pt.—epipterygoid; Ex.—exoccipital; F.—

frontal; F.J.—foramen jugale; F.O. fenestra

ovalis; I.P.—interparietal (postparietal); J.—

jugal; L.—lacrimal; Mx.—maxilla; N.—nasal;

P.—parietal; Pal.—palatine; P.F.—pretrontal;

P.Mx.—premaxilla; P.O.—postorbital; P.Oc.—

paroccipital; Po.F.—postfrontal; Pr.Ot.—

proédtic; P.S.—parasphenoid; Pt.—pterygoid;

P.V.—prevomer Q.—dquadrate; Q.J.—quad-

ratojugal; S.A.—surangular; S.Mx.—septo-

maxilla; S.Oc.—supraoccipital; Sq.—squa-

mosal; St.—stapes; Tab.—tabular; Tr.—

transversum.

The figures are orthoprojections taken with

a pantograph and, except where specifically

stated, they are of the specimens as preserved

without correction of any distortion. Recon-

structed parts and uncertain features are

shown in broken line. All figures drawn by

the author.

THE PRISTEROGNATHID THEROCEPHALIANS We

the 4th incisor. The bulge of the maxilla above the canine is rugosely pitted,

but no nutritive foramina can be seen. The septomaxillary foramen is of

moderate size (6 mm.).

S.A.M. 4341 (Fig. 1c). Stinkfontein, Prince Albert. Coll. Haughton.

This is a weathered snout bleached white. Each premaxilla has the

stumps of 5 incisors preserved; in cross section the 5th is only slightly smaller

than the 4th; in the right maxilla imperfect crowns of 4 postcanines are

preserved; the 3rd and 4th are the smallest and the 2nd the largest

maxillary tooth.

The septomaxilla has a large facial exposure and a stout spur directed

anteriorly into the nostril. There is a fairly large oval septomaxillary foramen

and in the maxillary edge, greatly overlapping the premaxilla, there is a

nutritive foramen. The maxillary bulge is rugosely sculptured. The nasals

broaden as they approach the nostril.

S.A.M. 9084 (Fig. 2a). Rietkuil, Beaufort West. Coll. Boonstra.

This specimen consists of the major part of a distorted skull, lacking the

intertemporal region, together with the ends of a humerus, radius and ulna.

The articulatory region has recently been described and figured (Boonstra,

HSS, lig.) 1).

‘ «Yale deoran

. mC S| gs cl) ese mm, ee

~ ee?

“Ms ecee*

Fic. 2.—a, Scymnosaurus ferox. S.A.M. 9084. (x #.)

Left side of skull and lower jaw with correction of the

distortion. b, Scymnosaurus major. S.A.M. 10556.

(x ¢-) Right side of preorbital part of skull.

72 ANNALS OF THE SOUTH AFRICAN MUSEUM

With the full length of the lower jaw preserved it is possible to estimate

the maximum length of the Scymnosaurus ferox skull viz. 375 mm. The

incisors are large and robust with the 5th only slightly smailer than its

predecessor and the sharp posterior edge bears moderately fine serrations.

The 3 postcanines are short stout teeth with the 3rd much smaller than the

anterior ones and the serrations on the posterior edge are as in the incisors

and the canine. The 5th incisor lies close to the canine, whereas in the

specimens mentioned above there is an appreciable diastema.

The preorbital depression has clearly demarcated borders.

The maxillary bulge is pitted and is perforated by a number of nutritive

foramina. The septomaxillary foramen is fairly large (6 mm.) and the

septomaxilla well developed but with less facial exposure than in

S.A.M. 4341. 7

SAM. 9005.

Fig. 3:') Seymnosaurus major. Type: | S.A NE.) Coen:

(x 4.) .@, Dorsal view. 0, Left side.

THE PRISTEROGNATHID THEROCEPHALIANS TS

The orbit lies partly in the anterior half of the skull.

The dentary is strong, with a straight ventral edge, a sloping mentum and a

weak symphysis.

See: 4347.

SCYMNOSAURUS MAJOR. Sp. Nov.

Holotype. S.A.M. 9005. Klein-Koedoeskop, Beaufort West. Coll.

Boonstra.

The well preserved anterior two thirds of a skull together with parts of

the pectoral and pelvic girdles, ends of humerus and radius and ulna.

There are 2 to 3 postcanines; mentum fairly upright and symphysis fairly

strong; maximum length of skull 450 to 474 mm. (as reconstructed); frontal

excluded from orbital border.

S.A.M. 9005 (Fig. 3 a and 0).

The estimated maximum length of the skull is 450 mm. The incisors are

large and strong (length of 3rd incisor 55 mm.); the length of the canine is

80 mm. (as reconstructed), whereas the two postcanines are small (15 mm.).

All the teeth have a finely serrated sharp posterior edge. The 5th incisor

lies close up to the canine, but the distance between the canine and the first

postcanine is large (30 mm. on the right side and 50 mm. on the left).

The preorbital depression is deep but with rounded margins.

The maxillary bulge has pronounced pits and radiating grooves. The

septomaxilla is well developed and the foramen fairly large (11 mm.). The

maxilla is high but short.

The nasals widen considerably in their anterior half.

The frontals are small, very narrow and well removed from the orbital

border. The prefrontals are large and the postfrontals fairly large.

The orbits are small and round (45 mm.) and lie partly in the anterior half

of the skull.

The mentum is fairly upright but the two rami are loosely joined at

the symphysis.

S.A.M. 10556 (Fig. 2b). Knoffelfontein, Beaufort West. Coll Boonstra.

This specimen consists of a weathered and bleached preorbital part of a

large skull (474 mm. as reconstructed). Only cross sections of the roots of

the teeth are preserved. The five incisors are practically of equal size.

74 ANNALS OF THE SOUTH AFRICAN MUSEUM

Anterior to the canine there is a distinct diastema and the edge of the

maxilla curves sharply downwards (the “‘step’’ of Watson). Posterior to

the canine there is a diastema of 27 mm., then follow three closely packed

small postcanines.

The preorbital depression forms an oblique groove from the orbit in the

direction of the canine.

The maxillary bulge is strongly pitted. The septomaxilla is well developed

with a strong intra-nostril spine. The septomaxillary foramen is very large

(19 mm.) and is kidney shaped with a spur from the septomaxilla dividing

it into two, in very much the same way as the medially directed spine divides

the nostril.

The maxilla is low and fairly long.

The mentum is upright and the symphysis quite strong.

SCYMNOSAURUS Sp. INCERTAE SEDIS

The following fragmentary specimens can be referred to the genus but

specific determination is uncertain,

S.A.M. 8999. Locality unknown. Coll. unknown. A weathered snout

fractured cleanly just behind the canines and from which little more

can be determined than that there were 5 large subequal incisors and

a fairly strong canine.

S.A.M. 11459. Buffelsvle1, Beaufort West. Coll. Boonstra and Marais.

A weathered snout in pieces, with the dental formula 1.5, c.1I., p.c.3+?

together with parts of a carpus and tarsus.

S.A.M. 11833. Lammerkraal, Prince Albert. Coll. Boonstra and Pienaar.

A weathered snout with I postcanine on the right side and 2 on

the: lett:

S.A.M. 11961. Dikbome, Laingsburg. Coll. Boonstra. A weathered

snout with dental formula 1.5, c.1, p.m ee

S.A.M. 9126. Voélfontein, Prince Albert. Coll. Boonstra. This consists

of a fairly well preserved snout with the teeth of the left side preserved

i.5, C.I, p.c.3-+. The teeth are very similar to those of the genotype.

Being only 2 the size of the genotype this snout may represent a small

species of Scymnosaurus.

THE PRISTEROGNATHID THEROCEPHALIANS 75

Genus Scymnosaurus: Measurements in Mms.

Sey ELOX.: S. major.

632 3430 434% 9084 9005 10556

prvee— Post.sq. edge’ 4 -. 3752 375? 38% 375 4560? 474?

Biwi Ant.Orb. border =. - —— — eT eMOg | 215 216?

Width of Snout over Canines . 105? 106 105 #00? I15 100

ici tmterorbital 9... — — Corr 72. =

Height of Snout at Post. edge

Gigeaminers...... eee OO LOO 4 TOs LOOn. pT XO go

Length of Upper Incisor

Series (65 75 65 70 85 60

ea, #1010) 70 60 70 75 60

Diastema C.-T... ] 9 ae 15 3 6 18

r a Oe 16 5 9 16

DMiastemay C2—P.C. . 14 = == 20 51 =

Ts, h ed aes 24 20 27 25

Length of P.C. Series oo == = 42 18 =

Bey 20 — AI 36 21 30

Ant.-Post.Diam. of Canine 1 22 a 25 24 3311 25

ig 22 20 20 a 26 25

Genus Glanosuchus Broom.

Broom, 1904, p. 85. Genotype G. macrops Broom.

Large Pristerognathids with dental formula 1.5+1, c.1, p.c.5; the first

5 incisors are large but the 6th is inconstant and when present very small;

the canine is large and strong; the postcanines are moderately to weakly

developed.

Skull large (Max. length 315-321 mm.); preorbital hollow fairly shallow,

not sharply demarcated and really not more than a groove stretching from

the orbit in the direction of the canine; septomaxilla and septomaxillary

foramen well developed; frontal enters orbital border; postfrontal quite well

developed; snout broader than high, slightly narrowed between orbit and

canines; orbits well in posterior half of skull.

Mandibular symphysis formed solely by dentaries, weak, unankylosed,

mentum sloping. |

Epipterygoid only slightly widened. Sagittal crest of parietals only

moderately high but fairly sharp.

Glanosuchus macrops Broom. (Figs. 4 and 5.)

Broom, 1904, p. 85. Holotype. S.A.M. 637. Knofloksfontein, Beaufort

West. Coll. Snyman.

A well preserved skull lacking only the posterolateral corners of the skull

and thus the suspensorial region.

76 ANNALS OF THE SOUTH AFRICAN MUSEUM

Broom’s description was based on the only partially cleaned skull. I

have cleaned up the outer surface, thus revealing the sutures and prepared

the palate and basicranium so that these features can now be described.

In Fig. 4 a, b and c the

structural, features Of (ihe

lateral and dorsal surfaces

are shown. The _ septo-

maxilla has a large facial

exposure, the spur project-

ing into the nostril is well

developed and the foramen

fairly large (6 mm.). The

maxilla is fairly high and

long with its dentigerous

border curving upwards

anterior to the posterior

edge (ot) \the jcanine: yiihe

preorbital hollow has a

fairly definite dorsal and

posterior rim but anteriorly

it flows into the general

maxillary surface in the

direction of the canine.

The maxillary bulge above

the canine is fairly smooth.

The nasals are widest

anteriorly. The frontal is

of medium size apparently

just entering the orbital f

border. In dorsal view the Fic. 4.—Glanosuchus macrops. Type. S.A.M.

occiput is seen to be deeply 637- (X4%-) @, Right side. 6, Left side.

5 c, Dorsal view. d, Ventral view.

concave. In outline the

epipterygoid is hourglass-shaped and although not much widened it is not

a columnar columella crani. Its base, resting on the quadrate ramus of the

pterygoid, has no great posterior process. Although the edge of the sagittal

crest is not preserved it is clear that the crest was not very high and

moderately sharp. The temporal fossa is short but roomy.

The ventral surface (Fig. 4d). With the dentaries in occlusion it has not

been possible to expose the choanal region. The prevomers posteriorly form

a truncated spatulate sheet of bone underlying the median edges of the

palatine and the anterior edge of the anterior pterygoidal ramus. The lateral

edge of the prevomer lies medial to an oblique ridge on the palatine. This

ridge is continued on the pterygoid where it is dentigerous (a cross section of

THE PRISTEROGNATHID THEROCEPHALIANS 77

at least one tooth is clearly visible). The palatine is a relatively small bone,

anteriorly bounding the choana and posteriorly forming the front margin of

the oval shaped large suborbital vacuity. Two oblique sutures, meeting in

an obtuse angle, separate the palatine from the prevomer and pterygoid.

The pterygoid is a typically tetraradiate bone; the anterior ramus with its

fellow, forms the lozenge-shaped middle part of the palate; anteriorly,

between the two diverging dentigerous ridges, a V shaped hollow extends on

to the prevomerine surface where it shallows and fades out; laterally the

anterior ramus forms the inner border of the suborbital vacuity; the posterior

part of the anterior ramus carries a prominent oblique ridge stretching from

near the median line to near the edge of the suborbital vacuity; this ridge is

dentigerous, carrying small sharp teeth irregularly arranged in two rows; in

the median line just posterior to these dentigerous ridges there appears to

be a small interpterygoidal slit.

The transverse pterygoidal ramus forms a strong bar which laterally

descends to the level of the lower dentary border and this distal end lies just

median to the angle of the dentary. The movement of the lower jaw is thus

suided and confined with little lateral play allowed. Against the lateral part

of the anterior face of the transverse pterygoidal bar lies the transversum;

from here the transversum ascends rapidly and curving forwards as a thin

process forms the lateral edge of the suborbital vacuity and meets the palatine.

The posterior ramus of the pterygoid with its fellow forms a median keel

which clasps the anterior end of the parasphenoidal rostrum.

The quadrate ramus of the pterygoid is not completely preserved and

one does thus not know whether it met the quadrate or not. On its upper

surface the quadrate ramus carries the base of the epipterygoid. I have not

been able to determine whether the epipterygoid has a posteriorly directed

process running along the quadrate ramus. In ventral view it is seen that

the edge of the quadrate ramus is only slightly concave and the epipterygoid

has a roomy cavum epiptericum median to it and the anterior part of the

prootic.

The parasphenoid is in ventral view seen as a sheet of bone underlying

the basisphenoidal tubera and narrowing anteriorly is clasped by the pterygoids

where these bones form a fairly deep median keel.

Ventrally the occipital condyle is formed by the strong basioccipital.

Postero-laterally to the basisphenoidal tuber lies the fenestra ovalis with the

proximal end of the stapes tm situ on both sides. The rim of the fenestra is

formed postero-laterally by the opisthotic, postero-medially by the

basioccipital, antero-medially by the basisphenoid and antero-dorsally by

the prodtic.

Only part of the occiput is preserved; above the foramen magnum the

occiput is deeply excavated; the posttemporal fenestrae are large; the

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

paroccipital (opisthotic) strong and only the dorsal edge of the interparietal

and tabular seen on the dorsal surface.

S.A.M. 11843. (Fig. 5a@ and b.) Lammerkraal, Prince Albert. Coll.

Boonstra. :

This good specimen consists of

the major part of a skull lacking

only the suspensorial region and

the lower edge of the dentary. It is

complementary to the holotype

skull in that it shows that the frontal

has a small entry into the orbital

border, | the «postiremtal ame

postorbital flank the parietal for

a short distance within the temporal

fossa; the sagittal crest of the

parietals is preserved as a not very

high but fairly sharp crista; the

temporal fossa is short but roomy.

But the postcanines are smaller and

more slender although in both

skulls they occupy the same space

(48) ;mm.).. ihe; yothameieen) 1s

preserved on the right side and

here also it is very much smaller

Fic. 5.—Glanosuchus macrops. S.A.M. than the anterior incisors. As in

11843. (X¢-)_ 4 Lateral view. b, the type all the teeth have a finely

Dorsal view. Left side incorporating .

features determined of the right side. serrated posterior border. The

occiput is less deeply concave than

in the type skull. The preorbital depressien is very shallow and the maxillary

surface smooth over the bulge where the long canine root is housed.

S.A.M. 11964. Locality unknown. Coll. unknown.

This is a weathered snout showing the upper dentition to be

1.5+1, C.I, p.c.2+?, with the 6th incisor very small.as in the two foregoing

specimens. Although appreciably smaller than the type the dentition agrees

well with that of the type so that I am referring it to the species macrops.

S.A‘: M. 903: *Seekoeigat, Prince’ /Albert)))Colly Mu Bless

A very much weathered snout, which on account of the presence of a

small 6th incisor can be included in the genus Glanosuchus.

Genus Ptomalestes. Gen. Nov.

Fairly large Pristerognathids with the dental formula i.5, c.1, p.c.6; the

incisors are subequal and fairly weak, the canine strong and the postcanines

small slender teeth well spaced.

THE PRISTEROGNATHID THEROCEPHALIANS 79

Skull fairly large (Max. Length 258 mm.); preorbital depression fairly deep

but without abrupt margins and shallowing in the direction of the canine;

frontal just entering orbital border; postfrontal quite well developed; snout

broader than high, only slightly narrowed between orbit and canine; orbits

well in posterior half of skull.

Mandibular symphysis formed solely by dentaries, weak, unankylosed,

mentum sloping.

Sagittal crest of parietals only moderately high and narrow.

Ptomalestes avidus Sp. Nov. (Fig. 6a—d.)

Genotype. S.A.M. 11942. Steenboksfontein, Laingsburg. Coll. Boonstra.

A fairly well preserved skull, but it has been subjected to a slight shear

and some weathering, together with some cervical vertebrae, part of the

pectoral girdle, humeri, radii and ulnae.

In lateral view (Fig. 6a)

the maxilla is seen to be

low and long and has a long

overlap on the premaxilla;

the small slender _post-

canines are _ irregularly

spaced and occupy 51 mm.

with a diastema of 12 mm.

to the canine; anterior to

the canine the maxillary

edge curves upwards

(‘‘step’’ of Watson) and

the diastema is 13 mm. on

_ the left and 15 mm. on the

Hight side. On the pre-

maxilla there are 5 incisors,

fairly small and all more

or less the same size. All

the teeth have serrated

sharp posterior edges.

The prefrontal is large

with a dorsal and a lateral

face.

The jugal has a stout

dorsal ramus forming the Fic. 6.—Ptomalestes avidus. ype: wovAeML:

. LEO4Z)) (SC 4) a, a. Wateral view.’ b;, Dorsal

lower and hind part of view. (G7) Vellital view. id, Occipitaly view:

the postorbital bar; the

posterior ramus extends moderately far backwards underlying the anterior

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

part of the anterior squamosal ramus forming the infratemporal arch; the

anterior jugal ramus is short. ,

The postorbital is of moderate size forming the postero-dorsal corner of

the orbital border and has only a short posterior tongue applied to the

lateral face of the parietal.

The squamosal has a deep descending process to hold and support the

quadrate and quadratojugal, but I have not been able to determine the

relations of these three bones as seen in lateral view.

The temporal fossa is short but roomy.

The angular has a fairly large external face with radiating ridges and

posteriorly a deep notch and a well developed reflected lamina with a thin

ventral edge. The surangular forms, in lateral view, the dorsal girderlike

border of the jaw between the dentary and the articular. The lower edge

of the dentary is straight and the posterior edge concave between the angle

and the strong coronoid process, which, extending into the temporal fossa,

stands out well dorsal to the surangular and the dentigerous border of

the dentary.

In dorsal view (Fig. 6b) the parietals are seen to form a moderately high,

fairly sharp sagittal crest. The frontal is fairly short, narrow and is practically

excluded from the orbital border. The postfrontal is quite well developed and

the prefrontal has a large dorsal surface. The occipital edge is fairly deeply

concave with only the dorsal edges of the interparietal and tabulars seen in

dorsal view, but the posttemporal arch does not sweep very much in posterior

direction. The temporal fossa is short but wide.

In ventral view (Fig. 6c) the dentaries are in position so that the anterior

and lateral portions of the palate are not exposed. The choanal openings are

short and the interchoanal bar formed by the prevomers stout. Posteriorly

the prevomers form a truncated spatulate sheet of bone underlying the

palatines and pterygoids. The palatine is of fair size with its outline

roughly that of a parallelogram; it carries an oblique ridge which is continued

on the pterygoid. The anterior pterygoidal rami form a median flat surface

laterally bounded by the above mentioned oblique ridges; no teeth can be

seen on these ridges; laterally the pterygoid bounds the large suborbital vacuity;

posteriorly the anterior ramus carries an eminence in which a number of

small teeth are implanted mainly in a curved row. The transverse pterygoidal

ramus forms a moderately strong bar inclined ventrally in lateral direction;

its corner does not descend so far as the ventral edge of the dentary where

this forms its obtuse angle. Posteriorly the pterygoid clasps the anterior

process of the parasphenoid and forms the lateral sheets of bone of the fairly

deep median keel. The quadrate ramus of the pterygoid has a fairly straight

lateral edge, well removed from the braincase, and meets the quadrate.

THE PRISTEROGNATHID THEROCEPHALIANS 81

I have not been able to determine the relations of the epipterygoid to the

quadrate ramus of the pterygoid.

The transversum is biramic; a transverse ramus is applied, as a vertical

sheet of bone, to the anterior face of the transverse pterygoidal ramus; a

longitudinal ramus forms a girder, ascending in the skull in anterior direction

to meet the palatine at its junction with the maxilla, and laterally bounding

the large suborbital vacuity.

The parasphenoid forms a fairly deep median keel anteriorly clasped

by the pterygoids, and presumably forms the underface of the basisphenoidal

tubera. The basisphenoidal tubera are situated well below the level of the

basioccipital condyle; their posterior borders are developed as a sharp ridge

forming the anterior part of the deep rim bounding the fenestra ovalis; the

medial part of the rim of the fenestra is formed by an equally sharp ridge

formed by the basioccipital confluent with that of the basisphenoid. Medially

to the deep rim of the fenestra ovalis the under surface of the basisphenoid

and basioccipital appears as a wide deep groove.

The strong opisthotic abutting against the basisphenoid and basioccipital

forms the outer and more dorsally situated part of the border of the fenestra

ovalis. Laterally the paroccipital bar abuts against the everted sheet of the

squamosal which forms the inner face of the auditory groove.

The lower jaw, being in articulation, covers most of the quadrate in

ventral view, but medially the inner rounded edge of the median

condyle can be seen and laterally the posterior edge of the lateral

quadratic condyle.

The sharp ventral edge of the reflected lamina of the angular is shown

in fig. 6c as is also the unankylosed mandibular symphysis.

In occipital view (Fig. 6d) it is evident that the occiput is low and that

the quadrate complex lies well below the basis cranu and that the squamosal

bulging outwards lateral to the roomy temporal fossa has not carried the

quadrate laterally with it. The limits of the interparietal, supraoccipital and

tabulars cannot all be determined, but where uncertain are probably as shown

in broken lines. The posttemporal fenestrae are large, the auditory groove

fairly shallow but above the foramen magnum there is a very deep circular

depression. The paroccipital bar is strong and the descending sheet of the

squamosal covers the greater part of the face of the quadrate. The deep rms

of the fenestra ovalis are clearly seen and are formed by the basisphenoid

and basioccipital.

S.A.M. 11460. Buffelsvlei, Beaufort West. Coll. Boonstra and Marais.

This specimen consists of an imperfect anterior half of a skull smaller

than that of the type. There are 5 incisors and 5 postcanines; the incisors

are fairly weak, the first 4 subequal and the 5th much smaller; the postcanines

2—Annals

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

are small and slender; on both sides the 3rd is missing; they occupy 20 mm.

on the left and 22 mm. on the right side; the diastema anterior to the canine

as well as that posterior to it measures 12 mm. On the assumption that its

smaller size and the differences in the dentition may be due to its being a

juvenile I am including this specimen in the species avidus, but with

corroborative evidence from a more fully preserved skull it may well prove

to be a smaller species specifically distinct from the genotype.

S.A.M. go12a. Klein-Koedoeskop, Beaufort West. Coll. Boonstra.

This specimen is an anterior weathered part of a snout in which the

incisors agree with those of the type.

Genus Pnsterosaurus Gen. Nov.

Fairly large Pristerognathids with the dental formula i.?, c.I, p.c.3 (in all

probability there were 6 incisors); the canine small and weak; the postcanines

also small and weak and closely packed; in all probability the incisors were

also small and weak.

Skull fairly large (Max. Length 225? to 255? mm.); preorbital hollow

shallow to very shallow without definite margins; frontal with small entry

into the orbital border; postfrontal well developed; snout broader than high,

not narrowed between orbit and canine; orbits in all probability in anterior

half of skull. Sagittal crest of parietals high and sharp.

Pristerosaurus microdon Sp. Nov. (Fig. 7.)

Genotype. S.A.M. 9083. Rietkuil, Beaufort West. Coll. Boonstra.

A well preserved skull lacking only the precanine portion of the snout.

In lateral view (Fig. 7a) the maxilla is seen to be short but high. The

canine is small, probably not more than 15 mm. long and antero-posterior

diameter 6 mm; the three small postcanines occupy 13 mm. on the right

side and 15 mm. on the left. The prefrontal is large with a rounded

ridge separating a dorsal from a lateral face. The jugal is a large triradiate

bone; the dorsal ramus forms the major part of the postorbital bar; the

posterior ramus extends far posteriorly and with the anterior ramus of the

squamosal which overlies it forms the infratemporal arch. (In this feature

this early Therocephalian exhibits a condition found in the later Cynodonts

and not found in the Gorgonopsians.) The anterior jugal ramus forms the

infraorbital bar with its extremity wedged in between the lacrimal and maxilla.

The postorbital forms the postero-dorsal part of the orbital margin and

curving towards the median line is applied to the temporal face of the sagittal

crest formed by the parietal, but does not extend far in posterior direction.

THE PRISI!EROGNATHID THEROCEPHALIANS 83

The squamosal has a deep descending process supporting the quadratojugal

and quadrate, but the displacement of the lower jaw has disturbed the

articulatory region so that the relations here are difficult to determine.

SAM.4083.

Fic. 7.—Pristerosaurus microdon. Type. S.A.M. 9083. (x 4.) a, Lateral view.

b, Dorsal view. Left temporal arch corrected on basis of the right side.

The angular has a large external face with a notch situated above the

reflected lamina; a curved ridge separates a dorsal hollow from the lower part

of the external surface.

The temporal fossa is long and roomy.

In dorsal view (Fig. 7b) the high sharp parietal crest is evident. The

frontal is long with a small entry into the orbital border and the prefrontal

has a large dorsal face separated from the lateral face by a rounded ridge.

The occiput is deeply concave with the posttemporal arch sweeping

far posteriorly. Above the foramen magnum the occiput is very deeply

excavated; the posttemporal fossae are small; the paroccipital is strong, with

its distal end everted where it meets the squamosal to form the inner boundary

of the deep auditory groove.

Little of the ventral surface could be prepared; the condyle is moderately

large and single; the basisphenoidal tubera are very strong (as in Trochosaurus)

and the fenestra ovalis has a prominent sharp ventral rim; the parasphenoidal

keel is deep and narrow. The quadrate complex is posteriorly overlapped by

a sheet of the squamosal descending far ventrally.

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

The specimen in the British Museum B.M.N.H. R.4100 which has been

referred to Lycosuchus and then to Scymnosaurus can obviously not be

included in either of these two genera and undoubtedly falls within the limits

set for the genus instituted here and constitutes a second species —

Pristerosaurus watsom (Broom).

Measurements. Glanosuchus, Ptomalestes and Pmsterosaurus

Pristero-

Glanosuchus. Ptomalestes. saurus.

637 11843 I1964 903 I1942 11460 9083

Pr. Mx:==B.Ovscondyle .: inet 310. .200)0) 70) tn See ee ER LON

Pr: Ni Post! Squsedge..° 8. 32m? 32 heya ah ee eee Save

Pr. Mx.2-Ant: Orb, bonder 9.) 44.172 \ a0 — i | aero Gm

Pr.Mx.—Pin.For. AND) SDAA EN N21 dm Z20%) eee = be Re Sel

Width of Snout over Canines 82 80 i 65 70 48?

Width of Snout over last P.C. 85 O57) nee 95 75 58

Width *Tmteronbitale 2) ts 52 54 a 44 — 40

Width Intertemporal over

Pinon) SiMe arc i OF ae 23 a 24 a 20

Height of Snout at Post. Edge

Of) Camimen Wye en ee ae 61 Se 55 47 55?

Height)atiP Ow bam © se 5.5 5. eS 527 — — Joven 45

Length of Upper Incisor

series |. 55 47 AD he 40 35 =

Lee No) 50 45 45 40 35 i

Diastema C.-T a 10 Ty th wae 13 r2 a

r 7 ke TO nO 15 13 sos

Diastema C= Pee: hy a he ee 12 12 2

r 16 18 15, 8 be 12 1 2

Meneth).or ue © Series Wy ee — SS = 20 16

Te Ao WGP get aie 51 22 15

Ant:-Post, Diam, of) .Canine 11) 20 18 Sp) aS 13 12 af

1 Me 17 TO Lh) as 15 12 =

Genus Thernoides Boonstra.

Boonstra, 1953, p. 58. Genotype. TZ. cyniscus Boonstra.

DIAGNOSIS

Fairly large Pristerognathids with the dental formula 1.6, c.1, p.c.6; the

incisors are subequal, moderately long and slender, the canine long and strong

and the postcanines well spaced, small, slender teeth.

THE PRISTEROGNATHID THEROCEPHALIANS 85

Skull fairly large (Max. Length 275 mm., as reconstructed); preorbital

depression fairly deep but without abrupt margins and shallowing in the

direction of the canine; frontal probably just entering orbital border; snout

as broad as high, only slightly narrowed behind canines; orbit well in

posterior half of skull.

Mandibular symphysis fairly weak, but mentum fairly upright.

Quadrate situated very low down in skull, far ventral of the occipital

condyle.

Therioides cyniscus Boonstra. (Fig. 8.)

Beonsird, 1953, Pp. 56. Holotype. S.A.M. 11883." Vindraersfontein,

Beaufort West. Coll. Boonstra.

A slightly distorted and somewhat weathered skull, together with part

of the pectoral girdle, humerus, radius, ulna and part of manus.

In lateral view (Fig. 8) the maxilla is seen to be fairly low but long with a

moderate overlap on the premaxilla; the small and slender postcanines are

evenly spaced, occupying 39 mm. on the left and 40 mm. on the right side

with a diastema of 10 mm. on the left and 9 mm. on the right side to the

canine; the canine is robust and 48 mm. long; no “‘step’’ anterior to canine,

and the diastema to the last incisor is small (10 mm. on left and 9 mm. on

right). On the premaxilla there are 6 incisors occupying 33 mm. on the

right and 5 incisors occupying 30 mm. on the left side; they are slender with

a mean length of 15 mm.; the 6th is weaker than the anterior teeth. All the

teeth have serrated sharp posterior edges. The septomaxilla has a fairly large

facial exposure and the septomaxillary foramen is large.

SAM. 96S. iy ae

Fic. 8.—Therioides cyniscus. Type. S.A.M. 11888. (x 4.)

Lateral view. Incorporating features of the right side.

The prefrontal is fairly large with a dorsal and a smaller lateral face.

The jugal bar is strong. The squamosal has a very deep descending process

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

carrying the quadrate very far down in the skull with the result that the

articulation is situated on the same level as the ventral border of the dentary.

I have recently described this area in detail (Boonstra, 1953).

The occiput is only partly preserved. The paroccipital is strong and its

ventro-median corner well developed and this, together with the basioccipital

and basisphenoid processes, forms the strong rim of the fenestra ovalis, which

is thus situated low down in the skull, well below the level of the occipital

condyle. The basisphenoidal tubera are well developed and the parasphenoidal

keel deep. The posttemporal fenestra is fairly large and the auditory groove -

well developed. The lateral sweep of the squamosals has not carried the

quadrates far laterally.

Genus Pristerognathus Seeley.

Seeley, 1895, p. 994. Genotype. P. polyodon Seeley.

DIAGNOSIS

: Laer . 6 3?

Pristerognathids with the dental formula oc €.2, eae

Pristerognathus polyodon, Seeley.

The holotype is in the British Museum (Natural History), B.M.N.H.

R. 2581, Tamboerfontein, Beaufort West, and consists of a poor weathered

snout.

This genus has only historical importance since the poor genotype shows

only the following diagnostic characters: dental formula, ae Ci, pod,

In the premaxilla the first incisor’s root has a circular cross section and it

lies close to its fellow; incisors 2-5 are ovate in section and the 6th is smaller

than the anterior teeth; they occupy 35 mm. In the dentary there are three

incisors decreasing in size in posterior direction; they are long and curved

with finely serrated posterior edges; they occupy 17 mm. The upper canine

has a diameter of 14 mm. and a serrated posterior edge. There is a step in

the maxillary border anterior to the canine. The symphysis, formed chiefly

if not entirely by the dentary, is unankylosed and the mentum is sloping.

The width of the snout is 55 mm. and the height 46 mm. The bulge of the

maxilla above the canine has a sub-crocodilian pitted surface.

Pristerognathus baim (Broom).

Broom, 1904, p. 87. Holotype. S.A.M. 583. Locality unknown. Coll.

unknown.

THE PRISTEROGNATHID THEROCEPHALIANS 87

A very poor weathered snout fragment.

There are roots of 6 upper and 3 lower incisors. The upper incisors occupy

34 mm. (measured round the curve); the 6th is smaller than the preceeding

teeth; the first smaller than the succeeding 3 and lies close to its fellow.

There is no reason for this fragment to bear a distinct specific name and I

regard bain; as a synonym of polyodon.

I propose that the generic name Pristerognathus be retained for Seeley’s

historic specimen and be used as a suitable label for all Pristerognathid

Therocephalian specimens in which no other diagnostic characters are shown

except the presence of 6 upper incisors occupying a space of 35 mm. For

specimens showing other additional characters other generic names should be

used as has already been done viz. Alopecognathus, Cynariognathus, etc.

The following list of specimens comprises a number of fragmentary and

weathered snouts which on the nature of the incisors can only be referred

to the genus Pristerognathus as characterised above.

Se ven 63r Koup, Coll. Joubert:

Ly 751 Seekoeigat, P.A. Coll. Du Plessis.

A 752 Seekoeigat, P.A. Coll. Du Plessis.

pews iiettontem, P.A. Coll. J. H. Whaits.

,, 1213 Unknown locality.

», 3432 Janwillemsfontein, P.A. Coll. Haughton & Whaits.

», 9015 Klein-Koedoeskop, B.W. Coll. Boonstra.

,, 9084a Rietkuil, B.W. Coll. Boonstra.

peerir Ore Vier, P2A: Coll. Boonstra.

», 11456 Buffelsvlei, B.W. Coll. Boonstra & Marais.

», 11586 Koedoeskop, B.W. Coll. Boonstra.

Menrrea2 7 Lammerkraal, P)A.~ Coll. Boonstra.

feito |, Veldmansnvier, P2A. Coll: Boonstra.

Pete a berdewater,. PA. Coll. Boonstra,

ge L872 is -

ite WORSE) Ws 4

ee ke 74 : in - if

PATIO TS % A Y 3

yt o76 ‘3

,, 11936 Bosluiskraal, Laingsburg. Coll. Boonstra,

,, 11950 Klein-Koedoeskop, B.W. Coll. Boonstra.

», 11957 Abrahamskraal, P.A. Coll. Le Roux.

,, 11960 Dikbome, Laingsburg. Coll. Boonstra.

,, 11963 Dikbome, Laingsburg. Coll. Boonstra.

,, 11965 Skoppelmaaikraal, Laingsburg. Coll. Botes.

#2210906... Seekoeigat,,,P.A...Coll. Bz) Bain.

py. EIGQO7), »Seekoecigat,..PAn,Colly io Baim:

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Alopecognathus Broom.

Broom, I9I5, p. 116. Genotype. A. angusticeps Broom.

The holotype of this species :s in the American Museum of Natural History.

A.M.N.H.5559. Coll. Whaits.

DIAGNOSIS

Moderate to fairly large Pristerognathids with the dental formula

1.6, C.I, p.c.4-5; the anterior incisors are fairly long, slender teeth (the ist

incisors, as in all Pristerognathids, are smaller than the 2nd to 5th) but the

6th is appreciably to very much smaller; the canine is long and strong; the

postcanines are small and fairly weak, not close set.

Skull moderate to fairly large (Max. Length 240? mm. to 276 mm.);

preorbital hollow varies from a fairly shallow hollow shallowing evenly in

the direction of the canines and without abrupt borders to a deep depression

with sharply demarcated borders especially anteriorly; septomaxilla with well

developed facial exposure and septomaxillary foramen well developed; frontal

with small entry into orbital border or just excluded from it; prefrontal large

with well marked dorsal and lateral face; postfrontal moderately to well

developed; snout wider than high, broader over last postcanines than over

canines; orbits just entering anterior half of skull.

Mandibular symphysis weak and mentum sloping very much.

Sagittal crest of parietals moderately high but with sharp edge.

Quadrate low down, but well above the level of the lower border of

the dentary.

Temporal fossa fairly long and wide.

Squamosal with everted lateral edge.

Alopecognathus angusticeps Broom.

The holotype in the American Museum has been redescribed by me some

years ago when studying the South African specimens sold by Dr. Broom

to the American Museum (Boonstra, 1935; p. 2).

DIAGNOSIS

There are 5-6 postcanines; maximum length of skull 275 mm.; preorbital

depression fairly shallow to deep, extending in direction of canine; postfrontal

well developed; squamosal laterally everted.

S.A.M. 9112 (Fig. 9). Stinkfontein, Prince Albert. Coll. Boonstra.

A fairly well preserved anterior three quarters of a skull.

From the figures and from the accompanying list of measurements it is

clear that this specimen from Stinkfontein is co-specific with the genotype,

THE PRISTEROGNATHID THEROCEPHALIANS 89

notwithstanding certain less important differences e.g. in our specimen there

are 6 postcanines occupying 40 mm., and the preorbital depression is deep.

Pit | es th SAM 4112.

Fic. 9.—Alopecognathus angusticeps. S.A.M. QIi2. (x

a, Dorsal view. b, Lateral view.

Cols

This specimen allows us to correct two points in my description (Boonstra,

1935) of the genotype viz. with the anterior part of the sagittal crest preserved

it is now clear that in my reconstruction the height of the crest should have

been much higher; the preserved frontonasal suture in this specimen shows

that the broken line indication in my figure of 1935 should have been further

forward so that the frontal is a long narrow bone and the nasal shorter than

I thought in 1935.

Alopecognathus angustioriceps (Boonstra).

Boonstra, 1953, p. 63. Holotype. S.A.M. 9342. (Fig. 10.) Kroonplaas,

Beaufort West. Coll. Boonstra.

A very good undistorted and practically complete skull.

go ANNALS OF THE SOUTH AFRICAN MUSEUM

DIAGNOSIS

There are 5 postcanines; maximum length of skull 252 mm.; preorbital

depression shallow, continued anteriorly as a shallow groove; postfrontal

small; squamosal with lateral bulge.

age

~~ ~

x” any we

Fic. 10.—Alopecognathus angustioriceps. Type. S.A.M. 9342. (x 4.)

a, Lateral view. 0b, Dorsal view. c, Ventral view.

In lateral view (Fig. 10a) the maxilla is seen to be fairly low and of

moderate length with a moderate overlap on the premaxilla; it carries 5 small

THE PRISTEROGNATHID THEROCEPHALIANS Qi

(1st very small) postcanines occupying 27 mm. on the left and 29 mm. on the

right side. The diastema between canine and the first postcanine is 10 mm.

on the left and rr mm. on the right. The canine is well developed with

a length of 30? mm. and antero-posterior diameter of 14 mm. Anterior to

the canine is a diastema of 12 mm. on the left and 14 mm. on the right. The

premaxilla carries 6 teeth, the anterior 5 are long and slender and the 6th

small and slender occupying 33 mm. on both sides.

The septomaxilla has a well developed facial surface and intra-nostril spur.

The prefrontal has a small lateral face. The jugal has a short dorsal ramus

and a well developed posterior ramus overlain by a moderately long anterior

ramus of the squamosal. The postorbital forms most of the postorbital bar.

The squamosal has a deep descending ramus supporting the quadrate complex.

I have recently figured and described the suspensorial region (Boonstra, 1953).

The dentary is large with a strong coronoid process; the symphysis is weak and

the mentum sloping.

In dorsal view (Fig. 10b) the long slender shape of the skull is evident;

the orbits just enter the anterior half of the skull; the temporal fossa is long

and roomy; the everted anterior ramus of the squamosal forming most of the

lower temporal arch is noteworthy (seen also in the genotype). The frontal is

just excluded from the orbital margin; both the frontal and nasal are narrow.

The prefrontal has a large dorsal face bounded laterally by a sharp ridge.

The postfrontal is small and the postorbital well developed but extending little

along the lateral face of the parietal. The parietals are narrow and form

a sharp sagittal crest which is, however, not high. The upper occipital edge is

very deeply concave.

In ventral view (Fig. 10c) it is seen that with the dentaries in position

it has not been found feasible to clean the anterior part of the palate. Just

anterior to the transverse rami of the pterygoids the pterygoids bear two

dentigerous ridges. The lateral corner of the transverse pterygoidal ramus

lies in the plane of the angle of the dentary but does not descend to the level

of the ventral edge of the dentary. Posteriorly the pterygoid clasps the

parasphenoidal rostrum which lies in the median line and the quadrate ramus,

with a slightly curved lateral edge, extends to the quadrate. The

parasphenoidal keel is deep. The basisphenoidal tuber, the medio-ventral

corner of the opisthotic and the antero-lateral corner of the basioccipital form

three distinct protuberances which constitute the tripartite rim bounding the

fenestra ovalis. The paroccipital bar is strong; the basioccipital condyle fairly

weak. The squamosal extends down far to cover the posterior face of the

quadrate complex. The squamosal sweeps far posteriorly to make the occiput

deeply concave, and the occipital condyle and the quadratic condyles are

situated far anterior to the posterior limits of the skull. The quadratic condyles

also lie in a plane well ventral to that of the occipital condyle.

Q2 ANNALS OF THE SOUTH AFRICAN MUSEUM

Measurements. Alopecognathus

A.M.N.H. S.A.M. S.A.M.

5299 OTT 739344

Prbix.—-B.O., condyle}, Lib Avapeeah Me iebe 234 — 215

Pr. Mx. Pesto! | etlge my man. amare Bee be: 275 260 252

Py. Mx¢ 2 AntOrbs: (border ie ee ne annie 140 138 [NG

PG. Mix POT. eal ok UN cage mer en ka 210 210 180

Width ‘of -Snout vover ‘Camimes in) 04) sa 55 56 50

Width vot ‘Snomitover jlasth biG. ja. ao ae 68 70 53

Width, TnterorbitalQe\ oe Weck se. i a ane 35 45 40

Width, Intertemporallover sransHor.. 10? 15 17,

Height of Snout at Post. edge of Canine . . 50 51 48

Height at PostiOrp. ban) iii. saurwai\ Sepia 45 Ae 45

Length of Upper Incisor Series Le 33 34? 33

I. a SE OS

Diastema “C2er IF r2 1 2

I. = = 14

Diastemra (CEC. Ih. 12 16 10

ite =o = II

Peneth, of vb ©) Series lig 31 40 27

r. wax 39 29

Ant.-Post.Diam. of Canine I. 12 15 13

ie = — 13

Genus Cynariognathus Broom.

Broom, 1931, p. 161. Genotype. C. platyrhinus Broom.

The holotype of this species is in the American Museum of Natural

History. A.M.N.H. 5502. Collected by J. H. Whaits and sold to the

American Museum by Dr. Broom.

DIAGNOSIS

Medium sized Pristerognathids with the dental formula i.5-6, c.1I, p.c.6-9;

the posterior incisors are smaller than the anterior ones; the canine is long

and strong; the postcanines are fairly strong and closely set.

Skull of medium size (Max. Length 260-290 mm.); preorbital depression

shallow; septomaxilla fairly small; frontal with a moderate entry into the

orbital border; prefrontal large with well marked dorsal and lateral face;

postfrontal moderately developed; snout broader than high; broader over

last postcanines than over canines; orbit in posterior half of skull; skull low

over postorbital bar; orbits high up in the skull. Premaxilla weak. Angle

THE PRISTEROGNATHID THEROCEPHALIANS 93

of dentary obtuse with posterior edge of dentary not deeply concave. Dentary

not shallow behind the canine.

Cynariognathus pauciondens. Sp. Nov. (Fig. II.)

Holotype. S.A.M. 11560a. Kroonplaas, Beaufort West. Coll. Boonstra.

A weathered snout with some parts of the postcranial skeleton.

DIAGNOSIS

Six closely packed postcanines occupying 27 mm. on the left and 28 mm.

| on the right side; breadth of snout over the canines 50 mm.; height of snout

at posterior edge of canine 40 mm.; preorbital length of skull 99 mm.

-ee

7? See .

Fic. 11.—Cynariognathus paucioridens. Type. S.A.M. 11560a. (x 4.)

a, Dorsal view. b, Lateral view.

This skull has a long low maxilla, the lacrimal is large and the prefrontal

fairly small; the dentary is fairly lightly built; the preorbital groove is fairly

deep. Little more can be determined from this specimen and I would have

hesitated to name it were it not that a second specimen, also with 6 closely

packed postcanines, proves that we have here a definitely new species of

Cynariognathus.

Q4 ANNALS OF THE SOUTH AFRICAN MUSEUM

S.A.M. 11586. Koedoeskop, Beaufort West. Coll. Boonstra.

The antorbital weathered half of a skull.

Here there are 6 closely packed, fairly weak postcanines occupying

28 mm. and there are apparently 5 incisors occupying 30 mm.; the height

of the snout is 40 mm. and the width 53 mm.; the antorbital length is

100? mm.

5.A.M. 1080. Fraserburg Rd., Prince Albert. Coll. J: H. Whaite:

A weathered antorbital part of a skull together with a complete hindfoot.

In this specimen there are 7 postcanines; they are closely set and robust

and occupy 26 mm. The height of the snout is 35 mm. and its breadth

48 mm.; the antorbital length as estimated is only 75 mm. The shorter snout

and the larger number of postcanines make inclusion in the above species

provisional.

Cynariognathus spp.

S.A.M. 3713. Bloukrans, Prince Albert. Coll. Haughton.

S.A.M. g088a. Klein-Koedoeskop, Beaufort West. Coll. Boonstra.

S.A.M. 11968. Locality and Collector unknown.

These three incomplete snouts are included in the genus Cynanognathus

as the height of the snouts is much smaller than the breadth and because

in the postcanine series the teeth are closely packed. But as the posterior

postcanines are not preserved the determination of the species remains

uncertain.

Genus Pristerognathotdes Gen. Nov.

Genotype. Alopecognathus minor Haughton.

DIAGNOSIS

Medium sized Pristerognathids with the dental formula 1.6, c.1, p.c.5-6;

the incisors are small to fairly strong teeth, with the 6th incisor only slightly

to much smaller than its predecessors; the canine is fairly long; the postcanines

are small, weak and well spaced.

Skull of medium size (Max. Length 222-287 mm.); preorbital depression

shallow to fairly shallow continuing as a groove in the direction of the canine;

septomaxilla fairly small; frontals with a fairly small entry into the orbital

border; prefrontal large with well marked dorsal and lateral face; postfrontal

moderately well developed; snout wider than high; broader over last postcanine

than over canines; orbits in posterior half of skull or just entering anterior

half; skull very low over postorbital bar; orbits high up in skull.

THE PRISTEROGNATHID THEROCEPHALIANS 95

Sagittal crest high and with sharp edge. Quadrate fairly low down, but

well above the level of the lower border of the dentary.

Temporal fossa fairly long and moderately wide.

Premaxilla weak.

Squamosal with lateral edge not everted.

Angle of dentary squarish with fairly deep concave posterior edge sweeping

| far posteriorly towards the coronoid process. Dentary shallow behind

_ lower canine.

Pristerognathoides minor (Haughton). (Fig. 12.)

Haughton, 1918, p. 180. Holotpye. S.A.M. 3415. Klipbank, Beaufort

West. Coll. Whaits.

A good skull lacking only the temporal arches.

| DIAGNOSIS

There are 4 postcanines; maximum length of skull 240? mm.; preorbital

| depression deep with very definite and abrupt margins; postfrontal large.

Since Haughton’s description I have carried the development of the skull

_ further, exposing more of the palate, and removed the matrix from the

| preorbital depression.

In lateral view (Fig. 12a) the maxilla is seen to be low and long with

_a long overlap on the premaxilla; on the right 4 postcanines are preserved but

there is a space between the rst and 2nd which may have housed an additional

tooth; the length of the series is 28 mm.; on the left 3 postcanines are preserved

with a gap between the 1st and 2nd; the length of the series is 26 mm.;

the postcanines are short but stout. Between the Ist postcanine and the

canine there is a diastema of 8 mm. on the left and 12 mm. on the right side.

|The canine is long and strong (diameter 11-12 and length 24? mm.).

Anterior to the canine there is a diastema of 10 mm. to the last incisor. There

are 6 incisors of which the 5th and 6th are shorter than the anterior ones

and the series occupies 32 mm. on both sides. All the teeth have finely

serrated posterior borders.

The septomaxilla has a fairly large facial exposure and a well developed

intra-nostril spur; the septomaxillary foramen is fairly large. The preorbital

depression is a deep hollow, especially on the right side, with a sharp rim,

especially along its dorsal and ventral borders; a low longitudinal ridge

divides the hollow into two equal parts on the right side, but on the left the

lower part is the larger; anteriorly the hollow ends abruptly and is not

continued in the direction of the canine.

The bulge of the maxilla above the canine is not rugosely pitted. The

anterior orbital border just enters the anterior half of the skull. The

96 ANNALS OF THE SOUTH AFRICAN MUSEUM

postfrontal is very well developed with a large entry into the orbital border;

whereas the postorbital appears to have only a very small entry into the

orbital border, forming the posterior part of the postorbital bar.

The edge of the sagittal crest is not preserved but was probably sharp and

fairly high with little flanking of the parietal by the postorbital.

S.AM.3LI5.

’

« &

tos SA ee ey

oe SD) oer Ce

f . vetted

Got ee:

frp ao

Pola a

Fic. 12.—Pristerognathoides minor. Type. S.A.M. 3415. (x #4.)

a, Lateral view. Incorporating features of the left side. b, Dorsal

view. c, Ventral view.

The articulatory region is not well shown since the bone has been stripped.

The angular has a deep notch and a small bone above the articular I have

THE PRISTEROGNATHID THEROCEPHALIANS 07

ident:fied as the quadratojugal. The squamosal has a deep descending

process posteriorly supporting the quadrate complex.

In dorsal view (Fig. 12b) the width across the temporal arches (as restored)

is great making a roomy although short temporal fossa, whereas the snout,

unconstricted behind the canines, is relatively narrow. The frontal is fairly

small and probably just excluded from the orbital border. The postfrontal is

very large and the postorbital weak, both with little posterior prolongation

along the lateral face of the parietals. The prefrontal has a well developed

| dorsal face separated from the smaller lateral face by a sharp ridge which

_ also forms the sharp dorsal rim of the preorbital depression. The nasals are

large, expanded anteriorly and posteriorly. The dorsal occipital edge is

deeply concave.

In ventral view (Fig. 12c) development has exposed all but the choanal

region of the palate. The posterior ends of the prevomers form a

shovel-shaped sheet of bone underlying the palatines and pterygoids. The

_ pterygoids have long anterior rami, which bear, in their posterior part, a

dentigerous ridge; the transverse pterygoidal ramus is not very strong or

_ wide and its anterior face is lined by a descending sheet of the transversum

which extends anteriorly as a girderlike ramus to form the outer border of

the large suborbital vacuity. There is a small interpterygoid vacuity. The

outer edge of the quadrate ramus of the pterygoid is slightly concave, thus

increasing the size of the temporal space. In the median line the parasphenoid

underlies the basisphenoid and anteriorly forms a deep keel. Surrounding

the fenestra ovalis there is a fairly strong rim formed by the basioccipital,

opisthotic and basisphenoid-parasphenoid. The paroccipital bar is strong and

its ventro-anterior corner abuts against the quadrate. Lateral to the quadrate

lies a small guadratojugal which does not enter the condylar surface. The

squamosal covers most of the posterior face of the quadrate complex.

S.A.M. 4332 (Fig. 13.) Wilgerbos, Prince Albert. Coll. Haughton.

A good skull of which some aspects of the internal structure have been

described by Haughton (Haughton, 1918).

The skull is very similar to that of the holotype of Pristerognathoides

minor. On the left side there are 5 postcanines occupying 29 mm. The

distance between canine and postcanines is 15 mm. on the left side and

between canine and postcanines is 15 mm. on the left side and between canine

and incisor the diastema is 9 mm. and the 6 incisors measure 30 mm.

The preorbital depression is fairly deep, but shallows in the direction

of the canine.

As is evident from the figures there are several other minor differences

in the proportions of some of the surface bones, but notwithstanding these

I am referring this skull to P. minor.

8—Annals

ANNALS OF THE SOUTH AFRICAN MUSEUM

SAM 4332.

oan WN. : ysis =

ie ee oe eS

Win se mi sae SO xP

at [! = Vj ‘ Zs ” se

Fic. 13.—Pristerognathoides minor. S.A.M. 4332. (x

a, Dorsal view. 0b, Lateral view.

3+)

S.A.M. 3435 (Fig. 14). Jakkalsfontein, Prince Albert. Coll. Rogers.

A well preserved antorbital part of a skull laterally compressed.

SAM. 3135.

Fic. 14.—Pnisterognathoides minor. S.A.M. 3435.

(x 4.) Lateral view.

Although the snout as preserved appears to be shorter and the ma il

consequently relatively higher and shorter than in the type, the dentition is”

very similar and the clearly demarcated preorbital depression has an abrupt

anterior border as in the type specimen. I am thus referring this snout to.

P. minor. a

S.A.M. 11891 (Fig. 15). Lammerkraal, Prince Albert. Coll. Boonstra

& Pienaar. : ki i i:

THE PRISTEROGNATHID THEROCEPHALIANS 99

A good skull lacking only the right postorbital and temporal arches.

In this skull there are on the right side roots of six postcanines. The snout

is broader than in the type and the preorbital depression is fairly shallow and

although there are other minor differences as can be seen from the figures

and the table of measurements I am referring it to the species minor.

SAM. 11541.

+ ee ey Canes aus oes

- Soe n8 abe oe =

7 & SOSA Soe ee

5 o> tA

pe ES, B

®*eae”

Fic. 15.—Pristerognathoides minor. S.A.M. 11891. (x 4.)

a, Dorsal view. 0b, Lateral view.

_ Pnisterognathoides roggeveldensis (Boonstra). (Fig. 16.)

Boonstra, 1953, p. 60. Holotype. S.A.M. 9356a. Roggekloof,

Sutherland. Coll. Walker.

A fair though somewhat distorted skull lacking the posterior part of the

skull roof.

DIAGNOSIS

Skull long and narrow; with five weak, well spaced postcanines; incisors

fairly strong with the 6th only slightly smaller than the 5th; orbit well in

posterior half of skull; squamosal extending well forward on lateral face of

parietal; prefrontal narrow.

In lateral view (Fig. 16b) the maxilla is seen to be long and high; the

septomaxilla has a moderate facial exposure; the orbit is small and situated

high up in the skull; the lacrimal large and the preorbital depression fairly

shallow. I have recently (Boonstra, 1953) described the articulatory region.

I00 ANNALS OF THE SOUTH AFRICAN MUSEUM

SAM. G35bQa.

Fic. 16.—Pristerognathoides roggeveldensis. Type. -S.A.M. 9356a. (x #)

a, Dorsal view. b, Lateral view.

In dorsal view (Fig. 16a) the snout appears long and narrow and the

temporal fossa long and narrow.

Pristerognathoides vanwyki (Broom). (Fig. 7)

Broom, 1925, p. 318. Holotype. S.A.M. 6533. Bloukrans, Prince

Albert. ‘Coll, Le“ Roux:

A weathered and distorted skull.

DIAGNOSIS

Skull moderately long and narrow; with six weak well spaced postcanines;

incisors fairly weak, with the sixth very slender; orbit just in posterior half

of skull; prefrontal large.

In lateral view (Fig. 17b) the maxilla is seen to be fairly short and high;

the septomaxilla has a good facial exposure; the orbit is small and situated

high up in the skull; the lacrimal is large and the preorbital depression shallow.

In dorsal view (Fig. 17a) the skull is fairly long and fairly narrow and

the temporal fossa fairly short and narrow.

S.A.M. 11893 (Fig. 18). Lammerkraal, Prince Albert. Coll. Pienaar.

THE PRISTEROGNATHID THEROCEPHALIANS

SAM 6533...

“eeee?” -c°

oe”

M@weooer’

Fic. 17.—Prsterognathoides vanwyki. Type. S.A.M.

a, Dorsal view. b, Lateral view.

6533= (Xx

Incorporating features of the left si

)

sue

SAM. 11893.

Fic. 18.—Pristerognathoides vanwyki. S.A.M. 11893.

a, Dorsal view.

(OS aya)

6, Lateral view.

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

This snout, presented to the Museum by Mr. J. Pienaar of Lammerkraal,

has only two postcanines preserved, but as this is probably only due to ©

postmortem loss and the other characters preserved agree fairly well with

those of vanwyki I am referring it to that species.

S.A. M: 11689, (Fig. 19). Prince Albert Road. Coll. Hesse:

SAM Hb $4. This dorso-ventrally crushed

skull has the postcanines badly

preserved but there were probably

6. Although the snout is longer

and there are differences of actual

size and proportions | am referring

it to vanwyki.

Fic. 19.—Pmnsterognathoides vanwyki. S.A.M. 11689. (x +4.) a, Dorsal view.

b, Lateral view. Dorso-ventral crushing corrected.

Pristerognathoides paruus Sp. Nov. (Fig. 20.)

Holotype. S.A.M. 3611. -Bloukrans, Prince Albert. © Coll. Scholtz

A weathered anterior two thirds of the skull.

DIAGNOSIS

Small low and fairly narrow skull; with 5 small well spaced postcanines;

incisors weak with the 5th weaker than the anterior ones and the 6th very

feeble, the diastema between incisors and canine great (15 mm.), and canine >

slender; prefrontal large; very low over postorbital arch.

The skull is small with the maximum length probably not more than

195 mm., the maxilla is long and low; the suborbital arch slender and the

dentary weak; the preorbital depression quite deep.

THE PRISTEROGNATHID THEROCEPHALIANS

5.AM, 360).

Fic. 20.—Pysterognathoides parvus Sp. Nov. Type.

MEASUREMENTS—Pristerognathoides

SE Aci 3 Om.

a, Dorsal view. 6b, Lateral view. Incorporating some features of the left side.

103

(X 3+)

3415 | 4332 | 3435 | 11891 | 9356a| 6533 | 11893 | 11689) 3611

Pr. Mx.—B.O. condyle oe ae Bo) 25) 200 — 200 —_ — = —= ee

Pr. Mx.—Post. Sq. edge St af <. |_240 218 — 215 255 2227 | 213'2-|\ 235 —

Pr. Mx.—Ant. Orb. border .. te Se |) GIS) 110 90 113 117 106 | 108? | 118 97

Pr. Mx.—Pin. For. .. &. we «|| 165 155 — | 160 — | 158 — | 1772) 148

Width of Snout over Canines as Bie 50 47 _ 55 52? | 45 50 52 45

Width of Snout over last P.C. me So 55 60 -- 60 — 52 =if/ — 59

Width Interorbital .. ope ae Ae 33 35 — 30 — 25 2} — 30 28

Width Intertemporal over Pin. For. nie 20 — — 16 _ 14 — 17 25

Height of Snout at Post. edge of Canine .. 40 45 40 39 53? 40 51 STi? 40

Height at P. Orb. bar =% “ aos Zor) +30 a 20 — — — 302) |) 30)?

Length of Upper Incisor Series ate E 32 33 _— — 35 30? 35 30 35

r 32 _ 30 35 — — 34 — 35

Diastema C.—I. an as ze 1. il 8 5 — 9 9 13 9 10

r il 11 6 5 — — 13 ~ 9

Diastema C.—P.C. .. ae ae 1. 8 15 12 15 16 10 11 12 10

Tt 11 Ui 9 14 10 10 10 11 8

Length of P.C. series .. he oh jail 33% 30 22 26 27 27 — _— 25

r 29 — 27 27 DON 135 a 3325) 21

Ant.-Post. Diam. of Canine .. Ns i 12 10 10 11 12 13 12 12 5

; Tr 11 10 9 7 12 12 12 11 6

I04 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Maraisaurus Boonstra.

Boonstra, 1953, p. 62. Genotype. M. parvus Boonstra.

DIAGNOSIS

Small Pristerognathids with dental formula unknown but probably

1.02 feelhar p.c.4%

Skull very small [Max. Length (as reconstructed) 162? mm.]; preorbital

depression very deep with sharp rim; frontal with moderate entry into orbital

border; postfrontal well developed; orbits large and just entering anterior

half of skull. Sagittal crest of parietals fairly low, but with sharp edge.

Maraisaurus parvus Boonstra (Fig. 21).

Genotype. S.A.M. 11944. Buffelsvlei, Beaufort West. Coll. Marais.

The posterior two thirds of a somewhat weathered skull lacking the snout.

Lateral view (Fig. 216). The articulatory region of the lower jaw has

recently been figured and described (Boonstra, 1953, p. 62). The prefrontal

has a sharp dorsal edge separating }

a small lateral from a larger dorsal ‘

face. The jugal is well developed

with an anterior ramus forming the ai

stout suborbital bar; a weaker SAMIIM44UL. vit

dorsal ramus forming the lower : od.

half of the orbital bar and a fairly

long posterior ramus, overlain by

an anterior ramus of the squamosal,

forming the anterior and much of

the ventral part of the lower

temporal arch. The postorbital is

small forming only a small part of

the orbital margin and posteriorly

extending as a weak tongue on to

the lateral face of the parietal.

The squamosal has a moderate

descending flange carrying the

quadrate complex but not extending

far ventrally so that the quadratic

condyle still lies well above the level

of the lower border of the dentary.

In dorsal view (Fig. 21a) it is

seen that the temporal fossa is

fairly long and wide, the inter- Fic. 21.—Maraisaurus parvus. Type.

temporal region narrow and the >”: cenit Boe ae et

sagittal crest low but sharp. The

frontal is long and has a fairly large entry into the orbital border: the

Vee tet

ay at

THE PRISTEROGNATHID THEROCEPHALIANS 105

postfrontal is well developed; the postorbital has only a small entry into the

orbital border and its posterior tongue weak and flanks the parietal for only

a short distance; the ridge on the prefrontal sharply separates the dorsal from

the lateral surface.

An Unidentified Pristerognathid.

S.A.M. 11959 (Fig. 22). Dikbome, Laingsburg. Coll. Boonstra.

Fragmentary weathered skull pieces.

The dental formula is 1.6, c.I, p-C-5. The incisors are robust, with the

6th strong, occupying 40 mm. on the left and 43 mm. on the right side.

There is no diastema between incisors and canine and the diastema between

canine and postcanines is very small (5 mm. on the right and 6 mm. on the

left side). The postcanines are irregular with the first, fourth and fifth fairly

weak but the third quite robust and occupying 36 mm. on the left and 45 mm.

on the right side. The articulatory region has recently been described

(Boonstra, 1953)-

This form appears to occupy a position intermediate between genera like

Scymnosaurus and Cynariognathus.

SAM, 11459.

Fic. 22.—An Unidentified Pristerognathid. S.A.M. 11959 a, Lateral view (x 4.)

b, Oblique view of the articular region. (x 3.) The quadratojugal is not preserved;

when present it rested on the ledge on the posterior surface of the quadrate.

DISCUSSION

As all recent authors derive the Therapsids from the Pelycosaurs it will

be of interest to compare the Pristerognathidae — the most primitive

Therocephalian family — with the Pelycosaurs. From Romer’s monograph

I have extracted a list of morphological features mentioned by him and am

comparing the same structures as identified in the Pristerognathids.

The Pristerognathids agree with the Pelycosaurs: in most forms the

frontal enters the supraorbital border, but in some this entry is small and in

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

some the frontal is secondarily excluded; a ridge separates a dorsal from a

lateral surface in the prefrontal, but the general dorsal surface is not sharply

separated from the lateral surface; otic notch closed at junction of squamosal

and tabular; step anterior to canine as in predaceous Pelycosaurs; quadrate

situated below plane of maxillary teeth; maxilla with its large canine crowds

out lacrimal to form a junction with the nasal as in the advanced Pelycosaurs;

squamosal covers most of posterior face of the quadrate; choanae well

forward; expansion of posterior end of prevomer; quadrate ramus of pterygoid

reaching quadrate; epipterygoid narrow, but not a typical columella cranu

and reaches parietal; quadrate in contact with paroccipital; angular notch

as in Sphenacodontidae.

The Pristerognathids differ from the Pelycosaurs: the parietal separates

the postorbital from the squamosal; there is no basal movable articulation;

the palate is not Rhynchocephalian-like; supratemporal absent; dorsal surface

not sharply demarcated from lateral surface and sides not steep; premaxilla

not extending posteriorly between nasals; parietal short but narrow; upper

edge of occiput not moved forward and occiput is thus not slanting and the

surface of the interparietal and tabular are not visible in dorsal view; condyle

and quadrates not in posterior position; lacrimal always with little anterior

extent; the postorbital bar is always slender, never developed as a plate

of bone between orbit and temporal fossa; maxilla never meets quadratojugal;

jugal though extending far posteriorly never meets the quadratojugal;

quadratojugal always very small; septomaxilla with large facial exposure;

only the anterior ramus of the pterygoid bearing teeth and no teeth on the

lateral pterygoidal flange nor any on the palatine and ectopterygoid; lateral

pterygoid flange never meeting the jugal; suborbital vacuity always present

and large; epipterygoid somewhat broader and reaching the parietal; angular

notch present and dentary always with a strong coronoid process.

It will furthermore be of interest to note in what characters the

Pristerognathids agree with the Cynodonts viz. dentary with large coronoid

process; sloping mentum; interparietal region narrow and sagittal crest;

postorbital does not meet squamosal; preparietal absent; posterior end of

prevomer spatulate and underlying palatines and pterygoids; jugal stretching

far posteriorly ventral to the anterior ramus of the overlying squamosal;

frontal sometimes excluded from orbital border as in Cynodonts.

REFERENCES

Boonstra, L. D. 1934. A Contribution to the Morphology of the Mammal-like

Reptiles of the Suborder Therocephalia. Ann. S. Afr. Mus. XXXI, pp. 215-267.

Boonstra, L. D. 1935. On Some South African Reptiles of the Suborder

Therocephalia preserved in the American Museum of Natural History. Am.

Mus, (Noy. (77 2.:) Pps ais b2:

Boonstra, L. D. 1953. The Lower Jaw Articulatory Region in some Pristerognathid

Therocephalians. Ann. S. Afr. Mus. XLII, 1, pp. 54-63.

_THE PRISTEROGNATHID THEROCEPHALIANS 107

Broom, R. 1903. On Some New Primitive Theriodonts in the South African Museum.

Ann. S. Afr. Mus. IV, pp. 147-158.

Broom, R. 1904. On Two New Therocephalian Reptiles (Glanosuchus macrops and

Pnsievopnatnus baini)..~ Trans. 5. Afr.’ Phil’ Soc. 15, pp. 85-88.

Broom, R. 1915. On Some New Carnivorous Therapsids in the Collection of the

British Museum. Proc. Zool. Soc., pp. 163-173.

Broom, R. 1915. Permian, Triassic and Jurassic Reptiles of South Africa. Bull.

ames vas. Nat. Hist. XXV, Il, pp. 105-164.

Broom, R. 1925. On some Carnivorous Therapsida. Rec. Alb. Mus. III, pp. 309-326.

Broom, R. 1931. Notices of some new Genera and Species of Karroo Fossil Reptiles.

Rec. Alb. Mus. IV, pp. 161-166.

HauGutTon, S. H. 1918. Some New Carnivorous Therapsida, with Notes upon the

Brain-Case in Certain Species. Ann. S. Afr. Mus. XII, p. 175-216.

SEELEY, H.G. 1895. On the Therosuchia. Phil. Trans. Roy. Soc. 185, pp. 987-1018.

8. The Cranial Structure of the Titanosuchian: Anteosaurus. By L. D.

Boonstra, D.Sc.

(With 22 text-figures.)

HISTORICAL

Although the Anteosaurus skull was first described by Broom as long ago

as 1910, the details of the cranial structure are still very inadequately known.

Watson in 1914 gave some details of the structure of the incomplete

skull in the British Museum and in 1921 the same author, giving details of

the snout, instituted the name Anteosaurus for the specimen hitherto thought

to be a Titanosuchus.

Broom in 1929 founded an additional species — A. minor — on a piece

of the skull roof.

Broili and Schréder in 1935 described certain skull fragments under the

name TJitanognathus lotzt.

In 1936 the present author figured and described a distorted skull of

A. minor that had been sold to the American Museum by Dr. Broom.

In the same year Broom described a good skull and lower jaw under the

name Dinosuchus vorstenr.

In 1948 the present author published a figure of a skull, which in 1952

was named A. abeli, and in 1953 a taxonomic account of the Titanosuchians

included a number of photos of Anteosaurus skulls in the South African

Museum.

MATERIAL

The present paper is based mainly on the large number of specimens in

the collection of the South African Museum, viz.

S.A.M. 2752. Vivier Siding, Beaufort West. Coll. Haughton & Whaits.

Posterior two thirds of the skull without basis cranii.

S.A.M. 4340. Leeurivier, Beaufort West. Coll. Haughton. A good skull,

though distorted by a simple shear, with part of the lower jaw.

S.A.M. 5621. Leeurivier, Beaufort West. Coll. Haughton. A snout and

part of the skull roof.

S.A.M. 9123. Voélfontein, Prince Albert. Coll. Boonstra. A weathered

skull.

S.A.M. 9139. Voélfontein, Prince Albert. Coll. Boonstra. A weathered

skull fragment.

108

CRANIAL STRUCTURE OF THE TITANOSUCHIAN I0Q

S.A.M. 9140. Voélfontein, Prince Albert. Coll. Boonstra. A partial

disarticulated skull.

S.A.M. 9329. Kruisvlei, Beaufort West. Coll. Boonstra. A good skull

and much of the lower jaw.

S.A.M. 11293. Boesmansrivier, Beaufort West. Coll. Boonstra. A good

weathered skull, slightly dorso-ventrally compressed and distorted,

with some bones of the occiput disarticulated.

S.A.M. 11296. Kruisrivier, Sutherland. Coll. Boonstra & Laurenson.

A very good skull and lower jaw, though somewhat distorted by a

simple shear.

S.A.M. 11302. Buffelsvlei, Beaufort West. Coll. Boonstra & Marais.

A fair, weathered skull and lower jaw.

S.A.M. 11492. Mynhardtskraal, Beaufort West. Coll. Boonstra. A

fairly complete skull without lower jaw.

S.A.M. 11576. Klein-Koedoeskop, Beaufort West. Coll. Boonstra. A

snout with fairly well preserved teeth.

S.A.M. 11577. Bulwater, Beaufort West. Coll. Boonstra & Truter. A

good practically undistorted three quarters of a well preserved

skull.

S.A.M. 11592. Dikbome, Laingsburg. Coll. Boonstra & Du Plessis.

A weathered skull, but with a good palatal region.

S.A.M. 11694. Koringplaas, Laingsburg. Coll. Boonstra & Du Plessis.

A very good undistorted skull without the lower jaw.

S.A.M. 11929. Kruisvlei, Beaufort West. Coll. Boonstra. The greater

part of a skull in intractable matrix.

S.A.M. 11946. Buffelsvlei, Beaufort West. Coll. Boonstra & Marais.

A nearly complete, good skull, slightly distorted by a simple shear.

S.A.M. 11949. Nuwefontein, Fraserburg. Coll. Boonstra & Jooste. A

partial skull, snout end occiput not in contact.

GENERAL SKULL FoRM

There is some difficulty in determining the correct skull form in

Anteosaurus, due to the post-mortem deformation the available material has

usually undergone. In only one skull (S.A.M. 11694) there appears to be

little disturbance of the original symmetry. In 8 the deformation is chiefly

due to dorso-ventral pressure, but accompanied by some measure of distortion.

In 5 specimens the skulls have been subjected to a shearing action — mostly

in the form of a simple shear. In these sheared specimens it is of interest

to note that they were all lying on their left sides. In only 1 specimen,

lying on its right side, the deformation was due almost wholly to compression

from side to side.

IIO ANNALS OF THE SOUTH AFRICAN MUSEUM

In the figures the legend indicates where an attempt has been made to

correct the effects of the deformation. The resulting correction made from the

dorsal and ventral aspect has in some cases not produced the same result and

a mean between the two results may indicate the correct condition, but not

necessarily so.

The Anteosaurus skull is large to very large (480-800 mm. max. length;

222-612 mm. max. width). The snout (with the mandible) is much higher

than broad and thus, notwithstanding the width over the temporal region,

the skull gives the impression of being high and narrow. This is in sharp

contrast to the other Titanosuchians and the Tapinocephalia. In those

specimens where the incisors are present the carnivorous nature is strongly

evident. The prominent boss-like development of the upper part of the

postorbital bar is striking, and in some specimens this rugged appearance

is further strengthened by the presence of bosses on the jugal and

angular and around the pineal foramen with a lesser or greater

amount of swelling of the frons. The great lateral and posterior sweep of

the temporal arches is characteristic; the temporal fossa is large and extends

far ventrally with a relatively narrow infra-temporal arch. The ventral

postero-lateral corner of the skull is not formed by the quadratojugal lying

on the surface, but, lying medially, gives an un-Deinocephalian-like

appearance to the Anteosaurus skull.

The orbits are of medium size and face anterolaterally; the nostrils are

non-terminal and lie laterally, whereas in all other Deinocephalians they are

directed much dorsally. The intertemporal region is fairly narrow, but

without any suggestion of a sagittal crest. The mentum of the lower jaw

is high and squarish. |

The anterior part of the dentigerous border of the upper jaw sweeps

sharply upwards, exposing the long intermeshing incisor teeth.

The lower jaw is hinged fairly far posteriorly.

a. The Skull in Lateral View (Figs. 1, 6, 8, 11, 1)

In lateral view the skull is roughly pearshaped in outline. The preorbital

portion is much longer than the postorbital part. The lateral direction of the

nostril, orbit and temporal opening is apparent. The temporal opening

extends far ventrally and the infratemporal bar is narrow. The quadrato-

jugal is not a bone of the surface but lies medially of the postero-lateral corner

of the skull. The side of the snout is fairly vertical.

The Premaxilla (P.M.) forms the anterior } of the upper edge of tne skull;

anteriorly it has a rounded curve to the dentigerous border, which curves

CRANIAL STRUCTURE OF THE TITANOSUCHIAN IIl

sharply upwards in antero-posterior direction; from the nostril a groove in

the surface of the bone’ runs anteriorly, lying above the curved

premaxilla-maxillary suture; on the lower border of the nostril the premaxilla

is separated from the septomaxilla by a narrow tongue of the maxilla; the

anterior 4 of the internarial bar is formed by a strong girder of the premaxilla,

but in the posterior half the nasal helps in forming the internarial bar;

‘ Bis ay ‘ 9

yO et

2° * Ar.

= o

-~e-<or? -.7

Fic. 1.—Anteosaurus abeli. Holotype. S.A.M. 11296. Kruisrivier, Sutherland.

Lateral view of the left side of the skull. (x 4.) The distortion due to a simple shear

corrected. In this and all the other figures, orthoprojections, obtained with a

pantograph, are given. In the lateral views the projection is on the median plane. In

the dorsal and ventral views the projection is on the plane in which the postcanines lie.

In the occipital views the projection is at right angles to the plane of the postcanine

alveolar border.

An.—angular; Ar.—articular; B.O.—basioccipital; B.S.—basisphenoid (sheathed by

parasphenoid); D.—dentary; E.O.—exoccipital; F.—frontal; I.P.—interparietal; J.—

jugal; L.—lacrimal; M.—maxilla; N.—nasal; P.—parietal; Pal.—palatine; P.M.—

premaxilla; P.O.—postorbital; P.O.c.—paroccipital; P.V.—prevomer; Po.F.—

postfrontal; Pr.F.—prefrontal; Pt.—pterygoid; Q.—dquadrate; Q.J.—quadratojugal;

S.A.—surangular; S.M.—septomaxilla; S.O.—supraoccipital; Sgq.—squamosal; St.—

stapes; IT.—tabular; Tr.—transversum.

posteriorly the premaxilla has a wedge-shaped prolongation into the nasal,

extending posteriorly of the nostril roughly for the length of the nostril, or

in some cases for double this length. The premaxilla is either edentulous or

carries a variable number of incisor teeth (1-5), with the number sometimes

varying in the two premaxillaries of the same skull. There is a considerable

variation in the length of the dentigerous border of the premaxilla, even

within what I consider to be the same species; this variation is concomitant

Ii2 ANNALS OF THE SOUTH AFRICAN MUSEUM

with both the number and size of the incisor teeth; in A. abeli, S.A.M. 4340,

the dentigerous edge on the left side with 5 incisors is 160 mm., whereas in

S.A.M. 11296 with 3 teeth it is 90 mm., and in the right size with 4 teeth

it is (ro) mam:

The Septomaxilla (S.M.) has a small facial exposure; dorsally it forms the

posterior part of the lower narial border; curving sharply inwards it extends

_ to near the median line to form the floor of the nostril. Anteriorly it does not

meet the premaxilla, being separated from this bone by a narrow tongue

of the maxilla. Posteriorly it extends as a short wedge in between the nasal

and maxilla. There is a small septomaxillary foramen.

The Nasal (N.) is in lateral view seen to be a long narrow bone with a

constricted waist. Anteriorly it forms the posterior border of the nostril, with

a dorsal prong extending in to the dorsal narial border and a ventral prong

extending in to the ventral narial border. Posteriorly it extends to about

halfway the swelling of the forehead, where it meets the frontal in a feebly

sigmoid suture.

The Maxilla (M.) is a large bone extending far posteriorly, where it

overlaps the jugal. In the smaller forms, e.g. A. cruentus (S.A.M. 11694),

it does not in its ventral part stretch so far posteriorly as in the larger forms,

so that in the former the suture with the jugal is oblique, whereas in the

latter it is vertical. On the surface of the maxilla there is a strong ridge

running obliquely from the orbit in the direction of the canine. In its

dorso-posterior part the maxilla has a triangular tongue which separates the

jugal from the lacrimal except for a short distance near the orbit. The

maxilla carries a variable number of teeth (5-8), often irregularly spaced.

The Frontal (F.) forms in lateral view the middle portion of the dorsal

edge of the skull. It forms the major portion of the fore-head swelling, as

this is varyingly developed in the different forms. The frontal has a small

tonguelike entry into the dorsal orbital border, except in S.A.M. 11296 where

the large postfrontal boss overlaps it.

The Prefrontal (Pr.F.) has in lateral view a curved dorsal edge and a

fairly straight ventral edge. Anteriorly it stretches as a wedge between the

nasal and maxilla, and posteriorly it forms the thickened anterodorsal part

of the orbital border. It forms the lateral part of the forehead swelling and,

where this is great, it forms an overhanging bulge.

The Lacrimal (L.) varies in size, due to the extent to which the maxilla

extends posteriorly. It is a roughly quadrilateral bone with a short suture

with the jugal and forming the anterior part of the orbital border. Its suture

with the prefrontal lies on a fairly sharp ridge.

The Jugal (J.) is quite a large bone notwithstanding the encroachment of

the maxilla anteriorly. It forms the lower orbital border and extends a little

on to the anterior as well as the posterior orbital border. It has only a small

CRANIAL STRUCTURE OF THE TITANOSUCHIAN EES

contact with the lacrimal. Entering the postorbital bar it is greatly overlapped

_by the postorbital. It has a long and strong posterior process which, in the

form of a sheet of bone, forms the inner surface of the zygomatic arch and

extends posteriorly to past the quadratojugal. The outer surface of the

process is overlapped by the zygomatic process of the squamosal. The

amount of this overlap varies in the different forms. In A. cruentus

(S.A.M. 11694) the outer surface of the jugal, below the orbit, is fairly

smooth, except for a hollow which is present in all forms immediately below

the orbital border, but in the larger forms there is a swelling, feeble in

A. abel, (S.A.M. 4340), low in A. abeli (S.A.M. 11296) and A. vorsten

(S.A.M. 11577) and strong and prominent in A. crassifrons (S.A.M. 11946

and S.A.M. 11302) and A. acutirostris (S.A.M. 9329) where it forms a

prominent overhanging outgrowth.

The Postfrontal (Po.F.) forms the large and prominent boss-like growth

overhanging the postero-dorsal orbital border. The size of this boss

determines the amount of its overgrowth over the surface of the frontal and

the size of the entry of the frontal tongue into the dorsal orbital border.

Ventrally the extent of the pachyostosis in the postfrontal determines the

degree to which it overlaps the postorbital on the postorbital bar so that the

position of the postfrontal-postorbital suture shows a considerable variation.

In the forms where the thickening of the frontal is not very great the

postfrontal forms part of the dorsal outline of the skull when observed in

lateral view; in the other forms it nearly reaches the dorsal edge or lies

well below it.

The Postorbital (P.O.) presents, in lateral view, two distinct surfaces.

The more lateral surface is exposed where the bone forms the fairly weak and

Narrow curved part of the postorbital bar overhung to a greater or lesser

extent by the bosslike postfrontal. In a more medial plane the postorbital

provides the sheet of bone which forms the lateral face of the intertemporal

region and forms the dorsal edge of the temporal fossa. There is a considerable

variation in the size and shape of this plate of bone. Posteriorly it meets the

upsweeping arm of the squamosal in a long suture. Ventrally its edge

overhangs the squamosal where this bone forms the inner face of the

temporal fossa.

The Parietal (P.) is in lateral view seen only where it presents the lateral

surface of the pineal boss and its long postero-lateral tongue intercalated

between the squamosal and tabular. It does not form the upper inner face

of the temporal fossa as it is here covered by the postorbital which forms

the inner face of the fossa.

The Tabular (T.) in lateral view is seen to form the postero-dorsal border

of the posttemporal arch and in some forms the tabular is also seen as it

4— Annals

II4 ANNALS OF THE SOUTH AFRICAN MUSEUM

descends on the posterior surface forming the prominent ridge limiting the

occiput proper.

The Squamosal (Sq.) is a large bone of intricate shape. In lateral view

it is seen to form the lateral part of the posttemporal arch and the greater

part of the inner surface of the temporal fossa, except for that part formed

by the postorbital, and the outer surface of most of the zygomatic arch.

Anteriorly it meets, within the temporal fossa, the postorbital, and lower

down the supraoccipital and paroccipital. In the angle between the zygomatic

process and the inner facing of the temporal fossa the squamosal also forms

the niche in which the upper and hinder faces of the quadrate are housed.

The Quadratojugal (Q.J.) is not preserved in most of the specimens, but

in the four skulls in which it is preserved, its lateral surface appears to be

triangular in outline and this does not lie on the surface of the postero-ventral

outer corner of the skull as it does in all other known Deinocephalia, but lies

in a plane medial to the subtemporal arch as in Therocephalians,

Gorgonopsians, and the higher Therapsids. The quadratojugal rests on a

ledge of the quadrate above the outer quadratic condyle. In these relations

it approaches very closely to the condition recently described by me in the

contemporary Pristerognathid Therocephalians. In one skull of A. cruentus

(S.A.M. 11694) there is a small notch in the squamosal lying lateral to the

quadratojugal.

Palatal bones. When the lower jaw is disarticulated the lateral surfaces

of some of the palatal bones are seen. The lateral surface of the lateral flange

of the pterygoid (Pt.), the quadrate ramus of the pterygoid and of the

anterior pterygoid process, the transversum and palatine are all visible. In

A. cruentus (S.A.M. 11694) the basisphenoid is also partly seen.

Noteworthy are the strong and deep pterygoidal flange and the prominent

dentigerous boss on the palatine.

bl The Skulliin Dorsal View {Figs"2, 7, 9; 10, 14;) 18; 21, 22)

In dorsal view the Anteosaurus skull presents in outline a pear-shaped

form, with the jugal bosses in some forms appearing as lateral protuberances,

in some cases low and in others prominent. In A. crassifrons the skull is

short and squat with a relatively broad snout. In the others the snout is

either narrow, fairly narrow or moderately wide. The forehead swelling

is low, moderate to very massive. In all the postfrontal boss on the dorsal

part of the postorbital bar is prominent. In A. acutirostris (S.A.M. 9329)

and A. crassifrons (S.A.M. 11946) the jugal boss protrudes strongly beyond

the general skull-outline; in A. abeli (S.A.M. 11296) and A. vorsten (S.A.M.

11577) moderately, and in the others not at all.

The orbits are in all cases only partly visible, being overhung by the

fore-head swelling; in A. cruentus, A. abeli and A. vorstert this overhang

CRANIAL STRUCTURE OF THE TITANOSUCHIAN EES

is least, in A. levops and A. acutirosiris fairly great and in A. crassifrons

very great.

Fic. 2.—Anteosaurus abeli. Holotype. S.A.M. 11296,

Kruisrivier, Sutherland. Dorsal View with distortion

due to a simple shear corrected. (x %-)

The temporal fossa is broad and long with the temporal arches flaring

greatly laterally and posteriorly. The occipital edge is deeply concave from

side to side. The intertemporal region is generally flat, but with a pineal

boss varying from low to prominent and massive. The intertemporal width is

small, medium to fairly broad with no suggestion of the development of a

sagittal crest. The pineal foramen is fairly small and situated near, to very

near, the occipital edge. The internarial bar is strong and massive.

The Premaxilla (P.M.). The two premaxillaries together form the rounded

anterior margin of the skull. Laterally they meet the maxilla in a curved

I16 ANNALS OF THE SOUTH AFRICAN MUSEUM

suture lying lateral to a groove in the premaxillary surface. The premaxilla

forms the anterior and most of the dorsal narial border. Together they form

a stout internarial bar, which, extending backwards for only a short distance,

is posteriorly abruptly truncated. This truncation is rather surprising since

the premaxilla-nasal suture lies in a groove which is continued to near the

fronto-nasal suture. This accounts for the error of observation on my part

as depicted in the figure of the specimen in the American Museum, which later

in this paper is made the type of a new species, A. minusculus.

The Septomaxilla (S.M.) is as already described in lateral view.

The Maxilla (M.), in addition to the features described when seen in lateral

view, shows clearly that the suture between it and the septomaxilla and nasal

lies along a curved ridge.

The Nasal (N.). The paired nasals are hourglass-shaped in outline, with

the truncated posterior end of the premaxillaries separating them in their

anterior half to a third. The posterior part of the nasals enters the fore-head

swelling to a greater or lesser extent depending on the development of this

thickening. Where the frontal swelling is least, the nasal extends furthest

posteriorly.

The Frontal (F.). The paired frontals form a more or less pronounced

cruciform figure with the narrow tongues entering the orbital borders

forming the cross member. In two cases this cross member is truncated and

on the surface does not enter the orbital border. The postfrontal boss

overflows on to the frontal surface to a varying extent. Posteriorly the

frontals meet the parietals in a suture running across the skull just anterior to

the pineal boss, but in A. crassifrons the pineal boss extends anteriorly as

a strong swelling on to the posterior part of the frontals. The frontals meet

the edge of the postorbitals posterior to the limits of the postfrontals.

The Prefrontal (Pr.F.). In dorsal view the extent to which the prefrontal

enters the fore-head swelling is indicated in the figures. Where this swelling

is least developed the overhang of the prefrontal over the orbit and the

lateral skull face immediately anterior to the orbit is least and the transition

from the dorsal to the lateral surface not very abrupt. Whereas in

A. crassifrons the swollen prefrontal all but obscures the lacrimal from

dorsal view.

The Lacrnmal (L.) is as described in lateral view, except that in dorsal

view it is overhung by the prefrontal swelling to a greater or lessor extent

depending on the amount of the pachyostosis of the prefrontal.

The Postfrontal (Po.F.). In dorsal view the knob-like bosses formed by

the greatly thickened postfrontals are a very prominent feature. In most

skulls this boss completely overhangs the part of the postorbital entering the

postorbital bar; in others the lower part of the postorbital above the suture

with the jugal is visible, and in three cases the boss leaves exposed, behind

CRANIAL STRUCTURE OF THE TITANOSUCHIAN G7

its posterior edge, the postorbital as it sweeps from the upper temporal face

down to the lower part of the postorbital bar.

The Parietal (P.) has a relatively small dorsal surface. The two bones

are fused except where they are pierced by the fairly small pineal foramen.

Round the pineal foramen the parietals are thickened to form a boss variable

in size and shape. In some skulls a distinct ring wall with sharp edges

surrounds the foramen, in others only a low mound is formed, whereas in

A. crassifrons a massive thickening extends into the frontals and in

A. vorsteri (S.A.M. 11577) the frontal surface, anterior to the low boss, is

slightly hollowed out in the form of a V shaped depression bounded by a

low ridge. In one specimen (S.A.M. 2752) the pineal boss overhangs the

occiput. The pineal foramen is near or very near the posterior edge of

the parietals.

Posteriorly each parietal sends out a long horn with its end wedged in

between the dorsal end of the tabular and the upper part of the squamosal.

The outer edge of the parietal horn forms a sharp ridge lying lateral to a

groove and thus sharply demarcating the intertemporal surface.

The Postorbital (P.O.). The posterior flange of the postorbital is a large

sheet of bone applied to the outer surface of the parietal and, extending far

back, it is met by the upsweeping squamosal. The ventral edge of the

postorbital flange overlaps and overhangs the antero-medially directed sheet

of the squamosal.

Little of that part of the postorbital which helps to make up the postorbital

bar is usually seen in dorsal view, being overhung by the greatly swollen

posttrontal.

The Jugal (J.). In dorsal view the boss of the jugal, in those forms

where it is developed, is shown very clearly as it determines the outline of

the skull in this region.

The Squamosal (Sq.) in dorsal view presents a zygomatic process covering

the latero-dorsal surface of the long zygomatic process of the jugal. From

this level the squamosal sweeps upwards to form the postero-lateral part of

the posttemporal arch, where it meets the postero-laterally sweeping horn

of the parietal and posteriorly abuts against the tabular. Within the temporal

fossa the squamosal forms its large internal surface; dorsally meeting the

postorbital and, at a lower level, sweeps antero-medially to meet the

supraoccipital and paroccipital. Between the zygomatic process and the

antero-medial process a niche in the anterior squamosal surface houses

the quadrate.

The Interparietal (1.P.), and Supraocctfital (S.O.). In dorsal view little

of the occiput is seen as this is vertical. The dorsal edge of the interparietal

and in some cases a bit of the surface of the supraoccipital can be seen, and

in A. acuttrostris the occipital condyle.

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

c. The Skull in Ventral View (Figs. 3, 13, 15, 16, 19)

In general terms it may be said that in the ventral aspect of the skull

two surfaces are shown meeting in an obtuse angle at the plane of the

transverse pterygoid flanges. Posterior to the very prominent transverse

pterygoid flanges the basis cranu lies in a more or less horizontal plane,

whereas anteriorly the palate is in anterior direction directed upwards. This

is accentuated by the upward retreat of the alveolar border of the

premaxillaries. The most prominent features in the ventral aspect of the

skull are: the strong and deep lateral flanges of the pterygoid, the strong

reniform dentigerous bosses of the palatines, the retreat of the alveolar border

era

Ae oe

DTT eR Wea et

z 22

Yo +2 ete ew? ’

--”

Fic, 3.—Anteosaurus vorsteri. Referred specimen. S.A.M. 11577,

Bulwater, Beaufort West. Ventral View. (x %-)

CRANIAL STRUCTURE OF THE TITANOSUCHIAN IIQ

of the premaxillaries, the medial shift of the quadratojugal away from the

lateral surface of the skull and the deeply concave occiput.

The Premaxilla (P.M.) has an alveolar border and a palatal process. The

alveolar border has retreated upwards. It carries a variable number of teeth —

5, 4, 3. The number of incisors in the two premaxillaries of the same skull

is, in some cases, unequal. Where no teeth roots are preserved there is a

groove in the alveolar surface which is either clearly separated into separate

alveoli, or not. The palatal process meets the prevomers which underly it,

and laterally meet the maxilla along a curved suture.

The Maxilla (M.). Ventrally the maxilla has, in addition to its alveolar

_edge, a medially directed tongue which separates the premaxilla from the

palatine and except in A. crassifrons makes contact with the prevomer.

_ Antero-medially of the upper canine the maxilla is deeply excavated to receive

the fang of the lower canine. The maxilla carries a large canine and a

variable number of postcanines. The latter are irregularly spaced and vary

_ in size and in number (5-8). The root cross-sections show the postcanines to

be rather feeble teeth of no great functional importance.

The Prevomer (P.V.). The pair of prevomers form a strong interchoanal

| bar, anteriorly underlying the palatal process of the premaxillaries and

_ posteriorly tapering in between the palatines to make a small contact with

the pterygoids. The interchoanal bar is, except anteriorly, deeply excavated

longitudinally and each prevomer has a well defined ridge on its lateral edge,

which forms the medial border of the slitlike choanae.

The Palatine (Pal.) meets ihe maxilla in a long suture lingually of the

_ alveolar border and curving upwards forms most of the lateral and posterior

_ border of the choana. Further back it carries a strong and prominent reniform

dentigerous boss. In most cases the fair-sized recurved teeth are implanted

in a single curve but in A. vorstert (S.A.M. 11577) there appears to be a double

row. Posterior to the boss there is a ridge continued on the pterygoid towards

the interpterygoidal slit.

The Transversunt (Tr.), composed of a sheet of bone lying antero-laterally

of the pterygoid flange, meets the palatine, maxilla and jugal and descending

along the front and side of the pterygoid flange forms the upper part of the

lateral and part of the anterior face of the strong lateral pterygoid flange.

There is no fenestra between the transversum and palatine.

The Pterygoid (Pt.) has the usual complex form of the Therapsids. The

anterior process is short, the. lateral flange deep and strong, the quadrate

process sweeps backwards with a straight outer edge and in the middle line

the pterygoids together form a fairly deep keel posterior to the interpterygoidal

slit. In the fork between the two quadrate rami the pterygoids meet the

basisphenoid (basi-parasphenoid) in a U-shaped suture with no development

of basipterygoid processes.

120 ANNALS OF THE SOUTH AFRICAN MUSEUM

In most skulls the posterior extremity of the quadrate ramus is not

adequately preserved and its relations with the quadrate uncertain. But in

a specimen which I believe to be an A. crassifrons (S.A.M. 11929) it would

appear that the quadrate ramus of the pterygoid is applied to the inner face

of the quadrate a little below the distal end of the stapes. If this observation

is correct then the condition is very similar to that in Jonkeria and in the

Tapinocephalids generally.

The Basisphenoid (B.S.) is apparently sheathed along its under surface

by the parasphenoid and presents a ventral surface cup-shaped in outline.

Just behind the median pterygoidal keel the median basisphenoidal surface

is excavated to form a broad groove, which is continued in the basioccipital

right up to the condyle. Lateral to the median groove the basisphenoid

develops a rounded ridge which broadens posteriorly until it is posteriorly

notched and develops a latero-posteriorly directed tongue, which forms the

antero-ventral border of the fenestra ovalis. The basioccipital-basisphenoidal

suture runs across the median groove from notch to notch. The

basisphenoidal-pterygoid suture runs along the lateral border of the rounded

basisphenoidal ridge.

The basisphenoid is a short bone, but in A. major (S.A.M. 11293) it is

nearly twice as long as in all the other skulls.

The Bastoccipital (B.O.) forms the greater part of the rounded condyle

with the exoccipitals forming the postero-lateral corners. In A. crassifrons and

A. acutirostris the exoccipitals are intimately fused to the basioccipital to form

a rounded condyle. Anterior to the condyle the under surface of the

basioccipital is in its median part hollowed out to form a wide groove which

is continued on to the basisphenoid. The vertical plate of the basioccipital

anterior to the condyle reported by Broom in A. vorstem is not shown in any

of the skulls I have examined and this incorrect statement can thus only be

due to an error in observation. Antero-laterally the corner of the basioccipital

is bent sharply downwards to form the strong medial border of the fenestra

ovalis. Postero-laterally the continuation of this prominent ridge round the

fenestra ovalis is formed by the antero-medial corner of the paroccipital,

which has a similarly bent down process. The basioccipital abuts against the

paroccipital in a diagonal line of contact.

The Exoccipital (E.O.), besides forming the postero-lateral corner of the

condyle, has a lateral process which overlaps the paroccipital and

supraoccipital, but is not very clearly shown in any of the skulls examined.

The Paroccipital (P.O.) forms a stout girderlike bone medially butting

against the basioccipital and distally applied to the inner face of the quadrate

and making contact with the descending sheet of the squamosal. The

dorso-lateral corner of the paroccipital bounds the post-temporal fossa, which

is anteriorly closed by the sheet of the squamosal forming the inner lining

CRANIAL STRUCTURE OF THE TITANOSUCHIAN UAE

of the temporal fossa. Dorsally the paroccipital lies against the lower edge

of the supraoccipital.

The Jugal (J.) in ventral view shows its inner surface. Its zygomatic

process is here clearly seen to extend far backwards and to form the inner

part of the arch, whose outer half is formed by the overlapping zygomatic

process of the squamosal.

The Squamosal (Sq.) shows its posterior surface as it sweeps downwards

{o overlap the posterior face of the quadrate and quadratojugal.

The Quadrate (Q.) and Quadratojugal (Q.J.). In most of the skulls

studiec the quadrate complex is missing. It would appear that it fits rather

loosely in the notch of the squamosal in which its upper end is housed.

There was apparently also little ankylosis in the joint between the paroccipital

and the quadrate. The quadraie and quadratojugal form an antero-posteriorly

flattened mass of bone carrying two moderate condyli on its lower edge,

which lies somewhat diagonally in the skull, with the outer condyle furthest

posteriorly. The quadratojugal is small and resting on a ledge above the

outer quadrate condyle is applied to the outer edge of the quadrate. It lies

well medial of the lateral edge of the skull and is not a bone of the lateral

surface as it is in all other known Deinocephalia.

d. The Skull in Occipital View (Figs. 4, 20)

The occiput is vertical, but deeply concave from side to side. The strong

tabular ridges bound the occiput proper laterally. Lateral to this ridge lies

the posterior surface of the posttemporal arch, and the groove which in higher

Therapsids functions as an auditory groove.

Fic. .4.—Anteosaurus laticeps. Sp. Nov. Holotype. S.A.M.

11592, Dikbome, Laingsburg. Occipital View. The _ possible

dorso-ventral compression not corrected. The dorsal part and

the quadrate-complex restored from all the other material. (x 4.)

I22 ANNALS OF THE SOUTH AFRICAN MUSEUM

The Squamosal (Sq.) is seen to form the lateral margin of the skull.

Dorsally the squamosal and tabular clasp the extremity of the parietal horn.

Ventrally the squamosal overlaps the quadrate and quadratojugal. Medially

it makes contact with the tabular in the ‘‘auditory groove’’, which lies lateral

to the strong tabular ridge. This ridge is ventrally continued on the squamosa!

as it fades out. This ridge thus differs materially from the ridge in Jonkena,

where the tabular, squamosal and paroccipital contribute to its formation.

The Tabular (T.) carries the strong ridge which forms the inner border

of the “‘auditory groove’’ and laterally limits the occiput proper. Externally

the tabular ridge appears to be sheathed by a thin bone, which I thought

may represent the supratemporal, but a number of cross sections in one skull

have yielded no evidence to substantiate this. From the ridge the tabular

extends medially as a sheet of bone to meet the interparietal. In its dorsal

part the occiput consists of 3 !ayers of bone viz. anteriorly there is a sheet of

the squamosal applied to a sheet of the parietal and this is posteriorly covered

by the tabular laterally and the interparietal medially.

The Interparietal (1.P.), as a large thin sheet of bone with a ridge in the

median line, forms the upper and middle part of the occipital surface.

The Supraoccipital (S.O.) lies below the interparietal and tabular and

forms the upper border of the foramen magnum and, laterally, the upper

border of the depression which represents the post-temporal fenestra now

anteriorly closed by the overgrowing sheet of the squamosal within the

temporal fossa. Lateral to the foramen magnum the supraoccipital is

overlapped by the exoccipital.

The Exocctpital (E.O.) is seen to form the dorso-lateral segment of the

condyle and its lateral flange overlaps the supraoccipital and paroccipitai

for a short distance.

The Paroccipital (P.O.) is a stout bar between the quadrate and the

basioccipital. In posterior view it is clearly shown how its proximo-ventral

corner is bent sharply downwards to form the posterior part of the prominent

rim of the fenestra ovalis. Similarly the downwardly directed processes of

the basioccipital and basisphenoid forming the median part of the rim of the

fenestra ovalis are clearly seen in occipital view. The median groove in the

ventral surface of the basioccivital and basisphenoid is also evident.

Stapes (St.) is only partly preserved in one of the skulls studied. It

appears to be a stout rod-lhke bone similar to that of other Deinocephalians

but longer.

The Quadrate (Q.) and Quadratojugal (Q.J.). In posterior view it is

very clear that the quadratojugal has shifted in medial direction as in the

higher Therapsids and is no longer a bone of the outer lateral surface of

the skull as it is in all other known Deinocephalia.

‘CRANIAL STRUCTURE OF THE TITANOSUCHIAN 123

The Pierygoid (Pt.). In occipital view the deep lateral flanges of the

pterygoids are well shown.

The Post-temporal Fenestra does not penetrate the occipital plate as it

is anteriorly closed by a sheet of the squamosal. In occipital view it is thus

only evident as a depression lying in the corner between the supraoccipital,

paroccipital and squamosal.

e. The Lower Jaw (Figs. 1, 5, 6)

Since the gquadrate has not shifted much in anterior direction the lower

jaw is long. The mandibular ramus is strong and heavy, particularly in its

anterior part. In all the larger forms there is a strong and prominent boss

on the anterior part of the angular. Unfortunately no lower jaw is preserved

in those forms without a jugal boss, but I think it likely that in these cases

no angular boss will be developed. In contradistinction to the condition

in other Deinocephalia there is in Anteosaurus some indication of the

development of a low coronoid process to the dentary.

In my material only the outer surface of the mandible is shown exposing

the dentary, angular, surangular and articular.

The Dentary (D.) is a massive bone. The mentum is high and fairly

upright and the symphysis strongly ankylosed. Posteriorly the dentary

curves upwards to form a low incipient coronoid process. The dentary

carries a strong canine and a variable number of incisors and postcanines.

The number of incisors varies from 2 to 4. The number of postcanines is

difficult to determine but seems to vary from 4 to 7.

The Angular (An.) is remarkable for the development on the outer surface

of a large egg-shaped swelling in its anterior part. Posterior to the boss the

angular has the typical Therapsid structure.

The Surangular (S.A.) is in outer view the typical curved girder-like bone

it is in Therapsids generally, but anteriorly it rises and meets the dentary

in the low coronoid process.

The Articular (Ar.) has only a small external face forming the extreme

postero-ventral corner of the lower jaw.

f. The Dentition (Figs. 1, 5, 6)

The dentition of Anteosaurus is clearly that of a specialised type of

carnivore. With the large canines, long intermeshing incisors, and feeble

postcanines it obviously did no chewing and very little shearing, but was

rather well adapted for grabbing and then tearing flesh from its victim, and

just before deglutition the lump of flesh was held by the recurved teeth

situated on the palatine bosses.

I24 ANNALS OF THE SOUTH AFRICAN MUSEUM

The Anteosaurus-incisors are quite distinct from those

in the other carnivorous Therapsids, as for example those

usually present in the Therocephalians Gorgonopsians and

Cynodonts. In the latter the upper incisors in occlusion

lie labially of the lower incisors, whereas in Anteosaurus

the two sets intermesh as do the herbivorous teeth of the

Tapinocephalia. To give the long incisors functioning

space the alveolar border of the premaxilla retreats and

the upper incisors are directed much anteriorly. The first

incisor is smaller than the second and with its fellow forms

a distinct pair, as is also the case in the Pristerognathid

Therocephalians. This pair passes in between the number ane : sO Ss coe

one pair of incisors of the dentary. When fully developed ferred specimen.

there are five upper and four lower incisors, but even in Pat ty

the same skull the number in the two halves is mostly Beaufort West.

different.‘ So!-we have; "in "S.A.M)’ 11576, 5 left’ upper, UEHOECIae” 3

4 right upper and in both sides 3 lower; in S.A.M. 11296, the plane of the

3 left upper, 4 mght upper and 2 lower on both sides; in Occt IME a)

S.A.M. 4340, 5 left upper, 4 right upper and 4 left lower; in S.A.M. 11694 no

uppers; in crassifrons no uppers; in S.A.M. 11577, 2 left and 1 right upper; in

S.A.M. 11492, 3 left and no right uppers. In those cases where no incisors

or incisor roots are preserved there is in the alveolar border a more or less

el ao OO ee alee ee ee Mit ie ee a oe

i et Ta ene eeer”?

Fic. 6.—Anteosavvus abeli. Paratype. S.A.M. 4340, Leeurivier, Beaufort West.

Lateral View. (x 4%.)

CRANIAL STRUCTURE OF THE TITANOSUCHIAN I25

distinct groove which in some shows more or less indefinite subdivision into

separate alveoli. Without sectioning it is difficult to determine whether some

forms were permanently edentulous (rather improbable) or whether the first

set has not yet erupted or whether the last set has been shed or whether an

antecedent set has been shed and a succeeding set not yet erupted.

In S.A.M. 11296 the left canine is on the point of being shed and the

cusp lying posterior to it may be the replacing canine as it does not have the

appearance of a postcanine.

The postcanines are not well shown and we usually only have the section

of the roots on which to base a count. In S.A.M. 11529 some crowns are

well preserved and here the last tooth is a fairly small tooth, 16 mm. long,

linguo-labially compressed with diameters 1r and 7 mm. In outline the

postcanine is fairly bluntly conical. The postcanines are usually irregularly

spaced. Their number sometimes varies in the two maxillaries of the same

skull. In the genus the postcanines vary in number from 4 to 8 in the

upper jaw.

Taxonomic

In 1921 Watson established the genus Anteosaurus on the grounds that a

skull in the British Museum (R. 3595) from Tamboerfontein differed from

all hitherto known Deinocephalians in the possession of only three incisor

teeth and in being the first known Deinocephalian skull in which the upper

part of the postorbital bar was strongly swollen to form a very prominent boss.

Subsequent finds have shown that in a series of skulls with the typical

boss the number of incisors varies and I propose that as diagnostic character

for the genus Anteosaurus we should consider only the presence of the typical

boss formed by the prefrontal bone.

Accepting this character as diagnostic we have to exclude from the genus

Anteosaurus the form named A. minor by Broom who says of it, “‘but differs

in having only a small thickening instead of a huge boss in the postfrontal

region’. For this unsatisfactory type, consisting of only the interorbital and

intertemporal! regions, nomenclatural procedure thus compels one to propose

a new generic name. I therefore propose that the skull fragment from

Merweville, British Museum (Natural History) R.5742, be known under the

generic name Pseudanteosaurus gen. nov., genotype Pseudanteosaurus

minor (Broom).

The specimen in the American Museum of Natural History stated by

Broom to be the topotype of the above has, however, well developed bulbous

swellings in the upper part of the postorbital bar and must be retained in

the genus Anteosaurus of which it then constitutes a new species to be named

and described later in this paper.

1260 ANNALS OF THE SOUTH AFRICAN MUSEUM

Broili and Schroéder’s genus Titanognathus was based on the following

diagnostic characters: ‘“Schadel mit schmaler und steil vom pramaxillaren

Kieferrand aufsteigender Schnauze, sehr gross. Praemaxillarer Kieferrand

gegentiber dem maxillaren stark in die Hohe gezogen, Symphysenregion des

Unterkiefers entsprechend erhoht gegentiber dem riickwartigen abschnitt des

Dentale. Zahnformel: ipa ly esas (OCles,

4(?) I Boiee

together with the additional inaterial now known has established that the

retreat of the premaxillary edge occurs in Anteosaurus and the above dental

formula is also within the limits of the genus. The generic name

Titanognathus thus becomes a synonym of Anteosaurus but we may continue

to regard this specimen as specifically distinct from the genotype A. magnificus

under the name A. Jotz: (Broili and Schroder).

Broom’s genus Dinosuchus was established on the following diagnostic

characters: presence of a dentigerous palatine boss, low position of temporal

arch, great width of occiput, large size, dental formula 1.4, c.1, p.c.5, huge

angular boss. In the large number of skulls now known to fall within the

limits set for the genus these above characters are present and any difference

in degree cannot be more than of specific value. Dinosuchus thus becomes a

synonym of Anteosaurus and the points in which Broom’s specimen differs

from the genotype warrant the retention of Broom’s specific name, vorstert.

Specific descriptions of the known and new species can now be given:

Restudy of the genotype

Anteosaurus magnificus Watson.

Broom, R. 1910. Tttanosuchus ferrox Owen (in errore).

Watson, D. M.S. 1914. Titanosuchus ferox Owen (in errore).

Watson, D. M. S. i921. Anteosaurus magnificus Watson.

Genotype. Incomplete skull. British Museum (Natural History) R. 3595.

Tamboerfontein, Beaufort West. Coll. Seeley.

In the type skull the snout was not in contact with the rest of the

skull and in his restoration Watson failed to realise that the premaxillary

edge curved upwards. This feature of the Anteosaurus skull was first reported

by Broili and Schréder and confirmed in al! subsequent specimens. Due to

the incompleteness of the type skull only an incomplete description can be

given for the species viz. :

Skull large; maximum length about 660 mm. Snout long, fairly high and

wide. Intersquamosal width not great (360? mm.). Postfrontal boss huge

and prominent. Jugal and angular bosses unknown. Fronto-naso-prefrontal

swelling fairly weak. Pineal boss prominent with sharp circular border,

situated very near occipital edge. Degree of upward inclination of

premaxillary edge uncertain. The occiput is fairly high, fairly wide, deeply

CRANIAL STRUCTURE OF THE TITANOSUCHIAN 127

concave with a great posterior sweep of the temporal arches and the upper

part of the temporal fossa roomy antero-posteriorly; the temporal arch rises

well above the plane of the intertemporal surface. Palate probably long.

Basts crani unknown. Condyle unknown. Watson gives the upper teeth

as 1.3, c.I, p.c. 8 on both sides, but Broom states that there may be 5 incisors.

Anteosaurus lotzi (Broili and Schroder).

Broili, F. and Schroder, J. 1935. Tttanognathus lotzi Broili and

Schroder.

Boonstra, L. D. 1953. Anteosaurus lotzi (Broili and Schréder).

Holotype. Skull fragments and some postcranial bones. Alte Akademie,

Miinchen. No. ?. Brakwater, Beaufort West. Coll. Schréder.

Se mle = = ol ele «

ba]

e -

Phat eo

. e

“em eevee?”

Fic. 7.—Anteosaurus abel. Referred specimen. S.A.M. 11949,

Nuwefontein (Roxana), Fraserburg. Dorsal view. This was the

first skull in which the truncated posterior limit of the

premaxillaries was determined. (x +.)

128 ANNALS OF THE SOUTH AFRICAN MUSEUM

As the type is rather unsatisfactory, consisting of only a fragmentary snout,

mandibular fragments, incomplete ilium, pubis and femur, only a very

incomplete diagnosis can be given viz. :

Skull size unknown, probably large. Snout length unknown, fairly high

and narrow. Intersquamosal width unknown. Postfrontal boss unknown.

Jugal boss unknown, angular boss strong. Fronto-naso-prefrontal swelling

unknown. Pineal boss unknown. Sharp upward inclination of premaxillary

edge, and Broili and Schroder record an accompanying step-up of the

precanine border of the dentary. The occiput is unknown. Palate unknown.

Basis cranu unknown. Condyle unknown. Broili and Schréder give the

iD I 6

dental formula i. pea.

4’ I Sain

Anteosaurus vorsteri (Broom).

Broom, R. 1936. Dinosuchus vorsteri Broom.

Boonstra, L. D. 1953. Anteosaurus vorsten (Broom).

Holotype. A good skull and part of the lower jaw. Transvaal Museum.

265. Stinkfontein, Prince Albert. Coll. Vorster, Botes and Broom.

Skull very large; maximum length 740 mm. Snout fairly long, high and

wide. Intersquamosal width very great (600 mm.). Size of postfrontal boss

unknown, probably fairly strong. Jugal boss unknown, angular boss very

strong. Fronto-naso-prefrontal swelling unknown, probably fairly weak.

Pineal boss unknown, near? occipital edge. Sharp upward inclination of

premaxillary edge. The occiput is high, wide, moderately deeply concave,

with a fairly great posterior sweep of the temporal arches and the upper

part of the temporal fossa roomy antero-posteriorly; the temporal arch not

rising above the plane of the intertemporal surface. Palate fairly long, with

deep lateral pterygoidal flanges not massive with sharp ventral edge. Basis

cranii short, with short basisphenoid. Exoccipitals fused with basioccipital

to form a rounded condyle. Broom gives the upper teeth as 1.4, c.I, p.c.5.

Referred Specimen. (Figs. 3, 8, 9.) A good skull, but lacking the

basis cranu. S.A.M. 11577. Bulwater, Beaufort West. Coll. Boonstra

and Truter.

Those areas of the outer skull surface lost by weathering in the type

specimen are here perfectly preserved and the description of this species can

thus be augmented as follows: postfrontal boss fairly strong, but not obscuring

the lower part of the postorbital in dorsal view; jugal boss low and small:

fronto-naso-prefrontal swelling weak, running evenly on to the anterior nasal

surface and with little overhanging of the sides of the skull; pineal boss low,

with rounded edges not near occipital edge, with hollow on frontal anterior

CRANIAL STRUCTURE OF THE TITANOSUCHIAN I29

to the boss; palatine bosses reniform, not circular as described by Broom;

in the left premaxilla stumps of two incisors are preserved and on the right

one; for the rest the alveolar border presents a matrix filled groove with little

indication of separate alveoli; on the left there appears to be room for 5 and on

the right for a total of 4 incisors; on the left alveolar border of the maxilla

Fic. 8.—Anteosaurus vorsteri. Referred specimen. S.A.M. 11577, Bulwater,

Beaufort West. Lateral view. (x }.)

cross-sections of 7 fairly regular postcanines are seen and on the right 6.

This specimen is of some historical interest in that it was known by the

local people to have been exposed when Seeley collected the well-known

Tamboerfontein specimen of Bradysaurus baim. Weathering is thus not

very rapid in the Tapinocephalus-zone of the Koup.

Anteosaurus abeli (Boonstra). (Figs. I, 2, 5, 6, 7.)

Boonstra, L. D. 1952. Anteosaurus abel (Boonstra).

Boonstra, L. D. 1953. Anteosaurus abel (Boonstra).

Holotype. A good skull and lower jaw. S.A.M. 11296. Kruisrivier,

Sutherland. Coll. Boonstra.

Skull large, maximum length 700 mm. Snout long, high, moderately

wide. Intersquamosal width fairly great (450? mm.). Postfrontal boss fairly

large and prominent. Jugal boss low, angular boss very strong.

Fronto-naso-prefrontal swelling strong, but only slightly overhanging the

sides of the skull. Pineal boss low with rounded edge, situated near occipital

5—Annals

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

edge. Sharp upward inclination of premaxillary edge. The occiput is high,

fairly narrow and deeply concave with a fairly great posterior sweep of the

temporal arches and the upper part of the temporal fossa roomy

antero-posteriorly; the temporal arch rises above the plane of the intertemporal

surface. Palate probably long. Basis cranu unknown. Condyle unknown.

In the type there are in the left upper jaw 3 incisors, 1 canine and probably

7 postcanines; whereas on the right side there are 4 incisors, I canine and

? postcanines. In the dentary there are 2 incisors on both sides.

Fic. 9.—Anteosaurus vorsteri. Referred specimen. S.A.M. 11577,

Bulwater, Beaufort West. Dorsal view. Symmetry restored on

the basis of the well preserved left side. (x %-)

Paratype. (Fig. 6.) A good skull and parts of the lower jaw.

S.A.M. 4340. Leeurivier, Beaufort West. Coll. Haughton.

CRANIAL STRUCTURE OF THE TITANOSUCHIAN I31

I have included this specimen in the species — abelt — although it differs

from the type in a number of points viz. there is only a faint indication of

a jugal boss; the frontal enters the supra-orbital border; the premaxilla has

a longer alveolar border and on the left carries 5 incisors — the first 4 being

very long and strong whereas no. 5 is small and weak; on the left dentary

there are 4 long and strong incisors intermeshing with the upper incisors.

S.A.M. 9123 from Voélfontein, Prince Albert, consists of the major part

of a skull which agrees in all essentials with the type skull. Here there are

on the left premaxilla 5 cross-sections of incisor roots; these represent

numbers 1-4 with a replacing incisor lying median to no. 4. On the night

side cross-sections of nos. 2, 3 and 4 are preserved. On the left there is

the root of a large canine, whereas on the right there is the empty alveolus

of the canine.

S.A.M. 11576 (Fig. 5) from Klein-Koedoeskop, Beaufort West, consists

of the anterior part of both jaws. The upper canines are large strongly

curved teeth. On the left premaxilla there are 5 incisors — the first 4 long

and strong and the fifth short but strong, whereas on the right there are

only the 4 anterior incisors. In both dentaries there are 4 incisors — the

first 3 long and strong and the fourth much smaller.

S.A.M. 5621, Leeurivier, Beaufort West, consists of two pieces not in

contact, representing most of the dorsal surface of a skull, which closely

resembles that of the type skull.

S.A.M. .11949 (Fig. 7), Nuwefontein (Roxana), Fraserburg, consists of

a snout and the dorsal part of the posterior part of the skull not in contact.

This specimen is weathered beautifully white and in the snout the sutures

are very clearly shown and it was in this specimen where I first noticed that

the premaxillaries were posteriorly abruptly truncated and not long and

tapering as in Jonkerta and the Tapinocephalids.

Anteosaurus acutirosins Sp. Nov. (Fig. 10.)

Boonstra, L. D. 1953. Anteosaurus abeli (Boonstra) in errore.

Holotype. A good skull and lower jaw. S.A.M. 9320, Kruisvlei, Beaufort

West. Coll. Boonstra.

Skull large, maximum length 675 mm. Snout long, high, narrow and

light. Intersquamosal width great (480? mm.). Postfrontal boss huge

and prominent. Jugal boss massive, angular boss massive. Fronto-naso-

prefrontal swelling very strong with a distinct step on to the anterior nasal

surface and laterally slightly overhanging the sides of the skull. Pineal boss

132 -ANNALS OF THE SOUTH AFRICAN MUSEUM

low with rounded edges extending to the occipital edge. Fairly strong upward

inclination of the premaxillary edge. . The occiput is high, fairly wide, very

deeply concave with a sharp and great posterior sweep of the temporal

arches; upper part of the temporal fossa roomy antero-posteriorly; the temporal

arch rises above the plane of the intertemporal surface. Palate apparently

long and narrow. Basis cranu fairly long. Exoccipitals fused with

basioccipital to form a rounded condyle, which is visible beyond the occipital

edge in dorsal view. On both sides there are 4 upper and 4 lower incisors,

I canine and 5?. postcanines. In both dentaries there are 4 incisors.

In this specimen the quadratojugal is clearly seen as a small bone resting

‘on a ledge above the outer quadratic condyle as in the Pristerognathid

‘Therocephalians.

Fic. 10.—Anteosaurus acutirostris. Sp. Nov. Holotype.

S.A.M. 9329, Kruisvlei, Beaufort West. Dorsal view.

Symmetry restored on the basis of the right side, which

has suffered a_ slight side-to-side compression. (xX ¢.)

CRANIAL STRUCTURE OF THE TITANOSUCHIAN Biss

_. Associated with the holotype skull of acutirostris there were a number of

skull fragments of other Anteosaurus skulls and in addition a large number

of bones of the postcranial skeleton of more than one individual. I hope to

be able to describe these bones of the postcranial skeleton in the near future.

In the meantime a general statement here will be useful.

As a whole the postcranial skeleton is lightly built without any trace of

the massiveness so typical of all the hitherto known South African

Deinocephalians. One’s first impression is that the skeletal bones could

belong to some large Therocephalian — associated, for instance, with a skull

not very much larger than that of Scymnosaurus ferox. This applies

particularly to the femur, which is a long bone with the proximal and distal

ends hardly expanded and the shaft long and slender.

Anteosaurus crassifrons Sp. Nov. (Figs. 11, 12, 13-

Holotype. A good skull, but distorted by a simple shear. S.A.M. 11946,

Buffelsvlei, Beaufort West. Coll. Boonstra and Marais.

Skull large, but short and quat; maximum length 570 mm. Snout short,

high and very wide. Intersquamosal width fairly small (330? mm.).

Postfrontal boss fairly massive and prominent. Jugal boss massive, angular

boss unknown. Fronto-naso-prefrontal swelling very massive with a very

distinct step on to the anterior nasal surface and laterally strongly overhanging

the sides of the skull. Pineal boss rounded, large and extending on to the

frontal. Very sharp upward inclination of the premaxillary edge. The

occiput is high, fairly wide, deeply concave, with a great posterior sweep of

Fic. 11.—Anteosaurus crassifrons. Sp. Nov. Holotype.

_S.A.M. 11946, Buffelsvlei, Beaufort West. Lateral view.

CxS)

134 ANNALS OF THE SOUTH AFRICAN MUSEUM

the temporal arches, and the upper part of the temporal fossa roomy in

antero-posterior direction; the temporal arch not rising above the plane of

the intertemporal surface. Palate very short with very massive lateral

pterygoidal flanges. Basis craniw short, with short basisphenoid. Exoccipitals

fused with basioccipital to form a rounded condyle. In both premaxillaries

no teeth are preserved but 5 matrix-filled alveoli are shown; in both maxillaries

5 postcanine roots of greatly varying diameter are preserved, numbers 2 and

5 being much smaller than the other 3.

S.A.M. 11302 from Buffelsvlei, Beaufort West, is a fairly complete but

weathered skull. Although a somewhat larger skull than that of the type, it

clearly belongs to the same species. It has the same massive

fronto-naso-prefrontal swelling with a distinct step anteriorly and laterally

Fic. 12.—Anteosaurus crassifrons. Sp. Nov.

Holotype. S.A.M. 11946, Buffelsvlei, Beaufort

West. Dorsal View. Symmetry affected by a

simple shear, restored on the basis of the least

affected left side. (x 3.)

overhanging the preorbital side-wall; the jugal boss is very massive. On the

right side there are stumps of 4 incisors and five or six postcanines; a well

CRANIAL STRUCTURE OF THE TITANOSUCHIAN 135

preserved crown of a postcanine is of fair size (25 x 14 x 8 mm.); it is a bluntly

conical tooth, labio-lingually compressed.

S.A.M. 11929 from an unknown locality, probably near Abrahamskraal,

Prince Albert, is an imperfect, weathered skull in extremely intractable

matrix, but showing a fairly good ventral surface. The quadrate shows its

articular condyles to lie obliquely in the skull with the outer one the more

posterior one as shown in reconstruction in all the figures in this paper, and

Fic. 13.—Anteosaurus crassifrons. Sp. Nov.

Holotype. S.A.M. 11946, Buffelsvlei, Beaufort

West. Ventral view. Symmetry restored on the

basis of the left half of the palate with the result

that it is narrower than it would have been in life.

On the right the quadrate-complex is indicated in

position. (x %.)

thus not as shown in Broom’s figure of Anteosaurus vorsteri. Part of the left

stapes can also be seen and this bone appears to be stout, firmly wedged in

the fenestra ovalis and abutting against the medial edge of the quadrate dorsal

to the extremity of the quadrate ramus of the pterygoid, which appears to

overlap the postero-median surface of the quadrate as in Jomkeria and the

Tapinocephalids. The basis cranii is short and wide, with a_ short

basisphenoid. The roots of five postcanines are preserved in the right maxilla.

136 ANNALS OF THE SOUTH AFRICAN MUSEUM

Anteosaurus major Sp. Nov. (Figs. 14, 15)

Boonstra, L. D. Anteosaurus abeli (Boonstra) in errore.

Holotype. A good skull, lacking the snout, but somewhat dorso-venirally

compressed. S.A.M. 11293. Boesmansrivier, Beaufort. West. Coll.

Boonstra.

Skull very large, maximum length 805? mm. Snout long, fairly broad

and high. Intersquamosal width very great (612 mm.). Postfrontal boss

Fic. 14.—Anteosaurus major. Sp. Nov. Holotype. S.A.M. 11293,

Boesmansrivier, Beaufort West. Dorsal view. Symmetry disturbed by

dorso-ventral crushing restored on the basis of the left side. ( x $.)

CRANIAL STRUCTURE OF THE TITANOSUCHIAN 137

only moderately strong and not very prominent, with postorbital forming the

postero-lateral part. Low and weak jugal boss, angular boss unknown.

Fronto-naso-prefrontal boss moderate, confluent with anterior nasal surface.

Pineal boss low, with rounded edges, situated some distance from the occipital

edge. Premaxilla unknown. The occiput is low and broad, moderately

deeply concave, sweep of temporal arches more laterally than posteriorly and

the upper part of the temporal fossa roomy antero-posteriorly; the temporal

arch rising above the level of the intertemporal surface. Palate fairly long

Fic. 15.—Anteosaurus major. Sp. Nov. Holotype. S.A.M. 11293,

Boesmansrivier, Beaufort West. Ventral view. Symmetry restored on

the basis of the left side, which is least disturbed. (x +.)

138 ANNALS OF THE SOUTH AFRICAN MUSEUM

and broad, with strong and deep lateral pterygoidal flanges but not so massive

as in crassifrons. Basis cranu long and basisphenoid much longer than in

any of the other species. Exoccipitals forming the dorso-lateral corners of

the condyle. No teeth are preserved.

Anteosaurus laticeps Sp. Nov. (Figs. 4, 16)

Holotype. An incomplete skull showing only the ventral and most of the

occipital surface. S.A.M. 11592. Dikbome, Laingsburg. Coll.

Boonstra and Du Plessis.

Skull large, maximum length 645? mm. Snout short and very broad.

Intersquamosal width great (522 mm.). Postfrontal boss unknown. Jugal and

angular bosses unknown. Fronto-naso-prefrontal region unknown. Pineal

region unknown. Premaxilla unknown. The occiput is low and very broad,

Fic. 16.—Anteosaurus laticeps. Sp... Nov. Holotype.

S.A.M. 11592, Dikbome, Laingsburg. Ventral view based

mainly on the right side. (x }-)

CRANIAL STRUCTURE OF THE TITANOSUCHIAN I39

shallowly concave; sweep ot temporal arches mostly laterally. Palate long

and very broad with only moderateiy strong lateral pterygoidal flanges.

Basis cranu short, with very short basisphenoid. Exoccipitals forming much

of the dorso-lateral corners of the condyle. No incisors are preserved, the

canines are very strong, slightly recurved teeth; in the mght maxilla stumps

of five postcanines can be made out, whereas on the left five teeth are

preserved with the probability of another two making a total of 7. The

postcanines are bluntly conical, but linguo-labially compressed teeth with the

crowns thus oval in crosssection (16 xX II xX 7 mm.).

Antéosaurus cruentus Sp. Nov. (Figs. 17, 18, 19, 20)

Boonstra, L. D. 1953. Anteosaurus minor (Broom) in errore.

Holotype. A good, undistorted skull, but lacking the lower jaw.

S.A.M. 11694, Koringplaas, Moordenaarskaroo, Laingsburg. Coll.

Boonstra and Du Plessis.

Skull moderately large, maximum length 565 mm. Snout long, narrow

and fairly low. Intersquamosal width not great (360 mm.). Postfrontal boss

Fic. 17.—Anteosaurus cruentus. Sp. Nov. Holotype. S.A.M. 11694,

Koringplaas, Laingsburg. Lateral view with the quadrate-complex

missing. (> $¢.)

fairly strong and prominent. No jugal boss, angular boss unknown.

Fronto-naso-prefrontal swelling small, passing evenly on to the anterior nasal

surface. Pineal boss prominent, with sharp circular border, situated very

near the occipital edge. Sharp upward inclination of the premaxillary edge.

The occiput is high and fairly broad, very deeply concave with a great

posterior sweep of the temporal arch and the upper part of the temporal

fossa roomy antereo-posteriorly; the temporal arch not rising above the very

I40 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 18.—Anteosaurus cruentus. S. Nov. Fic. 19.—Anteosaurus cruentus. Sp. Nov.

Holotype. S.A.M. 11694, Koringplaas, Holotype. S.A.M. 11694, Koringplaas,

Laingsburg. Dorsal view. (x %.-) Laingsburg. Ventral view. (x ¢-)

narrow intertemporal surface.

Palate long, with fairly robust

lateral pterygoidal flanges. Basis

cranu long, but the basisphenoid

is short. The exoccipitals form-

ing a large part of the dorso-

lateral corners of the condyle.

No trace of incisors is preserved;

the alveolar face of the pre-

maxillaries shows a matrix filled

groove divided in its posterior

part into distinct adveoli; there

Po.F.

i Fic. 20.—Anteosaurus cruentus. Sp. Nov.

appears to be room for 5 incisors Holotype. S.A.M. 11694, Koringplaas,

when developed. On the right Taingshurg. Occipital var" aaanae:

the canine root is followed by roots of 6 postcanines, but on the left only

four roots with a possible fifth can be seen. |

CRANIAL STRUCTURE OF THE TITANOSUCHIAN I4I

S.A.M. 9140 from Voélfontein, Prince Albert, is an imperfect disarticulated

skull agreeing fairly well with the type, but is of some interest in that it

shows a number of the roofbones as separate elements with exposed sutural

faces.

Anteosaurus levops Sp. Nov. (Fig. 21)

Holotype. A weathered skull, without the lower jaw. S.A.M. 11492.

Mynhardtskraal, Beaufort West. Coll. Boonstra.

Skull fairly small, maximum length 485 mm. Snout fairly short, lightly

built, narrow and low. Intersquamosal width relatively large (415 mm.).

Postfrontal boss strong and prominent. No jugal boss; angular boss unknown.

Fronto-naso-prefrontal swelling strong with a distinct step onto the anterior

nasal surface and laterally slightly overhanging the sides of the skull. Pineal

boss apparently prominent, reaching the occipital edge. Upward inclination

Fic. 21.—Anteosaurus levops. Sp. Nov. Holotype.

S.A.M. 11492, Mynhardtskraal, Beaufort West.

Dorsal view. Temporal fossae not cleared of matrix.

| (X &-)

of the premaxillary edge moderate. The occiput is fairly low and broad;

deeply concave, not vertical; strong postero-lateral sweep of the temporal

‘arches and the upper part of the temporal fossa shortened in antero-posterior

direction; the temporal arch rising above the plane of the narrow intertemporal

142 ANNALS OF THE SOUTH AFRICAN MUSEUM

surface. Palate long and narrow. JBasis cranu fairly short. Condyle

unknown. In the right premaxilla parts of the crowns of 3 incisors are

preserved, but on the left there is a matrix filled groove with no sign of

any teeth.

Anteosaurus minusculus Sp. Nov.

Boonstra, L. D. 1936. Anteosaurus minor (Broom) in errore.

Holotype. A distorted skull with the greater part of the arches and most

of the palate missing.

American Museum of Natural History, No. 2224. Vanderbylskraal?,

Beaufort West. Coll. Broom.

This specimen in the American Museum is stated by Broom to be the

topotype of the skull fragment in the British Museum (Natural History)

R.5742.

In my paper on the Titanosuchids in the American Museum I attempted

a reconstruction of the dorsal aspect of the skull and showed the premaxilla

as a long posteriorly tapering bone. With our present knowledge of the

truncated posterior end of the premaxilla in all species of Anteousaurus this

was obviously an error in observation. As this skull has well developed

postfrontal bosses it cannot beiong to Pseudanteosaurus minor (Broom) and

I propose to regard it as a new species of Anteosaurus under the specific

name — minusculus sp. nov.

The skull is fairly small with a maximum length of 480 mm.; the snout

is long, broad and fairly high; the intersquamosal width was probably small

(225? mm.); the postfrontal boss quite strong and prominent; there is no

jugal boss; the fronto-naso-prefrontal swelling is weak; the occiput is vertical,

very deeply concave with a great posterior sweep of the temporal arches

and the upper part of the temporal fossa is roomy in antero-posterior direction.

Anteosaurus sp. (Fig. 22)

S.A.M. 2752, Viviers Siding, Beaufort West. Coll. Hanghton and Whaits.

In this specimen we have only the posterior two thirds of the upper surface

of a skull. Although it cannot be included in any of the above described

species I am not naming it. The skull differs from all the described forms

in that the prominent mound-like pineal boss overhangs the occipital surface;

the occiput is very deeply concave from side to side with the two horns of

the parietal directed nearly wholly in posterior direction; in the posterior

CRANIAL STRUCTURE OF THE TITANOSUCHIAN 143

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Siding, Beaufort West. Dorsal View. (x #4.)

part of the frontals there is a step bringing this part of the surface down to a

lower level than that of the anterior part of the frontals. The postorbitals

form a small part of the posterior surface of the boss, whose surface is in

other species wholly formed by the postfrontal. These bosses are only

moderately strong and prominent. There is very little swelling in the

fronto-naso-prefrontal region, but notwithstanding this the prefrontal is so

developed to exclude the frontal from entering the supraorbital border.

DISCUSSION

From the above account it is clear that Anteosaurus is a genus of the

Deinocephalia quite distinct from all the known South African genera. It

may be characterised as follows: skull large, with prominent postfrontal

bosses; temporal fossa with large dorso-ventral diameter and deep

antero-ventral bay; infratemporal bar narrow; quadratojugal no longer lying

ANNALS OF THE SOUTH AFRICAN MUSEUM

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CRANIAL STRUCTURE OF THE TITANOSUCHIAN I45

on the surface on the latero-postero-ventral corner of the skull but shifted

medially as in the higher Therapsids; alveolar border of the premaxilla inclined

upwards to give working space to the long, simple, pointed incisors; a short

series of postcanine teeth, rather small and irregular; deeply concave occiput;

ridge forming the outer border of the occiput proper formed solely by the

tabular and squamosal; postorbital covering the lateral face of the parietal

and with a long contact with the squamosal; fronto-naso-prefrontal region

swollen to a greater or lesser extent; with bosses on the jugal and angular

in the larger species; number of incisors variable; great posterior overlap

of the maxilla over the jugal.

If the Titanosachia are defined as Deinocephalians with a carnivorous

dentition, with quadrate not displaced far anteriorly, with concave occiput,

with spacious temporal fossa, then Anteosaurus is undoubtedly a

Titanosuchian.

The only other S.A. Titanosuchian genus in which the skull is adequately

known is Jonkena.

Anteosaurus differs from Jonkeria in a number of important points: in

Anteosaurus the quadratojugal is no longer a surface bone, but has shifted

medially (in one species of Jonkeria viz. vanderbyli, there is an indication

that here also there is a similar tendency); in Anteosaurus the infratemporal

bar is narrow (again J. vanderbyli shows a similar tendency); in Anteosaurus

the temporal fossa extends far ventrally and has in addition an anterior bay

of the fossa extending to under the postorbital bar; the postfrontal develops

a large boss in Anteosaurus, whereas this bone is a small element in Jonkeria;

large jugal and angular bosses are unknown in Jonkerna and _ the

fronto-naso-prefrontal region remains unswollen; the pineal foramen is

situated near the occipital edge in Anteosaurus, whereas in Jonkeria it is

situated in the plane of the postorbital bar; in Jonkeria the premaxillary edge

does not curve sharply upwards; no reniform palatine boss bearing teeth is

known in Jonkeria; the lateral pterygoidal flanges are much weaker in

Jonkena; in Jonkerna the posterior sheet of the postorbital does not cover

the whole lateral face of the parietal; in Jonkeria the jugal is a much smaller

bone and does not flare out laterally as it does so characteristically in

Anteosaurus, nor does it extend so far posteriorly lying along the inner face

of the squamosal; in Jonkeria the postcanines form a long series, whereas

this is short in Anteosaurus; the posterior process of the premaxilla in

Jonkeria, as in the Tapinocephalians, is much longer than in Anteosaurus,

in Jonkeria the ridge on the occiput is formed by the paroccipital, squamosal

and tabular, whereas in Anteosaurus the paroccipital does not enter into it

at all; in Anteosaurus the quadrate rami of the pterygoid curve much outwards

as they approach the quadrate, whereas in Jonkeria they lie nearly parallel

to the median line; in Anteosaurus the stapes is a much longer bone; in

6—Annals

146 ANNALS OF THE SOUTH AFRICAN MUSEUM

Anteosaurus the maxilla has a much greater overlap over the jugal. These

differences, together with others not listed here, to my mind show that these

two genera lie on lines of development sufficiently divergent to warrant our

placing them in different families, which I propose to name the Anteosauridae

and the Jonkeridae.

It is thus clear that the Anteosauridae, although retaining a number of

primitive characters, have advanced farther in some points of structure than

their contemporaries the Jonkeridae.

COMPARISON WITH OTHER THERAPSIDS

In South Africa the Therapsids are first encountered in_ the

Tapinocephalus-zone. With its complex of monoclinal folds it has up to the

present not been possible to establish from what level within the zone the

various known Therapsid finds have come. Until then we are forced to

consider the assemblage of forms from this zone as being contemporaneous.

The thus contemporary Therapsids from the Tapinocephalus-zone are:

Anningiamorpha, Dromasauria, Anomodontia, Gorgonopsia, Therocephalia

and Deinocephalia. These are all present as well established groups clearly

distinct from each other.

The Anningiamorphs are not very well known, but appear to be a group

in which a large number of primitive characters have persisted.

The Dromasaurians are only known from four specimens and combine

a number of primitive characters with some rather specialised.

The Anomodonts, although but poorly represented in this zone compared

to the great diversity developed in the younger rocks of the Karroo, are

already quite specialised when they are first encountered. No group with

such a remarkable edentulous premaxilla can be anything but firmly set on

an independent line of development.

The Gorgonopsians, with but a few forms present in the Tapinocephalus-

zone, blossomed exceedingly in later ages, but the oldest known species are

already definite Gorgonopsians with a habitus well established and clearly

their own and the subsequent developments in no way exceeded the limitations

inherent in these early forms from the Tapimocephalus-zone.

The Therocephalians of the Tapinocephalus-zone are a virile suborder

of the Therapsids already represented by a large number of species, which,

representing different lines of development, can be placed in a number of

different families. Having already, by the beginning of the Tapinocephalus-

zone times, split into a number of families it is clear that the tempo of

development within the suborder during the antecedent ages must have been

greater than in the Gorgonopsians. This was probably due to a greater

lability in the original stock. As their successors — not only the higher

Therocephalian families but also their off-spring the Cynodonts, Bauriamorphs

CRANIAL STRUCTURE OF THE TITANOSUCHIAN 147

and Ictidosaurians, of the later Karroo Beds — show, this lability or greater

potentiality for further development beyond the confines of the suborder

was maintained.

The Deinocephalians, when first encountered in the Tapinocephaius-zone,

were already at the end of their tether. They are represented by the

end-products of their particular line of development which culminated in the

3 specialised groups — Titanosuchia, Tapinocephalia and Styracocephalia.

CHARACTERS OF A POSTULATED PRIMITIVE THERAPSID ANCESTOR

If we postulate a primitive Therapsid ancestor, common to all the

Therapsids, situated on a morphological level somewhere between that of

the early Pelycosaurs and the first Therapsids we would expect it to have

the following characters: intertemporal region broad and flat; preparietal

absent; premaxilla without long posterior process intercalated between the

nasals; the postfrontal well developed; the postorbital covering the lateral

face of the parietal and meeting the squamosal; the pineal foramen well in

advance of the occipital edge i.e. the upper occipital edge has not yet migrated

anteriorly; the premaxilla below the nostril would still be shallow; the

zygomatic arch shallow; the temporal fossa relatively small with the

squamosal not bowing out laterally or posteriorly; the quadratojugal still

lying on the outer surface i.e. not yet migrated internal to the squamosal; the

occiput low, fairly narrow, vertical, with its upper border not moved

anteriorly, not deeply concave from side to side; the paroccipital just

beginning to strengthen and to support the quadrate; the quadrate would still

be large with its condyle lying fairly far ventrally, but still in the posterior

position i.e. in line with the occipital condyle; the premaxilla would be

dentigerous and with hardly any palatal face; the maxillary teeth would form

a long series and there would probably already be a specialised canine;

choanae and nares anteriorly situated; no suborbital opening; the lateral

pterygoidal processes would still be situated fairly far back and be fairly weak;

the basisphenoid-pterygoid joint would no longer be movable and the contact of

the basisphenoid with the pterygoid shifted posteriorly; the basisphenoid

processes flattening; the quadrate ramus of the pterygoid reduced in height; the

parasphenoidal rostrum would still be visible in the posterior part of the

interpterygoidal slit; there would be no coronoid process to the dentary; the

lower jaw long and the angular notch and reflected lamina would be beginning

to develop.

Tabulating the characters of the Therapsids of the Tapinocephalus-zone

on the basis of the above list one gets the following result:

The Anningiamorpha are not sufficiently well known to make a count of

the characters corresponding with those in the above list, but those preserved

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

point to this group as approaching the postulated primitive Therapsid condition

most closely. |

The Dromasauria, as far as they are known, appear to occupy the

second place.

Then come the three groups of the Deinocephalia — which, although

clearly distinct from each other — due to particular specialisation — seem

to stand on more or less the same morphological level.

Somewhat further removed from the postulated primitive Therapsid

condition are the Dicynodonts, and then come the Gorgonopsians, followed

by the Therocephalians in which the rate of development away from the

primitive Therapsid condition has been the greatest.

Finally, Anteosaurus has advanced beyond the primitive Therapsid stage

in the following characters: the premaxilla, below the nostril has increased

in depth and then the alveolar edge has curved upwards; the temporal fossa

has become very roomy and the squamosal is bowed out strongly both

laterally and posteriorly; the quadratojugal has migrated from the outer

surface to lie medial of the zygoma; the occiput has greatly increased its surface

by becoming both broad and high; the paroccipital process has become very

strong and supports the quadrate firmly; the length of the postcanine series

has decreased and the long incisors have become intermeshing teeth; the

lateral pterygoidal processes are situated well forward and are very strongly

developed; the basisphenoid-pterygoid region has advanced as in higher

Therapsids, but there are no strong basisphenoidal tubera and lastly we have

the characteristic pachyostosis with the development of the peculiar

postfrontal, jugal and angular besses.

REFERENCES

Boonstra, L. D. 1936. The Cranial Morphology of Some Titanosuchid Deino-

cephalians. Bull. Am. Mus. Nat. Hist. 72, 3, 99-116.

Boonstra, L. D. 1948. Miljoene Jare Gelede in die Karoo. Voortrekkerpers,

Johannesburg.

Boonstra, L. D. 1952. ’n Nuwe TJitanosuchiérsoort (Anteosaurus abelt). Tydskrif

vir Wetenskap en Kumns. 12, 1, 150-151.

Boonstra, L. D. 1953. A Suggested Clarification of the Taxonomic Status of the

South Atrican’ Titanosuchians,, Anny S) Air Mus, 42)) a) meres:

Broiri, F. AND SCHRGODER, J. 1935. Ein Dinocephalen-Rest aus den wunteren

Beaufort-Schichten. Sitzb. Bay. Akad. Wiss. Sonderdruck. 93-114.

Broom, R. 1910. Observations on some Specimens of South African Fossil Reptiles

preserved in the British Museum. Trans. Roy. Soc. S. Afr. 2, 1, 19-25.

Broom, R. 1929. On the Carnivorous Mammal-like Reptiles of the Family

Titanosuchidae. Ann. Trans. Mus. 13, 1, 9-36.

Broom, R. 1936. On some New Genera and Species of Karroo Fossil Reptiles, with

Notes on some others. Ann. Trans. Mus. 18, 4, 349-386.

Broom, R. 1936. On the Structure of the Skull in a New Type of Deinocephalian

Reptile. Proc. Zool. Soc. 733-742.

Watson, D. M. S. 1914. The Deinocephalia, an Order of Mammal-like Reptiles

Proc. Zool. Soc. 749-786. ,

Watson, D. M.S. 1921. The Basis of Classification of the Theriodontia. Proc. Zool.

Soc. 35-98.

9g. The Smallest Titanosuchid yet recovered from the Karroo. By oe

Boonstra, D.Sc.

(With Plate XVIII and 5 text-figures)

In the collection of the South African Museum there is a specimen

(S.A.M. 4323) collected by Haughton on the Merweville Commonage in 1917.

This had been entered in the register as a Gorgonopsian, presumably because

of its small size. The specimen as preserved consists of the anterior third

of a small skull, the major part of a manus, a tarsus, a nearly complete

femur, a radius, a fibula, part of the head of the humerus, a coracoid, a

series of caudal vertebrae and some other fragments. This is the first specimen

of a South African Deinocephalian in which most of the bones of the fore-

and hindfoot have been found in articulation.

THE SKULL (fig. 1)

In the accompanying figure the lateral aspect of the snout is given,

with the missing part of the skull indicated by broken lines. The snout is

very similar in general build to that known in the large Anteosaurus, although

in size it is less than 4 of Anteosaurus abel. The nostril is not terminal; the

alveolar border, anterior to the canine, sweeps sharply upwards; this reduction

of the premaxilla creates the space necessary for the large anteriorly directed

anterior incisors, whose function has become that of snatching, piercing and

tearing teeth (cf. mechanical grab); the five upper incisors, increasing rapidly

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Fic. 1.—Micranteosaurus parvus Gen. et Sp. Nov. Lateral

view of anterior third of the skull with the missing posterior

two thirds in broken lines based on the structure of

Anteosaurus. S.A.M. 4323. Commonage, Merweville,

Beaufort West District. (x 4.)

149

150 ANNALS OF THE SOUTH AFRICAN MUSEUM

in size from number 5 to number 1, intermesh with the four incisors of the

dentary; the lower incisors also increase rapidly in size from number 4 to

number 1, but they are directed upwards and very much less anteriorly than

are the upper incisors; the posterior border of the incisors forms a sharp

cutting edge with fairly fine serrations; the upper incisors occupy 55 mm. on

the left and 57 mm. on the right side (measured over the curve).

Between the last incisor and the canine there is a diastema of 5 mm. on

the left and 4 mm. on the right side. The canine is a strong curved tooth

with a length of 35 mm. and at the base of the crown the antero-posterior

diameter is I4 mm.; its posterior edge, though sharp, is not serrated.

Between the canine and the first postcanine there is a diastema of 27 mm.

on the left and 23 mm. on the right side; on the right the first and second

postcanines are in part preserved, whereas on the left the first, third and

fourth are in part preserved; the postcanines are small stubby teeth apparently

irregularly spaced and functionally unimportant.

The dentary is strong with the alveolar border housing the incisors bent

downwards to increase the space necessary for the very long anterior incisors;

the symphysis is strong; the mentum squarish with ventrally a “‘gonial’’ or

“‘digastric’’ tubercle.

The premaxilla extends some distance posteriorly between the nasals.

The limits of the septomaxilla are not very clear, but the bone is small and

apparently forms the ventral border of the nostril as shown in the figure.

The maxilla is not swollen above the canine so that it would appear that

the root of the canine is not strong. Anteriorly the maxilla has a large overlap

over the premaxilla.

Femur (fig. 2a)

Except for its distal end the femur is fairly well preserved. It is a long

slender bone with both its proximal and distal end unexpanded and the shaft

long and slender. A twist in the shaft places the distal end at night angles

to the proximal end. The proximal facet is directed appreciably anteriorly.

No external trochanter is differentiated on the posterior edge of the bone

and the area for the insertion of the ilio-femoralis is narrow. Anteriorly a

ridge separates this area from the area for the insertion of the

pubo-ischio-femoralis internus. The posterior condyle has no epicondylar

widening and the facet for the fibula is terminal. The intercondylar fossa is

shallow and so is the popliteal fossa. The intertrochanteric fossa is shallow,

and with no clearly differentiated external and internal trochanters developed,

it is not clearly demarcated either anteriorly or posteriorly. There is thus little

left of the primitive Y midges.

It is thus evident that this femur differs greatly from that hitherto described

in any South African Deinocephalian where the bone is usually a short stout

SMALLEST TITANOSUCHID YET RECOVERED FROM KARROO I5I

element with short wide shaft and greatly expanded distal and proximal ends.

Although superficially resembling the femur of Therocephalians and

Gorgonopsians, it differs strikingly in the absence of a differentiated external

trochanter and in the rotation of the ends on the shaft.

a b d

Fic. 2.—Micranteosaurus parvus Gen. et

Sp. Nov. S.A.M. 4323, Commonage,

Merweville, Beaufort West District.

a, Worsal view of left femur. (x 4.) 0,

Ventral view of fibula. (x 4.) c, Dorsa

view of left coracoid. (x 4.) 4d, Dorsal

view of radius. (x #.)

Fiputa (fig. 20)

The fibula is a lightly built long and slender bone with its proximal end

strongly expanded and with its articulating facet for the femur terminal.

Whereas the proximal end is flattened, the distal facet is broadly oval in

outline.

Pes. (plate XVIIIa and fig. 3)

When this specimen came under my notice it had already been partly

cleared of matrix and the parts glued together. Before preparing it further

I embedded the whole in plaster. In the accompanying plate I give a

photograph of the pes after preparation and still in the plaster bed. From

this it is evident that the two proximal tarsals joined by matrix were rotated

as a unit through 180° when joined by glue to the distal part of the pes in

152 ANNALS OF THE SOUTH AFRICAN MUSEUM

the original preparation. In Fig. 3 the proximal tarsal elements are shown

right side up. |

The intermedium is an ovoid bone with its outer border concave and facing

a similar concavity of the fibulare, thus creating a passage for an artery.

Fic. 3.—Micranteosaurus parvus

Gen.,.,et) Spx j Nowy 1;S A. Meat 4g2ee

Commonage, Merweville, Beaufort

West District. Semi-diagrammatic

restoration of the right pes seen in

dorsal view > about 2?. C—

centrale; F—fibulare; I—inter-

medium.

The fibulare is a dorso-ventrally flattened bone, but thickened both |

proximally and distally to form articulating facets for the fibula and centrale

and 4th distale respectively.

The single centrale is a small pebble-like bone. |

Four distalia are preserved, but a fifth must also have been present. |

Five metacarpals are preserved as shown in the illustrations; they are |

dorso-ventrally flattened bones, constricted in the waist and with their distal _

ends more expanded than the proximal ends. No. 1 differs but little from

No. 2, but the 4th and especially the 5th are much reduced.

Phalanges. In the first two digits no phalanges are preserved, and in the

3rd and. 4th only the proximal end of the first phalanges are present; in the |

little toe there is a fairly long phalanx and an ungual phalanx preserved.

SMALLEST TITANOSUCHID YET RECOVERED FROM KARROO 153

The tarsal formula is thus 2, I, 5, and the phalangeal formula

Bae 3°, 4? 2

The pes is thus still closely related to that of the Pelycosaurians.

CoRACOID (fig. 2c)

The left coracoid is preserved. It is roughly circular in outline; thickened

laterally where it carries an articular facet to form the lower part of the

glenoid articulation; medially it forms a fairly thin sheet of bone with a

concave upper and a convex under surface. Anteriorly it has a free edge and

is not suturally united to the procoracoid, as is the case in most Therapsids

including the Deinocephalians such as Jonkeria and Moschops. This free

coracoid is, hower, encountered in a number of Deinocephalians, e.g.

Stvuthiocephalus, Tapinocephalus, Pelosuchus, etc.

Rapius (fig. 2d)

The, radius is much shorter than the fibula. It is a fairly slender bone

somewhat flattened dorso-ventrally; its proximal end is expanded to about

twice the width of the distal end, but the shaft has no waist-like constriction.

Manus (plate XVIIIb and fig. 4)

When the manus came to my notice it had also been partially prepared,

but all the constituent bones were still joined to each other by matrix. I also

embedded the whole in a block of plaster before continuing the preparation.

As is evident from the photograph, the first digit lies extended, the second

and third folded inwards, in the fourth digit the phalanges have been displaced

and the fifth lies extended. In the proximal row of the tarsus the intermedium

and ulnare have been displaced medially (anteriorly). .

In Fig. 4 I give a restoration of the dorsal aspect of the manus. In the

proximal row of the carpus there are a radiale, intermedium and ulnare. The

radiale is roughly rectangular in outline; dorso-ventrally compressed; it is

thickened distally and proximally to form facets for the distals (1 & 2) and

radius respectively, with both the dorsal and ventral surfaces’ concave

antero-posteriorly. The intermedium is a fairly small thin flat bone.

The ulnare is the longest bone of the proximal row. It is a stout bone,

proximally thick and knob-like, with a large convex proximal facet for the

ulna; anteriorly it is weaker, with a flat distal articular facet for the distals

(4 and 5); just behind the distal face the bone has a constricted waist.

The centrale is not preserved, but was probably a fair sized bone lying

anterior to the intermedium and between the distal ends of the radiale and

ulnare, articulating distally with two distals (2 and 3).

- There are 5 distal carpals. The first is a small pebble-like bone; the

second is the largest of the distals, its dorsal surface is excavated with ridges

154 ANNALS OF THE SOUTH AFRICAN MUSEUM

on the preaxial, distal and proximal edges; the third also has an excavated

dorsal surface; the fourth and fifth are pebble-like.

Fic. 4.—Micranteosaurus parvus

Gens eto Sp Nov: /SiAGME ©4323;

Commonage, Merweville, Beaufort -

West District. Semi-diagrammatic

restoration of right manus seen in

dorsal view x plus-minus 4. C.—

centrale; I—intermedium; Re—

tadiale; Ue—ulnare.

As a whole the carpus is very Pelycosaur-like, approaching that of

Ophiacodon fairly closely, except that there is only one centrale.

The metacarpals are somewhat dorso-ventrally flattened bones in general

rod-like with slightly constricted waists; the decrease in size from the first

to the fourth is fairly evenly graded, but the fifth is much reduced; the first

metacarpal is distinguished by being much broader than the others and thus

relatively more flattened with expanded ends.

In the first digit the first phalanx is broad proximally, with a waist situated

in the anterior half; the distal end is much narrower than the proximal; the

second phalanx is much smaller, also broader proximally than distally, with

the waist nearly in the middle; anterior to the second roughly hourglass-shaped

phalanx there is preserved the proximal end of the ungual phalanx, so that

in the first digit there are 3 segments. This is most unusual, but there is no

doubt that such is the case. Even without the preserved proximal end of

the ungual phalanx the count would also be 3, for the second phalanx with

its hourglass-shape could not possibly be an ungual phalanx.

SMALLEST TITANOSUCHID YET RECOVERED FROM KARROO 155

In the second digit there are also 3 segments with each of the constituent

phalanges closely resembling those of the first digit.

The third digit has four segments.

The phalanges of the fourth digit have been displaced and all but one lost.

What I believe to be the second phalanx lies above the displaced fourth

metacarpal. The fourth digit probably had 4 segments as is the case in

the third digit.

The fifth digit is much reduced with a short hourglass-shaped first phalanx

and a fairly high, narrow and long ungual phalanx.

The carpal formula is thus 3, 1, 5, and the phalangeal formula

3, 3, 4, 4?, 2, and in structure thus near that of the Pelycosaurs.

A MoscHopip CarpPus (fig. 5)

For comparison I am including here a figure of a carpus, which, on the

humerus, I have identified as a Moschopid fairly near Moschops. This

specimen, S.A.M. 9157, collected by me at Wolwefontein, Prince Albert,

consists of a good humerus, radius and ulna, with

elements of the carpus still joined by matrix to the

distal end of the epipodial in natural articulation.

In the proximal row there are four bones, viz.

radiale, intermedium, ulnare and pisiforme. Only

the proximal part of the radiale is preserved, but

it would appear that the complete bone was a strong

rounded element. The intermedium is_ nearly

circular in outline, with its dorso-ventra] diameter

about equal to the antero-posterior diameter, and it

is thus not a thin plate-like bone. The ulnare is a

large bone, subcircular in outline; a longitudinal

ridge divides its dorsal surface into two faces, of

which the postaxial one is the larger; it is a much

thinner bone than the radiale and intermedium, and

its under surface is deeply concave.

Postaxially there lies a pebble-like pisiforme.

In between the ulnare and the radiale there was

Fic.

5-—An_ undeter-

mined Moschopid.

S.A.M. 9157, Wolwe-

fontein, Prince Albert

District. Part of right

fore-imb x 2. C—

centrale; I—inter-

medium; P—pisiforme;

R—radius; Re—

radiale; U—ulna; Ue—

ulnare.

a fair sized centrale; the actual bone has been lost,

but an impression in the matrix on the preaxial

surface of the ulnare indicates its position. None of

the five distals is preserved.

This Moschopid carpus is thus structurally close

to that of the Pelycosaurs, but the shape of the

individual bones is quite different, whereas in Micranteosaurus the individual

bones very closely resemble those of the Pelycosaurs in shape.

156 ANNALS OF THE SOUTH AFRICAN MUSEUM

TAXONOMIC

Although we know nothing about the posterior two thirds of the skull, the

snout is sufficiently characteristic to enable us to establish its affinities. The

dorsally curving alveolar border of the premaxilla, the long simple

intermeshing incisors, the short postcanine series; the fairly narrow, high

snout, and the position of the nostril show that this small Titanosuchid is

structurally nearly akin to the large Anteosaurus. Because of this great

difference in size I propose that it be considered a new genus, under the

name Micranteosaurus parvus Gen. et Sp. Nov. With Anteosaurus it is to be

included in the Titanosuchid family Anteosauridae.

RET LARLY Sie AD A EB) ORE We. See Rone eee WM TIAY ep

Pate XVIII. Micranteosaurus parvus Gen. et Sp. Nov. S.A.M. 4323, Commonage,

Merweville, Beaufort West District. a, Pes. The proximal segment shows the ventral

surface of the intermedium and fibulare as these two bones have been turned

upside-down in the original preparation. This error is corrected in the restoration

(Fig. 3) x about 3 nat. size. b, Manus in dorsal view x about */ Daw.~SiZe.

Ann. S. Afr. Mus., Vol. XLII Plate xvi

a SY

10. Paranteosaurus, Gen. Nov.: A Titanosuchian Reptile. By L. D.

Boonstra, D.Sc.

(W:th 2 text-figures)

In 1940 I collected parts of a skull, a proximal end of a femur and a

vertebra of a Jitanosuchian on the farm Mynhardtskraal, Beaufort West

(S.A.M. 11485). The cranial material consists of a weathered pre-orbital part

and the dorsal cranial roof without actual contact. The snout on development

yielded a good palatal exposure (Fig. 1), and in the other part the interorbital

region and the structure of the postorbital bars could be determined (Fig. 2).

Fic. 1.—Paranteosaurus primus Gen et Sp. Nov.

Orthoprojection of the Ventral View of the skull of

the holotype, S.A.M. 11485, Mynhardtskraal,

Beaufort West. (x }.)

157

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

The palate as shown in Fig. 1 agrees very closely in structure to that of

the Titanosuchian genus Anteosaurus. As in Anteosaurus the alveolar border

of the premaxilla does not lie in the same plane as the maxillary border,

but curving upwards makes an obtuse angle just anterior to the canine; on

both sides only the root of the last incisor is preserved, with anteriorly a

matrix-filled groove with little indication of a division into separate alveoli;

in the left maxilla there is in this groove room for two incisors and in the

right possibly for three; the incisor count thus falls within the limits set for

Anteosaurus,; the canines have only the roots preserved, but these indicate

that the canines were directed much more anteriorly than in Anteosaurus, as is

usual in Anteosaurus, five irregularly spaced small postcanines are present;

these are small labio-lingually flattened conical teeth; in a horizontal fracture

through the right maxilla a replacing root is seen lying linguo-anteriorly to

the third postcanine. The crescentic dentigerous boss on the palatine and the

relations of the premaxilla, prevomer, maxilla, palatine, transversum and

the pterygoid are as in Anteosaurus.

a Os

®eseaee*

“e

Fic, 2.—Paranteosaurus primus Gen. et Sp. Nov.

Orthoprojection of the dorsal view of the skull of

the holotype, S.A.M. 11485, Mynhardtskraal,

Beaufort West. (x }.)

PARANTEOSAURUS I59

On the basis of the palate alone there is nothing to exclude the specimen

from the genus Anteosaurus. Similarly the outer surface of the snout (Fig. 2)

| is very similar to that of Anteosaurus, although the snout is relatively lower.

! But in the structure of the skull roof and particularly of the postorbital

bar our specimen differs very markedly from Anteosaurus. This marked

difference is due to the small size of the postfrontal, which is here a small bone

forming the dorso-posterior orbital margin, whereas in Anteosaurus the strong

pachyostotic development of the postfrontal has resulted in this element

overflowing on to the surface of the frontal and overgrowing much of the

postorbital and forming the prominent boss so characteristic of Anteosaurus.

An intermediary stage between the condition of the postfrontal in this specimen

and in Anteosaurus is shown by the Russian genus Titanophoneus. The

postfrontal in Pseudanteosaurus with its posterior tongue-like prolongation is

of quite a different nature. |

No features in this skull, which may be thought to indicate youthfulness,

are not also encountered in some species of Anteosaurus. I thus conclude that

the smallness of the postfrontal and the light build of the postorbital bar do

not represent a growth-stage in the individual, but are in fact a retention of

an older morphological stage of the family Anteosauridae.

TAXONOMIC

For this new form of the Anteosauridae I propose the name Paranteosaurus

primus Gen. et Sp. Nov. It may be characterised as follows:

A medium-sized Anteosaurid (max. length probably about 570 mm.), with

small postfrontal not extending posteriorly, lightly built postorbital bar,

without any sign of a boss-like development in the upper part of the postorbital

bar, dental formula i.3?-4?, c.1, p.c.5.

REFERENCES

Boonstra, L. D. 1954. The Cranial Structure of the Titanosuchian: Anteosaurus.

Ann. S. Afr. Mus. 42, 2, 106-148.

EFREMOV, J. A. 1940. Preliminary Description of the New Permian and Triassic

etrapodatrom U'S,S.R- Acad. Se. U'S:S.R., 10;)2, 1-140.

ANNALS

OF THE

SOUTH AFRICAN MUSEUM

VOLUME XIII

Descriptions of the Palaeontological Material acquired by the South African

Museum and the Geological Survey of South Africa.

PART III, containing:—

Ld:

12.

13.

14.

Some fossil Mammals in the South African Museum collections. By

i. B.S. Cooke, D.Sc: (With map and Plate XIX.)

Fossil Suiformes from Hopefield. By R. SINGER and E. N. KEEN.

(With Plates XX—XXIV and one text-figure.)

Struthiocephalellus: A new Deinocephalian. By L. D. Boonstra,

D.Sc. (With 3 text-figures.)

The Girdles and Limbs of the South African Deinocephalia. By

L. D. Boonstra, D.Sc. (With 6 diagrams, 108 text-figures and

Plate XXV.) .

ISSUED JANUARY 1955. PRICE {1 tos. od.

PRINTED FOR THE

TRUSTEES OF THE SOUTH AFRICAN MUSEUM

AND THE

CEOLGGICAL SURVEY. OF. SOUTH AFRICA

BY THE NATIONAL COMMERCIAL PRINTERS’ LIMITED,

JAN VAN RIEBEECK STREET, ELSIES RIVER

Ir. Some Fossil Mammals in the South African Museum Collections.* By

H B25. Cooks, D.Sc., F.ReS:S-Atr. Johannesburg.

(With map and Plate XIX.)

ABSTRACT

The collections of the South African Museum at Cape Town include the

first fossil mammal to be discovered in Southern Africa, Homotoceras bainit.

There are also some two hundred fossil teeth and bones of Quaternary

mammals from twenty-four sites, eleven of which have not previously been

recorded. Few of the specimens have been mentioned in published literature

unless they represented new species, and lists of faunal assemblages exist only

for two of the sites— Taung and Florisbad. The present account records

all the species identified in the collections and lists the assemblages for

each locality.

INTRODUCTION

The fossil collections of the South African Museum at Cape Town include

some two hundred bones and teeth of Quaternary mammals of which only

very few specimens have been mentioned in published literature. Twenty-

four sites are represented, eleven of these being new records. Of the thirteen

known localities, faunal assemblages have been listed only for two — Taung

and Florisbad — while the remaining eleven sites have been recorded merely

by the mention or description of isolated specimens. It is the purpose of

this account to provide as complete a record as possible for the assemblages

for each locality. The accompanying map shows the wide distribution

of the sites.

I. Modder River, O.F.S.

One of the most striking of the Quaternary mammalian specimens dis-

played in the South African Museum is the frontlet and gigantic horn cores

of an extinct buffalo, “‘Bubalus’’ bainii. The remains were recovered from

a depth of forty feet in alluvial deposits of the Modder River, in the Orange

* Read before Section D of the South African Association for the Advancement of

Science at its Jubilee Congress in Cape Town, July, 1952.

VOL. XLII. PART III.

ANNALS OF THE SOUTH AFRICAN MUSEUM

162

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FOSSIL MAMMALS IN S.A. MUSEUM COLLECTIONS 163

Free State in 1839. The remarkable civil engineer and naturalist, Andrew

Geddes Bain, saw the specimen and persuaded the actual finder, Mr. Martin

Smith, to send it to the Geological Society of London. Its subsequent history

is uncertain but it was displayed in the South African Museum and was seen

by the eminent English palaeontologist, Dr. Seeley, who described it in

r89r under the name Bubalus bainu.*

Several other specimens of this giant buffalo have since been found in

South Africa and three related species have been described from North

Africa (1951), East Africa (1933) and the Anglo-Egyptian Sudan (1949). In

describing the Sudan specimen, the late Miss D. M. A. Bate (1949, 1951)

created a new genus Homovoceras for the African long-horned buffaloes and

showed that this genus is allied to the African Syncerus and is very distinct

from the Asiatic Bubalus. The South African form is accordingly now

Homotoceras bain (Seeley).

Another fossil is recorded as coming from the banks of the Modder River

“half way between Kimberley and Bloemfontein’’. It comprises an imperfect

frontlet and partial horn core of a large hartebeest, described by Broom in

1909 and named Bubalis [sic] priscus. If Broom’s generic reference is

correct it should now be placed in Alcelaphus as A. priscus, but it is possible

that it might fall within the genus Peloroceras. ,

2. Delport’s Hope, Vaal River

The fauna from this site has been listed by the writer (1949) from

material in the McGregor Memorial Museum, Kimberley. The South African

Museum collection includes a number of bone fragments from which only

the proximal end of a right metacarpal can be identified as belonging to a

large bovid, possibly a kudu.

3. Sheppard Island, Vaal River

When material from this site was listed by the writer in 1949, it was

thought that some of the specimens collected by van Riet Lowe in 1928

had been destroyed in a fire at the University of the Witwatersrand in 1931.

Six of these have now been found in the S.A. Museum collections. The

species represented are Equus sanduithi, E. capensis, E. burchel, Syncerus

caffer, and possibly, a kudu.

4. Taungt

In 1920 Haughton described seven small skulls of baboons from the

limestone deposit at Taung but the full paper was not published. The name

* See Appendix, p. 168.

t Union of S. Afr. Rep. Form and Spelling of Geographical Proper Names, 1939, recom-

mended ‘‘Taung’’’ instead of the previous ‘‘Taungs’’.

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

Papio antiquus was proposed and was printed in an abstract (1924). In

1926 J. H. S. Gear collected additional material and distinguished two

species, the larger of which he named Papio africanus and the smaller

P. izodi, presumably regarding Haughton’s name as a nomen nudum. In

1940 the generic reference was transferred by Broom to Parapapio and

Gear’s views were upheld. In 1948, however, Broom re-examined Haughton’s

original specimens (which are in the S.A. Museum) and concluded that

only the larger species was represented and that the original specific name

antiquus must be held to ante-date africanus. The neotype skull of the

species is numbered S.A.M. 5356 and the best jaw is S.A.M. 5357. The

generic reference remains amended and the correct designation is now

Parapapio antiquus (Haughton).

5. Florisbad (Hagenstad Salt Pan)

The S.A. Museum collections include a selection of specimens from the

spring deposit at Florisbad first described briefly by Broom (1913) and later

by Dreyer and Lyle (1931), whose list is complete but subject to several

specific amendments. The assemblage includes the following species: Pedetes

hagenstadi, Aonyx robustus, Diceros bicornis, Equus quagga, E. lylet, E.

capensis, Hippopotamus amphibius, Phacochoerus helmei, P. compactus, P.

aethiopicus, Homotoceras bainu, Pelea capreolus, Pelorocerus helmei, Conno-

chaetes antiquus (2), Damaliscus albifrons, Taurotragus oryx, Strepsiceros

strepsiceros, Gazella bondi (?), Antidorcas marsupialis, Cephalophus sp.,

Syluicapra grimmia.

6. Hoogstede, Tarkastad District

Five specimens from the farm Hoogstede, west of Queenstown, are lightly

mineralised and are probably surface finds. One specimen is a tooth of

a. domestic ox. The remainder represent a lion, Equus burchelh and

Pelorocerus sp., possibly P. broomi.

7. Lwartkops, Sundays River

A small fragment of an upper molar of Hippopotamus sp. is the only

fossil mammal from this locality, which has yielded much invertebrate

material to other collections.

8. Victoria West

From the farm Jakkalsfontein near Victoria West comes an upper molar

of Equus burcheli. There is an upper premolar of Equus capensis from an

unknown locality in the same area.

FOSSIL MAMMALS IN S.A. MUSEUM COLLECTIONS 165

9. Brakfontein, Three Sisters

A single specimen from the farm Brakfontein, near Three Sisters, is a

fragment of a left lower jaw of Diceros bicornis.

to. Beaufort West

A lightly mineralised milk molar of Equus cf. burchelli comes from the

farm Little England near Beaufort West.

II. Cango Caves, Oudtshoorn

Six specimens from the Cango Caves are referable to Procavia capensis,

Connochaetes sp. and Tragelaphus cf. scriptus.

12. Linkerhandsgat and Noottgedacht, Stanford

A few miles north-east of Stanford is an unusual occurrence of chalky

limestone and calcified sands containing bones and teeth of mammals. Three

specimens from Linkerhandsgat are identified as Crocuta sp., Thos mesomelas

and Alcelaphus cf. caama. From the adjoining farm Nooitgedacht, there

are three fragments representing Redunca arundium and Aepyceros melampus.

A larger private collection and material collected by the writer will be dealt

with at some length in a separate paper.

13. Hawston

The sand-dunes along the coast near Stanford and Hawston have yielded

almost unmineralised teeth of the African elephant and of the black rhinoceros.

14. Skildegat Cave, Fish Hoek

This cave, also known as Peers’ Cave, was excavated twenty-five years

ago by B. and V. Peers and it is probable that the eight teeth from this

locality which are in the S.A. Museum collections were recovered during the

excavations. A bovid incisor cannot be determined generically but the

remaining specimens belong to Equus zebra and E. capensis.

15. Kalk Bay

This area yielded three molars of Hippopotamus MR and part of

a lower molar of Loxodonta africana.

16. Yzerplaats, Maitland District

The type series of teeth of Equus capensis, described by Broom in 1909

and figured in 1928, was contained in a block of limestone washed up on the

beach at Yzerplaats. :

166 ANNALS OF THE SOUTH AFRICAN MUSEUM

17. Bloembosch, Darling District

The dune-covered farm Bloembosch, north of Yzerplaats, has furnished

nearly fifty specimens, including bones, imperfect jaws and isolated teeth.

Equus capensis is abundant, the collection including the plesiotype upper

left second molar referred to Broom’s species by Haughton (1932). Other

equine material is not specifically identifiable except for an unmineralised

lower jaw of a young Equus zebra. Diceros bicornis occurs. The only

carnivore is Cyvrocuta crocuta. Artiodactyls include Hippopotamus cf.

amphibius, Guwiraffa camelopardalis, Homotoceras bainu, Syncerus caffer,

Connochaetes sp., Hippotragus cf. niger. The two lower mandibles which

the writer (Cooke 1947) named Hippotragus problematicus, are also in the

collection; it was suggested that these jaws might represent the almost

unknown H. leucophaeus but Dr. Broom informed the writer shortly before

his death that he had found material of the Blue Buck during his last

European-American tour and that the Bloembosch fossil was clearly distinct.

His notes and drawings have not so far been found.

The recently discovered site at Elandsfontein, near Hopefield, which is

being studied by the University of Cape Town, has yielded abundant fossil

material of a character similar to the Bloembosch specimens;* it has also

provided a fossil human cranium.

18. Saldanha Bay

The only species represented by the five isolated teeth from this locality

is Equus capensis. One of the specimens is the isolated upper fourth premolar

described by Broom (1913) as ‘‘from Darling’’, and another is the plesiotype

left lower premolar described by Haughton (1932).

19. Hoedpjesbaa

The limestone quarries at Hoedjiesbaai, near Saldanha Bay, have furnished

sixteen very nice specimens, mostly partial jaws with teeth. The five species

are Procavia cf. capensis, Thos mesomelas, Arctocephalus pusillus, Suricata

sp. (or possibly Cynictis), and Raphicerus campesiris. This is believed to be

the first record of the Cape sea lion in the fossil state.

20. Geelwal Karoo, Van Rhynsdorp District

This site has provided a partial lower jaw of Thos mesomelas.

21. Near Springbok, Namaqualand

A site 40 miles east of Springbok yielded the type series of cheek teeth

described by Haughton (1932) as Notohipparion namaquense. The original

illustration exaggerates the breadth of the crowns as the plane of drawing

is parallel to the rather oblique grinding surface.

* See: Following article by R. Singer and E. N. Keen.

FOSSIL MAMMALS IN S.A. MUSEUM COLLECTIONS 167

22. Bogenfels, S.W.A.

The collections include an incomplete lower jaw from Bogenfels, S.W.A.,

labelled “‘“Propalaeonyx africanus Stromer’’. It comes, presumably, from

the so-called Miocene beds.

23. Kalk Plateau, S.W.A.

Two equine teeth, apparently of Equus burchelli, are recorded as coming

from a well on the Kalk Plateau east of Marienthal in S.W.A.

fa Wsaros, S.W.A.

The type series of upper and lower cheek teeth of Equus sandwithi

(Haughton 1932), was associated with a number of other specimens not

previously mentioned. It is now possible to reconstruct all the cheek teeth,

though not of a single individual. One tooth of Equus capensis is also

represented in the assemblage, and the bovid genera Connochaetes and

Strepsiceros are present but the species cannot be determined.

25. Other Locahties

The collection includes a lime-encrusted upper premolar of Equus cf.

zebra from Broken Hill, Northern Rhodesia. There is a piece of. grey

limestone from the vicinity of Mt. Lemagrut (near the famous Olduvai

gorge), with part of the left lower jaw of a rhinoceros exposed.

26. Unknown Localities

Twenty specimens are without locality records. The species represented

are Equus burcheli, E. kuhni, E. capensis, Hippopotamus sp., Damaliscus

sp., Connochaetes sp. and Phacochoerus sp.

REFERENCES

Bate, D. M. A. 1949. A new African fossil long-horned buffalo. Ann. Mag. Nat.

Hist. (12), Il, pp. 396-308.

Bate, D. M. A. 1951. The mammals from Singa and Abu Hugar. In Fossil Mammals

of Africa, II, pp. 1-28. British Museum (Nat. Hist.), London.

Broom, R. 1909. On a large extinct species of Bubalis. Ann. S. Afr. Mus. VII,

PP: 27, 280.

Broom, R. 1909. On the evidence of a large horse recently extinct in South Africa.

IOI: BP. Zo... .2o2.

Broom, R. 1913. Man contemporaneous with extinct animals. Ann. S. Afr. Mus.

1, pp. 13-16:

Broom, R. 1928. On some new mammals from the Diamond gravels of the Kimberley

District. Ann. S. Afr. Mus. XXII, pp. 439-444. .

Broom, R. 1940. The South African Pleistocene Cercopithecid Apes. Ann. Transv.

Mus. XX, pp. 89-100.

Broom, R. 1948. Some South African Pliocene and Pleistocene Mammals. Ann.

Transv. Mus. XXI, pp. 1-38.

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

Broom, R. and ScuHepers, G. W. H. 1946. The South African Fossil Ape-men;

the Australopithecinae. Tvansv. Mus. Mem. No. 2, pp. 1-272.

Cooke, H. B. S. 1947. Some fossil hippotragine antelopes from South Africa.

So. Aly. J. Sct. SLIT, pp. i226-237.

Cooke, H. B. S. 1949. Fossil mammals of the Vaal River basin. Geol. Surv. Mem.

XXXV (3), pp. I-109g.

DREYER, T. F. and Lyre, A. 1931. New fossil mammals and man from South Africa.

Grey Univ. College Dept. Zool., Bloemfontein, pp. 1-60.

Gear, J. H. S. 1926. A preliminary account of the baboon remains from Taungs.

S. Agr oY set: OL pp ig3aa-7a9.

HaucutTon, S. H. 1924. Papio antiquus n.sp. Trans. Roy. Soc. S. Afr. XII. Min.

& Proce!" May)1920)"'p. (EX VEE.

Haucuton, S. H. 1932. The fossil Equidae of South Africa. Ann. S. Afr. Mus.

XXVIII, pp. 407-427.

SEELEY, H. G. 1891. On Bubalus bainii (Seeley). Geol. Mag. VIII, pp. 199-202.

APPENDIX

Bubalus bainii

The date when the specimen was received at the Museum is not recorded. Two

entries in the Account Book for 1856 may refer to this specimen: ‘‘March. To Ford

repairing horns, £1 15s.’’ Ford had already repaired the models of natives, presumably

made of plaster, thus having a knowledge of plaster-work, which would be suitable

for the repair of fossils. ‘‘September. To G. West, paint for fossil head, 3s. 6d.”’

The skull was on exhibition over the entrance to the old Museum (now the

west wing of the South African Library) in 1860, when Layard compiled his Catalogue:

“Immediately overhead is a magnificent fossil frontlet of an extinct bovine from

Thaba ’Nchu.’’ (Layard, E. L. Catalogue of the South African Museum, Part 1,

Mammals, p. 7. Cape Town, 1862. Preface dated 1st January, 1861, publication

delayed until 1862. See Annual Report S.A. Mus. for 1862.) It was in the same

position when Seeley saw it.

The locality Thaba ’Nchu is not mentioned in Bain’s references to this fossil

Bovine. Bain did not visit the eastern part of the Orange Free State until 1845.

Thaba ’Nchu, however, does lie within the catchment area of the sources of the

Modder River. lLayard’s entry seems to be the only record of the locality, unless

possibly a more precise site can be traced by research in the Deeds Office for Mr.

Martin Smith’s farm —dif he possessed one.

In Dr. Cooke’s map the site of the locality (No. 1) is placed south of Kimberley

in the western part of the Orange Free State; but if Layard was correct it should

be placed in the eastern part between Bloemfontein and the border of Basutoland.

Seeley (p. 201) gave two reasons for considering this type specimen as the

one referred to by Bain in 1839. The second reason seems to be acceptable. In his

letter to Sir H. de la Beche in 1844, read to the Geological Society in 1845, and

printed (abridged) in Trans. Geol. Soc. Lond. (2), VII, 4, 1856, Bain said that

Martin Smith’s fossil was ‘‘in Cape Town’’. The South African Museum possesses

Bain’s own copy of this part of the Transactions, in which two words have been

added by Bain (p. 59) to make the sentence read: ‘‘This fossil is now in the Cape

Town Museum.’’

Seeley’s first reason is less acceptable because there may be two interpretations.

Thomas Bain’s words ‘‘his father’s fossil’? may mean either the fossil which his

father induced Martin Smith to send to Cape Town, or an example which A. G. Bain

himself found.

It is, therefore, appropriate to reproduce here (pl. XIX) Bain’s MS. drawing

and description (from memory) of a specimen of this bovine which he himself found

on Mr. G. Southey’s farm near Graaff-Reinet. The original sketch is in the S. Afr.

Museum library — but what happened to the specimen?

See: Lister, M. H. Journals of Andrew Geddes Bain. Van Riebeeck Soc. Publ.

No. 30, p. 230. Cape Town, 1949. Also Seeley, H. G., 1891 (op. te

PVATE GES

MiSs, Wl SIUIUE

Annes. Afr.

12. Fossil Suiformes from Hopefield.* By R. SinGER and E. N. Keen.

Anatomy Department, University of Cape Town.

(With Plates XX—XXIV and 1 text-figure.)

INTRODUCTION

The fossil site on the farm ‘‘Elandsfontein’’ situated 10 miles south-west

of Hopefield, 90 miles north of Cape Town, was located by one of us (R.S.)

in May, 1951.f On numerous subsequent visits various members of the

University of Cape Town staff, Professor M. R. Drennan, Messrs. J. A.

Mabbutt, K. Jolly and the authors, have collected highly fossilized bones and

stone implements from the surface of the site. The range of specimens already

identified includes a large number of extinct and existing mammals, the

proportions of which suggest an early Upper Pleistocene period, with the

possibility of extension into the late Middle Pleistocene.

In January, 1953, Jolly, on a field trip with Singer, discovered pieces of

the greater portion of the calvarium of an hominid. On three subsequent

expeditions Jolly and Singer retrieved additional portions of the cranium

which has since been reconstructed (Drennan, 1953; Singer, 1954). Fluorine

tests, carried out through the courtesy of Dr. K. P. Oakley of the British

Museum, reveal that this Saldanha Skull and the Mesochoerus fossils (described

in this paper) were apparently contemporaneous.

Elandsfontein lies in the sandy veld between the Sout River and the

Langebaan-Saldanha Lagoon. The site is 300 feet above sea-level and is

divided into wind-scoured kloofs or depressions by sand-dunes which are

either drifting or, where covered by vegetation, stationary. Ridges of

ferricrete cut diagonally across the length of the site, and in places the dunes

are capped by massive calcrete mounds or flat boulders of partly silicified

surface limestone. Softer, cellular calcretes are found in certain places at

~ * Simpson (1945) has given reasons for placing the Hippopotamidae and the Smidae

into the separate infraorders ANCODONTA and SUINA; both are, however, included

in the suborder SUIFORMES.

+ The existence of fossil-bearing sites in the Darling-Hopefield district has been known

for years (Broom, R., Annals S. Afr. Mus. VII, p. 281, 1909, and XII, p. 13, 1913;

Cooke, H. B. S., XLII, p. 161, 1955), but hitherto no thorough investigation of

the area had been made.

279 ANNALS GF THE SOUTH AFRICAN MUSEUM

the lowest parts of the depressions. The tortuous courses of the ferricrete

ridges indicate that they are the indurated lower flanks of old sand-dunes

now stripped bare of the sand walls. This ferruginization is usually associated

with moist ground conditions, a fairly high stable water-table and an

abundance of vegetable acids in the soil. It seems that this fossil site was at

one time a large vlei or lagoon along the edge of which animals roamed.

The rich collections of stone implements indicate the presence of Man on

the site from a late stage of the Chelles-Acheul (Stellenbosch) Culture until

the period when the Bush races were developing their culture. The occupation

is not a continuous one. A large number of the implements exemplify the

transition from the South African Earlier to the Middle Stone Age.

DESCRIPTION OF MATERIAL

Fam. HIPPOPOTAMIDAE

Hippopotamus amphibius Linnaeus 1758

The collection of fossil mammalian remains from the Hopefield site

includes several fragments of jaws and teeth which are identified as hippo-

potamus remains. Many of the specimens derive from young animals and

show deciduous or unerupted permanent teeth. Six of the permanent molar

teeth are complete and can be compared in size with the limits stated in

Cooke’s (1949) survey of all the South African fossil material recovered up

to that date. None of the measurements fall outside the wide limits which

are characteristic of this species. There are therefore no grounds for

separating the remains from the existing species Hippopotamus amphibius,

fossil specimens of which have been found in several other South African

localities.

Fam. SUIDAE

Genus Mesochoerus, Shaw and Cooke, 1941

Mesochoerus lategant, sp. nov.

Dragnosis: A Mesochoerus resembling M. olduvaiensis (Leakey, 1942) in

the structure of the third lower molars. The second lower molars have a

well developed anterior median pillar. The crowns of the third lower molars

are longer by 10% and narrower by 10%, and the height of the unwom

pillars is greater by 10% than in M. olduvaiensis. The pillars of the upper

molars are obliquely arranged, those on the labial side being placed slightly

in front of their lingual partners. The fourth premolars are molariform.

Type: The complete upper and lower cheek dentition of a single animal,

recovered in seven fragments numbered S6, S7, S8, S14, S15, $16, S17 in

the Hopefield collection in the Anatomy Dept., University of Cape Town.

(Plate XX.) Specimens $7, S14, S16 and S17, constituting the right dentition.

FOSSIL SUIFORMES FROM HOPEFIELD rE

have been donated to the S.A. Museum where they have received catalogue

number I1I712.

Among the fossil specimens collected from the Hopefield site are several

upper and lower teeth and partial jaws which appear to belong to a single

species of large suid. All the fossils at Hopefield have been more or less

damaged by the movement of the sand in which they lie. For this reason

recovery of an intact jaw is uncommon, and the study is necessarily confined

to the characters of the individual teeth, which are often found lying free

and not close to any other teeth or suid remains.

The collection consists of teeth and jaws from ten individuals and includes

a complete upper and lower molar dentition of a single animal (S6: left

M: Mz M:;; S15 and S8: left M° M* M’; S16 and S17: right M: Mz M, P, P;;

S14 and $7: right M* M* M’ P*). Three other pairs of left and nght lower

third molars (S2 and $3, S12 and S13, Sg and Sga) appeared to derive from

a single animal in each case, and one pair of right and left upper third molars

(Str and S1o) similarly resembled each other. These upper molars could

not, however, be made to occlude satisfactorily with any of the lowers. Five

individuals are represented by isolated teeth (Sr, mght M:; S4, nght M’;

S5, left M° and M’; S2o, left M* and M’; and Sat, right M,).

The various teeth resemble each other sufficiently closely to support the

conclusion that only one species is represented.

LOWER CHEEK TEETH

Intervals: 4/5 3/4 D3 V2

Lateral Sth 4th 3rd 2nd Ist

Pillars: pair pair pair pair pair

Fic. 1.

Third lower molars: ten specimens from six individuals. A typical specimen

in moderate wear is illustrated in Fig. 1, which also shows the convention

by which the lateral and median pillars are identified. The crowns of these

172 ANNALS OF THE SOUTH AFRICAN MUSEUM

teeth are each made up of five pairs of lateral pillars arranged symmetrically

(Si, S2, S6, Pl. XXI). Each pair appears to meet in the median line of

the tooth early in wear, but becomes separated by median pillars in later

wear. The usual number of median pillars is nine. A single median pillar

is found in front of the first lateral pair, sometimes in the median line,

sometimes leaning to the labial side of the tooth. There are two median

pillars in the 1/2, 2/3 and 3/4 intervals. These may be of equal size, or

the posterior one may be larger than the anterior. A single median pillar

is usual in the 4/5 interval, and another in the posterior median position,

i.e. at the posterior extremity of the tooth. A few of the specimens show

duplication of the median pillars in the 4/5 and posterior positions.

The tooth erupts in such a way that the first three pairs of lateral

pillars rapidly come into wear one after another, beginning in front (Pl. XX,

B). Wear commences in these anterior pillars before the fourth and fifth

pairs erupt. Later on growth and subsequent wear proceed at a faster rate

in the posterior part of the tooth, so that in late wear the tooth may present

an appearance of relatively even attrition. The upper third molars erupt in

a similar way.

In general the pattern of the worn surface of the lower tooth is simpler,

less complex and less folded than that of the corresponding upper tooth.

There is a cingulum at the crown-root junction in all specimens, but in

some it is very poorly developed (S21, Pl. XXII). In many of the teeth

the crown is partially covered with a thick layer of cement (S2r, Pl. XXII).

Roots are very well developed along the whole length of the tooth, even at

the stage when the posterior pillars are neither fully developed nor erupted

(S21, Pl. XXII); at this stage the length of the root may exceed the height

of the crown. The roots supporting the anterior three pairs of lateral pillars

are more or less separated from their neighbours, and straddle a wide dental

canal (S16, Pl. XXII, D). In some cases the tips of these roots are distorted

in quite irregular ways (So, Pl. XXIII, B). Extra median rootlets are

occasionally present in both upper and lower third molars, and vary from a

minute nodule to a spicule as long as the neighbouring lateral roots (Sq,

S14, Pl. XXIV). The curvature of the roots is backwards and to the labial side;

the labial roots are always larger than those on the lingual side of the tooth.

The roots of the posterior two pairs of lateral pillars tend to be fused together

to form flat plates arranged antero-posteriorly. Towards the back of the

tooth these plates converge, and fuse together in the median line of the tooth,

under the posterior median pillar (cf. Pl. XXIV, D, S4).

173

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174 ANNALS OF THE SOUTH AFRICAN MUSEUM

Second lower molars: three specimens from two individuals. The crowns

of these teeth are made up of two symmetrically placed pairs of lateral pillars,

a single well developed anterior median pillar, two median pillars in the

interval between the lateral pairs, and a fairly large posterior median pillar.

The anterior median pillar is eccentrically placed on the labial side of the

tooth, and the posterior median pillar on the lingual side of the median line

(S6, Pl. XXI, B). In the two specimens in which the pattern is clear there

are small accessory enamel nodules behind each labial lateral pillar and a

small row of nodules at the extreme anterior part of the tooth, in front of

the anterior median pillar (S17, Pl. XXII, E).

There are four well developed separate roots, which appear to diverge

whether the tooth is viewed end-on or from the side. The general curvature

of the root is backwards and to the labial side (S17, Pl. XXII, F). The

posterior roots are larger than the anterior.

Dimensions of Crown.

Greatest Greatest Occlusal Length of

Length. Breadth. Breadth. Roots.

$3 (R) 24 20 — _

S6 (L) 29 18 16 —

$17 (R) 30 20 14 29

Mean 28 19 15

Mesochoerus olduvaiensis

Holotype 28 20 17 —_—

Paratype 29 ZU 18 —

Mesochoerus paiceae

Neotype 32 19 18 20-25

TaBLE II: Dimensions of Second Lower Molars in mm.,

and comparison with the dimensions reported

for M. olduvaiensis and M. paiceae.

First lower molars: two specimens from different individuals. Both these

teeth are extremely worn (S6, Pl. XXI, B; S17, Pl. XXII, E). The second

and third molars belonging to each of these first molars have been found, and

it can be seen that at the time when the first molar was worn down to the

roots, the posterior part of the third molar was incompletely developed and

unerupted (S6, Pl. XXI, B). The degree of wear of these teeth precludes

a determination of the enamel pattern in the earlier stages of wear. In each

there remain two fairly large convoluted pillars, one in front of the other,

with small islands of enamel in the centre of the outlines, indicating an

originally more complex pattern from which the present appearances were

derived. |

There are four roots, as in the second molar, diverging in both views,

the posterior roots being more massive than the anterior (S17, Pl. XXII, F;

56) Pl x)

FOSSIL SUIFORMES FROM HOPEFIELD 175

The greatest length, greatest breadth, and occlusal breadth of the crowns

of the two teeth are 20, 14 and 12 mm. (S17); 20, 14 and 14 mm. (S6),

respectively. One of the roots exceeded 20 mm. in length.

Fourth lower premolar. There is one partly broken representative of this

tooth, and it is molariform in type (S17, Pl. XXII, E, F). The crown

structure is poorly defined, but seems to be made up largely of two pillars

arranged antero-posteriorly. The greatest length of the crown is 16 mm.,

and the greatest breadth 12 mm. Two roots are disposed antero-posteriorly,

and appear to have exceeded 14 mm. in length. The posterior root is partially

divided into labial and lingual parts near its tip.

Third lower premolar. Only the remains of the roots of one of these

teeth were found, embedded in the fragment of jaw bearing a second and

first molar (S17, Pl. XXII, E, F). Two roots appear to be arranged antero-

posteriorly, and one of these is estimated to have exceeded 22 mm. in length.

The length of the lower molar series in the type specimen was 120 mm. on

the left side and 121 mm. on the right side.

UPPER CHEEK TEETH

When the upper teeth are compared with the lower teeth, certain differences

are sufficiently marked to be summarised as follows: The upper cheek teeth

are broader than the lower, and show more complex and folded enamel

patterns; the structure of the third upper molars is less regular, and there

is a tendency in all the teeth for the arrangement of the columns to be

oblique, the columns on the labial side of the tooth being placed slightly

in front of the level of their lingual companions (Pls. XXIII and XXIV).

Third upper molars: seven specimens from five individuals. As already

mentioned, the crown structure of these teeth is less regular than in the case

of the lower third molars (S20, Pl. XXIV, A). The first two, and in some

cases three, pairs of lateral pillars can be compared with the corresponding

structures in the lower teeth, although they are obliquely placed, with the

labial pillars ahead of the lingual partners. At least three regular lateral

pillars are found on the lingual side of all the specimens, but there may be

only two such regular lateral pillars on the labial side. Behind this regular

part of the tooth is found a variable number (5—7) of smaller pillars,

irregularly arranged in such a way as to defy division into median and lateral

pillars. The posterior extremity of the tooth may consist of a single pillar,

or of two pillars of approximately equal size. In the anterior median position

is a pillar either frankly divided into a central triangular mass with two

176 ANNALS OF THE SOUTH AFRICAN MUSEUM

small outriders, one on each side, or showing a clear tendency to be divided

in this way. In the 1/2 and 2/3 intervals there is usually a single smaller

median pillar; occasionally this is duplicated. A curious feature of the lateral

pillars is a groove which appears on the outer side of the pillars low down,

near the crown/root junction. This is well seen on the lingual side of the

specimen S15 (Pl. XX, B). The third upper molars erupt in much the same

way as the corresponding lower teeth (see p. 172).

As in the case of the corresponding lower teeth, the upper third molars

have well developed roots along their whole length. The roots supporting

the first two lateral pillars are usually separate, and may be divided so that

two rootlets support one pillar. Behind this the roots are more or less

fused (S4, Pl. XXIV, D), sometimes into plates similar to those seen in the

lower teeth. The roots diverge widely when seen end-on, and their general

curvature is backwards (S5, Pl. XXIII, B).

Second upper molars: four specimens from three individuals. The funda-

mental structure of the crown is two pairs of lateral pillars, an anterior median

pillar, a single central pillar, and a posterior median pillar. The enamel

bounding these pillars is considerably folded and the picture is complicated

by the presence of additional small enamel nodules. The maximum number

of these is eight, one on each side of the anterior and posterior median

pillars and two between each pair of lateral pillars. The disposition of the

pillars shows the same slant as is seen in the third molars, with the labial

side in advance. The anterior median pillar is situated more or less on the

lingual side of the tooth (S20, Pl. XXIV, A). In later wear the posterior

median pillar tends to fuse with the posterior lateral pillar on the lingual

side (S20, Pl. XXIV, A).

Four roots are present, the posterior pair being larger than the anterior.

The general curvature of the roots is backwards (S5, Pl. XXIII, B).

Dimensions of Crown.

Greatest Greatest Occlusal Height of unworn Length of

Length. Breadth. Breadth. pillars above root. Roots.

3rd Posterior Anterior. Posterior.

pillar. group.

Third Upper Molars

S4 (R) 65 25 22 29 22 34 25

$5 (L) 64 25 20 36 30 34 2g

S10(R) 67 26 23 — 26 — —

S1r1(L) 70 26 Sie. — 27 — —

S14(R) 70 24 <F 32 32 29 a

S15(L) 70 26 — 36 35 26 —

S20(L) 63 28 23 — —_ a =

Second Upper Molars

$5 (L) 29 22 19 20+

S7 (R) 29 22 ky, 29

S8 (L) 29 pips 17 =

S20(L) 32 24 22 274

TasBLE III: Dimensions of Second and Third Upper Molars in mm,

FOSSIL SUIFORMES FROM HOPEFIELD 177

First upper molars: two specimens from the same individual. As in the

case of the corresponding lower teeth, both specimens exhibit advanced wear.

Only one of the two is in a sufficiently early stage of wear to allow analysis

of the original enamel pattern. Specimen $7 (Pl. XXIII, D) suggests that

in early wear a first molar resembles a second, on a smaller scale; as wear

advances fusion develops between the anterior median pillar and the two

adjacent lateral pillars, and between the central and posterior median pillars

and the two posterior lateral pillars. In advanced wear the appearance is

of two rather large nondescript enamel figures arranged antero-posteriorly.

At this stage there is little difference in the appearance of two rather large

nondescript enamel figures arranged antero-posteriorly. At this stage there is

little difference in the appearance of the worn surfaces of the upper and lower

first molars. ,

In neither specimen were the roots clearly defined, but there are indications

that they are four in number, as in the case of the second molars and the

first lower molars.

The greatest length, greatest breadth, and occlusal breadth of the two

teeth are 20, 18 and 15 mm. ($7); 21, 20 and 17 mm. (S8), respectively.

Fourth upper premolar (S7, Pl. XXIII, C, D). A single specimen of this

tooth was recovered and it is distinctly molariform in appearance. There

are two lateral pillars, and an anterior and posterior median complex, each

consisting of a reasonably large mass with several additional minute enamel

nodules. Greatest length 8 mm., greatest breadth 9 mm., occlusal

breadth 8 mm.

There appear to have been two labial roots and one lingual root. The

anterior labial root is smaller than the posterior, and is 18 mm. long.

DISCUSSION

There is no difficulty in identifying the genus to which these teeth belong

as Mesochoerus, first defined by Shaw and Cooke in 1941. These authors

listed as diagnostic characters of the third lower molar (type specimen) of

this genus a relatively low crown, columns clearly separated from one another,

a cingulum demarcating crown from root, and large brachyodont roots.

The teeth here described agree with these criteria in each particular.

Apart from the specimen originally named Notochoerus paiceae by Broom,

to which species Shaw and Cooke referred further material when renaming

the genus, only Leakey (1942, 1943) has described additional species of

Mesochoerus. The first was Mesochoerus olduvaiensis, described on two

mandibular fragments, each with a third and second molar. These fragments

were recovered from Beds I and II at Oldoway, horizons regarded as Middle

178 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pleistocene. The second was Mesochoerus heseloni, described from seven

mandibular and two maxillary fragments found at Omo, Abyssinia. Here the

fossil beds were stated to be of Lower Pleistocene age.* ©

The differences between the present species and M. paiceae or M. heseloni

are considerable. M. heseloni has third molars which are shorter and lower

crowned than in the present species. The two described specimens of Ms;

of M. paiceae are not only shorter and higher crowned than the specimens

described here but are also less complex, having four pairs of lateral columns

as against five.

There remains M. olduvaiensis. The general structure and appearance of

the teeth do not differentiate them from those described by Leakey (1942).

There are five pairs of lateral pillars in M; and the teeth are not as hypsodont

as those of M. paiceae. The three specimens of M. here described have a

well developed anterior median pillar, whereas both teeth described by Leakey

lacked this feature. This characteristic of M. was included by Leakey in

the diagnosis of M. olduvaiensis, so that a difference of structure is apparent.

Its taxonomic value may, however, be doubted.

It is only when the measurements shown in Table I are examined in detail

that the differences between Leakey’s and the present species become clear.

Despite the length of M:. which exceeds that of Leakey’s species by about

10%, the teeth are on the average 10% narrower. The difference in length/

breadth index becomes correspondingly striking (mean of 3.4 against 2.7 in

M. olduvaiensis). With the increase in length has gone a certain increase

in the height of the crown so that the length/height index does not differ

markedly.

It is clear that these measurements must be held to differentiate the

present species from M. olduvaiensis. The name Mesochoerus lategani is

suggested in honour of Mr. S. P. Lategan, on whose farm the specimens

were discovered.

The upper cheek teeth of M. olduvaiensis have not been recorded. It is

of interest to see that the posterior part of the upper third molars described

here show the same irregularity of structure as is found in the upper third

molars of Mesochoerus heseloni. On the other hand neither Leakey’s (1943)

nor Arambourg’s (1947) illustrations of the occlusal surface of the upper third

molar of M. heseloni show any trace of the obliquity in the arrangement of

the pillars which is so characteristic of the upper teeth found at Hopefield

(Pls. XXIII and XXIV).

The geological period to which these fossil specimens should be referred

is difficult to determine, but the total faunal assemblage from Hopefield

* Arambourg (1947) allots this species to the genus Omochoerus which he created, and

argues that the genus Mesochoerus may have to be merged with Hylochoerus.

FOSSIL SUIFORMES FROM HOPEFIELD 179

suggests an early Upper Pleistocene period. The possibility that specimens

of a Middle Pleistocene period have been mingled with later material cannot

at this stage be altogether excluded. The results of fluorine estimations on

a wide range of specimens do not so far support the idea that fossil remains

of widely differing age have been confused by sand movement on the site.

Leakey (1953) comments that in East Africa no specimens of Mesochoerus

have been recovered from Upper Pleistocene deposits, and that if the evidence

for an Upper Pleistocene dating at Hopefield is correct, the presence of the

Mesochoerus represents survival of a genus which had become extinct further

north. It is also noteworthy that the Mesochoerus teeth and jaws are the

only suid specimens so far recovered from the site, which again suggests the

late survival of an isolated species.

_ SUMMARY

Fossil Hippopotamus and suid remains from MHopefield are described. The

single suid species represented is named Mesochoerus lategani sp. nov. The early

Upper Pleistocene age which must provisionally be attached to the specimens is

contrasted with the Middle Pleistocene date allotted to the type specimens of M.

olduvaiensis, and the Lower Pleistocene dating of M. heseloni.

ACKNOWLEDGEMENTS

We are indebted to Dr. H. B.S. Cooke, University of the Witwatersrand, for

generous advice and assistance. Mr. G. McManus, Surgery Department, University

of Cape Town, kindly photographed the specimens.

The Univ ersity of Cape Town is grateful to three donors who wish to remain

anonymous and who sponsored the work at Hopefield.

Publication of this paper was assisted by a grant-in-aid made by the Council of

the University of Cape Town.

REFERENCES

ARAMBOURG, C. 1947. Mission Scientifique de L’Omo, Tome I, Fascicule II:

Paléontologie.

Cooke, H. B. S. 1949. The Fossil Suina of South Africa, Trans. Roy. Soc. S. Africa,

92, 1-44.

Drennan, M. R. 1953. A preliminary note on the Saldanha skull, S.A. J. Sci., 5e,

7-11. ;

Leakey, L. S. B. 1942. Fossil Suidae from Oldoway, Journal of the East African

Natural History Society, 16, 178-196.

LeakEY, L. S. B. 1943. New fossil Suidae from Shungura, Omo, one of the East

African Natural History Society, 17, 45-61.

LEAKEY, L. S. B. 1953. Persona] communication.

MIDDLETON-SHAW, j-C.-angs Coors 4B: S: 1941.) New ossil Pig remains from

the Vaal River Gravels, Trans. Roy. Soc., S. Africa, 28, 293-299.

Smmpson, G. G. 1945. The Principles of Classification and a Classification of Mammals,

Bull. Am. Mus. Nat. History, 85, 1-350.

SINGER, R. 1954. The Saldanha Skull from Hopefield, South Africa. Amer. J. Phys.

Anthrop., September (in the press).

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Nittes. Alr. Mus., Vol. XLII

. Pirate XX. A. Labial (lateral) view of the left upper and lower jaw

| fragments of the type specimen, in occlusion (S6, S8, S15).

i B. Lingual (medial) view of the same.

ANON, S, Ase, WOOS,, Woll, XIU

PLaTE XXI. A, B. Occlusal views of type specimens shown in Plate XIX

(S8 + S25 upper, SO owen). ©) Dy Occlusal views or right a(S) iancdamlek:

(S2) lower third molars. S2 is in more advanced wear and is broken posteriorly.

Pirate XXII. A, B, C. Lingual, labial and occlusal views, respectively, of

right lower third molar. D. Anterior view of right lower third molar (S16).

E, F. Occlusal and lingual views, respectively, of right lower second molar,

first molar, and fourth premolar (S17).

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PLATE XXIII. A, B. Occlusal and labial views, respectively, of left upper

third and second molars ($5). C, D. Labial and occlusal views, respectively,

of right upper second molar, first molar, and fourth premolar (S7).

Annes. Atr. Mus: Vol, XHil

oo 2345 |

PLaTE XXIV. A. Occlusal view of left upper third and second

molars (S20). B. Posterior view of left lower third molar (Sg) to

show deformity of roots which have been painted black.

C. Anterior view of right upper third molar (S14). D. Oblique

posterior view of right upper third molar (Sq). E. Anterior view

of right upper third molar (S4).

Le) Do BOONSERA:: B.sc:

Struthiocephalellus: A new Deinocephalian

The Girdles and Limbs of the South African Deinocephalia

13. Struthiocephalellus: A New Deinocephalian. By Lizuwe Dirk

Boonstra, D.Sc.

(With 3 text-figures.)

In the collection of the South African Museum there is a specimen

(S.A.M. 5006) collected by Haughton in 1916 on Abrahamskraal, Prince

Albert, which according to Rossouw (4) is Low Tapinocephalus zone. This

specimen consists of much of the skeleton of a small Tapinocephalian. There

is preserved: much of a distorted and weathered skull in two parts, viz. a

maxillary and an occipital part not in contact; parts of the vertebral column,

part of a scapula, most of a humerus, the proximal end of an ulna; the

greater part of a distorted pelvis, a femur and the proximal end of a tibia,

together with other fragments.

The bones of the girdles and limbs are being described in a forthcoming

paper on these elements based on all the Deinocephalian material in the

Museum.

The skull (Figs. 1-2) is about half the size of that of Struthiocephalus

whaitst. The pachyostosis is weak so that the postorbital bar is relatively

slender and the posttemporal opening roomy. The occiput is fairly upright

and the quadrate not shifted very far anteriorly. The snout is moderately high.

Owing to the distortion and the lack of contact between the maxillary

and occipital pieces the two figures I give here are reconstructions and

represent orthoprojections of the dorsal and ventral surfaces projected on

to the plane in which the alveolar borders of the maxillaries lie.

A comparison of these figures with those given for Struthiocephalus whaitsi

(2) shows that the present skull, apart from its being only a half as large,

is very similar in general structure. The chief points of difference are:

Little pachyostosis, no fronto-nasal boss, slender postorbital bar, more upright

occiput and less forward displacement of the quadrate.

These points of difference might very well be thought to merely indicate

juvenility in the present skull. However, in our collection I (2) have

described a juvenile skull (S.A.M. 11493) of Struthiocephalus whaitst which

is just as long as that of two adult skulls, but its youth is indicated by the

fact that the teeth are just beginning to erupt. Now in the present skull

(S.A.M. 5006) there is a full set of sixteen teeth which would tend to show

that although so much smaller it cannot be younger than the juvenile skull

(S.A.M. 11493) of Struthiocephalus.

Unfortunately no crowns of any of the anterior teeth are preserved, but

judging from the nature of the roots it does not appear probable that in the

STRUTHIOCEPHALELLUS 181

anterior teeth the crowns had the

usual Tapinocephalian structure

consisting of talon and horse-shoe-

shaped attrition surface, but

rather that they were simple

pointed teeth.

Fig. 1.

IP.

Dorsal aspect of the skull of Struthio-

cephalellus parvus. S.A.M. 5006.

Abrahamskraal, Prince Albert. (x 3.)

This as well as Fig. 2, is a recon-

structed projection by pantograph on

to the plane in which the maxillary

alveolar borders lie. F .—frontal;

I.P.—interparietal; J.—jugal; L.—

lacrimal; M.—maxilla; N.—nasal;

P. — parietal; Po.F. — postfrontal;

Pr.F.—prefrontal; P.M.—premaxilla;

P.O. — postorbital; S.M. — septo-

maxilla; Sq.—squamosal; T.—tabular.

Ventral aspect of the skull and the

first three cervicals of Stvuthiocepha-

lellus parvus. (x 3.) At.C.—atlantal

centrum; At.I.—atlantal intercentrum;

Ax.C.—axial centrum; Ax.].—axial

intercentrum; B.O. — basioccipital;

B.S.—basisphenoid; Diap.—diapophy-

sis; ]1.—third intercentrum; Pal.—

palatine; Par.—parapophysis; P.Oc.—

paroccipital; Pt.—pterygoid; P.V.—

prevomer (vomer); O.—quadrate;

Q.J.—quadratojugal; Tr.—transver-

sum; 3 C.—centrum of third vertebra.

182 ANNALS OF THE SOUTH AFRICAN MUSEUM

The crowns in some of the posterior teeth are imperfectly preserved.

Here the crowns appear spatulate in outline and labio-lingually compressed.

In one or two of the rear crowns there appears to have been a stronger

central cusp with a weaker posterior and anterior cusp, strongly reminiscent

of those known in Agnosaurus (1) and the Russian Rhopalodon.

In view of the condition in Agnosaurus the possibility that the spatulate

rear teeth may represent “‘milk’’ teeth cannot be excluded.

A series of three cervical vertebrae is preserved in articulation with

the occipital condyle.

The cervical vertebrae (Figs. 2 and 3) are very similar to those of

Moschops and Mochognathus (3) and to those of the Synapsids generally.

The proatlas is a stout bone; in lateral view its outline is that of a

shallow segment; anteriorly it has a ventral facet for articulation with the

Fig. 3.

A. Lateral view of the first three cervical vertebrae of

Struthiocephalellus parvus. (x 4.) At.A.—atlantal

arch; Pr.At.—proatlas.

B. Lateral view of left and right atlantal ribs. ( x 3.)

STRUTHIOCEPHALELLUS 183

exoccipital and a similar facet posteriorly articulates with the atlantal

prezygapophysis.

The atlas is temnospondylous with a paired neural arch resting on a

large atlantal intercentrum and an odontoid-like pleurocentrum. The arch

is of complex shape; dorso-posteriorly a postzygapophysis articulates with

the prezygapophysis of the axis; dorso-anteriorly a prezygapophysis gives

articulation to the proatlas; postero-laterally there is a strong diapophysial

process carrying a facet for the articulation of the tuberculum of the atlantal

rib; near its antero-ventral edge there is a small vertebrarterialis foramen;

dorsally the two halves do not meet to form a spine.

The atlantal intercentrum is large; it carries a posteriorly directed

parapophysial facet on its postero-lateral edge for the capitulum of the

atlantal rib.

The atlantal pleurocentrum is mostly covered by the atlantal rib, but

is probably as that described in Moschops and Mochognathus.

The axis has the two halves of the neural arch fused to each other and

to the pleurocentrum to form a typical holospondylous vertebra.

The spine is comb-shaped; anteriorly an upwardly directed prezygapo-

physial facet articulates with the postzygapophysis of the atlantal arch;

postero-dorsally horizontal upwardly facing postzygapophyses articulate with

the prezygapophysis of the succeeding vertebra; well below the junction

of arch and centrum there is situated a strong process carrying the diapo-

physial facet for the tuberculum of the axial rib, whose capitulum articulates

with a parapophysial facet situated on the postero-lateral corner of the

axial intercentrum.

The axial centrum is laterally excavated below the diapophysis and

ventrally has a sharp keel.

The third intercentrum is smaller than its predecessors.

The third cervical has a bluntly pointed spine and its centrum is deeply

excavated below the diapophysis so that ventrally it shows a sharp keel.

The atlantal rib (Fig. 3 B) is preserved on both sides; it is a greatly

flattened small bone with a weak shaft, a greatly expanded leaflike tuber-

culum and a much weaker capitulum.

This specimen in size comes very near to Moschosaurus and Agnosaurus

and agrees with these two forms in having the pachyostosis little developed,

the quadrate not greatly forwardly displaced and the skull height not

greatly reduced. In the structure of its palate and the dorsal cranial surface

it shows considerable similarity to the much larger Struthiocephalus from

which it differs however in the nature of its dentition and in the structure

of the girdles and limb-bones.

184 ANNALS OF THE SOUTH AFRICAN MUSEUM

For this form I propose the name—

Struthiocephalellus parvus. Gen. et Sp. Nov. Holotype: S.A.M.

5006. Skull, vertebrae, girdle- and limb-bones. Abrahamskraal,

Prince Albert. Low Tapinocephalus zone.

REFERENCES

zr. Boonstra, L. D. Agnosaurus Gen. Nov.: ’n Nuwe Geslag van die Deinocephaliérs.

Tydsk. Wet. Kuns. 12, 2, 242-245. 1952.

2. Boonstra, L. D. The Cranial Morphology and Taxonomy of the Tapinocephalid

genus Struthiocephalus. Ann. S. Afr. Mus. 42, I, 32-53. 1953.

3. GreGory, W. K. The Skeleton of Moschops capensis Broom, a Dinocephalian from

the Permian of South Africa. Bull. Amer. Mus. Nat. Hist. 61, 179-251. 19206.

4. Rossouw, P. J. en DE VILLiERS, J. Die Geologie van die Gebied Merweville,

Kaapprovinsie. Toel. Blad. 198. Geol. Opname, Unie S. Afr., I-71. 1952.

14. The Girdles and Limbs of the South African Deinocephalia. By LIEUWE

Dirk Boonstra, D.Sc.

(With 6 diagrams, 108 text-figures and Plate XXV.)

t. INTRODUCTION

In this communication I intend describing the pectoral and pelvic girdles

and the fore- and hind-limbs of the large number of Deinocephalian specimens

housed in the South African Museum in Cape Town. The nature of the

material determines the method of treatment, which will in this case have

to assume, in the taxonomic part, the form of an illustrated catalogue.

In all, the Museum Register contains 106 separate specimens in which

some part or other of the girdles or limb-bones is preserved. In a very large

number of cases these postcranial elements are not associated with any

cranial material and identification is thus rendered difficult in view of the

fact that authors in the past have usually made skulls and jaws ‘‘types’’.

Moreover, in many cases the specimens consist of isolated individual bones

which, being unassociated with other elements of the postcranial skeleton,

makes the task of identification no easier.

All the specimens were collected in the Tapinocephalus zone of the Lower

Beaufort; nearly all in the area known as the Koup, which includes parts

of the districts Beaufort West, Prince Albert and Laingsburg, with only a

couple from the area north of the Nieueveld Escarpment in the districts of

Sutherland and Fraserburg.

Rossouw (26) has recently subdivided the Tapinocephalus zone into three

subdivisions, and it is of interest to record that the specimens to be

described here come mostly from the lower subdivision.

Since the Museum’s first acquisition in 1904 the following collectors have

contributed to the collection of Deinocephalian girdle- and limb-bones: Cloete,

Oakley, le Roux, du Plessis, Cairncross, Maddison, Whaits, Haughton,

van der Byl, Boonstra, Avenant and Marais. Of these, Haughton with 18

and Boonstra with 65 specimens are the more important collectors.

2. HISTORICAL AND MORPHOLOGICAL

A. The Pectoral Girdle (Diagram 1)

In 1889 Seeley (28) figured a coracoid and incorrectly interpreted it as

the pubis of Titanosuchus. Broom (8) in 1905 mentioned parts of the shoulder

186 ANNALS OF THE SOUTH AFRICAN MUSEUM

girdle of Pelosuchus. In 1914 Watson (29) figured the scapulocoracoid of

Phocosaurus and described the scapula, coracoid and cleithrum of Titano-

suchus. In the same year Broom (11) figured and described the shoulder

girdle of Moschops, and on the same material Gregory and Camp (17), Romer

(25) and Gregory (18) based their figures and descriptions. In 1914 Broom

(10) also published a photograph of a Titanosuchian pectoral girdle (A.M.N.H.

5611) under the name Tapinocephalus atherstonei in error, and this scapulo-

coracoid was later figured by

Gregory (18) under the name

“‘Tapinocephalus’’ to indicate his

Be Oe Menus eek lack of confidence in Broom’s iden-

+o ssc. tification. In 1915 the pectoral

girdle of Struthiocephalus was shown

in Haughton’s (20) photograph of

the mounted specimen in the South

African Museum. Broom (14) in

1929 figured and described the

pectoral girdle of Jonkena trucu-

lenta. In 1931 von Huene, (21)

figured an interclavicle, which he

thought was that of a Titanosuchid.

In 1940 Byrne (16) described and

figured parts of the girdle of

Moschoides. Although pectoral ele-

ments are mentioned in connection

with some of the other described

Deinocephalians, they were neither

figured nor described.

In the South African Museum

collection the pectoral girdle is well

Semi-diagrammatic sketch of a Titano- represented. The scapula is pre-

suchian pectoral girdle in lateral view. oa rell : i :

Cl.—cleithrum; Cla.—clavicula: Cor.— S€rved wholly or in part in 36 speci-

coracoid; Gl.—glenoid; I.Cl.—interclavi- mens, the coracoid in 16, the

cula; P.Cor.—precoracoid; Sc.—scapula; ees ; i

S.C.F.—foramen supracoracoideus; S.Sc. precoracoid in 12, the interclavicle

—suprascapula; Tr.—mound, ridge or in 15, the clavicle in 5, whereas the

tubercle from which the scapular head eth . .

of thes a ericens wonleinates. cleithrum is only preserved incom-

pletely in I specimen.

The Deinocephalian pectoral girdle is composed of 11 bones — one unpaired

bone and five pairs — as is the case in all primitive reptiles. These are the

interclavicula, a pair of scapulae, coracoids, precoracoids, cleithra and

claviculae. With which in life a cartilaginous sternum was probably associated.

These eleven bones together form a U-shaped framework encasing the thorax

and is supported semi-slung by the paired fore-limbs.

Diag. 1.

GIRDLES AND LIMBS OF DEINOCEPHALIA 187

Ventrally the interclavicle, bisected by the median line, forms the con-

necting link between the two limbs of the U. Each limb is formed by the

more or less vertically disposed scapulo-coracoid, which consists of a dorsal

bladelike scapula and a ventral, antero-posteriorly elongated, coracoidal plate

formed by the large precoracoid and a smaller coracoid. The lower edge

of the coracoidal plate rests on the stem of the interclavicle, which sometimes

carries a strong median ridge against which the coracoids abut.

On the posterior surface of the scapulo-coracoid, at the junction of the

scapula and the coracoid, lies the simple, antero-posteriorly shortened,

glenoid cavity, which receives the shortened oval head of the humerus. The

glenoid has a dorsal facet lying in the scapula and this faces ventro-

posteriorly but sometimes also slightly externally, and a ventral facet on

the coracoid facing dorso-posteriorly and also much externally. The nature

of the glenoid cavity makes the humerus lie with its distal end somewhat or

fairly much lower than its proximal end. Moreover, it limits the anterior

disposition of the humerus.

On the posterior border of the scapula, some distance above the glenoid,

lies a mound, or ridge or a prominent tubercle, from which the scapular

head of the triceps originates. Along its anterior border the scapula is flanked

by the elongated cleithrum, which in the Tapinocephalia is, where known,

a rudimentary splintlike bone, but in the Titanosuchia is still functional as

a fairly strong elongated element with a spatulate dorsal end projecting

above the upper edge of the scapula, to which a cartilaginous suprascapula

was attached in life. Below the cleithrum and lying along the anterior edge

of the scapula and precoracoid is the long platelike clavicle, which in the

Titanosuchia, and to a lesser extent in the other forms, dorsally overlaps

the lower end of the stem of the cleithrum, fitting into a groove in this

bone. In all the Deinocephalia the clavicle, where known, has its widely

spatulate ventral end clasping the upturned antero-lateral external surface

of the interclavicle, which develops a more or less deep groove for the

reception of the clavicle.

In the pectoral girdle the two coracoids and the scapula are more or less

firmly joined by sutures to each other, with the other elements, the dermal

bones, more loosely applied to them. Ridges and grooves limit the move-

ments of the elements inter se and strong ligaments holding them together

allow a fair amount of movement. There is considerable freedom of movement

between the two halves allowing for difference of disposition when one

fore-limb is disposed anteriorly and the other posteriorly.

As a whole the pectoral girdle is thus a massive structure of overlapping

and supporting bones, united by suture or ligament, forming a strong but

188 ANNALS OF THE SOUTH AFRICAN MUSEUM

The girdle is strongest in the Titanosuchia, where quick forceful move-

ments are requisite for their carnivorous habits, weaker in the Tapinocephalia,

whose habits were more leisurely and more aquatic, and in the Anteosauria it

is weak, with the distinctive tricipital tubercle on the scapula, and a more

crawling habit.

The Deinocephalian pectoral girdle shows definite advances beyond the

morphological stage achieved by the Pelycosaurs. The more important

advances are: the loss of the supraglenoid buttress and the supraglenoid

foramen and the development in its stead of a mound, or ridge, or tubercle

for the insertion of the scapular head of the triceps muscle; the antero-

posterior shortening of the glenoid cavity, entailing the loss of the intricate

““screw-shape’’ and the formation of a simple glenoid with a dorsal facet on

the scapula and a ventral facet on the coracoid, and the glenoid, as a whole,

facing ventro-postero-externally; the total exclusion of the precoracoid from

the articulation in most forms, whereas in the others it forms no more than

a small part of the anterior rim of the glenoid; the reduction in size and

functional importance of the cleithrum (greatest in the Tapinocephalia, least

in the Titanosuchia and probably also in the Anteosauria); the increase in

size of the precoracoid, particularly antero-dorsally for the m. supracoracoideus,

and concomitantly an extension of the area for the origin of the m. scapulo-

humeralis; the loss of the dorsal process of the coracoid from which the

coracoidal head of the triceps originated primitively; the development of a

very strong interclavicle with a strong spatulate anterior end bent sharply

upwards; the clavicle is a flat bone of considerable size.

The Deinocephalian pectoral girdle is less advanced than that of some

of the higher Therapsids in that: no acromion is developed and there is no

indication of the development of an incipient spina scapularis.

B. The Fore-limb (Diagrams 2 and 3)

HUMERUS

The first Deinocephalian humerus to be figured was one from Varsfontein,

which Owen (23) in error referred to Pareiasaurus, and Lydekker (22)

provisionally included in Tapfinocephalus. In 1889 Seeley (28) figured a

specimen from Koedoeskop as that of Titanosuchus, since it was said to be

associated with the type skull fragments. Broom (7) in his description of

Scapanodon mentions three humeri as possibly associated with the jaw

fragments, but does not figure them. Then in 1909 Broom (9g) made a

humerus the type of Eccasaurus priscus and erroneously stated that it was

from the Ecca Beds, whereas in fact, the farm Sandvlakte has outcrops of

the Tapinocephalus zone of the Lower Beaufort Beds. Eccasaurus was

GIRDLES AND LIMBS OF DEINOCEPHALIA 189

finally by Broom (15) considered to be one of the herbivorous Tapinocephalia,

whereas, as will appear later, it is one of the carnivorous Anteosauria. In

1914 Broom (11) figured the humerus of Moschops and published (10) a

Dieg. 2.

P.lat. OPC.

a b c

Semi-diagrammatic sketch of a Jonkerid humerus. a. Dorsal

view. b. Ventral view. c. Proximal view. A.D.V.L.—anterior

dorso-ventral line; B.F.—bicipital fossa; Cap.—caput humeralis;

D.P.C.—delto-pectoral crest; Ect.Ep.—ectepicondyle (radial);

Ect.F.—ectepicondylar foramen; Ent.Ep. —entepicondyle

(ulnar); Ent.F.—entepicondylar foramen; L.M.L.—latero-

medial line (separates the proximal and distal dorsal surfaces

of the humerus); O.F.—fossa for the olecranon; P.Lat.—

processus lateralis; P.Med.—processus medialis; R.—radial

condyle or capitellum; U.—ulnar condyle.

photograph of the fore-limb of a Titanosuchian, which was erroneously

referred to Tapinocephalus atherstonet. This humerus was subsequently

figured by Gregory (18) without the ectepicondylar foramen being indicated.

I located this foramen when at the American Museum in 1935 thus proving

it to be Titanosuchian and not Tapinocephalian. In 1914 Watson (29),

reviewing the Deinocephalian material in the British Museum (N.H.), figured

an imperfect humerus of Pnigalion oweni, and also located an ent- and

ect-epicondylar foramen in Seeley’s humerus of Titanosuchus. The humerus

of Struthiocephalus is poorly shown in Haughton’s (20) photograph and

Broom’s (15) reconstruction. In his comparative study Romer (25) figured

the humerus of Moschops, as did Gregory (18). In 1928 Broom (13), dis-

cussing the confusion with regard to the British Museum material, figured

a humerus, which he regarded as that of Tapinocephalus atherstonet, whereas,

having two epicondylar foramina, it must be that of some Titanosuchian.

The following year Broom (14) figured the humerus of Jonkeria crassus and

briefly described, without a figure, a humerus identified as Jonkeria sp. and

gave photographs of the humerus of Dinosphageus ( = Jonkeria) haughtont,

I90 ANNALS OF THE SOUTH AFRICAN MUSEUM

and stated that there is a humerus associated with the type skull of Jonkeria

pugnax. Von Huene (21) figured a humerus identified simply as Deino-

cephalian, but clearly belonging to the genus Jonkeria. Finally Byrne (16)

figured the humerus of Moschoides.

In the collection of the South African Museum 35 specimens have the

humerus adequately preserved and a number where only unsatisfactory ends

are preserved.

The humerus in the three divisions of the Deinocephalia, where it is

known (Tapinocephalia, Titanosuchia, Anteosauria), shows certain distinctive

characters but these are overshadowed by the features they have in common.

The Deinocephalian humerus is a fairly to very short bone, massive in

the majority of forms, with greatly expanded ends and a short to very short,

strong to very strong shaft. The proximal and distal expansions make an

angle relative to each other because of a twisting on the shaft. The amount

of this “‘twist’’ varies from about 10° to over 40°, but the usual twist is

about 15-25° and postmorten deformation usually accounts for the extremes.

The proximal surface of the humerus has in its median part the caput

humeralis, which is oval in outline — sometimes narrowly and in other cases

broadly oval. Its edges overhang the dorsal and ventral faces of the humerus

to a varying degree. The face of the caput is only slightly convex, and, in

life, it must have had a thick cartilaginous pad in order to fit the glenoid

and make a relatively efficient shoulder joint. Postaxially the caput merges

into the processus medialis, which in the Tapinocephalia and Anteosauria

lies in about the same plane as the caput, but in the Titanosuchia the

processus medialis lies in a plane slightly (Titanosuchidae) or much

(Jonkeridea) distally of that in which the caput lies. Preaxially the caput

flows into the processus lateralis, which in its turn is distally continued as

the deltopectoral crest (D.P.C.). The processus lateralis usually forms the

most proximal corner of the humerus and it limits the anterior disposition

of the limb.

The distal end of the humerus differs considerably in the Tapinocephalia

and Anteosauria on the one hand and the Titanosuchia on the other hand.

In the former the capitellum or radial condyle is weak, not sharply modelled,

transversely elongate and with little ventral swelling and its face does not

extend in proximal direction along the ventral face of the bone; it is thus

more distally situated and directed. Whereas in the Titanosuchia the

capitellum is strongly bulbous, rounded and it extends moderately to very

far proximally along the ventral face, in some cases right up to the plane

in which the entepicondylar foramen has its ventral opening; its modelling

is strong with sharp and prominent edges in some specimens; thus, although

extending to the distal end of the bone, the capitellum is more ventrally

situated and directed. The ulnar condyle is relatively weak and not

GIRDLES AND LIMBS OF DEINOCEPHALIA I9Q1l

distinctly demarcated from the radial condyle and its mostly rounded but

sometimes slightly trochlear surface lies distally and only just extends on

to the dorsal surface; with, in some cases, a shallow trochlear fossa on the

dorsal surface to receive the olecranon of the ulna when the epipodial is

semi-extended.

Neither of the two epicondyles stretch distally of the articulatory surfaces.

In distal view both epicondylar ends are very little thickened in the Tapino-

cephalia and Anteosauria, whereas in the Titanosuchia they are thick to very

thick, indicating an important difference in the development of the flexors,

extensors and supinators.

The proximo-dorsal surface is roughly triangular in outline, ay tHe

base situated proximally and the apex merging into the dorsal surface of the

shaft. This surface is divided by the anterior dorso-ventral line (A.D.V.L.)

into two parts. Anteriorly (preaxially) to this line lies the triangular upper

face formed by the strong development of the delto-pectoral crest (D.P.C.).

This large area (greatest in the Titanosuchia) is indicative of the presence of

a very strong m. deltoideus. The ventro-distal corner of the delto-pectoral

crest, to which was inserted the m. pectoralis, lies much further distally

in the Jonkeridae than in the Tapinocephalia and, in particular, the Anteo-

sauria. Both m. deltoideus and m. pectoralis were apparently much stronger

in both groups of the Titanosuchia than in the Tapinocephalia and

Anteosauria.

On the other part of the proximo-dorsal surface there lies the fairly weak

oblique latero-medial line (L.M.L.), which is strongest in the Titanosuchia.

The separate muscle-scars for the scapulo-humeralis, latissimus dorsi and

the triceps on this surface are usually not clearly demarcated. But in the

plane of the shaft a thickening on the L.M.L. is developed for the attachment

of the lateral humeral head of the triceps and in some Titanosuchians a

strong moundlike tubercle is developed here.

The dorso-distal surface is also roughly triangular with the apex directed

proximally and dying out on the surface of the shaft. Distally this surface

is moderately to very greatly expanded by the development of the two

epicondylar flanges. Proximal to the distal edge of the bone, formed by

the edge of the radial and ulnar condyles, there is a shallow triangular

depression lying between the dorsal surfaces of the two epicondylar flanges.

The anterior (preaxial) border of this hollow is sharper than the posterior

(postaxial) border and is a continuation of the L.M.L. Anterior to this ridge

lies the confluent ectepicondylar (radial epicondyle) and supinator flange,

which in the Titanosuchia is pierced by the rounded ectepicondylar foramen,

either vertically or obliquely. Distally this flange has a thickened truncated

end, to which the extensor muscles were attached. The supinator part of

this flange has a thin, sharp edge — thinner in the Tapinocephalia and

Anteosauria than in the Titanosuchia. In the Titanosuchia the ectepicondylar

IgQ2 ANNALS OF THE SOUTH AFRICAN MUSEUM

foramen pierces this flange either quite near its edge where it is thinnest, or

deeper in through much thicker bone. In the first case the perforation is at

right angles to the surface, whereas in the latter case the perforation is

oblique, with the ventral exit situated further distally than the dorsal entry.

The entepicondylar flange is thicker than the confluent supinator and

ectepicondylar flange. To its thick truncated distal end the strong flexors

were attached. The slitlike entry of the entepicondylar foramen really lies

somewhat proximal to the epicondylar flange proper on the dorso-postaxial

or the postaxial surface of the shaft. The foramen perforates the bone

obliquely so that its ventral exit lies further distally as well as ventrally.

The proximo-ventral surface of the humerus consists of a large hollowed

out triangular surface (the bicipital fossa), which is bounded anteriorly by

the large delto-pectoral crest, the proximal edge, formed by the processus

lateralis, the caput humeralis and the processus medialis, and posteriorly by

the thick posterior edge, to which the median head of the triceps was

attached. In this view the delto-pectoral crest is seen to extend ventrally,

from its origin at the processus lateralis, as an inclined strong sheet of bone

terminating ventro-distally in the thickened corner for the insertion of the

m. pectoralis. The pectoralis-corner is never knoblike as in the Pareiasaurs,

but is thicker in the Titanosuchia than in the Tapinocephalia and Anteosauria.

In Jonkeria parva the D.P.C. is particularly massive and short. In the

Titanosuchia, with one exception, the pectoralis-corner is situated very far

distally, whereas in the Tapinocephalia its distal extension is moderate to

small, and in the Anteosauria small. From this corner the pectoral crest

subsides into the surface of the shaft in postero-distal direction, abruptly

in the Titanosuchia, but gently in the case of the Tapinocephalia and the

Anteosauria.

The ventro-distal surface of the humerus is broad to very broad and

consists chiefly of the capitellum or radial condyle which is large and massive

in the Titanosuchia, but in the Tapinocephalia and Anteosauria it is a much

smaller feature. The capitellum lies mostly in the pre-axial half of the distal

surface and in the Titanosuchia it forms a large bulbous mass of bone

stretching far to very far in proximal direction nearly to or even beyond the

plane in which the entepicondylar foramen lies. It is well modelled with

sharp borders in the Titanosuchia, whereas in the Tapinocephalia and

Anteosauria the modelling is weaker and it is not bulbously swollen and

never stretches far in proximal direction. The ulnar articulatory surface

is not clearly demarcated from the radial condyle and simply forms the

rounded part of the distal edge of the humerus. The lateral portions of the

ventro-distal surface form the under surfaces of the ect- and entepicondylar

flanges. The ectepicondylar flange is thin and has its edge bent downwards.

In the Tapinocephalia and Anteosauria its upper end is separated from the

surface of the shaft by a groove, and it is, in the absence of the ectepicondylar

GIRDLES AND LIMBS OF DEINOCEPHALIA 193

foramen in these groups, along this groove that the radial nerve ran. The

entepicondylar flange is thick. The ventral opening of the entepicondylar

foramen, through which passed the medial artery and nerve, is in the Tapino-

cephalia and Anteosauria a fairly narrow slit, but it is more oval in the

Titanosuchia. In the Tapinocephalia and Anteosauria the entepicondylar

foramen is always a considerable distance from the proximal edge of the

radial condyle, whereas in the Titanosuchia the radial condyle in some forms

extends so far in proximal direction that the foramen lies close to the proximo-

postaxial edge of the condyle.

THE SHOULDER JOINT

From the description of the glenoid cavity and the shape of the caput

humeralis it is clear that the shoulder joint was not a very efficient structure.

A large amount of cartilage and strong ligaments were necessary to make

the joint function at all well. Even so, any great forward disposition. of the

humerus must have taken the head of the humerus out of the socket. It

would appear that in the Titanosuchia the humerus lay more horizontally

than in the Tapinocephalia, where it was inclined obliquely downward towards

the elbow.

RADIUS

The radius is not very well known.

are preserved.

Diag. 3.

In this collection only 7 good radii

Hitherto the radius has been described in Moschops by

Broom (11) and Gregory (18). Of the radius

of Tapinocephalus Broom (13) in 1928 says

“the radius is broad and flat’’ and agrees

with that of Moschops but is more massive.

In 1929 Broom (14) described a radius of

Jonkeria sp. and figured a right radius of

“probably Titanosuchus ferox’’. In 1931

von Huene (21) gave a figure of a

Deinocephalian radius. Byrne (16) figured

the bone in Moschoides and I (4) figured the

radius of a Moschopid in 1954.

A Titanosuchian radius and

ulna in dorsal view. B.F.—

flange for the insertion of

the biceps; C.P.—coronoid

process; in.—facet for inter-

medium; L.S.F.—lip to sig-

moid fossa; R.—radius;

ta.—facet for radiale; Sty.—

styloid flange; U.—ulna;

Ue.—facet for the ulnare.

In the larger Titanosuchians the radius is

a large stout bone, but in some of the smaller

forms it is relatively slender (length 190-318

mm.). The two ends are expanded, especially

the proximal one in the larger forms. Both

the ends and the shaft are oval in

section. The proximal end is flat or

194 ANNALS OF THE SOUTH AFRICAN MUSEUM

‘becomes convex towards the postaxial border, which abuts against the rim

‘of the sigmoid facet of the ulna (but in Keratocephalus the whole proximal

face is convex). On its postaxial corner there is a weak to strong flange

(bicipital), whose edge fits into the hollowed surface of the ulna below the

rim of the sigmoid facet. A similar flange is developed on the distal postaxial

surface, where the radius fits against the ulna. The pre- and postaxial faces

are deeply concave in the Titanosuchia, but the shaft is little constricted

in Keratocephalus and some Moschopids. The preaxial edge is fairly sharp,

whereas the postaxial edge is rounded. The distal end is fairly flat or quite

strongly convex, and in some Titanosuchia shows a well developed facet for

the intermedium and a lesser one for the radiale.

ULNA

In 1876 Owen (23) noticed as Paretasaurus an ulna, which Seeley figured

and, as Lydekker (22), considering it to be of Tapinocephalus, remarks

“referred it in the abstract to Titanosuchus’’.’ This is undoubtedly a

Titanosuchian ulna notwithstanding that Broom (14) also considered it to

be of Tapinocephalus. The supposedly Titanosuchus ulna figured by Seeley

(28) is really a tibia of a Pareiasaurian. The Moschops ulna was figured

by Broom (11) and refigured by Gregory (18), and in Broom’s (10) photo

of the ‘“‘Tapinocephalus’’ fore-limb a large Titanosuchian ulna is shown. In

1914 Watson (29) mentioned the ulna of Phocosaurus and Broom (13)

mentions an ulna of Tapinocephalus and figured (14) part of an ulna of

Jonkeria crassus and describes the ulna of Jonkena sp. In 1931 von Huene

(21) figured an imperfect ulna of a Deinocephalian. Byrne (16) figured

the bone of Moschoides and I (4) figured a Moschopid ulna in 1954.

In our collection there are 15 ulnae, some without the distal end. In no

case is there a pair and in only two cases have we a radius associated with

an ulna.

The Deinocephalian ulna is fairly to very massive and short (length

258-402 mm.). Its proximal end is greatly expanded; the shaft is broad but

flattened; the postaxial face is fairly straight, but the preaxial face is deeply

concave; the distal end is only slightly more expanded than the shaft.

Proximally the lateral corner is developed into a strong olecranon which is

rugosely striated for the reception of the strong triceps muscle. Medially

lies the large sigmoid face for articulation with the humerus; this surface is

only shallowly concave longitudinally and it is continued on to the strong

coronoid process; dorsally to the coronoid process the sigmoid face ends, in

its medial part, in a concave rim, to which the outer edge of the head of

the radius is applied, so that the sigmoid face is continuous with the proximal

face of the radius to form a conjoined articulatory facet for the capitellum

GIRDLES AND LIMBS OF DEINOCEPHALIA IQ5

of the humerus. The sigmoid face has a more or less distinct ridge running

proximo-distally so that two faces, inclined at an obtuse angle to each other,

can be distinguished; the dorsal part articulates with the humerus when

the limb is directed forwards, and the ventral part comes in play when the

epipodium is rotated to the posterior position. The dorsal lip of the sigmoid

face forms a more or less prominent rim, which is continued medially by

the sharp edge of the head of the ulna. Lateral to this rim the dorsal surface

of the ulna bears a longitudinal groove fading away distally on the shaft;

on the ventral face there is a similar though deeper groove. The distal end

is indistinctly divided into three facets; the central and larger facet is for

the ulnare, the preaxial facet for the intermedium and the postaxial one for

the weak pisiforme. Proximally of this facet for the pisiforme the postaxial

edge of the ulna develops a sharp ridge, which apparently has the same

function as a styloid process.

THE ELBow JOINT

For all major movements the two bones of the epipodial move as a unit

and the articulatory face formed by the ulna and radius is in the shape

of a fairly large arc whose surface moves over the distal end of the humerus

in flexion and extension. When the radius is rotated on its long axis the

ulnar facet slides backwards or forwards against the distal face of the humerus.

Extension is limited by the olecranon, for which little provision is made in

the form of a shallow trochlear fossa.

MANUS

In the Deinocephalia the manus is insufficiently known. In the Moschopids

the carpal formula is 3, 2, 5 Byrne (16) or 4, 1, 5 Boonstra (4) and the

phalangeal formula 2, 3, 3, 3, 3 Byrne (16). In the Anteosauria I (4)

recorded, in one specimen, a carpal formula 3, I, 5 and the most unusual

phalangeal formula 3, 3, 4, 4?, 2.

In the structure of the fore-limb the Deinocephalians show certain important

advances beyond the Pelycosaur condition: in the humerus the proximal

face has lost the elongated straplike articulatory face and the articulation

is concentrated in a medially situated caput; the twist of the proximal and

distal ends relative to each other is strongly reduced; the epicondyles are

much reduced in size and there is never a separate supinator crest; the loss

of the ectepicondylar foramen in all but the Titanosuchia; the distal position

of the condyles makes a more upright disposition of the epipodial possible;

the sigmoid face of the ulna forming a more widely open arc makes a greater

196 ANNALS OF THE SOUTH AFRICAN MUSEUM

extension of the epipodial possible; the greatly laterally expanded delto-

pectoral crest makes the area for the origin of the m. brachialis much greater.

C. The Pelvic Girdle (Diagram 4)

Owen (23), in 1876, was the first to examine elements of the pelvic

girdle of a South African Deinocephalian. But, due to the confused labelling

of the materials sent to the British Museum (N.H.), he mistakenly considered

these to be Pareiasaurian, and also erroneously identified an ischium and

pubis (B.M. 43525r) as parts of the pectoral girdle and humerus.

These pelvic elements were re-examined by Seeley (28) and made the

type of a new genus, Phocosaurus. Lydekker (22) provisionally included this

specimen in the genus Tapinocephalus, and Watson (29) refigured the pelvis.

In 1914 Broom (11) figured and described the pelvis of Moschops. This

Moschops-material was re-examined by Gregory and Camp (17), Romer (25)

and Gregory (18) and their restorations of the pelvis differ considerably from

that of Broom.

Meanwhile Haughton (20) had published a photo of the mounted skeleton

of Struthiocephalus showing the pelvis and on this Broom (15) based his

reconstruction.

Broom (13) in 1928 figured an ilium and ischium under the name

Tapinocephalus atherstonei, whereas they are in fact of a Titanosuchian.

In the following year Broom (14) described an ilium of Jonkeria sp. and

another under the name Dinosphageus haughtom,; also an ischium under

the name Phoneosuchus angusticeps. Both these have since been included in

the genus Jonkena.

In 1931 von Huene (21) figured and described the ilium of Keratocephalus

and Broili and Schréder (6) an incomplete ilium and pubis of Titanognathus

lotzt, which has since been included in the genus Anteosaurus. Finally

Byrne (16) gave a brief account of the imperfect pelvis of Moschoides.

In our collection the pelvic girdle is represented in 18 specimens. In only

six cases parts of the pectoral girdle are associated with parts of the pelvic

girdle of the same specimen.

The pelvis is much lower than the pectoral girdle and the antero-posterior

length of the pubo-ischiadic plate is also less than that of the coracoidal plate.

The three bones of each half are mostly not firmly united at the sutures

and in immature specimens the centre of the acetabulum may still be

cartilaginous.

GIRDLES AND LIMBS OF DEINOCEPHALIA 197

The two halves, meeting in the median line, are weakly united, and in

immature specimens a large lacuna is present, which may persist as a small

opening even in older specimens; but in life would be closed with cartilage.

The junction of the pubes, parti-

Diag. 4. cularly, is very weak, due to the

eversion of the antero-ventral pubic

Glu. edge. The ichial symphysis is

stronger, but here the posterior median

edges also diverge and no strong sym-

physial carina is developed.

The attachment to the vertebral

column is by a pair of very stout main

sacral ribs, assisted by three posterior

ribs, which are very much weaker and

decrease in functional importance in

posterior direction, and by a long

slender anterior lumbo-sacral rb

usually with a feeble attachment to the

everted anterior iliac process. The main

sacral rib lies above and in the same

plane as the acetabulum — it is

thus not pre-acetabular but supra-

The pelvis of a Titanosuchian in bul

lateral view. Ac.—acetabulum; aeChipia®,

A.Pr.—anterior iliac process, more The acetabulum is large and

or less everted (lower in most eal ; heel eihs

Tapinocephalia); Glu.—area of roughly circular in outline; a ree

origin of the ilio-femoralis (gluteus);

I.F.R.—ilio-fibularis ridge (this lies

more horizontally in most Tapino-

cephalia); I1.—ilium (lower in most

Tapinocephalia); Is.—ischium (re-

stored); P.—pubis; P.F.—pubic

foramen; P.Pr.—posterior iliac

process (lower and lying more

horizontally in most Tapino-

cephalia); P.T.—tuberculum pubis;

S.A.B.—supra-acetabular buttress;

S.A.N.—supra-acetabular notch.

bones help in its formation, with the

iliac part constituting a half to two

thirds. The acetabulum faces mainly

outwards except in the Anteosauria

where it is directed considerably ven-

trally. It is fairly shallow but with

strong raised rims for the attachment

of the joint capsule and ligaments.

Dorsally a very strong buttress,

stronger in the Titanosuchia than in the Tapinocephalia, overhangs the

acetabulum (except in the Anteosauria), and this receives the main thrust

of the femur. In its dorso-posterior corner, just posterior to the edge of

the buttress, les the large supra-acetabular notch.

The ilium is constricted above its acetabular part to form a neck or shaft;

above this the blade is antero-posteriorly elongated, with a fairly long anterior

iliac process and a shorter and weaker posterior process. In the Titanosuchia

the supra-acetabular part of the ilium is high, whereas in the Tapinocephalia

it is low to moderate. This also applies to both the posterior and the anterior.

198 ANNALS OF THE SOUTH AFRICAN MUSEUM

processes. The anterior process is everted, weakly in most Tapinocephalia

and strongly in most Titanosuchia. The posterior process extends further

posteriorly in the Tapinocephalia than in the Titanosuchia, and in the

former lies more horizontally and in the latter more upright. On the

posterior edge of the posterior process an everted or folded-over ridge is

developed, presumably for the origin of the ilio-fibularis muscle. This ridge

lies horizontally in nearly all the Tapinocephalia, where it forms a sharp

to rounded postero-ventral edge, whereas it lies much vertically in the

Titanosuchia, where it forms a rounded to swollen thickening anterior to

the posterior edge of the ilium.

The upper iliac edge overhangs the area of origin of the m. ilio-femoralis

(gluteus) in some Tapinocephalia. The gluteal area is shallow in the

Tapinocephalia, but in some Titanosuchians it is deeply concave antero-

posteriorly.

In dorsal view the outline of the upper iliac surface is roughly triangular.

The apex, lying medially, is at a lower level than the outer iliac edge, which

is fairly straight in the Tapinocephalia, but concave in the Titanosuchia.

The apex represents a great thickening on the middle part of the inner

surface of the ilium. The massive main sacral rib abuts against this thickening

— more particularly to its anterior face. This sacral rib extends ventrally

along the inner iliac face to a plane just dorsal to that in which the outer

buttress lies. Posterior to the attachment of the main sacral rib there are

three irregular depressions for the reception of the three weak posterior ribs,

whose upper surfaces lie in the same plane as the upper surface of the main

rib, but as they are weak the ventral extent of their contact with the ilium

is small. On the inner face of the anterior iliac process there is an ill-defined

area for the reception of the end of the lumbo-sacral rib. The inner face

of the ilium, dorsal to the sacral ribs, afforded an area for the attachment

of the axial muscles.

The ischium is only adequately known in the Tapinocephalia. Its antero-

posterior length is fairly small. Except for its acetabular portion it is fairly

lightly built. It forms the postero-ventral segment of the stout acetabular

rim. From its junction with the ilium its postero-dorsal edge is thick but

posteriorly rapidly decreases to a thin edge, without the development of an

ischial tuberosity. Its ventral edge is curved so that posteriorly it is separated

from its fellow by a triangular incisure. Similarly the anterior ends also

do not meet, but here the opening between the ischia and pubes was closed

by cartilage. The central part forms a fairly strong symphysial face. The

ischia meet at an angle to form a V-shaped pelvic exit, and no carina is

formed along the median line.

GIRDLES AND LIMBS OF DEINOCEPHALIA 199

The pubis has its anterior corner strongly everted. From this corner to

the acetabulum the upper border of the pubis is thickened to form a stout

pubic ridge from which the pubo-tibialis muscle originated. The corner or

the tuberculum pubis, for the insertion of Poupart’s ligament, is strong.

Below the pubic ridge there lies the oval pubic foramen. Owing to its

eversion only the posterior part of the ventro-median pubic edge meets its

fellow to form a very weak line of contact, and the antero-posterior length

of the pubis is small. On the inner face it shows a quite distinct area from

which the pubo-ischio-femoralis internus originated.

In the pelvis the Deinocephalia have advanced beyond the Pelycosaurian

stage in that the pubo-ischiadic plate has been greatly reduced in length;

the strong development of the anterior iliac process and the reduction of the

posterior iliac process; in the development of a large circular acetabulum;

and increased height and a greater width to the pelvic outlet.

It most obviously differs from the higher Therapsids in the retention

of the simple pubic foramen and in the shortness of its pubo-ischiadic plate.

D. The Hind-limb (Diagrams 5 and 6)

THE FEMUR

The first Deinocephalian femur from the Karroo to be described was a

proximal half from Varsfontein (B.M. 43525t) first noticed by Owen (23)

in 1876 under the name Paretasaurus bombidens and figured by him (24) in

1880. Lydekker (22) referred this specimen to Tapinocephalus, and it will

here be included in Phocosaurus.

Seeley (28) in 1889 figured (Pl. 19) a femur from Koedoeskop (B.M.

49368), which he considered to be associated, correctly I believe, with the

type of Owen’s Titanosuchus ferox. Broom (8) in his description of Pelosuchus

mentions a femur, which he (15) figured in 1932. In 1914 Broom (11) figured

a femur associated with the type material of Moschops, and figures of femora

from this material were subsequently published by Gregory and Camp (17),

Romer (25), Gregory (18) and Broom (15).

The femur of Pnigalion was figured by Watson (29) in 1914, and in the

following year Broom (12) mentions that femora are associated with the

type of jaws of Moschognathus and these were figured by Gregory (18). The

femur of Struthiocephalus is shown in Haughton’s (20) photo of the mounted

skeleton. In 1928 Broom (13) figured a femur he thought to be of Tapino-

cephalus, but which in the sequel is shown to be of Jonkeria, and in the next

publication (14) a femur of Jonkeria sp. In 1931 von Huene (21) figured

the femur of Keratocephalus and Broili and Schréder (6) figured a distal end

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

of the Anteosaurian, Titanognathus. Byrne (16) gave figures of the femur

of Moschoides and I (4) of Micranteosaurus.

Diag. 5.

The femur of a Titanosuchid. a. Dorsal view. b. Ventral

view. Cap.—caput femoris; Ect.C.—ectocondyle; Ent.C.—

entocondyle; Ex.Tr.—external trochanter; F.T.—origin of

the femoro-tibialis; Fib.—facet for articulation of fibula;

Gast.—origin of gastrocnemius; Intc.F.—shallow inter-

condylar fossa; Int.F.—intertrochanteric fossa; Int.S.—

intercondylar sulcus; Int.Tr.—internal trochanter; Is.Tr.—

insertion of ischio-trochantericus; Pop.—popliteal fossa.

In the collection of the South African Museum there are 37 femora of

which some are only represented by one of the two ends. Although there

are about an equal number of femora and humeri it is noteworthy that

only in 8 specimens there are preserved both a humerus and a femur of

the same individual.

The femur of the Anteosauria differs so markedly from that of the

Tapinocephalia and the Titanosuchia that in this general account of the

Deinocephalian femur it is not considered, but is treated of separately in the

systematic part of this paper.

The femur, always longer than the humerus, is a massive bone with

very little of a shaft but with expanded ends. The proximal end is always

wider than the distal end. There is a very little ‘‘twist’’ on the shaft so

that the two ends lie in nearly the same plane, but the distal pre-axial condyle

does lie a little more ventrally than the distal post-axial condyle. The post-

axial condyle also lies a little further distally (where it lies much distally

this appears to be due to postmortem deformation or is pathological). The

shaft is always much broader than thick. In dorsal view the pre-axial border

is more concave than the post-axial border. In pre-axial view the ventral

border is more concave than the dorsal and the long axis is straight.

GIRDLES AND LIMBS OF DEINOCEPHALIA 201

Proximally the caput femoris is terminal, but somewhat preaxially

directed; it is antero-posteriorly elongated, thick preaxially, where it has an

abrupt edge; but postaxially it tapers more or less gently towards the external

trochanter from which it is sometimes separated by a slight notch; in its

preaxial part the caput is moderately convex. The caput is much smaller

than the acetabulum — its dorso-ventral diameter being about a third to

a quarter of that of the acetabulum.

The external trochanter lies far postaxially to form the proximo-postaxial

corner of the femur; from here the postaxial edge of the bone sweeps inwards

with a fairly straight edge to the shaft and then more or less sharply outwards

again to form the outer border of the ectocondyle. On the dorsal surface

just preaxial to the external trochanter some femora have a slight pit, where

the ischio-trochantericus muscle was presumably inserted.

_ The internal trochanter (sometimes not evident) is a rounded or oval

tubercle, sometimes very prominent, lying on the ventral surface, well away

from the preaxial border so that the intertrochanteric fossa lies well postaxially.

The distal articulatory facets for the tibia lie far distally and face only

little ventrally; the outer facet is slightly larger than the inner and lies a

little further distally. The two facets are separated by a deep and wide

intercondylar sulcus, which proximally has a sharp mndge separating it from

the popliteal fossa, and distally passes over into a shallow intercondylar

notch; a rounded depression in the proximal part of the intercondylar sulcus

presumably housed the quadriceps tendon.

The dorsal surface of the femur shows few prominent features; the

intercondylar fossa is very shallow; from the outer condyle an oblique ridge

runs in the direction of the caput femoris; between this ridge and the

external trochanter the ilio-femoralis muscle has a large area of insertion;

the femoro-tibialis originated from the middle part of this mdge.

On the postaxial face of the ectocondyle lies the weakly modelled surface

to which the bipartite head of the fibula articulated.

The Deinocephalian femur has advanced beyond the Pelycosaur stage

by the breaking up of the Y system of ridges; lack of a 4th trochanter;

decreased ‘“‘twist’’ on the shaft; the distal position of the condylar facets,

but with the ectocondyle nearly in the same plane as the entocondyle and

with a larger articulatory face. The widening of the whole bone and the

flattening of the shaft is apparently a secondary approach to the Pareiasaur

condition. It differs from the higher Therapsids in the great width and

the lack of a long distinct shaft and the caput is not turned antero-dorsally.

THE HIp-JOINT

The Deinocephalian hip-joint is a very ill-fitting joint. The elongated

caput femoris requires for its rotation on its long axis the very large circular

202 ANNALS OF THE SOUTH AFRICAN MUSEUM

acetabulum in which it fits very insecurely so that the strong iliac buttress is

required to prevent its dislocation when the thrust is transmitted. The

femur in transmitting the thrust would le obliquely with its distal end

considerably lower than the caput and it would have great, but not very

secure, possibility for antero-posterior movement. A very strong capsule

and ligaments would be required to strengthen the joint.

THE TIBIA

Owen (23) mentions a tibia (B.M. R.1707) referred by him to Pareiasaurus

bombidens. This tibia was figured by Seeley (28) who said “it may be

new or it may be Parevasaurus or Tapinocephalus’’. Considered by Lydekker

(22) as Tapinocephalus, it probably really is Titanosuchus. Two other tibiae

(one 47100) are mentioned by Lydekker as probably Titanosuchus or

allied form.

The tibia of Moschops was figured by

Broom (11) and refigured by Gregory (18).

In 1914 Watson (29) mentions a tibia as part

of the type material of Phocosaurus and

stated that the tibia of Titanosuchus ‘‘does

not differ essentially from that of the

Tapinocephaloids’’. In Haughton’s photo-

graph (20) a tibia of Struthiocephalus can

be seen.

Diag. 6.

In 1928 Broom (13) figured a tibia,

which I believe to be Jonkeri, as that of

Tapinocephalus atherstonei, and, in_ the

following year, he (14) figured the tibia of

fibula in dorsal view.

Cn.B.—cnemial boss; Cn.G. Jonkeria truculenta and noticed that of

—cnemial groove; Cn.R.— : :

coal anal Here eat Jonkeria sp. In 1931 von Huene (21) figured

for the outer femoral con- the tibia of Keratocephalus and Byrne (16)

dyle; Ent.C.—facet for the

inner femoral condyle; Fem.

—proximal facet on fibula

| Besa GPR 5 iN ER CRI In the collection of the South African

surface of the ectocondyle; Museum the tibia is represented by I7 more

fi.facet for the fibulare; :

FM UN eer eel LN Sa or less complete bones. It is remarkable that

medium. so many more tibiae than radii should be pre-

served. Only in one case both right and left

tibia of the same animal are present. In seven cases we have a tibia with

a fibula in the same specimen.

in 1940 that of Moschovdes.

GIRDLES AND LIMBS OF DEINOCEPHALIA 203

The tibia is in size medium to large and fairly to very massive with

strongly expanded ends and a robust shaft. The length varies from 180-355

mm. On its dorsal surface there is a very strong cnemial protuberance for

the insertion of the tendon of the femoro-tibialis and associated muscles.

This boss extends distally as a rounded ridge, which fades out on the shaft

before reaching the middle of the bone. Postaxially of the cnemial ridge lies

a fairly shallow groove bounded further postaxially by a weak ridge lying

parallel to, but is much weaker than the cnemial ridge. The proximal

expansion flares out preaxially as does also the distal expansion so that the

preaxial border is deeply concave. The proximal expansion flares out less

postaxially, but the distal expansion flares out abruptly postaxially.

The proximal surface is essentially triangular in outline with the apex

formed by the massive cnemial boss. Above the cnemial eminence the

surface is convex, but this flattens out towards the ventral corners, with

the formation of a low indistinct ridge separating these two flattened faces.

The preaxial face is greater than the postaxial one and they correspond

to the two condylar faces of the femur. In a tibia of Struthocephalellus

a high ridge separates two concave faces.

The distal surface is strongly convex for the two-thirds of the surface

lying preaxially; the postaxial part is flattened and tends to be concave.

The ventral surface is separated from the postaxial face by a strong ridge

and is furthermore indistinctly divided into two parts by a ridge running

obliquely, with its distal end lying preaxially; the upper part receives the

long ventral muscles and the lower the short ventral muscles.

THE FIBULA

The fibula has hitherto been described by Seeley (28) in Titanosuchus

and this identification is probably correct, although Broom (14) does not

think the bone is a fibula at all; in Moschops by Broom (11) and refigured

by Gregory (18); seen in Haughton’s (20) photograph of Struthiocephalus;

in errore by Broom (13) for Tapinocephalus and (14) in Jonkeria truculenta

and Jonkena sp. and finally by Byrne (16) in Moschoides.

In the collection described here there are 16 fibulae, of which only a

couple are not very well preserved.

Although the weakest of the major limb-bones the Deinocephalian fibula

is still a stout bone (length 220-345 mm.). The proximal expansion is

great and moderately to very massive; the distal end is somewhat less

expanded, but still quite strong; the postaxial edge is rounded and is

in outline fairly straight, whereas the preaxial border is deeply concave

and this edge is sharp up to near the distal end where it becomes rounded,

with a sharp ridge lying more ventrally; the shaft is long, fairly wide but

204 ANNALS OF THE SOUTH AFRICAN MUSEUM

flattened, and there is very little ‘‘twist’’ on the shaft, but the distal end

is bent down preaxially so that the two ends do not lie in the same plane.

The proximal facet is convex in section and occupies the whole of the

proximal end; although terminal it faces ventrally; in outline it is reniform

with the hilus due to a hollowing out of the ventral edge; a thickening

on the medio-dorsal surface near the head was for the insertion of the

ilio-fibularis muscle.

The distal facet is broad but flattened, convex, thicker preaxially and

facing more preaxially; the tilted preaxial part articulating with the inter-

medium and the postaxial part with the fibulare.

THE KNEE JOINT

The thrust of the hind foot is along the long axis of the tibia and is

transmitted to the two femoral facets meeting the proximal end of the

tibia in an efficient joint. As the articulatory surfaces on the femur lie

much distally and curve round dorsally the lower leg can be greatly

extended to bring the long axes of the tibia and femur nearly in line.

Rotation of the tibia on its long axis is well allowed for with the fibula

sliding over the postaxial corner cf the femur and distally carrying the foot

with it. In the Titanosuchia the amount of the extension possible is thus

much greater in the hind-epipodial than in the fore-epipodial.

THE PES

The Deinocephalian pes is but poorly known; in the Titanosuchia the

proximal row of tarsals consists of an intermedium and fibulare; in the

Moschopids (16) the tarsal formula is 2, 1, 4 and the phalangeal formula

apparently 2, 3, 3, 3, 3; in a single Anteosaurian I (4) recorded a tarsal

formula’ 2,1, 5 and ‘a phalangeal formula 2? ;°37)>3" 7 ary

SYSTEMATIC DESCRIPTIONS

Deinocephaha

Sub-ordinal Characters of the Girdles and Limbs.

The pectoral girdle is structurally Therapsid, but comparatively large

and massive and without an acromion; the cleithrum is splintlike or well

developed; the interclavicle with its anterior spatulate end turned upwards;

the three scapulo-coracoid elements not firmly ankylosed; there is no

ossified sternum.

a a =

GIRDLES AND LIMBS OF DEINOCEPHALIA 205

The humerus is structurally Therapsid but comparatively massive and

with either one or two epicondylar foramina; the radial facet is either much

distally or much ventrally situated.

The ulna has a massive olecranon.

The pelvic girdle is structurally Therapsid but large and massive; the

pubo-ischiadic plate is short; the acetabulum large; there is only a simple

pubic foramen; in most groups there is developed a distinctive ridge on

the posterior edge of the posterior iliac process for the origin of the m. ilio-

fibularis; the anterior pubic edge is everted; the symphisis is weak.

The femur, of typically Therapsid structure, is either short and massive

or long and slender.

Tapinocephaha

Infra-ordinal Characters of the Girdles and Limbs.

The pectoral girdle is fairly light to moderately massive; the scapula

is fairly low to low (310-530 mm.) with the upper part of its blade fairly

narrow to fairly broad (120-270 mm.); the scapular head of the triceps

originates from a low mound; the internal opening of the foramen supra-

coracoideus does not open into or is connected by a groove to the subscapular

groove; the glenoidal facet of the scapula is small or of medium size and

faces little or not at all externally. —

The coracoidal plate is fairly short or long antero-posteriorly; the pre-

coracoid is fairly large, but fairly weak with a thin anterior border and a

fairly strong dorso-posterior apex; the precoracoid excluded from the glenoid

or just enters the rim; its outer face is only moderately convex.

The coracoid is small to medium, light to fairly strong with a fairly large

glenoidal facet facing dorso-posteriorly and also much externally. Except

near the glenoid the three bones of the scapulo-coracoid are weakly united

suturally.

The clavicle is a medium-sized flattened bone with expanded ends and a

slight waist.

The cleithrum is a weak splintlike bone.

The interclavicle is a moderately light bone, moderately expanded

anteriorly with a long slender to fairly slender stem. A shallow groove

receives the lower end of the clavicle.

The humerus is fairly light to moderately massive; it varies in size from

small to large (length 276-588 mm.), but is never short and squat, and some-

times long and fairly slender; there is never an ectepicondylar foramen;

the capitellum is weak to moderately strong and usually poorly modelled

and it never extends proximally along the ventral face but remains situated

206 ANNALS OF THE SOUTH AFRICAN MUSEUM

much distally; the processus medialis always lies far proximally, nearly

in the same plane as the caput; the processus lateralis always les far

proximally; the proximal expansion is moderately great to great; the epi-

condyles are moderately to greatly expanded; the shaft is short to long,

narrow to very broad; the ventral opening of the entepicondylar foramen

is always slitlike.

The ulna is small and light to moderate and fairly strong, or is short,

massive and squat.

The radius is long and slender or fairly short with strongly expanded ends.

In the manus the carpal formula is 4, I-2, 5, and the phalangeal

formula, 18.2, 2). 2818.2

The pelvis has a fairly long pubo-ischiadic plate; the iliac blade is low

to only moderately high; the ilio-fibularis ridge has a fairly sharp edge and

is only moderately strong and usually lies nearly horizontally; the anterior

iliac process is only slightly to moderately everted; the posterior iliac process

is fairly long and low; the outer iliac face is only slightly concave in antero-

posterior direction. |

The femur varies from fairly small to fairly large (length 276-468 mm.),

light, moderate to fairly robust, sometimes short and squat; the internal

trochanter undeveloped or forming and elongated tubercle situated well

away from the preaxial border; the width over the external trochanter is

small (95 mm.) to great (256 mm.), and without a notch.

The tibia is massive and short (232 mm. in length) or slender or

moderate (length 180-270 mm.).

The fibula is light and slender (length 222-288 mm.).

In the pes the tarsal formula is 2, 1, 4, and the phalangeal formula

2, 3, 3, 3) 3:

Tapinocephahdae

Family Characters of the Girdles and Limbs.

The pectoral girdle is only moderately massive; the scapula is low, with

its posterior edge fairly straight.

The humerus is fairly massive but fairly long.

The ulna is moderate and fairly long or it is massive and squat.

The iliac blade is low with both the anterior and posterior processes

long and low; the ilio-fibularis ridge is strong and prominent.

The femur is either fairly massive and fairly long or massive and squat.

The above statement on the characters of the girdles and limbs of the

Tapinocephalidae is based on very inadequate material, as will become

evident below.

GIRDLES AND LIMBS OF DEINOCEPHALIA 207

Genus Tapinocephalus Owen (Figs. 1 a-b and 2)

In the collection of the South African Museum there is only one specimen

in which there is an identifiable skull associated with some parts of the

postcranial skeleton. This skull (S.A.M. 2344) is associated with two

humeri and a scapula (S.A.M. 3355).

Scapulae. a. Tapinocephalus atherstonei, S.A.M. 3355. Lateral view. (x 3.)

b. Tapinocephalus atherstonei, S.A.M. 3355. Posterior view. (x %.) c. Keratocephalus

moloch, S.A.M. 11937. Lateral view. (x %.)

Note: All the figures in this paper are orthoprojections and not perspective drawings.

In the girdles the figures labelled ‘‘lateral view’’ are of the bones in natural position

projected on to the median plane, “‘anterior’’ and ‘‘posterior’’ are on to the one

‘

plane at right angles to the median plane and ‘‘dorsal’’ and “‘ventral’’ on to the

other plane at right angles to the median plane.

The scapula (Fig. I a-b) is relatively low (530 mm.), with a broad

blade (as reconstructed the width in its upper part is 270 mm.); the tricipital

ANNALS OF THE SOUTH AFRICAN MUSEUM

208

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GIRDLES AND LIMBS OF DEINOCEPHALIA 209

ridge forms a strong but low mound; the internal opening of the foramen

supracoracoideus does not open into the subscapular groove, which is very

shallow indeed; the glenoidal facet is small and faces postero-ventrally

and but very little externally; the upper part of the blade is fairly straight

with little curvature to fit round the thorax.

The humerus (Fig. 2). Associated with the skull from Uitkyk (S.A.M.

2344), described by Haughton and correctly referred to Tapinocephalus, there

is a fairly complete left humerus and a weathered proximal half of the

right humerus (S.A.M. 3355). This association enables us to clear up the

confusion which exists in regard to the Tapinocephalus humerus. The story

of this confusion is as follows: In the first instance Owen (23) mistakenly

referred the humerus B.M. 43525p from Varsfontein, Prince Albert, to

Pareiasaurus. Lydekker (22), in correcting this obviously mistaken identi-

fication, himself erroneously referred it to Tapinocephalus. Broom (12),

accepting Lydekker’s identification, subsequently referred a specimen which

he sold to the American Museum (A.M.N.H. 5611) to Tapinocephalus.

Later Broom (13) figured a humerus (collection not stated) and states that

this represents the true humerus of Tapinocephalus and at the same time

refers B.M. 43525 to Titanosuchus. To make the confusion more confounded

Broom then contradicts Seeley’s (28) identification of B.M. 49369 as the

humerus of Titanosuchus and refers this humerus to Tapinocephalus.

The Uitkyk humerus (S.A.M. 3355) associated with a Tapinocephalus

skull, has no ectepicondylar foramen. In this feature it agrees with the

known humeri of all the other members of the infra-order Tapinocephalia.

Whereas all the humeri known to be associated with skulls classified as

belonging to the infra-order Titanosuchia do have a well-developed ect-

epicondylar foramen. Thus quite apart from other features still to be

considered below, the humeri A.M.N.H. 5611, B.M. 49369 and Broom

1928 (no collection or number given) must all belong to some genus of

the Titanosuchia. B.M. 49369 should be left where Seeley put it, viz. in

the genus Jitanosuchus and A.M.N.H. 5611 in Jonkeria and Broom’s 1928

humerus in Scapanodon and B.M. 43525 is the humerus of Phocosaurus.

The humerus of Tapfinocephalus may be described as follows: very large

(length 520-590 mm.); not very massive; proximal expansion great (width

276 mm.); distal expansion moderate (width 246-282 mm.); shaft fairly

long and moderately thick (diams. 105 xX 90 mm.); the delto-pectoral crest

long, strong, terminating well proximal of the ventral opening of the

entepicondylar foramen; caput oval in outline and lying in the same plane

as the processus medialis and the processus lateralis; the processus medialis

thus lies far proximally; the capitellum (radial condyle) is not well modelled,

is weak, and lies much distally with its ventral part lying well distal of the

plane in which the entepicondylar foramen opens; twist on shaft fairly

210 ANNALS OF THE SOUTH AFRICAN MUSEUM

great (35°—~40°); the lateral median line is weak and the antero-dorsal

ventral line is fairly weak; the entepicondyle is not greatly expanded and

the foramen enters ventro-postaxially and leaves in a narrow ventral slit;

the ectepicondyle is moderately expanded to form a thin unperforated.

curved plate with a sharp edge.

No other bones of the girdles or limbs in Tapinocephalus are known

with any certainty, but below it will be seen that certain elements could

probably be those of Tapinocephalus.

Tapinocephalus atherstoner Owen

The specific description is as for the genus.

Referred specimens in the S.A.M. collection:

S.A.M. 3355. An imperfect left.scapula (Fig: 1 \.a-b)jaay game. leit

humerus (Fig. 2 a-d) and the weathered proximal end of the right

humerus, associated with a good skull. Uitkyk, Beaufort West.

Low Tapinocephalus zone. Coll. Haughton. |

S.A.M. 5007. An isolated good left humerus (Fig. 2e). Wilgerfontein,

Prince Albert. Low Tapinocephalus zone. Coll. Haughton.

Genus Phocosaurus Seeley

This description is based mainly on the Varsfontein material in the

British Museum (B.M. 43525), incorporating the accounts of Seeley (27),

Lydekker (22) and Watson (20).

The pectoral girdle is large and massive; the scapula probably fairly

low (lower than reconstructed by Watson, Fig. 11, which is given x 4,

whereas it is in fact x 4); the scapular blade is fairly broad (240 mm.); the

tricipital ridge is low; the glenoidal facet faces well externally. The

precoracoid is inadequately known; ‘‘foramen opens into a distinct pit

on the visceral surface’ (Watson). The coracoid is massive with the facet

facing much externally.

The humerus (Fig. 3) is large and massive (length 492 mm.); proximal

(276 mm.) and distal (306 mm.) expansions large; shaft short and wide

(diams. 168 X 96 mm.); the delto-pectoral crest very long and nearly

reaching the plane of the ventral opening of the entepicondylar foramen;

the caput is widely oval and lies in the same plane as the processus medialis

and the processus lateralis, which also lies far proximally; the radial condyle

is strong, fairly thick and well modelled, but situated much distally and not

extending much proximally along the ventral face and thus well distal of

the plane in which the entepicondylar foramen opens; the twist on the

2i1

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212 ANNALS OF THE SOUTH AFRICAN MUSEUM

shaft moderate (25°); the L.M.L. is fairly distinct and the A.D.V.L. is

not prominent; the entepicondyle is strongly developed as a thick plate of

bone and the ventral opening of the foramen is slitlike and situated well

away from the edge of the bone; the ectepicondyle is developed as a greatly

flaring thin sheet of curved bone

The ulna (see Seeley’s Pl. 22) is short (320 mm.); the dorsal lip to the

sigmoid face is fairly strong; the styloid ridge is prominent; width over

coronoid process is moderate (204 mm.).

The pelvis (see Seeley, Pl. 21, Lydekker, Fig. 17 and Watson, Fig. a

The pubo-ischiadic plate is short (as reconstructed + 80% of the total

height of the pelvis); the ilium appears to have been fairly low in its supra-

acetabular part and fairly long antero-posteriorly; the ilio-fibularis ridge on

the posterior iliac process lies fairly horizontally and appears to have been

thickly rounded; the antero-ventral edge of the pubis is strongly everted

with the tuberculum pubis confluent with the thickened antero-ventral edge

curving in towards the median line.

The femur (see Owen (24), Pl. 17, Fig. 8) is only represented by a

proximal half. The proximal expansion is fairly broad (225 mm.); the

preaxial face is fairly deeply concave with the caput directed well pre-

axially; caput fairly massive; external trochanter not distinctly separated

from the proximal face; the internal trochanter lies well in and is developed

as a prominent oval tubercle; the shaft is fairly slender (diams. 112 xX

72, mm.).

Phocosaurus megischion Seeley

The specific description is as for the genus.

Type: B.M. 43525. Incomplete coraco-scapulae, humeri, mght ulna,

incomplete pelvis and the proximal half of the mght femur. Vars-

fontein, Prince Albert. Middle Tapinocephalus zone. Coll.

Atherstone.

Referred specimens in the S.A.M. collection:

S.A.M. 11300. (Figs. 4 and 5.) A right humerus, showing a pathological

lesion and an incomplete ilium. Deesweesfontein, Laingsburg. Low

Tapinocephalus zone. Coll. Boonstra.

S.A.M. 11988. Proximal half of an isolated left femur. Boeteka.

Beaufort West. High Tapinocephalus zone. Coll. Boonstra.

Genus Keratocephalus v. Huene

Although there are four skulls of Keratocephalus in the collection of the

South African Museum there are only a few bones of the girdles and limbs,

213

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214 ANNALS OF THE SOUTH AFRICAN MUSEUM

but, together with von Huene’s account of some bones in Tiibingen, the

following incomplete description can be given.

Of the pectoral girdle only a piece of a scapula (Fig. 1c) is known and

this is very similar to the corresponding part of the scapula of Tapinocephalus.

In the collection there are two imperfect ulnae (Fig. 6a). The ulna is

massive and squat (length + 350 mm.); the styloid ridge is prominent;

broad over the shaft and very broad over the coronoid process.

The only known radius

Fig. 5. (Fig. 6 b-d) is massive and

: squat (length 240 mm.); the

ventral face is concave with a

strong longitudinal ridge; the

flange (for the insertion of the

biceps) on. the proximo-

postaxial corner is strong; the

proximal facet is convex.

With no pelvis in this col-

lection I am _ extracting the

characteristic features from von |

Huene’s (21) account.

The supra-acetabular part

of the ilium is low and very

long (the height is only 52%

of its antero-posterior length);

the anterior process of the iliac

blade is very long and low and

Ilium of Phocosaurus megischion. (x %.) is fairly strongly everted; the

S-A.M. 11300. Lateral view. low posterior process is fairly

long, but much shorter than the anterior process, with the ilio-fibularis

ridge sharp and prominent and lying horizontally; the dorsal iliac edge is

not folded over laterally; on the inner face of the anterior process there is

a large face to receive a rib lying anterior to the main sacral mb.

The femur (see v. Huene’s Figs.) is short (400 mm.) and broad; the

width over the external trochanter is 256 mm.; the preaxial face is deeply

concave with the caput much preaxially directed; the external trochanter is

not demarcated from the proximal face, which extends far outwards from

the caput; the internal trochanter developed as a prominent tubercle;

the shaft is short and wide (diams. 132 x 84 mm.); the distal facets

are terminal.

With only one crushed tibia in our collection (Fig. 6 e-f) and the one

figured by von Huene, the Keratocephalus tibia appears to be massive

GIRDLES AND LIMBS OF DEINOCEPHALIA 215

but short (length 232-276 mm.) with a broad shaft and a massive cnemial

eminence.

Keratocephalus moloch. (x %.) a. Ulna of S.A.M. 11937. Dorsal view. b. Radius

of S.A.M. 11937. Dorsal view. c. Radius of S.A.M. 11937. Ventral view. d. Radius

of S.A.M. 11937 Posterior view. e. Tibia of S.A.M. 8946. Dorsal view. f. Tibia

of S.A.M. 8946. Posterior view.

Keratocephalus moloch von Huene

The specific description is as for the genus.

Type: Tiibingen. An ilium, femur and tibia, associated with parts of

the skull. Abrahamskraal, Prince Albert. Low Tapinocephalus zone.

Coll. v. Huene.

Referred specimens in the S.A.M. collection:

S.A.M. 8946. An imperfect left ulna and a crushed left tibia (Fig. 6

e-f), associated with a skull. Mynhardtskraal, Beaufort West. Low

Tapinocephalus zone. Coll. Boonstra.

216 ANNALS OF THE SOUTH AFRICAN MUSEUM

S.A.M. 11937. An imperfect right scapula (Fig. 1c), an imperfect left

ulna (Fig. 6a) and a good nght radius (Fig. 6 b-d), associated with

a skull. Buffelsvlei, Beaufort West. Low Tapincephalus zone. Coll.

Boonstra and Marais.

Genus Pelosuchus Broom

A part of the scapula preserved indicates a fairly close similarity to the

scapula of Keratocephalus, as does also the incomplete weathered coracoid.

The femur (Fig. 7) is fairly j

short (420 mm.) and broad; the Fig. 7.

width over the external tro-

chanter is 215 mm.; the preaxial

face is deeply concave with the

caput directed much preaxially;

the external trochanter does not

appear to be demarcated by a

notch from the proximal face;

the internal trochanter lies near

the middle of the bone far away

from the preaxial border and is

a prominent and strong tubercle;

the shaft is fairly strong and

broad (diams. 132 x 80 mm.);

the distal facets are terminal.

A distorted tibia appears to be

stout and short with a strong

cnemial eminence.

Pelosuchus priscus Broom

The specific description is as Femur of Pelosuchus priscus. (x #.)

S.A.M. 918. a. Ventral view. b. Anterior

for the genus. view. Note: In the femur ‘‘dorsal’’ and

Type: S.A.M. gre. “Part or “‘ventral’’ are projections on the plane

. in which the ventral faces of the condyles

= 4 scapula, coracoid, : lie and “‘anterior’’ on to the axial plane

fairly good femur (Fig. 7) at right angles to the above.

and distorted tibia, associated with a weathered dentary. Bokfontein,

Prince Albert. Middle? Tapinocephalus zone. Coll. Cairncross.

Generically Undetermined Specimens:

S.A.M. 2753. A middle portion of a humerus (Fig. 12a) and the

distal end of a femur. Viviers Siding, Beaufort West. Mid Tapino-

cephalus zone. Coll. Haughton and Whaits.

S.A.M. 9097. An isolated femur (Fig. 8). This is probably a femur

of Tapinocephalus. Merweville Commonage. Low Tapinocephalus

zone. Coll. Boonstra.

GIRDLES AND LIMBS OF DEINOCEPHALIA 217

Above: Femur, probably

of Tapinocephalus. (x &-)

S.A.M. 9097. a. Dorsal

view. b. Ventral view.

c. Anterior view.

Left: Interclavicle of ?

Tapinocephalus. (xX &-)

S.A.M. 9153. a. Ventral

view. b. Lateral view.

218 ANNALS OF THE SOUTH AFRICAN MUSEUM

S.A.M. 9153. An interclavicula (Fig. 9) and a distorted femur (Fig. 10).

The interclavicle has a long slender stem and the femur a ridgelike

internal trochanter. These may represent a genus lying between

Tapinocephalus and Struthiocephalus. Jacobskraal, Prince Albert.

Low Tapinocephalus zone. Coll. Boonstra.

Fig. 10,

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7.5.4 8

Femur of ? Tapinocephalus. (x #@.) S.A.M. 9153. (Slightly

distorted.) a. Dorsal view. b. Ventral view. c. Anterior view.

S.A.M. 9164. The middle portion of a humerus. Wakkerstroom, Prince

Albert. Low Tapinocephalus zone. Coll. Boonstra.

S.A.M. 11303. An isolated imperfect femur. Buffelsvlei, Beaufort West.

Low Tapinocephalus zone. Coll. Boonstra.

S.A.M. 11306. An isolated good femur (Fig. 11), which agrees well

with the proximal end of the femur (B.M. 43525) which I am

including in the type material of Phocosaurus.

The femur is quite robust and fairly long (498 mm.); the width over

the external trochanter is moderate (234 mm.); the preaxial face is concave,

with the caput directed well preaxially; the caput is massive (164 x II4 mm.);

the shaft is fairly long and narrow (diams. 120 x 86 mm.); the tibial facets are

GIRDLES AND LIMBS OF DEINOCEPHALIA 219

directed much distally and the postaxial epicondyle lies further distally than

the preaxial epicondyle; the femoro-tibialis ridge is fairly strong and the

area for the insertion of the ilio-femoralis is narrow.

Boesmansrivier, Beaufort West. Mid Tapinocephalus zone. Coll.

Boonstra. |

Femur of ? Phocosaurus. (x 3.) S.A.M. 11306. a. Dorsal view.

b. Ventral view. c. Anterior view.

S.A.M. 11702. An isolated distal end of a humerus (Fig. 12 b-d) which

agrees fairly well with that of Tapinocephalus. Elandsberg, Suther-

land. Low? Tapinocephalus zone. Coll. Boonstra.

S.A.M. 11993. An isolated good left humerus (Fig. 13) with a long

delto-pectoral crest and a well-modelled radial condyle as in Phoco-

saurus, but otherwise much like the humerus of Tapinocephalus.

Locality and collector unknown.

S.A.M. 11997. An incomplete ilium (Fig. 14), distal end of a femur

and the distal end of a radius, associated with a fairly good but as yet

unprepared skull.

ANNALS OF THE SOUTH AFRICAN MUSEUM

220

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GIRDLES AND LIMBS OF DEINOCEPHALIA 221

The ilium has most of its blade missing, but on the lower edge of the

posterior iliac process there is a strong ilio-fibularis ridge lying horizontally;

the acetabulum is very large but shallow.

In its unprepared state the skull cannot be definitely identified; it is

certainly not that of Keratocephalus and in shape differs from that of

Tapinocephalus and may thus very possibly be near Phocosaurus.

Locality and collector unknown.

Fig. 15.

Humerus of ? Tapinocephalus.

(x é-) S.A.M. 11993. Ventral Ilium of ? Phocosaurus. (x %.)

view. S.A.M. 11997. Lateral view.

Struthtocephalidae

Family Characters of the Girdles and Limbs.

With much more material at our disposal than in the case of the

Tapinocephalidae a much fuller account can be given of the girdles and

limbs of the Struthiocephalidae.

The pectoral girdle is small to medium sized and light to only fairly

massive.

222 ANNALS OF THE SOUTH AFRICAN MUSEUM

The scapula is small and low (310 mm.) to fairly large and high

(480-500 mm.) with the upper part of the blade expanded (120-230 mm.);

its posterior border is fairly concave; the tricipital ridge is not very strongly

developed; the subscapular groove is shallow and confluent with a groove

lying in the visceral face of the precoracoid, dorsally of the inner opening

of the supracoracoid canal; the glenoidal facet of the scapula is of medium

size, concave and with a fairly well-moulded and raised external rim and

it faces postero-ventrally but not externally.

The coracoidal plate is long antero-posteriorly. The precoracoid is fairly

large and consists of a fairly thin plate of bone, but is thickened at the

apex, which lies dorso-posteriorly of the outer opening of the supracoracoid

foramen; its outer face is only very slightly convex; the supracoracoid canal

is not directed very obliquely and its inner opening still lies in the precoracoid

near the precoracoid-scapular suture, but has a groove dorsally confluent

with the subscapular groove. The precoracoid is excluded from the glenoid

and on the apex it carries a ridge which limits the anterior movement

of the humerus.

The coracoid is a fairly strong element with a fairly large glenoidal facet

which faces dorso-posteriorly and also externally.

No cleithrum is preserved but in all probability was a splintlike bone

as in Moschops.

A pair of clavicles is preserved in Struthiocephalus whaitsi, but as is

evident from Fig. 17 they have suffered from distortion, so that the right

one appears to be a much squatter bone than the left one. The left

clavicle, which I believe shows the natural form more truly, is a fairly

light bone with an expanded lower end, which is applied to the outer

face of the upturned interclavicular antero-lateral corner; the upper end

has its anterior edge thickened and posteriorly there is a thin flange, which

is applied to the outer face of the scapula; internally a ridge limits the

posterior movement of the clavicle over the scapula.

The interclavicle is a strong bone with a fairly thick and long stem,

which is, however, only moderately wide; the anterior spatulate end is

considerably expanded and curves strongly upwards; the moderately

hollowed surface for the reception of the clavicle is dorso-posteriorly bounded

by the thickened raised edge of the bone; the upper surface of the stem

has no median ridge for the lower edge of the coracoidal plate to abut

against.

The humerus of this family varies considerably; in some cases it is

fairly short and broad and in others long and fairly slender; in length it

varies from small to fairly large (276?-475 mm.); the proximal expansion

is moderate to large (156-270 mm.); the distal expansion is moderate to very

large (174-288 mm.); the shaft is short to long and narrow to very broad

GIRDLES AND LIMBS OF DEINOCEPHALIA 223

(width 84-125 mm.); the delto-pectoral crest is short to fairly long and

it terminates, in all cases, well proximal to the plane in which the ventral

opening of the entepicondylar foramen lies; the caput is narrowly to

moderately narrowly oval; both the processus medialis and the processus

lateralis lie far proximally, more or less in the same plane as the caput;

the radial condyle is weak, not well modelled, and does not extend much

proximally along the ventral surface and thus always lies well distal of

the plane in which the entepicondylar foramen lies; the twist on the shaft

is small to moderate (8°—-20°) and the one case in which it is 50°, this is

undoubtedly due to postmortem distortion; the L.M.L. is weak to fairly

strong; the A.D.V.L. is moderately to well developed; the entepicondyle is

weak to moderate and the foramen slitlike; the ectepicondyle is little. to

greatly expanded.

The ulna is very inadequately known and is apparently fairly light and

relatively long (258-366? mm.).

The radius is only known from a couple of ends.

The pelvis, hitherto unknown except for Haughton’s (20) photograph

of the mounted skeleton of Struthiocephalus and Broom’s (15) restoration

sketch based on it, is represented in the collection by three fairly well

preserved specimens and a fourth with two iliae. The three bones of the

pelvis are weakly united and the symphysis between the two halves is weak.

The pubo-ischiadic plate is relatively fairly long (the antero-posterior

length is 9g0-102% of the height ot the pelvis); the supra-acetabular part of

the ilium is low (45-65% of its antero-posterior length) and long antero-

posteriorly; the anterior process of the ilium is long and low and only

slightly everted; the posterior process of the ilium is not much shorter than

the anterior process, but is much lower, with its posterior edge lying nearly

horizontally and partially everted to form a weak ridge (for the insertion of

the m. ilio-fibularis), which also lies nearly horizontally; the dorsal iliac

edge is folded over laterally to overhang and limit the gluteal area; on the:

inner face of the anterior iliac process there is a strong attachment of a mb

anterior to the main sacral rib. In the pubis the upper part of the anterior

edge is strongly everted to form an elongated tuberculum pubis, which ‘is:

ventrally demarcated from the slightly thickened antero-ventral edge by a’

distinct step; in the median line the middle part of the pubis meets its

fellow in a very weak pubic symphysis; the middle part of the ventral’

edge of the ischium is thickened to form a large sutural face for a fairly

strong ischial symphysis; between the pubic and ischial symphyses there is

a large quadrangular fenestra, in life filled with cartilage.

The femur of the Struthiocephalidae shows considerable variation: small:

(length 276 mm.), light and slender or medium sized (length 408-440 mm.},:

224 ANNALS OF THE SOUTH AFRICAN MUSEUM

fairly robust and broad or fairly long (468 mm.), fairly light and fairly

slender; the width over the external trochanter is small (95 mm.) to great

(210 mm.); the preaxial face is moderately to strongly concave with the

caput only slightly or moderately preaxially directed; the external trochanter

is not separated from the proximal face by any notch; the internal trochanter

is undeveloped or forms a strong elongated tubercle, situated well away

from the preaxial border; the shaft is fairly long to short and fairly slender

to broad (55-125 mm.); the tibial facets are much distally directed; the

m. femoro-tibialis originates on a fairly strong ridge; the ilio-femoralis area

of insertion is narrow or broad.

The tibia is small (length 180 mm.) and slender to medium sized (length

240-270 mm.) and fairly slender with a well-developed cnemial ridge or

eminence, and cnemial groove.

The fibula is light, fairly short to long and slender (228-288 mm.).

Genus Struthiocephalus Haughton

The pectoral girdle is of medium size and only fairly massive (total

height as projected on to the median plane 696 mm.). The scapula is fairly

massive and high (480 mm.); its posterior border is deeply concave. As

the family description is based mainly on specimens of this genus refer

back for additional characters of the pectoral girdle.

The humerus is preserved in two specimens with skulls associated, but

in one, in addition to being juvenile, the humerus has undergone considerable

postmortem distortion and in the other proximo-distal compression has

shortened the bone with concomitant widening.

The humerus is fairly large (length 393-430 mm.) and fairly massive;

the proximal expansion is large (270 mm.) and the distal expansion very

large (288 mm.); the shaft is fairly long to short, moderately thick (diams.

102 X 60 mm.) to robust (diams. 126 x 60 mm.); the delto-pectoral crest

is of medium length, terminating fairly far proximal of the ventral opening

of the entepicondylar foramen; the caput is narrowly oval and lies slightly

proximal to the processus medialis; the twist on the shaft is moderate (20°;

in the deformed S.A.M. 11493 it is 50°); the L.M.L. is fairly strong; the

A.D.V.L. is well developed; the entepicondyle is fairly strong with the

foramen entering dorso-postaxially and leaving as a ventral slit; the ect-

epicondyle is greatly expanded as a thin sheet of bone.

The ulna and radius are inadequately known.

The pelvis is as described for the family; it may be noted here that in

both specimens known, but in particular in the juvenile one, the ilium has

not grown out to meet the other two bones in the central part of the

GIRDLES AND LIMBS OF DEINOCEPHALIA 225

acetabulum, whereas the three bones meet in the pelvis of Struthtocephatellus

which on account of its small size might have been considered a not fully

grown Siruthiocephalus.

Fig. 15.

A.M. 3012. Lateral view. (

b. Coraco-scapula of Struthiocephalus whaitsi. S.A.M. 11493. Lateral view. (

a. Pectoral girdle of Struthiocephalus whaitsi. S.

The femur is of medium size (length 440 mm.) and fairly broad and

robust; the width over the external trochanter is 204 mm.; the preaxial

face is moderately concave and the caput only slightly preaxially directed;

the internal trochanter forms a fairly strong ridgelike tubercle; the shaft

~weeecoeo ®*

~»

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 16.

Interclaviclae of Struthiocephalus whaitsi. (yx 4.)

a. S.A.M. 3012. Ventral view. b. S.A.M. 3012.

Lateral view. c. S.A.M. 11579. Ventral view.

d. S.A.M. 11579. Lateral view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 227

short and broad (diams. 125 xX 50 mm.); as preserved the postaxial tibial

facet lies much distally; the ilio-femoralis area of insertion is broad.

The tibia appears to be fairly short (240 mm.).

The fibula is fairly short (230 mm.) but fairly robust with a deeply

concave preaxial border.

Struthiocephalus whaitsi Haughton

The specific description is as for the genus.

Referred specimens in the S.A.M. collection:

S.A.M. 3012. A nearly complete and fairly good pectoral girdle (Figs.

15a, 16 a-b and 17 a-d), a good humerus (Fig. 18), a fairly good

pelvis (Fig. 19), two somewhat crushed femora (Fig. 20), a fair

tibia (Fig. 21) and fibula (Fig. 22), associated with a fairly good

but crushed skull. Abrahamskraal, Prince Albert. Low Tapino-

cephalus zone. Coll. Haughton.

a b Cc d e

Claviculae. (x 4.) a. Struthiocephalus whaitsi. S.A.M. 3012. Lateral view of

right clavicle. b. Struthiocephalus whaitsi. S.A.M. 3012. Inner view of right

clavicle. c. Struthiocephalus whaitsi S.A.M. 3012. Lateral view of left clavicle.

d. Struthiocephalus whaitst. S.A.M. 3012. Inner view of left clavicle.

e. Moschops capensis. S.A.M. 11972. Inner view of left clavicle.

ANNALS OF THE SOUTH AFRICAN MUSEUM

228

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GIRDLES AND LIMBS OF DEINOCEPHALIA 229

S.A.M. 11493. A fair coracoidal plate (Fig. 15b), a distorted humerus

(Fig. 23) and a good pelvis (Figs. 24 and 25), associated with a

fairly good skull. Juvenile. Mynhardtskraal, Beaufort West. Low

Tapinocephalus zone. Coll. Boonstra.

Fig. 19.

Struthiocephalus whaitsi. S.A.M. 3012. (x %.) a. Pelvis in lateral view.

b. Ischium in inner view. c. Ilium in dorsal view.

S.A.M. 11572. A fair interclavicle, most of both coracoidal plates and

the proximal ends of both scapulae (Fig. 26). Klein-Koedoeskop,

Beaufort West. Low Tapinocephalus zone. Coll. Boonstra.

S.A.M. 11579. A good proximal two-thirds of a humerus, a good

precoracoid and a good interclavicle (Fig. 16 c-d). Buffelsvlei,

Beaufort West. Low Tapinocephalus zone. Coll. Boonstra.

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

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Femora of Struthiocephalus whaitsi. S.A.M. 3012. (x @.) a. Left femur in

dorsal view. b. Right femur in ventral view. c. Left femur in anterior

view. d. Right femur in anterior view.

a b Cc Fibula of Struthiocephalus wha

ae : we S.A.M. 3012. (x %.) a. Dag

Tibia of Struthiocephalus whaitsi. S.A.M. 3072.. (x 4:) view. b. Ventral view. c. Anteri

a. Dorsal view. b. Ventral view. c. Posterior view. view. 4

231

GIRDLES AND LIMBS OF DEINOCEPHALIA

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232 ANNALS OF THE SOUTH AFRICAN MUSEUM

Struthiocephalus whaitsi. Juvenile. S.A.M. 11493. (x 4.)

a. Ilium in dorsal view. b. Pelvis in lateral view. c. Pelvis

in anterior view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 233

S.A.M. 11939. A weathered humerus, ulna (Fig. 27 a-b) and femur

(Fig. 27 d-f), associated with a jaw fragment. Dikbome, Laingsburg.

Low Tapinocephalus zone. Coll. Boonstra.

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Pelvis of Struthiocephalus whaitsi. Juvenile. S.A.M.

11493. (x %.) Ventral view.

Specifically undetermined specimen:

The following specimen approaches sufficiently close to warrant pro-

visional inclusion in the genus, but not in the species — whattsz.

S.A.M. 11941. <A good interclavicle (Fig. 28). Droéfontein, Prince

Albert. Low ? Tapinocephalus zone. Coll. Boonstra.

ANNALS OF THE SOUTH AFRICAN MUSEUM

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GIRDLES AND LIMBS OF DEINOCEPHALIA 235

Fig. 27.

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a. Ulna of Struthiocephalus whaitsi. S.A.M. 11939. Dorsal view. (x $.)

b. Ulna of Struthiocephalus whaitsi. S.A.M. 11939. Anterior view. (x 4.)

c. Ulna of Struthiocephalus parvus. S.A.M. 5006. JDorsal view. ( x é.)

d. Femur of Struthiocephalus whaitsi. S.A.M. 11939. Dorsal view. (x #.)

e. Femur of Struthiocephalus whaitsi. S.A.M. 11939. Ventral view. (x 3.)

f. Femur of Struthiocephalus whaitsi. S.A.M. 11939. Anterior view. (x é.)

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

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Interclavicle of Struthiocephalus sp. S.A.M. 11941.

(x 4.) a. Ventral view. b. Lateral view.

Genus Struthiocephaloides Boonstra

The pectoral girdle is only known from a single scapula; the scapula

is fairly large and high (500 mm.); the upper part of the blade is expanded

(210 mm.) and the posterior face is not deeply concave. No other part

of the girdles and limbs is known.

Struthiocephaloides cavifrons Boonstra

Specific description as for the genus.

Type: S.A.M. 5607. A left scapula (Fig. 29 a-b), associated with a

good skull. Lammerkraal, Prince Albert. High Tapinocephalus zone.

Coll. Haughton.

GIRDLES AND LIMBS OF DEINOCEPHALIA 237

Fig. 29.

= eco @

Scapulae. (x %.) a. Struthiocephaloides cavifrons. S.A.M. 5607. Lateral view.

b. Struthiocephaloides cavifrons. S.A.M. 5607. Posterior view. c. Struthio-

cephalellus parvus. S.A.M. 5006. Lateral view.

Genus Struthiocephalellus Boonstra

The pectoral girdle is only known from an imperfect scapula and frag-

ments of the coracoidal plate; the scapula is small (height 310? mm.) and

the width of the blade is 210? mm.; the posterior face of the scapula is only

slightly concave.

The humerus is small (length 276? mm.) and is lightly built; the proximal

expansion is fairly great (156 mm.); the distal expansion is relatively great

(174 mm.); the shaft is fairly long and fairly wide (diams. 66 x 48 mm.);

the delto-pectoral crest is only moderately long and is fairly weak and

terminates well proximal of the plane in which the ventral opening of the

entepicondylar foramen lies; the relations of caput, processus medialis and

lateralis are unknown; the radial condyle is weak (thickness 48 mm.),

distally situated and curving round on to the dorsal surface; the twist on the

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

shaft is moderate (20° ?); the L.M.L. is weak and the A.D.V.L. fairly

strong; the entepicondyle is expanded as a thin sheet of bone and the foramen

enters ventro-postaxially and leaves as a ventral slit well away from the

edge of the bone; the ectepicondyle is expanded as a thin curved sheet

of bone. me

The ulna has only the proximal part preserved; it is lightly built; proximal

to the coronoid process a sharp longitudinal ridge separates a groove from

the sigmoid surface; the ventral surface is deeply excavated longitudinally.

The pelvis has a relatively long pubo-ischiadic plate (97% of the height

of the pelvis); the supra-acetabular part of the ilium is low (126 mm.) and

long (200? mm. as restored) and the height is thus 63% of the length;

although neither the anterior nor posterior iliac processes are fully preserved

both appear to have been long and fairly low. The pubis is anteriorly

slightly everted with a prominent tuberculum pubis, clearly demarcated from

the medially directed antero-ventral edge of the pubis; the pubic symphysis

is fairly long but is weak. The ischium is long with a long but weak

symphysis; the postero-dorsal edge of the ischium is strongly thickened.

The femur is small, slender and fairly lightly built (length 276 mm.);

the width over the external trochanter is small (95 mm.); the preaxial face

is only slightly concave and the caput, which is relatively weak (55X25 mm.),

is directed much dorsally as well as preaxially — there is thus a twist on

the shaft; the external trochanter is indistinctly developed and the internal

trochanter not developed at all; the shaft is long and fairly narrow and

thin (diams. 55 xX 48 mm.); the tibial facets face ventro-distally with the

postaxial one lying further distally; the femoro-tibialis mdge is moderately

strong and the ilio-femoralis area of insertion very narrow.

The tibia has its proximal face divided into two distinct concave facets

by a sharp and strong dorso-ventral ridge; a prominent cnemial protuberance

is present, continued distally as a sharp cnemial ridge, with a deep groove

lying postaxially of this ridge; all these features are well and sharply

modelled.

Struthocephalellus parvus Boonstra

Type: S.A.M. 5006. A partial scapula (Fig. 29c), a fairly good

humerus (Fig. 30), the proximal end of an ulna (Fig. 27c), a fairly

good pelvis (Fig. 31), a good femur (Fig. 32) and the proximal end

of a tibia (Fig. 33), associated with a fair skull. Abrahamskraal,

Prince Albert. Low Tapinocephalus zone. Coll. Haughton.

GIRDLES AND LIMBS OF DEINOCEPHALIA 239

Generically undetermined specimens:

S.A.M. 8947. This specimen, which consists of two fairly good femora

(Fig. 34 a-c), a good tibia (Fig. 34 d-f) and a good fibula (Fig.

34 g-i), appears to be of a Struthiocephalid, larger and of somewhat

heavier build than Struthiocephalus whaits:. The tibia is note-

worthy for its very strong cnemial protuberance. Mynhardtskraal,

Beaufort West. Low Tapinocephalus zone. Coll. Boonstra.

Fig. 30.

Humerus of Struthiocephalellus parvus. S.A.M. 5006.

(x 3.) a. Ventral view. b. Dorsal view. c. Anterior view.

Fig. 3], The following four specimens are of

Struthiocephalids which are much longer

in the limb than Struthiocephalus.

S.A.M. 3614. A_ good isolated

humerus (Fig. 35 b-e). This

humerus is relatively long and

slender (length 480 mm.); both

the proximal (width 192 mm.) and

the distal (width 198 mm.) expan-

sion is small; the shaft is fairly

long and slender (diams. 95 x 78

mm.); the twist on the shaft is

small (8°); the capitellum is very

weakly modelled and situated far

é distally; both epicondyles are weak

(x 4) and the entry of the entepicon-

parvus. S.A.M. 5006.

Lateral view.

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

dylar foramen is clearly visible in dorsal view. Abrahamskraal,

Prince Albert. Low Tapinocephalus zone. Coll. Haughton.

Fig. 32.

Right: Femur of Struthio-

cephalellus parvus. S.A.M.

5006. (x ¢-) a. Dorsal

view. b. Ventral view.

c. Anterior view.

Left: Tibia of Stvuthio-

céphalellus parvus. S.A.M.

5006. (Sees) Ca orsal

view. b. ~Ventral view.

c. Posterior view.

S.A.M. 5009. A fair humerus with the proximal end not in contact with

the shaft (Fig. 35a). Wolwefontein, Prince Albert. Low Tapino-

cephalus zone. Coll. Haughton.

Although the delto-pectoral crest is shorter than in S.A.M. 3614 and

the processus medialis situated more nearly in the same plane as the caput

these two specimens probably are of the same species.

S.A.M. 4349. Two fairly good iliae (Fig. 36 a-b), a good femur (Fig.

36 c-e), the proximal halves of both ulnae (Fig. 36f) and of both

tibiae and a fibula (Fig. 36g).

GIRDLES AND LIMBS OF DEINOCEPHALIA 241

| Fig. 34.

| 2? Struthiocephalus. S.A.M. 8947. (> }.) a. Femur in dorsal view.

b. Femur in ventral view. c. Femur in anterior view. d. Tibia in

\ dorsal view. e. Tibia in ventral view. f. Tibia in posterior view.

| g. Fibula in dorsal view. h. Fibula in ventral view. i. Fibula in

anterior view.

ANNALS OF THE SOUTH AFRICAN MUSEUM

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GIRDLES AND LIMBS OF DEINOCEPHALIA 243

Fig. 36.

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i An unnamed Struthiocephalid. S.A.M. 4349. (xX 4.) a. Ilium in dorsal view.

' b. Ilium in lateral view. c. Femur in dorsal view. d. Femur in ventral view.

Le ° . ° . : . e .

e. Femur in anterior view. f. Ulna in dorsal view. g. Fibula in ventral view.

3—Annals

244

ANNALS OF THE SOUTH AFRICAN MUSEUM

2° Sou

scare

An unnamed Struthiocephalid. S.A.M. 9008 Ose 3.) “ave Femur

in dorsal view. b. Femur in ventral view. c. Femur in anterior

view. d. Tibia in dorsal view. e. Tibia in ventral view. f. Tibia

in posterior view. g. Fibula in ventral view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 245

The ilium is low (supra-acetabular height 210 mm.), with both the

anterior and the posterior process long and low (antero-posterior length

335? mm.); with the height thus 62% of the length; the anterior process is

slightly everted and on its inner face receives a rib lying anterior to the

main sacral rib; the origin of the m. ilio-fibularis hes on the everted

horizontal ridge on the posterior iliac process; the dorsal edge of the ilium

is folded over the upper limit of the gluteal area. It is thus apparent that

this ilium is very similar to that of Struthiocephalus.

Scapulae of two indetermined Struthiocephalids. (x %.) a. S.A.M. 10197. Lateral

Whew... De) SoA M. 101974 . Posterior, view.- c..'S.A.M. 211700. Lateral ..view.

d: S.A.) 11700. ~ Posterior "view.

The femur is long (468 mm.) and slender; the width over the external

trochanter apparently small (Ig0? mm.); the preaxial face is quite strongly

concave and the caput directed well preaxially; the internal trochanter is

246 ANNALS OF THE SOUTH AFRICAN MUSEUM

situated fairly near the preaxial border; the shaft is long and narrow (diams.

102 X 66 mm.); the tibial facets are directed well distally with the postaxial

one situated furthest distally.

The ulna is apparently long with a strong longitudinal ridge and a deep

excavation on its dorsal face. The tibia and fibula are both long and

slender bones.

Leeurivier, Beaufort West. Low Tapinocephalus zone. Coll. Haughton.

S.A.M. 9008. A good femur (Fig. 37 a-c) and tibia (Fig. 37 d-f) and

the proximal end of a fibula (Fig. 37g). Klein-Koedoeskop, Beaufort

West. Low Tapinocephalus zone. Coll. Boonstra.

Fig. 39.

Femur of uncertain affinity. S.A.M. 11880. (x 2.)

a. Dorsal view. b. Ventral view. c. Anterior view.

This specimen is co-specific with S.A.M. 4349. These two specimens

thus represent, together with S.A.M. 3614 and S.A.M. 5009, a Struthio-

cephalid with an ilium very similar to that of Struthiocephalus but with

much longer pro- and epipodials.

The following two specimens both come from the Nieueveld, where the

fauna of the Tapinocephalus zone is very poorly known.

S.A.M. 1to197. An isolated well-preserved scapula and coracoid (Fig.

38 a-b). Grootfontein, Fraserburg. Low? Tapinocephalus zone.

Coll. Boonstra.

GIRDLES AND LIMBS OF DEINOCEPHALIA 247

S.A.M. 11700. <A good isolated scapula (Fig. 38 c-d). Elandsberg,

Sutherland. Low? Tapinocephalus zone. Coll. Boonstra.

These two scapulae are clearly Struthiocephalid and they may yet prove

to be of one or other of the forms as yet only known from skulls (e.g.

Struthionops, Riebeeckosaurus or the other species of Struthiocephatus).

S.A.M. 11880. An isolated femur (Fig. 39). The affinities of this bone

are very doubtful; the tibial condyles are practically terminal and

the internal trochanter situated well away from the preaxial border.

Arthurskraal, Beaufort West. High? Tapinocephalus zone. Coll.

Geol. Survey.

Moschopidae

Family Characters of the Girdles and Limbs.

The girdles and limbs of the Moschopidae are only moderately well

represented in our collection. There are a good number of pectoral elements

of Moschops, but of the pelvic girdle there is only a single ilium, and the

pro- and epipodials reasonably represented. In drawing up the descriptions

I have drawn upon the accounts of Broom (11), Romer (25) and Gregory

(18) of the Moschops material in the American Museum.

The pectoral girdle is lightly to fairly lightly built. The scapula is

fairly low (400 mm.) to fairly high (440 mm.) or high (540 mm.) with the

upper part of the blade broad (215 mm.) to fairly narrow (156-190 mm.);

the posterior border is fairly straight; the tricipital ridge is moundlike, low

and not very prominent; the canal for the supracoracoideus does not open

directly into the subscapular groove, which, however, crosses the suture

to meet the internal opening of the canal; the glenoidal facet of the scapula

faces ventro-posteriorly, but not externally in Avenantia.

The coracoidal plate is not very long antero-posteriorly. The precoracoid

is fairly large, but is mainly composed of a thin plate of bone which has

its anterior edge only slightly thickened and its outer face moderately convex;

the supra-coracoidal canal penetrates the bone very obliquely, so that its

internal opening lies just below the precoracoid-scapular suture, with its

upper rim notched and with a groove running into the subscapular groove;

but in Avenantia the canal crosses the suture to open into the subscapular

groove. The postero-dorsal apex of the precoracoid sometimes only just

enters the anterior corner of the glenoid and here forms a slight depression

in the ridge bounding the lateral edge of the glenoid.

The coracoid is small and light, but with a well-developed glenoidal facet,

which faces postero-dorsally, but also well externally.

248 ANNALS OF THE SOUTH AFRICAN MUSEUM

No cleithrum is preserved in any of my specimens, but if present could

only have been a splintlike bone as found by Broom (11) in one of the

Spitskop specimens.

Moschopid ulnae. (x %.) a, a’. ? Moschops. S.A.M.

9124A in dorsal and anterior views. b, b’*. ? Moschops.

S.A.M. 9157, in dorsal and anterior views. c, c’. Moschops

capensis. S.A.M, 11292, in dorsal and anterior views.

Of the clavicle only one dorsal and two ventral ends are preserved.

Dorsally the posterior expansion overlying the scapula is relatively better

developed than in Struthiocephalus and the Titanosuchia.

GIRDLES AND LIMBS OF DEINOCEPHALIA 249

The interclavicle is a fairly large and strong bone, with a relatively long

stem expanded posteriorly; anteriorly the spatulate end is not greatly

expanded and the articulatory surface for the reception of the lower end

of the clavicle is only slightly excavated, with the development of a slight

groove in only one specimen. In my material the anterior end is not bent

up as sharply as in the Spitskop material. There is no median ridge on

the dorsal surface of the stem as in some Titanosuchia.

The humerus is moderately long (345-444 mm.); the proximal expansion

is moderate to fairly large (170-240 mm.); the distal expansion is moderate

(165°-240 mm.); the shaft is fairly long and moderately broad (width

78-91 mm.); the delto-pectoral crest is fairly short and its distal end lies

well proximal of the ventral opening of the entepicondylar foramen; the

caput is narrowly oval; both the processus medialis and the processus

lateralis lie far proximally, more or less in the same plane as the caput; the

radial condyle is fairly weak, not well modelled and it lies far distally and

does not extend along the ventral face, and thus lies well distally of the

entepicondylar foramen, but does curve a little round on to the dorsal face;

the twist on the shaft is small to moderate (10-20°); the L.M.L. is moderately

strong; the A.D.V.L. is distinct; the entepicondyle is moderately to fairly

greatly expanded to form a thin curved sheet of bone, and the foramen

is slitlike with its entrance visible in dorsal view on the dorso-postaxial

border; the ectepicondyle is strongly expanded as a thin sheet of bone with

ventrally, in some humeri, a well-marked groove functioning as does the

ectepicondylar foramen in the Titanosuchids for the passage of the radial

nerve.

The ulna is fairly light to moderate (length 275-300 mm.); the dorsal lip

to the sigmoid face is weak to fairly strong; the styloid ridge is moderately

strong.

The radius is relatively long (186-204 mm.); it is a slender nearly feature-

less bone or is fairly robust, with a fairly strong proximo-postaxial flange;

the ventral face is longitudinally moderately concave; the proximal facet is

slightly convex.

The manus, fully known in only one specimen, has the carpal formula

3-4, I-2, 5, and the phalangeal formula 2, 3, 3, 3, 3.

The pelvis. As our collection contains only a single incomplete Moschopid

ilium of the genus Avenantia, I am extracting the characteristic features

for this group from the descriptions by Broom (11), Romer (25), Gregory

(18) and Byrne (16).

The pubo-ischiadic plate is short (70-80% of the height of the pelvis);

the supra-acetabular part of the ilium is high and fairly short (height

73%-75% of its antero-posterior length); the anterior process of the iliac

250 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 41.

Interclaviculae of Moschops capensis. (x }.) a. S.A.M. 11972, in

ventral view. b. S.A.M. 11972, in lateral view. c. S.A.M. 119722,

in ventral view. d. S.A.M. 11975, in ventral view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 251

blade is fairly long and high and diverges fairly strongly laterally; the

posterior process is somewhat shorter and appreciably to greatly lower than

the anterior process, and its ventro-posterior edge lies fairly horizontally

and this is slightly everted, but without forming a definite ridge for the

ilio-fibularis; the dorsal iliac edge is slightly folded over laterally; there is

apparently no face for the attachment of a rib lying anteriorly of the main

sacral rib on the inner face of the anterior iliac process.

CIP ea a

° o beg G

_— ar CT Tae af

ne Sb ded oe TL

icitee 9

wii? ehee

re ik

fet gabe

Jasce

Femur of Moschops capensis. S.A.M. 11974.

(3¢3-) 0a. Dorsal view, . b. Ventral, view.

c. Anterior view.

In the pubis the anterior edge is everted and ends antero-externally in a

low tuberculum pubis; the pubes barely meet in their posterior part in a

median symphysis; the two ischia meet below in a deep symphysis forming

a median keel.

The femur of the Moschopids is a medium-sized, fairly robust bone

(length 310-414 mm.); the width over the external trochanter is fairly small

to fairly great (I20?-205 mm.); there is no notch separating the external

trochanter from the proximal surface; the preaxial face is deeply concave

and the caput preaxially directed; the internal trochanter is undeveloped;

252 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 43.

Humeri of Moscops capensis. (x %.) a. S.A.M. 11976. Ventral view. b. S.A.M.

11976. Dorsal view. c. S.A.M. 11976. Anterior view. d. S.A.M. 11976. Proximal

view. €.)) SAM nora. Ventral view. fo SSHAL ME) oe ra7s Dorsal view.

g. S.A.M. 11975. Anterior view. h. S.A.M. 11975. Proximal view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 253

the shaft is fairly long, fairly to moderately broad (84-110 mm.); the condyles

are much distally directed, with the external condyle situated slightly or

considerably further distally than the internal condyle; the ilio-tibialis ridge

is fairly strong and the area for the insertion of the ilio-femoralis quite broad.

The tibia is medium-sized (length 210-270 mm.); cnemial ridge and

groove fairly well developed.

The fibula is lightly built, slender and relatively long (222-258 mm.).

The pes is only fully known in one species; here the tarsal formula is

2, I, 4 and the phalangeal formula 2, 3, 3, 3, 3.

a eae Fig. 44.

ra te

t ae SU a tlay Oe te

% ' no N

i. é :

6 4

\ {

\ (

Scapulo-coracoids of Moschops capensis. (x %.) a. S.A.M. 11976a. Lateral view.

b. S-ALM, -21976b. Lateral view. c. ‘S:A.M. r197o0b. Posterior view.

254 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Moschops Broom

The pectoral girdle is lightly built; the scapula is fairly high (440-515

mm.) with the upper part of the blade fairly narrow (174-190 mm.); the

glenoidal facet of the scapula faces externally as well as ventro-posteriorly.

The femur is a medium-sized fairly robust bone (length 310-390 mm.). For

other characters see the family description.

Moschops capensis Broom

The specific description is as for the genus.

Topotypes: A.M.N.H. 5551-5557. Remains of seven or eight skeletons,

associated with skulls. Spitskop, Laingsburg. Low? Tapinocephalus

zone. Coll. Whaits and Broom.

Scapulo-coracoid of Moschops capensis. (x %.) S.A.M.

r19g76e. a. Lateral view. b. Posterior view.

Referred specimens in the S.A.M. collection:

S.A.M. 11292. An ulna (Fig. 4oc), associated with a skull 11291.

GIRDLES AND LIMBS OF DEINOCEPHALIA 255

S.A.M. 11971. A distorted humerus, associated with a distorted skull.

S.A.M. 11972. Two interclaviculae (Fig. 41 a-c) and two distal and one

proximal end of the clavicle (Fig. 17e), found near a skull, 11972.

S.A.M. 11974. A femur (Fig. 42), associated with a skull.

S.A.M. 11975. Part of a scapula, a humerus (Fig. 43 e-h) and an

incomplete interclavicula (Fig. 41d).

S.A.M. 11976. A good humerus (Fig. 43 a-d).

S.A.M. 11976 a, b, c, d ande. Five coraco-scapulae (Figs. 44 and 45).

Fig. 46.

Scapula of Moschops capensis. S.A.M. 11977a. (x #%.) a. Lateral view. b. Posterior

view. Coraco-scapula of Moschops capensis. S.A.M. 11977b. (x %.) c. Lateral

view. d. Posterior view.

256 ANNALS OF THE SOUTH AFRICAN MUSEUM

' .§.A.M. 11977 a-d. A scapula (Fig. 46a, b) and an incomplete coraco- |

scapula (Fig. 46c, d), two fibulae (Fig. 47) and three tibiae (Fig. 48),

together with a number of incomplete and weathered girdle and |

limb-bones. 1

S.A.M. 11978. An isolated coracoid. |

d e f

Fibulae of Moschops capensis. (xX &-)

S.A.M. 11977c. Dorsal view. Tibiae of Moschops capensis. S.A.M. 11977. (X #:|

b. S.A.M. 11977c. Ventral view. a. Normal left tibia in dorsal view. b. Normal lef

c. S.A.M. 11977c. Anterior view. tibia in ventral view. c. Normal left tibia i}

d.° S: ACM. ‘r19e77da" ) Dorsal waew,: posterior view. d. Deformed left tibia in dorsa|

e. S.A.M. 11977d. Ventral view. view. e. Crushed right tibia in dorsal view}

f. S.A.M. 11977d. Anterior view. f. Crushed right tibia in posterior view.

Son PETE .

GIRDLES AND LIMBS OF DEINOCEPHALIA 257

All the above specimens were found in an area of about 500 sq. yards,

associated with the remains of at least twenty Moschops skulls, mostly badly

weathered.

Kruisvlei, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra.

Fig. 49.

Scapula of Avenantia kruisvleiensis. S.A.M. 9167. (x §.) a. Lateral

view. b. Internal view. c. Posterior view.

Genus Avenantia Boonstra

The girdles and limbs are very inadequately known. The scapula is

low (400 mm.), and apparently with a very broad blade (width 210 mm.);

the glenoidal facet faces ventro-posteriorly and not externally at all; the

supracoracoid canal crosses the suture into the scapula to open into the

subscapular groove. The ilium is incompletely preserved, but its blade is

fairly low, and both the anterior and posterior processes are long and low.

On the other hand the femur is very Moschops-like; it is a moderately

robust bone with a length of 378 mm.; 168 mm. wide over the external

trochanter; the preaxial face is deeply concave and the caput directed well

preaxially; the shaft is fairly slender (diams. 96 X 60 mm.); the condyles

face ventro-distally and the postaxial condyle is situated much further

distally than the preaxial condyle.

258 ANNALS OF THE SOUTH AFRICAN MUSEUM

Avenantia kruisvleiensis Boonstra

The specific description is as for the genus.

Type: S.A.M. 9167. A scapula, a weathered ilium (Fig. 50) and a

good femur (Fig. 51), associated

with a fairly good skull. Kruis- Fig. 50.

vlei, Beaufort West. Low |

Tapinocephalus —_ zone. Coll. |

Boonstra.

Genus Moschognathus Broom

There is no specimen in the South

African Museum referable to this genus

and in the only known specimen only

some features of the pelvis and femur

are known. These point to a large

measure of similarity with the cor-

responding bones in Moschops (see eek Bi? aah

Gregory’s (18) Plate). Right ilium of Avenantia Rruis-

vietensis. S.AIM. oxb7. Px z.) |

Lateral view.

Femur of Avenantia kruisvleiensis. S.A.M. 9167. (x

a. Dorsal view. b. Ventral view. c. Anterior view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 259

Moschugnathus whaits: Broom

Type: A.M.N.H. 5602. Part of the pelvis, two femora and partial

vertebral column, associated with parts of a skull and lower jaw.

Beaufort West District. ° Tapinocephalus zone. Coll. Whaits.

Genus Pnigalion Watson

In our collection there is no specimen that can be referred to Puigalon.

In the only known specimen in the British Museum, Watson (29) has

associated with the skull a humerus, femora and a tibia. The humerus is

medium sized and fairly robust (length 455? mm.) with a greatly expanded

proximal end (width 246 mm.); the shaft is fairly long and broad (breadth

102 mm.); the strong delto-pectoral crest terminates well proximal of the

entepicondylar foramen; the ectepicondyle apparently greatly expanded as

a thin sheet of bone.

The femur figured by Watson is a stout and fairly squat bone (length

420 mm.); width over the external trochanter 216 mm. and the width of

the shaft 144 mm.). Another femur, consisting of distal and proximal ends

not in contact, represents a long fairly slender bone with a preaxially

directed head. I took the following measurements in 1934: width across

external trochanter 205 mm., across the epicondyles 170 mm. and the

diameters of the shaft 110 X 70 mm. It would thus appear that Watson’s’

figure is that of a dorso-ventrally crushed femur, which thus appears much

broader than it was in life and in that case it would not be so very different

from the Moschops femur.

Pnigahon owem Watson

The specific description is as for the genus.

Type: B.M. R.3596 (and R.3606). A humerus, femora and tibia,

considered to be associated with a partial skull. De Cypher, Beaufort

West. Low? Tapinocephalus zone. Coll. unknown?

Genus Moschoides Byrne

As there is no specimen referable to this genus in this collection I am

extracting the following from Byrne’s description :

‘“‘The scapula is a massive bone, thickened greatly along its posterior

edge. It forms the upper part of the glenoid cavity. The coracoid, which

forms the remainder of the glenoid, is a rather small element, but bears a

260 ANNALS OF THE SOUTH AFRICAN MUSEUM

well-developed coracoid process. The precoracoid is much larger, consider-

ably thinner, and is moderately convex. It is thickened only in the glenoidal

corner where, also, it is pierced by a large supra-coracoid foramen. The

clavicles are moderately expanded elements somewhat constricted in their

middle portions. The humerus is a powerfully developed element. There is

a strong delto-pectoral crest, a somewhat constricted shaft, and two

moderately developed distal condyles. Apparently there is only an ent-

epicondylar foramen.

Both radius and ulna are conspicuously flattened antero-posteriorly. The

ulna, the longer bone, carries a strong olecranon process.

The carpus consists of three rather large proximal elements, two central

elements, and five rather small distal carpals. The metacarpals are

unusually small with the exception of the fifth which is expanded and

platelike. The phalangeal formula is 2, 3, 3, 3, 3; the first phalanx in each

digit being unusually short.

The pelvic girdle is characterised particularly by the large anterior

process of the ilium. The acetabulum is roughly circular in outline.

The femur is a long element and is considerably flattened antero-

posteriorly. It bears a well-defined head and two sharply demarked distal

condyles. The lesser trochanter is not evident but the great trochanter is

well developed.

The tibia is very short. While greatly constricted along its shaft, it

flares broadly at either end.

The tarsus is represented by a massive astragalus, an expanded platelike

calcaneum, a rather small navicular, and four distal tarsals, the first three

of which are quite small. It would seem that a good deal of cartilage must

have been present in life to complete the ankle structure.

The metatarsals and the digits are similar in size and shape to the

corresponding elements in the forefoot. The distal portion of the hindfoot

is much weaker, however, in comparison with the forefoot.’’

Moschoides romen Byrne

Specific description is as for the genus.

Type: Walker Museum. No. ? The pectoral girdle, humerus, radius,

ulna and manus; a fair pelvic girdle, femur, tibia, part of the fibula

and the pes. Hottentotsrivier, Beaufort West. High? Tapinocephalus

zone. Coll. Romer and Miller.

Generically undetermined specimens: |

S.A.M. 1057. An isolated good femur (Fig. 52).

GIRDLES AND LIMBS OF DEINOCEPHALIA 2601

Fig. 52.

Above: Femur of a Moschopid

intermediate between Moschops and

Pnigalion. SPAPIVIR SS 1057. >< ai)

a. Dorsal view. b. Ventral view.

c. Anterior view.

Left: Scapula of a Moschopid. Gen.

Indet. S.A.M. 9003. (x 4.)

a. Lateral view. b. Posterior view.

262 ANNALS OF THE SOUTH AFRICAN MUSEUM

This is a medium sized (length 414 mm.) fairly robust bone (width over

the external trochanter 180 mm.); the shaft is fairly long and moderately

broad (diams. 102 X 72 mm.); both condyles lie in practically the same

plane; the ilio-tibialis ridge is strong. This femur appears to lie about

midway between that of Moschops and Pnigalion.

Letjiesbos, Beaufort West. Mid Tapinocephalus zone. Coll. Maddison.

Fig. 54.

Above: Humerus of ? Mos-

chops., S.A.M. 0157.7 (X-3-)

a. Ventral view. b. Dorsal

view. c. Anterior view.

Left: Radius of ? Moschops.

S.A.M. | 915750" ( p@teaen

Dorsal view. b. Ventral

view. c. Posterior view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 263

S.A.M. 9003. A good isolated scapula (Fig. 53). This scapula is high

(540 mm.) with a fairly narrow blade (width 156 mm.) and with its

posterior border fairly straight and the glenoid not facing externally.

Klein-Koedoeskop, Beaufort West. Low Tapinocephalus zone. Coll.

Boonstra.

S.A.M. g101. The distal end of a fibula. Rietfontein, Prince Albert.

Low Tapinocephalus zone. Coll. Boonstra.

S.A.M. 9124A. A good isolated ulna (Fig. 40 a, a’). Very similar to

the ulna of Moschops capensis. Voélfontein, Prince Albert. Low

Tapmocephalus zone. Coll. Boonstra. 3

S.A.M. 9157. A good humerus (Fig. 54), radius (Fig. 55), ulna (Fig.

40 b, b’) and part of the carpus.

I (4) have recently described the epipodial and the carpus. The humerus

is of medium size (length 384 mm.); the proximal expansion is fairly large

(210 mm.) and the distal expansion is moderate (212 mm.); the shaft is

fairly wide and short (diams. 91 x 66 mm.); although the ulna is very

similar to that of Moschops the radius is a much more slender bone. This

specimen thus lies near Moschobs.

Wolwefontein, Prince Albert. Low Tapinocephalus zone. Coll. Boonstra.

S.A.M. 11581. An isolated flattened femur (Fig. 56). This small femur

(length 372 mm.) is most probably of a juvenile Moschops. Buffels-

vlei, Beaufort West. Low Tafinocephalus zone. Coll. Boonstra.

S.A.M. 11987. An isolated weathered distal two-thirds of a humerus.

The groove on the under surface of the ectepicondyle for the passage

of the radial nerve is well developed. Buffelsvlei, Beaufort West.

Low Tapinocephalus zone. Coll. Boonstra.

S.A.M. 11995. An isolated radius (Fig. 57) with a well-developed

proximo-postaxial flange. Stouter than S.A.M. 9157. Sutherland

District. Low? Tapinocephalus zone. Coll. Boonstra.

S.A.M. 3308. An isolated good fibula (Fig. 58). With a length of 252

mm. and its slender build it comes very near that of Moschobps.

Uitkyk, Beaufort West. Low Tapinocephalus zone. Coll. Haughton.

264 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 57.

Above: Radius of ? Moschops. S.A.M. 11995.

(x ¢.) a. Dorsal view. b. Ventral view.

c. Posterior view.

Left: Femur of ? Moschops. S.A.M. 19581.

(x%.) Ventral view.

ao Job Cc

Fibula of ? Moschops. S.A.M. 3308.

(x %.) a. Dorsal view. b. Ventral view.

c. Anterior view.

Titanosuchia

Infra-ordinal Characters of the Girdles and Limbs.

The pectoral girdle is large and massive; the scapula is fairly to very

high (530-690 mm.) with the upper part of its blade fairly to very broad

GIRDLES AND LIMBS OF DEINOCEPHALIA 265

(220-426 mm.); the scapular head of the triceps originates from a fairly to

very prominent ridge; the internal opening of the foramen suprecoracoideus

opens into the deep subscapular groove, i.e. it crosses the suture between

the precoracoid and the scapula; the glenoidal facet of the scapula is large,

facing ventro-posteriorly and not, or but little, externally. The coracoidal

plate is of great antero-posterior extent. The precoracoid is large and strong,

with its anterior border thickened, and its greatly thickened dorso-posterior

apex forms at most the anterior depressed rim of the glenoid; the foramen

supracoracoideus penetrates the bone very obliquely, so that internally it

forms a groove in the sutural face crossing to open into the subscapular

groove; the outer precoracoidal face is only moderately convex. The coracoid

is massive with its large glenoidal facet facing dorso-posteriorly and also

much externally.

These three bones have greatly widened sutural faces near the glenoid,

but distally from this corner the bones become abruptly thinner along the

sutural lines.

The clavicula is a medium-sized to large bone; flat with expanded ends

and a constricted waist.

The cleithrum is a strong element with its upper end expanded and

projecting beyond the upper scapular border.

The interclavicle is massive to very massive with a greatly widened

stem; broadly spatulate anteriorly, and here curving sharply upwards. In

some specimens a ridge along the median line on the dorsal surface of the

stem limits the movement of the coracoidal plates. A deep groove receives

the posterior edge of the lower end of the clavicle.

The humerus is always massive, but varies in size from fairly small, short

and squat to very large and long (312-575 mm.); an ectepicondylar foramen

is always present; the radial condyle is always strong, thick and well

modelled and extends far proximally along the ventral face, in some cases

nearly reaching the base of the delto-pectoral crest; the processus medialis

lies either nearly in the same plane as the caput or in a more distal plane,

i.e. it is proximally or not proximally situated; the processus lateralis is

always situated far proximally; the proximal expansion is relatively

moderately or very great; the epicondyles are greatly expanded; the shaft

is either fairly long or very short, fairly slender or very broad; the ventral

opening of the entepicondylar foramen is slitlike or broadly oval; except

in one case, the delto-pectoral crest is long; the ectepicondylar foramen

is always small and circular.

The ulna is large and massive (length 320-402 mm.).

The radius is large and massive (length 282-318 mm.).

The manus is unknown.

266 ANNALS OF THE SOUTH AFRICAN MUSEUM

The pelvis has a short pubo-ischiadic plate; the iliac blade is high; the

ilio-fibularis ridge is rounded and strong and usually lies vertically; the

supra-acetabular buttress is strong with a well-developed notch; the aceta-

bulum faces outwards; the anterior iliac process is moderately to strongly

everted; the posterior iliac process is fairly short and high; the iliac blade

is concave in antero-posterior direction.

The femur is large and massive (length 498-605 mm.) with the internal

trochanter developed as a strong rounded tubercle, situated well away from

the preaxial border; the width over the external trochanter, which sometimes

has a notch separating it from the proximal face, is great (234-306 mm.).

The tibia is large and massive with strongly expanded ends (length

255-355 mm.).

The fibula is large and stout with a deeply concave anterior face (length

235-345 mm.).

The pes is incompletely known.

Titanosuchidae

Family Characters of the Girdles and Limbs.

The Titanosuchid pectoral girdle is only adequately known in one genus.

The pectoral girdle is large and massive; the height is 860-890 mm., and

the length of the coracoidal plate is 560-580 mm., which is thus 62-67%

of the height. The scapula is high (680 mm.) and the upper end of the

blade is broad to very broad (372-426 mm.); the scapular head of the triceps

is attached to a sharp ridge or a prominent mound.

The precoracoid is large and massive (340 X 342 mm.).

The coracoid is of medium size but massive (250 X 252 mm.).

The clavicle is a large bone (length 505 mm.); it is medio-laterally

flattened, with expanded dorsal and ventral ends, but the waist not greatly

constricted; the ventral spatulate end curves inwards to fit over the outer

face of the upturned antero-lateral corner of the interclavicle; the dorsal end

has its anterior end greatly thickened and is produced dorsally as a short,

strong process, which presumably fits into a groove on the lower end of the

cleithrum, which is however not preserved.

The interclavicle is a large and massive bone; the length is 570 mm.;

width over anterior expansion 455-545 mm.; width over posterior end of

stem 215-310 mm. and over the waist of the stem 155-180 mm.; the lateral

horns have a thickened postero-lateral edge, anterior to which there is a

fairly deep groove to house the ventro-posterior edge of the spatulate end

of the clavicle.

GIRDLES AND LIMBS OF DEINOCEPHALIA 267

The humerus is long and large (length 480-575 mm.); the radial condyle,

although strong and extending far along the ventral face, does not reach the

plane of the entepicondylar foramen, and lies well distal of the base of the

delto-pectoral crest; the processus medialis always lies far proximally —

nearly in the same plane as the caput; the proximal expansion is moderately

great (240?-310 mm.); the shaft is fairly long and relatively not very broad;

the ventral opening of the entepicondylar foramen slitlike or oval.

The ulna has a short massive shaft and a long sigmoid face with a

rounded lip.

The radius is a long robust bone.

In the massive pelvis the upper edge of the ilium does not slope strongly

downwards in posterior direction; the anterior process is moderately everted;

the ilio-fibularis ridge is massive; the ilium forms much the greater part

of the acetabulum.

The femur is very large (length 564-605 mm.); the width over the

external trochanter is great (275-306 mm.); the preaxial face moderately to

deeply concave.

The tibia and fibula large and strong.

Genus Titanosuchus Owen

The pectoral girdle poorly known; the coracoid is massive with the

glenoidal facet facing much externally.

The humerus is very large (length 530 mm.); the shaft is short and wide

(130 mm.); the width over the proximal expansion is 308 mm.; the delto-

pectoral crest is fairly short and its extremity is knoblike; the processus

medialis is situated well proximally; the capitellum is strong and extends

far proximally along the ventral face of the bone; the opening of the

entepicondylar foramen ventrally is slitlike; the ectepicondyle is developed

as a fairly thin sheet of bone and the foramen is situated well away from

the edge of the bone.

The epipodial bones and the manus are unknown.

The pelvic girdle is unknown.

The femur is very long (605 mm.); the width over the external trochanter

is 275 mm.; the shaft is fairly narrow (135 mm.); the preaxial face is not

very deeply concave, but the caput is preaxially directed.

The fibula is large and massive (295 mm.).

The tibia and pes are not known.

268 ANNALS OF THE SOUTH AFRICAN MUSEUM

Titanosuchus ferox Owen

The specific diagnosis is as for the genus (see Seeley’s (28) plates).

B.M. 49367, coracoid; 49369, humerus; 49368, femur; and 49367b, fibula;

all said to be associated with the type cranial material. Koedoeskop,

Beaufort West. Mid Tapinocephalus zone. Coll. T. Bain.

Referred specimens in the S.A.M. collection:

S.A.M. 739. A fairly good isolated femur, with a length of 570 mm.

and shaft diameters of 157 XxX 65 mm. _ Beaufort West District.

Tapinocephalus zone. Coll. Oakley.

S.A.M. 11491. A good isolated right femur (Fig. 59) with a length of

540 mm.; 294 mm. over the external trochanter, which is separated

from the proximal face by a distinct notch; the shaft diameters are

144 X 84mm. Mynhardtskraal, Beaufort West. Low Tapinocephalus

zone. Coll. Boonstra.

Fig. 59.

Femur of Titanosuchus ferox. S.A.M. 11491. (x %.) a. Dorsal view. b. Ventral view.

c. Anterior view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 269

Genus Scapanodon Broom

All that is known of the pectoral girdle is a single incomplete scapula

which shows that the scapular head of the triceps originated from a sharp

ridge on the posterior face of the scapula just above the glenoid.

The humerus is fairly long to long (480?-522 mm.); the proximal

expansion is not very great (240-264? mm.); the distal expansion fairly

great (282-290? mm.); the shaft is fairly short to long and not very broad

(diams. 114-118 X I10-12I mm.); the delto-pectoral crest is long with its

extremity extending to fairly near the entepicondylar foramen or terminating

well proximally of this plane; the caput is oval and fairly massive and lies

in nearly the same plane as the processus medialis, which is thus well

proximally situated; the twist on the shaft is great (40°); the L.M.L. is

strong, forming a pronounced bulge on the dorsal surface of the shaft; the

A.D.V.L. is strong and sharp; the entepicondyle is developed as a moderately

thick, but not greatly flaring flange of bone, with the foramen ventrally

showing a slitlike opening not very far removed from the edge of the bone;

the ectepicondyle is a greatly flaring thin sheet of bone pierced vertically

by the foramen lying well removed from the edge of the bone.

The epipodial bones of the forelimb and the manus are not known.

The pelvic girdle is massive and high; the pubo-ischiadic plate is short

(80% of the height of the pelvis as restored); the supra-acetabular part of

the ilium is high and relatively short (the height is 74% of the antero-

posterior length); the anterior iliac process, although not fully preserved,

appears to have been fairly short, but fairly high, and it is only moderately

everted; the posterior iliac process is short and has its posterior edge directed

much upwards and this edge is folded over to form a massive ilio-fibularis

ridge, which forms a prominent feature on the outer iliac face; anterior to

this ridge the iliac blade is deeply concave where the gluteal muscle is

attached; the inner face of the anterior process is irregularly pitted for the

reception of a rib lying anterior to the main sacral rib; the iliac portion of

the acetabulum is very great, forming over two thirds of the acetabulum.

The antero-ventral edge of the pubis is strongly everted with the tuber-

culum pubis confluent with the strongly thickened outwardly directed antero-

ventral edge, which thins as it proceeds towards the median line. The

pubic symphysis must have been short and weak. The pubic foramen

is a large oval opening.

Only the upper half of the ischium is preserved; it forms only a small

part of the acetabulum; its postero-dorsal edge is greatly thickened and it

would appear that the ischial symphysis was weak.

The femur is massive and long (564 mm.); its proximal expansion is

great, with the width over the external trochanter, which has a notch

270 ANNALS OF THE SOUTH AFRICAN MUSEUM

separating it from the proximal face, 306 mm.; the preaxial face is fairly

strongly concave with the caput moderately thick (134 mm.) and directed

moderately preaxially; the shaft is fairly long but very broad (162 mm.);

the preaxial condyle is much weaker than the postaxial condyle, which also

lies further distally and is very massive (174 mm. thick); the femoro-tibialis

ridge is strong, but not strongly bulging on the dorsal surface of the shaft.

No tibia, fibula or pes is known.

Scapanodon duplessis: Broom

Of the girdle and limb-bones only the humerus is known. The humerus

is long (520-522 mm.); the delto-pectoral crest with a swollen end terminates

well proximally of the entepicondylar foramen.

S.A.M. 772 and 773. Two fairly good left humeri (Fig. 60 b and c) said

to be associated with the poor type jaws. Seekoeigat, Prince Albert.

High Tapinocephalus zone. Coll. du Plessis.

Fig. 60.

Aaa es ea w A Sw

Humeri in ventral view.

(x 3.) a. Scapanodon septemfontis. S.A.M. 5001.

b. Scapanodon duplessisi. S.A

.M. 772. c. Scapanodon duplessisi. S.A.M. 773.

—e—B_

GIRDLES AND LIMBS OF DEINOCEPHALIA 271

Scapanodon septemfontis Sp. Nov.

I am making the specimen (S.A.M. 5001), which Broom (13) erroneously

described as being referable to Tapinocephalus atherstone1, the type of a

new species of Scapanodon.

Nothing is known of the pectoral girdle.

The humerus is large and fairly long (480? mm.); the delto-pectoral

crest terminates far distally, but still well away from the ventral opening of

the entepicondylar foramen.

Fig. 61.

OS PES SS

. Cis = . 3

ee ee.

Oe tes

ai woe

. -

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Pelvis of Scapanodon septemfontis. S.A.M. 5001. (x ¢@.) Lateral view.

272 ANNALS OF THE SOUTH AFRICAN MUSEUM

The rest of the forelimb is unknown.

The pelvis and femur are as described in the generic diagnosis.

Type: S.A.M. 5001. An incomplete humerus (Fig. 60a), a nearly

complete pelvis (Fig. 61) and a good femur (Fig. 62). Sewefontein,

Prince Albert. Low Tapinocephalus zone. Coll. Haughton.

Referred specimens in the S.A.M. collection:

S.A.M. 1203. A pubis. Letjiesbos, Beaufort West. Mid Tapinocephalus

zone. Coll. Maddison.

S.A.M. 11578. Part of a scapula and the major part of a humerus.

Aasvoélbos, Beaufort West. Low Tapinocephalus zone. Coll.

Boonstra.

Femur of Scapanodon septemfontis. S.A.M. 5001. (x %.) a. Dorsal view. b. Ventral

view. c. Anterior view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 273

Se Dn a

“0 414,°"

“ere

Precoracoid of Parascapanodon avifontis. S.A.M. 9127. (x #%.) a. Outer view.

b. Outline of sutural face for the scapula. c. Outline of sutural face for the coracoid.

d. Inner view. e. Lateral view (as projected on to the median plane).

274 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Parascapanodon Gen. Nov.

The generic characters of the pectoral girdle are as described for the

family.

The humerus is very large and massive; the length is 575 mm. and the

ends greatly expanded (proximal 310? mm., distal 312 mm.); the shaft is

fairly long but very robust (diams. 144 x 142 mm.); the delto-pectoral

crest is long, but terminates well proximal of the plane in which the ent-

epicondylar foramen lies; the caput is very massive, but short; the processus

medialis lies just a little distally of the plane in which the caput lies; the

capitellum is very strong and massive and extends far along the ventral

face, but does not reach the plane of the entepicondylar foramen; the

““twist’” on the shaft is large (40°); the L.M.L. is strong, with a massive

swelling on the dorsal surface of the shaft; both epicondyles are strongly

developed; the ventral opening of the entepicondylar foramen is large and

broadly oval, and the ectepicondylar foramen is small and situated well

away from the edge of the bone.

Right clavicle of Parascapanodon avifontis. S.A.M. 9127. (x %.) a. Outer

view. b. Inner view. c. Anterior view.

"MOTA JOMOJUY “9 “MOTA

: eusA *q “MaIA [esIog “e (‘fF X) L216 “W'S ‘syuofian uopoundvaspang JO Inuls 7

275

GIRDLES AND LIMBS OF DEINOCEPHALIA

4—Annals

276 ANNALS OF THE SOUTH AFRICAN MUSEUM

The ulna is a large and massive bone (length 372 mm., width over the

coronoid process is 200 mm.); the sigmoid face is long, with its ventral part

broadly rounded; the coronoid process is situated far distally and the shaft

is massive, broad and short.

The radius is a long robust bone (length 294 mm.) with a strong proximo-

postaxial flange.

No pelvis is known.

The femur is very long and massive (length 595 mm.); very broad over

the external trochanter (300 mm.); the preaxial face is deeply concave, with

the caput much preaxially directed and massive (diams. 215 x 167 mm.);

the external trochanter is indistinctly separated by a notch from the proximal

face; the shaft is fairly long and broad (breadth 150 mm.); wide over the

massive distal facets; the area of origin of the femoro-tibialis forms a strong

bulging ridge.

The tibia is large and massive (length 330-355 mm.); the cnemial

eminence very massive, continued distally as a strong ridge, with a deep

groove lying postaxially.

The fibula is large and stout (length 330-345 mm.).

Parascapanodon avtfontis Sp. Nov.

The specific characters are as for the genus.

Fig. 66.

Left fibula of Parascapanodon avifontis. S.A.M. 9127.

(x ¢.) a. Dorsal view. b. Ventral view. c. Anterior view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 277

Pm es Se “~ 5

“ae Us te be eh ee le

Scapulo-coracoid of Parascapanodon avifontis. S.A.M. 9106. (x $.)

Lateral view.

278 ANNALS OF THE SOUTH AFRICAN MUSEUM

Type: S.A.M. 9127. A very good precoracoid (Fig. 63), a good clavicle

(Fig. 64), a good femur (Fig. 65), a well-preserved fibula (Fig. 66),

associated with parts of a large skull. Voélfontein, Prince Albert.

Low Tapinocephalus zone. Coll. Boonstra.

Paratypes: Although lacking cranial parts, and direct comparison with

the type skull parts thus impossible, I have used the following specimens

in drawing up the generic description of Parascapanodon:

S.A.M. 9106. Parts of the scapula, coracoid and precoracoid (Pig. - 67).

Veldmansrivier, Prince Albert. Low T apinocephalus zone. Coll.

Boonstra.

S.A.M. 11488. Parts of the precoracoid, coracoid and _interclavicle

(Fig. 68) and a tibia (Fig. 69 a-c) and fibula (Fig. 69 d-f). Voélfontein,

Prince Albert. Low 7 apinocephalus zone. Coll. Boonstra.

Fig. 68.

Interclavicle, precoracoid and coracoid of Parascapanodon avifontis in ventral view.

S.A.M. 11488. (x 4.)

GIRDLES AND LIMBS OF DEINOCEPHALIA 279

Fig. 69.

tT ee _

Parascapanodon avifontis. S.A.M. 11488. (x §.) a. Right tibia in dorsal

view. b. Right tibia in ventral view. c. Right tibia in posterior view.

d. Right fibula in dorsal view. e. Right fibula in ventral view. f. Right

fibula in anterior view.

These two specimens are included in this species on the ground of the

great similarity of their precoracoids to that of the type.

280 ANNALS OF THE SOUTH AFRICAN MUSEUM

On the similarity of its fibula to that of the type I have also included:

S.A.M. 9163. A good fibula (Fig. 70 a-c) and a fair ulna (Fig. 70 d-e).

Wakkerstroom, Prince Albert. Low T apinocephalus zone. Coll.

Boonstra.

Fig. 70.

. Loc cosmos ‘

oot bras

Pe oye F 8 ee

a s, tee

26s,

Oleh =

Parascapanodon avifontis. S.A.M. 9163. (x #.) a. Left

fibula in dorsal view. b. Left fibula in ventral view.

c. Left fibula in anterior view. d. Left ulna in dorsal

view. e. Left ulna in anterior view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 281

On the similarity of its tibia to that of S.A.M. 11488 I have also

included :

S.A.M. 11299. A good tibia (Fig. 71 a-c) and a good radius (Fig. 71 d-f).

Boesmansrivier, Beaufort West. Mid Tapinocephalus zone. Coll.

Boonstra.

Parascapanodon avifontis. S.A.M. 11299. (xX @-) a. Right tibia in dorsal

view. b. Right tibia in ventral view. c. Right tibia in posterior view.

d. Right radius in dorsal view. e. Right radius in ventral view. f. Right

radius in posterior view.

ANNALS OF THE SOUTH AFRICAN MUSEUM

282

‘MOTA [PUITXOIG *9

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‘MOIA TesIOg ‘q

‘syuofian uopoundvosvavg JO snzreuInyzT

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Vide he

GIRDLES AND LIMBS OF DEINOCEPHALIA

Scapulo-coracoid of Parascapanodon. S.A.M. goto.

Lateral view.

283

284 ANNALS OF THE SOUTH AFRICAN MUSEUM

I am including the following isolated humerus as a paratype:

S.A.M. 11881. <A fairly good isolated humerus (Fig. 72). Bloukrans,

Prince Albert. Mid Tapinocephalus zone. Coll. Boonstra.

This very large and massive humerus can only be associated with such

a large skull and massive femur as that of Parascapanodon.

Referred specimens in the S.A.M. collection:

S.A.M. go10. A scapula and coracoid (Fig. 73). Klein-Koedoeskop,

Beaufort West. Low Tapinocephalus zone. Coll. Boonstra.

The coracoid in this specimen agrees very well with that of the paratype

S.A.M. 9106 and I am thus referring it to this genus although it has a much

wider scapular blade.

S.A.M. 11938. An isolated tibia. Steenboksfontein, Laingsburg. Low

Tapinocephalus zone. Coll. Boonstra.

S.A.M. 11989. An isolated good fibula (Fig. 74). Koedoeskop, Beaufort

West. Mid Tapinocephalus zone. Coll. Boonstra.

Fibula of Pavascapanodon. S.A.M. 11989.

(x %.) .a. Dorsal view. b. Ventral view.

c. Anterior view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 285

Jonkeridae

Family Characters of the Girdles and Limbs.

The pectoral girdle is large and massive; the height is 650-780 mm. and

the length of the coracoidal plate is 385-534 mm., which is thus 59-68%

of the height. The scapula is fairly high to high (530-665 mm.) and the

upper end of the blade is fairly broad (220-360 mm.); the scapular head

of the triceps is attached to a low or a prominent mound.

The precoracoid is fairly large, but not very massive (315-330 X

235-270 mm.).

The coracoid is small to medium-sized, but massive (180-200 xX 220-

282 mm.).

The clavicle is a large bone (420?-520? mm.) with a greatly constricted

waist.

The cleithrum is strong (length 450? mm.).

The interclavicle is a large and massive bone; the length is 396-558 mm.;

width over anterior expansion 292-456 mm.; width over posterior end of

the stem 210-252 mm. and over the waist of the stem 96-180 mm.; the

lateral horns with thickened postero-lateral edge and a deep groove to

receive the clavicle.

The humerus is relatively short and small to fairly large (length 312-510

mm.); the capitellum is very strong and extends far along the ventral face

to reach the plane of the entepicondylar foramen, and sometimes even the

base of the delto-pectoral crest; the processus medialis always lies well

distally — in a plane well distal of that of the caput; the proximal expansion

is relatively great (222-366 mm.); the shaft is short and broad to very broad

(breadth 84-168 mm.); the ventral opening of the entepicondylar foramen

is large and oval.

The ulna has a long, fairly massive shaft, and the radius is a long

robust bone.

In the pelvis the upper edge of the ilium slopes strongly downwards

in posterior direction; the anterior process is strongly everted; the ilio-

fibularis ridge is fairly light to moderately massive; the ilium does not

form the greater part of the acetabulum.

The femur is not large (length 504 mm.); the width over the external

trochanter fairly large (264 mm.).

The tibia and fibula fairly large but fairly slender.

Genus Jonkeria van Hoepen

The generic characters are as for the family.

286 ANNALS OF THE SOUTH AFRICAN MUSEUM

Jonkena truculenta van Hoepen

In our collection there are no specimens of this species with elements

of the girdles and limbs preserved. The following specific characterisation

is thus drawn from Broom’s (14) account.

Scapula fairly high (530 mm.) and the blade fairly wide (220 mm.);

the glenoidal facet faces somewhat externally. The precoracoid just enters

the glenoid; it is high (315 mm.) and fairly long (235 mm.); the anterior

edge is thin. The coracoid is 200 mm. high and 220? mm. long. The

cleithrum is, according to Broom, small and probably fairly similar to that

of Moschops. Length of clavicle + 520 mm. Interclavicle has a probable

length of 520 mm., width anteriorly 360 mm., width of stem 120-240 mm.

Humerus, ulna, radius, manus, pelvis and femur unknown.

Length of tibia 248 mm., and of fibula 240 mm.

The intermedium is strong and the fibulare a thin bone.

Type: Transv. Mus. 212. Right scapulo-coracoid, right and part of left

clavicle, part of the right cleithrum, part of interclavicle, right tibia,

fibula, intermedium and fibulare, associated with a good skull.

Abrahamskraal, Prince Albert. Low Tapinocephalus zone. Coll.

van Hoepen.

Jonkena angusticeps (Broom)

In our collection there is no specimen referable to this species. Broom’s

(14) figure of the ischium of the type is very misleading. With our increased

knowledge it is now evident that of the ischium figured by Broom nearly

half of the bone, viz. that part forming the contact with the pubis, is missing

and what Broom figured was the dorso-posterior part only.

Type: A.M.N.H. 5633. A very imperfect ischium, associated with a

good lower jaw. Abrahamskraal, Prince Albert. Low Tapmocephalus

zone. Coll. van der Byl.

Jonkeria crassus (Broom)

With no specimen in our collection the following characterisation is

extracted from Broom’s (14) account.

The scapula is simply mentioned but not described by Broom.

The humerus is relatively short (440 mm.), very massive, with greatly

expanded ends (proximal 345 mm. and distal 322 mm.); the shaft is very

short and broad (135 mm.); the delto-pectoral crest is very long and powerful

and nearly reaches the plane of the entepicondylar foramen; the caput is

GIRDLES AND LIMBS OF DEINOCEPHALIA 287

widely oval; the processus medialis lies well distal of the plane of the caput;

the capitellum is strong, thick and well modelled and extends very far

proximally along the ventral surface right up to the plane of the entepicondylar

foramen; the L.M.L. is strong with a strong swelling on the dorsal surface

of the shaft; the entepicondyle is very strongly developed as a thick flange,

with the ventral opening of the foramen large and broadly oval, and it lies

well away from the edge of the bone, nearly under the base of the delto-

pectoral crest; the ectepicondyle is developed as a thick sheet of bone,

pierced obliquely by the small rounded foramen lying near the edge of

the bone.

The ulna is massive; the width over the coronoid process is 217 mm.

The ilium is not figured or described by Broom.

Type: A.M.N.H. 5577. Scapula, humerus, part of ulna and ilium,

associated with the dentaries. Kruidfontein, Prince Albert. Low

Tapinocephalus zone. Coll. van der Byl.

Jonkena haughtom (Broom)

With about a dozen specimens in the collection a fairly full characterisation

of this species can be given.

The scapula is high to very high (590-665 mm.), with the upper part

of the blade broad to very broad (300-360 mm.); the tricipital ridge is fairly

to quite prominent; the glenoidal facet of the scapula faces mainly postero-

ventrally and but slightly outwards; the precoracoid forms the anterior part

of the rim of the glenoid. The massive coracoid has a large glenoidal facet

facing much externally.

The clavicle is represented by the upper half of the nght clavicle; the

upper end is moderately expanded (less than in Parascapanodon), with its

antero-dorsal corner produced as a process, which overlies the lower end of

the cleithrum, fitting into a groove in that bone, it has a strongly constricted

waist and its lower end is probably well expanded.

The cleithrum is a long (440? mm.) fairly slender bone, with its dorsal

end spatulate; in its lower half the posterior two-thirds of the outer face is

grooved for the reception of the clavicle; the inner face is excavated along

its posterior edge and in this recess the edge of the scapula fits. The

cleithrum extends dorsally of the upper edge of the scapula.

The interclavicle is not preserved.

The humerus is massive but fairly short (456 mm.); the proximal

expansion is very great (width 324 mm.) and the distal end is also wide

(306 mm.); the shaft is very short and very broad (diams. 168 X I14 mm.);

288 ANNALS OF THE SOUTH AFRICAN MUSEUM

the delto-pectoral crest is very long and nearly reaches the plane of the

entepicondylar foramen; the caput is oval in outline; the processus medialis

lies in a plane well distally of the plane in which the caput lies; the processus

lateralis forms the most proximal point of the humerus; the capitellum is

strong, thick, well modelled, and it extends proximally along the ventral

face to reach the plane in which the entepicondylar foramen lies; the twist

on the shaft is moderate (10°); the L.M.L. is well developed with a strong

swelling on the dorsal surface of the shaft; the A.D.V.L. is strongly developed;

the entepicondyle is strongly developed as a thick plate of bone, with the

ventral opening of the foramen large and broadly oval, and it is situated

far away from the edge of the bone; the ectepicondyle is developed as an

outflaring thick sheet of bone pierced obliquely by a small foramen IyANg

well away from the edge of the bone.

The ulna is large and massive (length 402 mm.), wide over the coronoid

process (222 mm.); the dorsal rim to the sigmoid face is strong and prominent.

The radius is a large (length 320 mm.) robust bone with a strong

proximo-postaxial flange for the biceps.

The manus is unknown.

In the pelvis the supra-acetabular part of the ilium is high (280 mm.)

and relatively short (380? mm.), with the height thus 73% of the length;

although not fully preserved the anterior iliac process appears to have been

fairly short, but fairly high and fairly strongly everted; the posterior iliac

process is short, with its postero-ventral edge folded over to form a fairly

low and fairly weak oblique ilio-fibularis ridge, which is only slightly

thickened in its upper part. The pubes are not preserved and both ischia

are incomplete but these appear to have been short.

Both the femur and tibia are too inadequately preserved to merit a

description.

The fibula is of moderate robustness and length (294 mm.); the distal

end is rather weak and flattened.

The pes is not preserved.

Type: S.A.M. 4343. An imperfect scapula, coracoid and precoracoid

(Fig. 75); an imperfect right clavicle (Fig. 76 c-d) and right cleithrum

(Fig. 76 a-b); a good right and an imperfect left humerus (Fig. 77);

a good left ulna (Fig. 78 a, a’); the distal end of a radius; an

incomplete pelvis (Fig. 79); an imperfect femur; the proximal end of a

tibia and a good fibula (Fig. 80), associated with a fairly good skull.

Welgemoed, Prince Albert. Low Tapinocephalus zone. Coll.

Haughton.

GIRDLES AND LIMBS OF DEINOCEPHALIA

fig. 75.

oF Paw = -= —

Scapulo-coracoid of Jonkeria haughton. S.A.M. 4343.

Lateral view.

(Xx

..)

289

ANNALS OF THE SOUTH AFRICAN MUSEUM

290

Fig. 76.

(e)

-e

Outer view of right

d. Inner view of right

Jonkeria haughtoni.

a. Outer

cleithrum.

cleithrum. c.

clavicula.

(xX é-)

clavicula.

291

"MOIA [VUNXOIG “Pp ‘MOTA IOLIoyUy “9 “MOTA

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GIRDLES AND LIMBS OF DEINOCEPHALIA

292 ANNALS OF THE SOUTH AFRICAN MUSEUM

LAPS

+f ~

"eae

tee:

Ulnae of Jonkeria haughtoni. (x %.) a and a’. Dorsal and anterior

views of S.A.M. 4343. b and b’. Dorsal and anterior views of

S.A.M. 11464. c and c’. Dorsal and anterior views of S.A.M. go02.

GIRDLES AND LIMBS OF DEINOCEPHALIA 293

eu,

os whats ©

see F

. *

Above: Jonkeria haughtom. S.A.M.

4343. (x#-) a. lium in dorsal

view. b. Ilium in posterior view.

c. Ilium in lateral view. d. Ischium

in lateral view.

Left: Right fibula of Jonkena

haughion. S.A.M. 4343. (x #4

a. Dorsal view. b. Ventral view.

c. Anterior view.

2904 ANNALS OF THE SOUTH AFRICAN MUSEUM

Referred specimens in the S.A.M. collection:

S.A.M. 4342. An isolated scapula (Fig. 81). lLeeurivier, Beaufort

West. Low Tapinocephalus zone. Coll. Haughton.

(

‘ i) c

(

ae (

~ i]

~ =

=~ ese -—- - ee

Scapula of Jonkeria haughtoni. S.A.M. 4342. (x #@.) a. Lateral view.

b. Posterior view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 295

S.A.M. gooz. An isolated scapula (Fig. 82) and a distorted ulna

(Fig. 78 c, c’). In outline the posterior border of the scapula has

a distinctive step also shown in S.A.M. 4342, and probably also in

the type where this edge is not preserved. Klein-Koedoeskop,

Beaufort West. Low Tapinocephalus zone. Coll. Boonstra.

Fig. 82.

ott

oe : b

Ue para

le» ee ete

eit’. Sari ar

recat’ yas

ee me eed, e*.

ore eye 8 8 8

y- .

= -

Men e Coe

Scapula of Jonkeria haughtoni. S.A.M. 9002. (x #@.) a. Lateral view.

b. Posterior view.

206

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 83.

Left: Scapula of Jonkeria haughtoni.

S.A.M. 19207. (x ¢.) Lateral view.

Below : Right radius of /Jonkena

haughtoni. S.A.M. 9145. (x @.) a. Dorsal

view. b. Ventral view. c. Posterior view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 297

S.A.M. 11297. An isolated scapula (Fig. 83). Boesmansrivier, Beaufort

West. Mid Tapinocephalus zone. Coll. Boonstra.

S.A.M. goog. An isolated precoracoid. This medium-sized bone,

composed of a thin sheet except at the apex, where the sutural faces

for the coracoid and scapula are fairly strong, is very similar to

that of J. truculenta but larger, and is probably that of J. haughtoni.

Klein-Koedoeskop, Beaufort West. Low Tapinocephalus zone. Coll.

Boonstra.

S.A.M. 9145. A good radius (Fig. 84) and an incomplete pubis probably

belong to this species. Seekoeivlei, Beaufort West. Low Tapino-

cephalus zone. Coll. Boonstra.

S.A.M. 9147. An isolated ulna with a damaged proximal end agrees

sufficiently well with that of the type to be included in this species.

Seekoeivlei, Beaufort West. Low Tapinocephalus zone. Coll.

Boonstra.

S.A.M. 11464. A very well preserved ulna (Fig. 78 b, b’); although

somewhat shorter than that of the type it can safely be included in

this species. Koedoeskop, Beaufort West. Mid Tapinocephalus zone.

Coll. Boonstra and Avenant.

Jonkenia ingens (Broom)

In the specimen described by Broom (14) as Jonkeria pugnax but by

me (1) considered a synonym of Jonwkena ingens Broom mentions the

scapulae but without figuring or describing them; and of the humerus he

states that it resembles that of Jenkeria crassus very closely.

In this collection there are four humeri, which in point of size can best

be tentatively referred to this large species of Jonkeria, rather than to any

other of the Titanosuchids.

The humerus in these specimens is large, massive, but not very long

(length 504-510 mm.); both ends are greatly expanded (proximal 330-336

mm., and distal 3122-324 mm.); the shaft is short and broad (diams. 130-156

x Ir0 mm.); the delto-pectoral crest is long and powerful, but terminates

well proximal of the plane in which the entepicondylar foramen lies; the

caput is large and widely oval; the processus medialis lies very far distally

of the plane in which the caput lies; the capitellum is strong, thick and very

well modelled and it extends far proximally along the ventral face, but does

not reach the plane of the entepicondylar foramen; the L.M.L. is strong;

the A.D.V.L. is not very sharp; the entepicondyle is strongly developed as

a thick flange of bone, with the ventral opening of the entepicondylar foramen

fairly large and oval, lying fairly near the edge of the bone well removed

from the base of the delto-pectoral crest; the ectepicondyle is a flaring sheet

298 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 85.

ast .

ee ‘ oe

Dna ex co

O10 COPEL Cis Sy

Cc.

Humeri of Jonkeria ?ingens. CX es) jal.) SHACME iazia3t

Ventral view. b. S.A.M. 738.

Ventral view. c. S.A.M. 738. Dorsal view. d. S.A.M. 738. Proximal view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 299

of bone pierced vertically by the small foramen lying well away from the

edge of the bone.

Referred specimens:

A.M.N.H. 5608. Much of two scapulae and a humerus, associated with

a skull. Kookfontein, Prince Albert. Mid Tapinocephalus zone.

Coll. van Wyk.

S.A.M. 738. A good isolated humerus (Fig. 85 b-d). Gamka River.

Low? Tapinocephalus zone. Coll. Cloete.

S.A.M. 3433. A good isolated humerus (Fig. 85a). Janwillemsfontein,

Prince Albert. Low Tapinocephalus zone. Coll. Haughton.

S.A.M. g006. An isolated proximal half of a humerus. Klein-

Koedoeskop, Beaufort West. Low Tapinocephalus zone. Coll.

Boonstra.

S.A.M. 11994. An incomplete isolated humerus with a good distal end.

Welgemoed, Prince Albert. Low Tapinocephalus zone. Coll.

Haughton.

S.A.M. 9348. An incomplete and distorted humerus and an ilium lacking

both the anterior and posterior processes (Fig. 86). The ilium is

larger and more strongly built than those of the other species of

Jonkeria and may provisionally be considered to represent that of

Jonkeria ingens. Mierfontein, Beaufort West. Mid Tapinocephalus

zone. Coll. Boonstra.

Fig. 86.

ee ae,

Ilium of Jonkeria ?ingens. S.A.M. 9348.

(x %.) Lateral view.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 87.

300

c .

Ca eer Yt

rh 2.

feuet

“Neg see

(x 8.) a. Pelvis in lateral view.

Jonkeria koupensis. S.A.M. 9004. :

b. Ilium in dorsal view. c. Pelvis in anterior view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 305

Jonkeria koupensis Sp. Nov.

This new species appears to be very closely related to J. haughtoni, but

the differences in the pelvis, as will be evident from the accompanying

characterisation, warrant treatment as a distinct form.

In the pelvis the pubo-ischiadic plate is probably short (83% of the

height as reconstructed); the supra-acetabular part of the ilium is high (264

mm.) and the antero-posterior length of the iliac blade is very short (282

mm.), so that the height is 93% of the length; the anterior iliac process is

fairly short, but appears shorter than it really is because of its strong eversion

antero-laterally, and is fairly high; the posterior process is short and fairly

low, with its postero-ventral edge moderately strongly folded over to form

a fairly strong ilio-fibularis ridge on the outer face; this ridge is directed

obliquely upwards, with its upper end not strongly bulbously thickened; a

slight groove on the inner face of the everted anterior iliac process indicates

the attachment of a rib anterior to the main sacral rib; antero-posteriorly the

outer face of the iliac blade is deeply concave.

The antero-ventral edge of the pubis is strongly everted, with the

tuberculum pubis confluent with the thickened part of the antero-ventral

edge, which stretches to the median line where the pubes meet, but do not

form a real symphysis.

The ischium is not preserved but probably is as reconstructed.

The ulna has the distal two-thirds preserved; it is a massive bone with

the dorsal lip of the sigmoid face developed into a massive swelling in its

preaxial part.

Fig 88. Type: S.A.M. 9004. A good pelvis,

; lacking only the ischia (Fig.

87) and the distal two-thirds of

the ulna. Klein-Koedoeskop,

Beaufort West. Low Tapino-

cephalus zone. Coll. Boonstra.

Referred specimen:

S.A.M. 11983. An isolated ilium

(Fig. 88). | Abrahamskraal,

Prince Albert. Low Tapino-

cephalus zone. Coll. Haughton.

Illium of Jonkeria koupensis. S.A.M.

11983. (x %.) Lateral view.

ANNALS OF THE SOUTH AFRICAN MUSEUM

302

"MaIA [esIogd “q

‘MOTA [PUIITXOIG "Pp ‘MOIA IOLIOJUY ‘9

“MOIA [eI]UDA “Be (°2 X)

‘6b16 “WV'S

‘pasvg visayuof Jo snxsWINn;T

GIRDLES AND LIMBS OF DEINOCEPHALIA 393

Jonkena parva Sp. Nov.

Although only a single isolated humerus is known, this bone is so

distinctive that I do not hesitate to make it the type of a new species of

Jonkena.

This humerus is the smallest Jonkeria humerus as yet known; its length

is only 312 mm., but it is a very massive element with both the proximal

(222 mm.) and the distal (252 mm.) ends very greatly expanded; the shaft

is very short and the bone is greatly constricted in the waist; the diameters

of the shaft are 84 x 78 mm.; the delto-pectoral crest is fairly short and it

terminates very far proximal of the plane in which the entepicondylar

foramen lies; it has a very massive ventral edge and it terminates as a very

thick knob. The caput is weak and its face is straplike, but it forms the

most proximal part of the bone; the processus lateralis lies more distal than

the caput; the processus medialis, as in all the Jonkerias, lies well distally

of the caput; the capitellum is very massive indeed and it extends very

far proximally along the ventral face, with its proximal border lying in a

plane proximal to that in which the entepicondylar foramen lies; posteriorly

of the well-modelled capitellum there is a deep groove in which the coronoid

process moved when the ulna was flexed; the twist on the shaft is great

(30°); the L.M.L. is distinct, with a large moundlike muscle scar on the

dorsal surface of the shaft; the A.D.V.L. is very strong and forms a

prominent ridge; the entepicondyle is strongly developed to form a greatly

outflaring thick sheet of bone; the ventral opening of the entepicondylar

foramen is oval and it lies well postaxially, near the edge of the bone; the

ectepicondyle is developed as a greatly flaring thin sheet of bone penetrated

in its thinner part, near the edge, by a small round ectepicondylar foramen.

Type: S.A.M. 9149. An isolated right humerus very well preserved

(Fig. 89). Saairivier, Prince Albert. Low Tafinocephalus zone.

Coll. Boonstra.

Jonkeria rossouwi Sp. Nov.

The specimen on which this new species is founded was found by Broom

(14) to be different from the then known species of Jonkeria, but Broom,

with a restraint unusual for him, did not name it. I am naming it Jonkeria

rossouw?t for Mr. P. J. Rossouw of the Geological Survey of the Union in

recognition of the importance of his stratigraphical work on the Tapino-

cephalus zone.

In this species the pectoral girdle is fairly large and fairly massive. The

scapula is fairly low (height 552 mm.) and the upper part of the blade is

greatly expanded (width 324 mm.); the tricipital bulge is very prominent;

the supra-glenoidal edge forms a strong raised rim; the internal opening of

304 ANNALS OF THE SOUTH AFRICAN MUSEUM

the supracoracoid foramen opens into the deep subscapular groove; the

glenoidal facet of the scapula faces ventro-posteriorly but not externally.

The precoracoid is long but low (somewhat affected by crushing); the foramen

pierces the bone very obliquely. The coracoid is small but massive, with

a large glenoidal facet facing well externally.

The interclavicle is massive but short (480? mm.) with the stem wide

posteriorly and with a narrowed waist anteriorly; the anterior spatulate end

curves upwards very sharply and has a deep groove on its outer antero-

Fig. 90.

Jonkeria vossouwi. S.A.M. 5014. (> %.) a. Scapulo-coracoid in lateral view

b. Scapula in posterior view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 305

lateral face for the reception of the postero-ventral edge of the clavicle; on

the dorsal surface of the stem there is a strong medial ridge against which

the precoracoids abut.

The clavicles and cleithrum are not known.

The humerus is fairly short (378 mm. in length), but massive with greatly

expanded proximal (330 mm.) and distal (276? mm.) ends; the shaft is very

short, thick and broad (132 x 84 mm.); the delto-pectoral crest is very long

and nearly reaches the plane in which the entepicondylar foramen lies;

the caput is broadly oval; the processus lateralis lies well proximally, in the

same plane as the caput, whereas the processus medialis lies well distally;

the capitellum is fairly massive and extends well along the ventral face

and nearly reaches the plane of the entepicondylar foramen; the ‘‘twist’’ on

the shaft is fairly small (15°); the L.M.L. is fairly strong with muscle scars

on the dorsal surface of the shaft; the A.D.V.L. is well developed; the

entepicondyle is strong with the ventral opening of the foramen large and

nearly round; the ectepicondyle forms a thick flange pierced by the small

foramen situated well away from the edge of the bone.

(The dorsal surface of the ectepicondyle shows a pathological lesion in

the form of an irregular deep excavation and this is accompanied by a

concomitant outgrowth of the olecranon of the ulna. In another humerus,

that of Phocosaurus S.A.M. 11300, the disto-ventral face of the capitellum is

deeply eroded with subsequent healing and the formation of a new but

concave instead of a convex articulatory facet. (Osteitis fibrosa?

Osteomalachia ? ))

The ulna has suffered pathological deformation so that the olecranon is

proximally prolonged as a thick outgrowth. The normal ulna would appear

to have had a more slender shaft and a weaker coronoid process than the

other known species of Jonkenia.

The radius is only slightly pathologically deformed. Its length is 312 mm.

and the flange on the proximo-postaxial corner is weak.

The manus is not known.

In the pelvis the supra-acetabular part of the ilium is high (288 mm.)

and relatively short (336 mm.) so that the height is 87% of the length; the

anterior iliac process is relatively short, but fairly high and strongly everted;

the posterior process is short and fairly low, with its postero-ventral edge

folded over strongly to form a strong vertical ridge, which is dorsally strongly

bulbous, and it projects strongly laterally; on the inner face of the anterior

iliac process no distinct facet is preserved for the attachment of a rib lying

anterior to the main sacral rib.

No pubis or ischium is preserved.

The femur is fairly long (504 mm.); fairly broad over the external

trochanter (264 mm.), which is not separated by a notch from the proximal

306 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 91.

oa:

=a:

~ .

“oe ae.

aan we

° aly

wit) ¢

om! ee

Interclaviculae. (x %.) a. Jonkeria rossouwi. S.A.M. 5014. Ventral view.

b. Jonkeria rossouwi. S.A.M. 5014. Lateral view. c. Jonkeria sp. indet.

S.A.M. 9124. Ventral view. d. Jonkeria sp. indet. S.A.M. 9124. Lateral view.

wr,

GIRDLES AND LIMBS OF DEINOCEPHALIA

jesioq “q

"MOIA [VWIXOIG “p

“MOIA [eIjUeA “eB (°2 X)

‘MOIA IOWSJUY “9 “AMOIA

‘troS “JuTW'S ‘“tanossos pisayuof JO SnisUIN}{

5—Annals

308

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 93:

Left radius of Jonkeria rossouwi. S.A.M. 5014. (x 2.)

a. Dorsal view. b. Ventral view. c. Posterior view.

Right ilium of Jonkeria rossouwi. S.A.M. 5014. (x #.) a. Dorsal view.

b. Posterior view. c. Lateral view. d. Anterior view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 309

face; the caput is fairly thick (114 mm.) and is directed much preaxially;

the shaft is broad but flat (diams. 156 x 84 mm.); the femoro-tibialis ridge

is fairly strong; the distal facets of the femur are small and directed much

distally, especially the ectepicondyle which lies far distally. (These features

of the distal end of the femur, differing as they do from the other Jonkerias,

appear to be due to some measure of pathological deformation.)

The tibia is fairly robust (length 300 mm.); the proximal face is inclined

much postaxially to correspond with the distally situated postaxial facet of

the femur.

The fibula is fairly slender and long (330 mm.).

The pes is not known.

Type: S.A.M. 5014. The left scapula and incomplete precoracoid and

coracoid (Fig. 90), an imperfect interclavicle (Fig. 91 a, b), a diseased

left humerus (Fig. 92), a deformed left ulna and a left radius (Fig.

93), a right ilium (Fig. 94), a left femur (Fig. 95), a left tibia (Fig. 96)

and a left fibula (Fig. 97). Abrahamskraal, Prince Albert. Low

Tapinocephalus zone. Coll. van der Byl.

Fig. 95.

Left femur of Jonkeria rvossouwi. S.A.M. 5014. (X é-)

b. Ventral view. c. Anterior view.

| 6—Annals

310 ANNALS OF THE SOUTH AFRICAN MUSEUM

Referred specimens:

S.A.M. 11979. An incomplete interclavicle (Fig. 98 a, b) and the

proximal ends of an ulna and fibula. Kruisvlei, Beaufort West.

Low Tapinocephalus zone. Coll. Boonstra.

S.A.M. 11982. A fairly good interclavicle (Fig. 98 c, d). Abrahamskraal,

Prince Albert. Low Tapinocephalus zone. Coll. Haughton.

Fig. 96.

Left tibia of Jonkeria rossouwi. S.A.M. 5014. (xX

view. b. Ventral view. c. Posterior view

Re eee]

je)

(2)

(op)

job)

—

Left fibula of Jonkeria rossouwi. S.A.M. 5014. (x @.)

a. Dorsal view. b. Ventral view. c. Anterior view.

art

GIRDLES AND LIMBS OF DEINOCEPHALIA

‘SG

Ventral

Ventral view.

c. S.A.M. 11982.

(SC at) ao Sra rT arg7 0:

11982 Lateral view.

11979. Lateral view.

d. S.A.M.

Interclaviculae of Jonkeria rossouwt.

bz SAM.

view.

312 ANNALS OF THE SOUTH AFRICAN MUSEUM

Jonkena Sp. Indet.

S.A.M. 9124. A good interclavicle (Fig. 91, c, d). Voélfontein, Prince

Albert. Low Tapinocephalus zone. Coll. Boonstra.

This interclavicle is larger and stronger than those of the oe species

of Jonkeria and may thus possibly be J. ingens.

S.A.M. 11984. An imperfect isolated ilium (Fig. 99) of an unknown

locality. It would appear to lie fairly close to the ilium of

J. haughtont.

Right ilium of Jonkeria sp. indet. S.A.M.

11984. (x %.) Lateral view.

S.A.M. 11886. A nearly complete interclavicle (Fig. Ioo c, d).

Vindraersfontein, Beaufort West. Low Tapinocephalus zone. Coll.

Boonstra.

This small bone with a narrow waist and the strongly expanded posterior

end of the stem is apparently of a small species of Jonkeria — possibly

of J. parva.

Titanosuchia Genera Indet.

5.A.M. 11571. A large isolated interclavicle with a strongly upturned

anterior end and a strong median keel on the dorsal face of the stem

(Fig. 100 a, b). It has the long stem of the Tapinocephalians but

the width is that usually seen in the Titanosuchians. Probably a

large Titanosuchid. Klein-Koedoeskop, Beaufort West. Low

Tapinocephalus zone. Coll. Boonstra.

GIRDLES AND LIMBS OF DEINOCEPHALIA S53

Re ice ete:

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SoS eee RIOR ROC

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Interclaviculae. (x %.) a. Titanosuchian gen. indet. S.A.M. 11571. Dorsal

view. b.

cf. parva.

Titanosuchian gen. indet. S.A.M. 11571. Lateral view. c. Jonkeria

S.A.M. 11886 Ventral view. d. Jonkeria cf. parva. S.A.M. 11886.

Lateral view.

314 ANNALS OF THE SOUTH AFRICAN MUSEUM

S.A.M. 11986. An isolated ilium (Fig. roz). Letjiesbos, Beaufort

West. Mid Tapinocephalus zone. Coll. Boonstra.

This ilium has on the outer face of the posterior iliac process neither the

horizontally placed ilio-fibularis ridge, usually seen in the Tapinocephalia,

nor the obliquely placed ridge, usual in the Titanosuchia. The area of

origin of the ilio-fibularis is rather a thickening of the postero-ventral edge

without any eversion or folding over. In outline this illum agrees more

with those of the Titanosuchia.

Ilium of a ? Titanosuchian. S.A.M. 11986. (x €.) \

a. Lateral view. b. Dorsal view.

Anteosauria

Infra-ordinal Characters of the Girdles and Limbs.

The pectoral girdle is light; the scapular head of the triceps originates

from a prominent tubercle; the outer face of the precoracoid is strongly

convex and the glenoidal facet on the coracoid is bipartite.

GIRDLES AND LIMBS OF DEINOCEPHALIA B15

The humerus is fairly light; without an ectepicondylar foramen; the radial

condyle is well modelled, but situated well distally; both the processus

lateralis and medialis lie far proximally; the delto-pectoral crest is very short

and is directed much postaxially.

In the manus the carpal formula is 3, I, 5, and the phalangeal formula

3) 3, 4, 42, 2.

The iliac blade is low and the acetabulum faces much ventrally; the

ilio-fibularis ridge lies horizontally; no supra-acetabular buttress is developed.

The femur is light, long and slender with the internal trochanter developed

as a ridge; the width over the external trochanter is very small.

In the pes the tarsal formula is 2, 1, 5 and the phalangeal formula

ora 4!» 2.

Anteosauridae

Family Characters of the Girdles and Limbs.

The Anteosaurid pectoral girdle is only imperfectly preserved in two

specimens; but, even so, it is evident that the Anteosaurid girdle differs

greatly from that of all the other Deinocephalia.

The girdle is of moderate size and is lightly built. No complete scapula

is known but it would appear that it was of moderate width and height

(455-475 mm. as reconstructed); instead of a tricipital mound or ridge, as

in all the other Deinocephalia, there is developed (for the origin of the

scapular head of the triceps) a prominent tubercle on the posterior edge of

the scapula 25-35 mm. above the glenoidal edge; the internal opening of the

foramen supracoracoideus opens into the fairly deep subscapular groove;

the glenoidal facet of the scapula is large and faces ventro-posteriorly, but

also much externally, and the outer and posterior margins are sharp,

prominent and well moulded.

The coracoidal plate is fairly long antero-posteriorly. The precoracoid is

relatively large; its dorso-posterior apex, just posterior to the outer opening

of the foramen supracoracoideus, forms a fairly long margin to the glenoid;

here the outer face of the precoracoid forms a decided depression or recess

to house the processus lateralis of the humerus, when this bone is in its

forward position; the foramen supracoracoideus penetrates the bone obliquely

so that it internally forms a groove in the sutural face and thus opens into

the subscapular groove; the outer precoracoidal face is strongly convex, so

that its lower edge is directed much medially.

The coracoid is small and light, but greatly thickened where it houses

the lower half of the glenoid; this facet is bipartite — the inner and posterior

moiety receives the head of the humerus in its posteriorly disposed position,

and the outer and anterior moiety receives the humerus when it is disposed

316 ANNALS OF THE SOUTH AFRICAN MUSEUM

anteriorly; the inner part of the facet is directed dorso-posteriorly and slightly

externally, whereas the outer part is directed much externally.

Nothing is known of the cleithrum, clavicula and interclavicula.

__ The humerus, known only in three specimens, is fairly lightly built and

fairly small to medium (length 300?-375 mm.); the proximal expansion is

fairly small (width 162-168 mm.); the distal expansion is somewhat greater

(width 200-205 mm.); the shaft is fairly long and slender (diams. 76-84 xX

48-65 mm.); the delto-pectoral crest is very short and terminates a consider-

able distance proximal to the ventral opening of the entepicondylar foramen,

and is directed much postaxially; the caput is narrowly oval; the processus

medialis lies in the same plane as the caput, whereas the processus lateralis

lies a little further proximally; the radial condyle is situated distally, but

extends a little along the ventral face and also curves on to the dorsal face;

the ‘“‘twist’’ on the shaft is large (40°); the L.M.L. is fairly definite and the

A.D.V.L. is sharp and prominent; the entepicondyle is fairly greatly

expanded, with the foramen entering dorso-postaxially and leaving as a

ventral slit; the ectepicondyle is well expanded as a thin plate of bone

housing on its ventral face a groove for the passage of the radial vessels.

The only radius known is a light, slender featureless bone and the only

known ulna (length 260 mm.) is crushed flat.

The manus in the only specimen known has the carpal formula 3, I, 5

and the phalangeal formula 3, 3, 4, 4?, 2.

The pelvis is only known from one imperfect ium. This, however,

shows that the Anteosaurids had an ilium of a type radically different from

any known in the South African Deinocephalia. It is without the strong

supra-acetabular buttress so typical of the Deinocephalia; instead, the

acetabulum is dorsally bounded by a continuous curved ridgelike margin

without a supra-acetabular notch; the thickly rounded posterior edge of the

fairly short posterior process, from which the ilio-fibularis originates, lies

horizontally as in the Tapinocephalia; the iliac blade was presumably short

and low.

In Fig. ro4 an orthoprojection, taken by pantograph, vertically on to

the iliac part of the acetabulum (as is the case in all the figures showing

the lateral aspect of the pelvis, i.e. a projection on to the median plane) is

shown. But I believe that in the naturally articulated skeleton the iliac

portion of the acetabulum, in this case, would be directed much more

ventrally so that the disposition of the femur in the Anteosaurids would be

radically different from the condition in all other Deinocephalia.

The femur of the Anteosaurids looks very different from that of the

other Deinocephalians; although not very long (210-420 mm.) it appears

very long because of its general slenderness and long shaft; the width over

GIRDLES AND LIMBS OF DEINOCEPHALIA a7

the external trochanter is small and varies from 47 to 120 mm.; the shaft

is narrow and has diameters ranging from 28 x 18 to 75 X 45 mm.; the

external trochanter is situated far proximally and nearly in the same plane

as the caput; the caput is short pre-postaxially and flows directly into the

external trochanter; with the preaxial border deeply concave and the postaxial

border convex the femur is a curved bone with the caput well preaxially

and somewhat dorsally directed; the internal trochanter forms a low ridgelike

tubercle situated well away from the preaxial face, with a resulting reduced

inter-trochanteric fossa; the femoral condyles face much distally with the

external one further distally than the internal one; the popliteal fossa is

elongated; the epicondyles, in one specimen, form a thin sheetlike expansion,

both pre- and postaxially; the femoro-tibialis ridge is weak and the area

for the insertion of the ilio-femoralis is narrow; there is a strong ridge for

the insertion of the pubo-ischio-femoralis internus.

The tibia is unknown.

The fibula is a lightly built long and slender bone with its proximal end

strongly expanded and with the facets for the femur nearly terminal.

Whereas the proximal end is flattened, the distal facet is broadly oval in

outline; the preaxial face is fairly straight, whereas in the other Deinocephalia

this is deeply concave.

The pes has the tarsal formula 2, I, 5 and the phalangeal formula

uneeriaim but probably 2?, 3°, 3°, 42, 2.

Anteosauridae

The Family Characters are as for the infra-order.

Genus Anteosaurus Watson

In this genus the girdles and limbs are very inadequately known. The

generic characters that can be determined from an incomplete scapula, an

incomplete and distorted humerus, three coracoids, an incomplete ilium, two

good and two incomplete femora are as follows: Tricipital tubercle elongated,

35 mm. above the edge of the glenoid; humerus fairly small (length 300°

mm.); the shaft is slender (diams. 60 x 48 mm.); the ilium is as described

for the infra-order; the femur is slender (length 402?-420 mm.); width

across the external trochanter 108-120 mm.; the epicondyles with moderate

or pronounced sheetlike expansions.

Anteosaurus acutirostris (Boonstra).

Femur 420 mm. in length, with flaring epicondyles.

S.A.M. 11977A. Two good femora (Fig. 102 a-c), associated with a good

skull (S.A.M. 9329). Kruisvlei, Beaufort West. Low Tapinocephalus

zone. Coll. Boonstra.

318

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 102.

Femora. (x %.-) a. Anteosaurus acutirostris. S.A.M. 11977A.

Dorsal view. b. Anteosaurus acutivositns. S.A.M. 11977A. Ventral

view. c. Anteosaurus acutivostris. S.A.M. 11977A. Anterior

view. d. Anteosaurus sp. SaA.Miin 2753 A. Ventral view.

e. Paranteosaurus primus. S.A.M. 11485. Ventral view. f. Anteo-

saurus sp. S.A.M. 5614. Ventral view.

GIRDLES AND LIMBS OF DEINOCEPHALIA 319

‘Anteosaurus lotz: (Broili and Schroder)

Alte Akademie. No. ?. Parts of peaalinats pubis and femur, associated

with some skull pieces. Brakleegte, Beaufort West. High? Tapino-

cephalus zone. Coll. Schroder.

Little can be said about these fragments, but if the pubis is correctly

determined and is as figured by Broili and Schroder, then this element is

radically different from the pubes of other Deinocephalia.

Anteosaurus spp.

_§.A.M. 2752. A partial distorted humerus and a coracoid, associated

with a skull. Viviers Siding, Beaufort West. Mid Tapinocephalus

zone. Coll. Haughton and Whaits.

Fig. 103.

Scapulo-coracoids. (xx %.) a. Anteosaurus sp. S.A.M. 5614. Lateral

view. b. Eccasaurus priscus. S.A.M. 11597. Lateral view.

320 ANNALS OF THE SOUTH AFRICAN MUSEUM

S.A.M. 2753A. Proximal and distal ends of a femur (Fig. 102d),

associated with a skull fragment. Vjiviers Siding, Beaufort West.

Mid Tapinocephalus zone. Coll. 2

Haughton and Whaits. Fig. 104.

S.A.M. 5614. Part of a scapula (Fig.

103a), an incomplete ilium (Fig. 104) wn ores

and the proximal end of a femur (Fig. ait, i iba ecg

1o2f), associated with two dentaries. ates eh alta

BSc S30 a ri :

bitte “G

te seis

Abrahamskraal, Prince Albert. Low

Tapinocephalus zone. Coll. Haughton.

S.A.M. 7396. An isolated coracoid. Abra-

hamskraal, Prince Albert. Low

Tapinocephalus zone. Coll. van der Byl.

S.A.M. 11887. An _ isolated coracoid.

Vindraersfontein, Beaufort West. Low Right ilium of Anteosaurus sp.

Tapinocephalus zone. Coll. Boonstra. SN

Genus Paranteosaurus Boonstra

Paranteosaurus primus Boonstra

S.A.M. 11485. The proximal end of a femur (Fig. 102e), associated with

a skull. Mynhardtskraal, Beaufort West. Low Tapinocephalus zone.

Coll. Boonstra.

This fragment shows no features by which it can be distinguished from

the femur of Anteosaurus.

Genus Micranteosaurus Boonstra

The femur is slender and curved (length 210 mm.); the width over the

external trochanter is 47 mm.; the shaft is long and narrow (diams. 28 x 18

mm.); a ‘‘twist’’ on the shaft causes the caput to be directed much dorsally;

the ectepicondyle (and probably also the entepicondyle) is without any sheet-

like expansion.

The fibula is very long (138 mm.) and slender.

The pes has the tarsal formula 2, I, 5 and the phalangeal formula

22,38) Heats 2

The radius (105 mm.) is much shorter than the fibula.

The manus has the carpal formula 3, I, 5 and the phalangeal formula

3, 3) 4, 4?, 2.

GIRDLES AND LIMBS OF DEINOCEPHALIA 320

Micranteosaurus parvus Boonstra

As I (4) have only recently described this form, the above generic

diagnosis of the limbs is extracted from that account, which consult for

the figures.

Type: S.A.M. 4323. A coracoid, radius, manus, femur, fibula and pes,

associated with a snout. Commonage of Merweville, Beaufort West.

Low Tapinocephalus zone. Coll. Haughton.

Genus Eccasaurus Broom

Notwithstanding its very distinctive humerus (which constitutes the type

specimen) Broom (9) included this genus in the Tapinocephalia and later

even referred a typical Tapinocephalian tooth to this genus.

Although Broom in his original description only mentions the humerus

there is in the South African Museum collection, associated with the type

humerus, some skull pieces, two imperfect and weathered femora, a flattened

radius?, some vertebrae, the distal end of a fibula, some other fragments

and a long slender simple pointed tooth, which is probably an incisor.

The generic characterisation below is based on the type supplemented by

features determined from a second specimen (S.A.M. 11597).

In the scapula the tricipital tubercle is oval in outline and is situated

25 mm. above the edge of the glenoid; the other features of the scapulo-

coracoid are as described above for the family. The humerus is 348-375

mm. long and the width of the shaft is 76-84 mm.; other features as for

the family.

The only known ulna is crushed dorso-ventrally; the length is 260 mm.

The pelvis is unknown.

The femur is 355 mm. in length and fairly slender; the width over the

external trochanter (120 mm.) is larger than in Anteosaurus; the epicondyles

are not laterally expanded as thin flanges.

There is thus no doubt that Eccasaurus with its long pointed incisor, its

humerus with the delto-pectoral crest very short, its slender femur, where

the external trochanter is situated so far proximally and so little laterally,

and in the referred specimen (referred because of the great similarity in

the humeri) the distinctive tricipital tubercle on the scapula, cannot possibly

be a Tapinocephalian but is in fact an Anteosaurian.

The tooth, which Broom has referred to Eccasaurus, does not belong to

this genus but is typically Tapinocephalian.

Eccasaurus priscus Broom

The specific characters are as for the genus.

322 ANNALS OF THE SOUTH AFRICAN MUSEUM

Type: S.A.M. 915. <A good humerus (Fig. 105), two imperfect femora.

(Fig. 106), distal ends of the two fibulae (Fig. 107a) and a flattened

?radius, associated with parts of the skull. Sandvlakte, Prince

Albert. Low? Tapinocephalus zone. Coll. Cairncross.

Fig. 105,

Humerus of Eccasaurus priscus. S.A.M. 915. (x #%.) a. Ventral view. b. Dorsal

view. c. Anterior view. d. Proximal view.

Referred specimen:

S.A.M. 11597. Part of the nmght coracoscapula (Fig. 103b), an incomplete

humerus and an ulna (Fig. 107b). Dikbome, Laingsburg. Low

Tapinocephalus zone. Coll. Boonstra.

? Anteosauridae Incertae Sedis

S.A.M. 11996. An isolated interclavicula (Fig. 108). Koedoeskop,

Beaufort West. Mid Tapinocephalus zone. Coll. Boonstra.

This interclavicle is small, with a short, fairly narrow stem, and the

anterior widely spatulate end is not curved up sharply. I am tentatively

referring this bone to the Anteosauridae as it cannot belong to any of the

other Deinocephalians, where this element has quite different features.

GIRDLES AND LIMBS OF DEINOCEPHALIA

ane cae

te 8 we

Fibula in ventral view. b. S.A.M.

Ulna in dorsal view.

Femur of FEccasaurus priscus. S.A.M. 915.

(x +4.) a. Ventral view. b. Anterior view.

-Interclavicula of an ? Anteosaurid. S.A.M.

mie904 (0 32) a.- Ventral waew.' b., Lateral

view.

323

Eccasaurus priscus. (x 4.) a. S.A.M. 915.

11597.

324 ANNALS OF THE SOUTH AFRICAN MUSEUM

CERTAIN PALAEOBIOLOGICAL CONCLUSIONS

From the above study it is abundantly clear that we still know very

little about the girdles and limbs of the South African Deinocephalia.

Complete skeletons are unknown and moderately complete skeletons are

extremely rare. In the herbivorous Tapinocephalia we have most specimens

in which a fair amount of the skeleton has been found in association; in all

about ten specimens (half of them of Moschopids) are known with a fair

amount of the skeleton preserved in association. For the rest the specific

and generic accounts are composite and based on a number of individuals,

each with some part of the skeleton preserved, supplementing each other.

In the carnivorous Titanosuchia only two specimens have a fair amount

of the skeleton of an individual animal preserved in association. In the

Anteosauria only a few bones in all are known and in the Styracocephalia

nothing whatever of the postcranial skeleton is known.

An inventory shows that parts of the girdles and limbs are known in

20 genera, and in Ir genera nothing at all is known of this part of the

skeleton.

Now it is of interest to note that articulated partial skeletons have been

more often found in the case of the Tapinocephalia than in the other infra-

orders. This denotes a difference in the vicissitudes of entombment apparently

largely due to differences in habitat. The two or three skeletons found in

isolation were probably entombed in much the same manner as were the

contemporary Pareiasaurs. In the two cases of mass-entombment of

Moschops herds the conditions were different. Broom considered that the

material of Moschops found by Whaits at Spitskop in the Moordenaarskaroo

represents a herd of males, females and young. In the Moschops-material

found by me at Kruisvlei in the Koup there are also males, females and

juveniles, but here they were found together with remains of at least two

individuals of the Anteosauria. There does not appear to be sufficient

evidence for concluding that the two Moschops-finds represent herds over-

whelmed by some catastrophy whilst peacefully feeding. It appears more

probable that, living together in a certain suitable area, the remains were

included in the sediment in a bed where deposition was taking place when

death, not necessarily catastrophically or even simultaneously, overtook

them. At Kruisvlei the bed in which the remains were found is not much

more than about a foot thick and is a fairly fine argillaceous sandstone and

the disposition of the various partial skeletons points to their having been

transported by water for a short distance at least.

The other Tapinocephalian, and by far the majority of the Titanosuchian

and Anteosaurian specimens, were found as isolated specimens consisting

GIRDLES AND LIMBS OF DEINOCEPHALIA 325

of only a few bones not articulated but entombed quite near each other.

The main reason for this scattering of the remains I believe to be due to

the activities of carrion-eaters and the effects of water transport from higher

to lower ground.

The above preliminary study of the locomotor apparatus in three of the

infra-orders of the Deinocephalia has revealed three main types of loco-

motion. Without going deeply into this subject, at this stage, I can briefly

indicate the broad differences that have become apparent.

In the Tapinocephalia the articulatory facets of the elbow-joint are weakly

modelled and must have contained considerable cartilage. In addition the

humeral condyles are situated much distally. This distal situation of the

humeral condyles may indicate a more upright stance of the fore-limb

when standing on firm ground. This is very probably the case in Moschops

and its close relations, but in the Tapinocephalids and in many of the

Struthiocephalids I am more inclined to think that the unfinished articulation

is more indicative of a life spent much in water — but by no means as

aquatic as the later Lystrosaurs. These reptiles living in the marshy parts

of the Karroo basin, often wallowing in the water, but coming on firmer

ground for part of their feeding would then, on death, be more likely to be

entombed as fairly complete cadavers.

In the Titanosuchia the radial condyle is well modelled and the elbow-

joint forms a much stronger and efficient hinge for terrestrial locomotion.

But, the condyle being situated more on the ventral face of the humerus,

this bone would in striding lie much more horizontally than in the Tapino-

cephalia. However, in the hind-limb, where the femoral condyles are placed

much distally, the femur would be approaching a semi-upright position and

the efficient joint with the tibia allows of the transmission of a powerful

thrust by the hind-limb. Moreover, the muscle scars on all the elements

of the stoutly built girdles and limb-bones are strong and indicate a muscular

action much more powerful than in the Tapinocephalia and capable of the

execution of sudden, rapid and forceful movements, which would be necessary

for these carnivores. Thus the Titanosuchia, although having the chest

slung fairly low between the fore-limbs, would be fairly active reptiles of

prey, living chiefly on firm ground with little of the wallowing partaken of

by the Tapinocephalia.

In the Anteosauria little is as yet known of the locomotor apparatus.

The nature of the hip-joint and the structure of the femur seem to point

to a crawling habit, showing, in some respects, much similarity to that of

the modern crocodiles, but the dentition seems to indicate that they were

possibly slinking carrion-eaters.

326

28.

29.

ANNALS OF THE SOUTH AFRICAN MUSEUM

REFERENCES

‘Boonstra, L: D. A Note on the Synonymy of the two Deinocephalians,

Dinophoneus ingens Broom and Jonkeria pugnax (Broom). S. Afr. Jour. Sci. 32,

329-331. 1935.

Boonstra, L. D. Some Features of the Cranial Morphology of the Tapinocephalid

Deinocephalians. Bull. Am. Mus. Nat. Hist. 72, 75-98. 1936.

Boonstra, L. D. The Cranial Structure of the Titanosuchian: Anteosaurus.

Ann. S. Afr. Mus. 42, 2, 108-148. 1954.

Boonstra, L. D. The Smallest Titanosuchid yet recovered from the Karroo.

Ann. S. Afr, Mus, 42, 2, 149-156. 1954.

Boonstra, L. D. SE EET OCE eS A New Deinocephalian. Ann. S. Afr. Mus.

42.3) 180- 184. 1955.

Broiti, F. and ScHR6DER, J. Ein Dinocephalen-Rest aus den unteren Beaufort-

Schichten. Sitzb. Bay. Akad. Wiss. Sonderdruck, 93-114. 1935.

Broom, R. Notice of a new Fossil Reptile (Scapanodon duplessisi) from the

ae Karroo beds of Prince Albert, Cape Colony. Rec. Alb. Mus. 1, 3, 182-

183. 1904,

Broom, R. Notice of some new Fossil Reptiles from the Karroo Beds of South

Afpica. 4 Rec), Alls; Mus, nisi 331-937). angas!

Broom, R. Notice of some New South African Fossil Amphibians and Reptiles.

Ann. S. Afr. Mus. 7, 3, 270-278. 1909.

. Broom, R. Further Observations on South African Fossil Reptiles. Am. Mus.

Jougy 147) 135-145. Tord.

. Broom, R. On the Origin of Mammals. Phil. Trans. Roy. Soc. B.206, 1-48. i

: Broom, R. Permian, Triassic and Jurassic Reptiles of South Africa. Bull.

Mus. Nat. Hist. 25, 2, 105-164. I915.

. Broom, R. On Tapinocephalus and Two Other Dinocephalians. Ann. S. Afr.

Mus. 22, 3, 427-438. 1928.

. Broom, R. On the Carnivorous Mammal-like Reptiles of the Family Titanosuchidae.

Ann. Transv. Mus. 13, I, 9-36. 1929.

- Broom, R. The Mammal-like Reptiles of South Africa. pp. 18-46. Witherby, 1932.

. Byrne, F. A Preliminary Report on a New Mammal-like Reptile from the

Permian of South Africa. Trans. Kans. Acad. Sci. 40, 221-231. 1940.

. GreGory, W. K. and Camp, C. Studies in Comparative Myology and Osteology

No, II. Bull. Am. Mus. Nat. Hist. 38, 15, 447-563. 1918.

. GREGORY, W. K. The Skeleton of Moschops capensis Broom, a Dinocephalian

Reptile from the Permian of South Africa. Bull. Am. Mus. ‘Nat. Hist. 50, 3.

179-251. 1926.

- Haucuton, S. H. On a Skull of Tapinocephalus atherstonei, Owen. Ann. S. Afr.

Mus. 12, I, 40-42. 1915.

: HAUGHTON, S. H., in Report of the South African Museum for 1915, Pp. 4- 1916.

. Huene, F. von. Beitrag zur Kenntnis der Fauna der siidafrikanischen Karroo-

formation. Geol. Pal. Abh. 18, 3, 159-227. 1931.

. LYDEKKER, R. Catalogue of the Fossil Reptilia and Amphabia in the British

Museum (Natural History) IV, 81-98. 1890.

. Owen, R. Descriptive and Illustrated Catalogue of the Fossil Reptilia of South

Africa in the Collection of the British Museum, 1-86. 1876.

. OweEN, R. Description of Parts of the Skeleton of an Anomodont Reptile (Platy-

podosaurus vobustus, Ow.) from the Trias of Graaff-Reinet, S. Africa. Quart.

Jour. Geol. Soc. 36, 414-425. 1880.

. Romer, A. S. The Locomotor Apparatus of Certain Primitive and Mammal-like

Reptiles. Bull. Am. Mus. Nat. Hist. 46, 10, 517-606. 1922.

. Rossouw, P. J. en DE ViILLieRS, J. Die Geologie van die Gebied Merweville,

Kaapprovinsie. Toel. Blad. 198. Geol. Opname, Unie S.Afr., 1-71. 1952.

SEELEY, H. G. Researches on the Structure, Organisation and Classification of

the Fossil Reptilia. II. On Pareiasaurus bombidens (Owen), and the Significance

of its Affinities to Amphibians, Reptiles and Mammals. Phil. Trans. Roy. Soc.

179, B. 59-109. 1888.

SEELEY, H. G. Researches on the Structure, Organisation and Classification of

the’ Fossil) Reptilia,’ Vl. On the Anomodont Reptilia and their Allies. Phil.

Trans. Roy. Soc. 180, B. 215-296. 1889.

Watson, D. M.S. The Deinocephalia, an Order of Mammal-like Reptiles. Proc.

Zool. Soc., 749-786. 1914.

‘ydq siog 9opeuoIseNy ;0JOUd

“royyne oy} Aq pelfepoyy “wnasny, UeorIpyY YyNOS ey} UT dnors PeyIQey ve UT

poyiqiyxe se iszwym snypydar01yjNATS JO [apouL azis-ofIT Ve JO Ydeisoyoyq— AXX ALVId

ANNALS

ORTH

SOUTH AFRICAN MUSEUM

VOLUME (XLIT

Descriptions of the Palaeontological Material acquired by the South

African Museum and the Geological Survey of South Africa.

PART IV, containing:—

15. Tertiary Nautiloids dredged near Cape of Good Hope. By A. K.

MILLER and W. M. Furnisu. (With Plate XXVI and two text-

figures. )

16. Phospkatic-Glauconitic Deposits off the West Coast of South Africa.

bye) ti. HAUGHTON, DSc:,. E.G:S:

17. Fossil Carnivora from Hopefield. By R. F. Ewer and R. SINGER.

(With Plates XXVII-XXXII and one text-figure.)

18. Further Fossil Suidae from Hopefield. By E. N. KEEN and R. SINGER.

(With Plates XXXIII-XXXV.)

: Title-page and Index to Volume.

ORD

wT ES |

ISSUED MARCH 1956. PRICE: 1os\od.

PRINTED FOR THE

TRUSTEES OF THE SOUTH AFRICAN MUSEUM

. AND FOR THE

CLOLOGICAL “SURVEY. OF ‘SOUTH. VAPRICA

BY THE NATIONAL COMMERCIAL PRINTERS LIMITED

JAN VAN RIEBEECK STREET, ELSIES RIVER

15. Tertiary Nautiloids dredged near Cape of Good Hope. By A. K. MILLER,

and W. M. FurRNIsH, State University of Iowa, lowa City, lowa.

(With Plate X XVI and 2 text-figures.)

Four Aturias from collections in the South African Museum at Cape Town

were recently sent to us on loan by Dr. Sidney H. Haughton, formerly of the

Union of South Africa Geological Survey. These specimens were secured

early in 1938 by Captain J. T. R. Gibson, skipper of a trawler belonging to

Messrs. Irwin and Johnson; he dredged them from a depth of 160 to 170

fathoms just west of Cape of Good Hope, between Slangkop Lighthouse and

Cape Point. A single species is represented by the four specimens, which

are internal molds composed of phosphate rock. Dr. Haughton has informed

us that similarly preserved Jlamellibranchs, gasteropods and some

mammalian remains were found associated.*

Representatives of Atuna, though almost invariably rare, are of very

widespread occurrence in the marine Tertiary. They range from the Eocene

to the Miocene, inclusive, and may have been found in slightly older strata.

In the Atlantic-Gulf coastal region of North America, where Tertiary deposits

are well developed and have been studied extensively, Aturias do not occur

below the base of the Eocene, and they are known from only one locality in

the Miocene. However, in westernmost North America, South America, and

Europe they seem to be less rare in the Miocene. After studying specimens

from many widely separated localities in various parts of the world, we are

convinced that typical representatives of the genus have a long range in

the Tertiary. Therefore, we can not be certain as to the precise age of the

specimens under consideration.

Fic. 1.—Atura lotzi Bohm, Suture.

*) The first record in print of the occurrence of Aéuria in the phosphatic-glauconitic

deposit off the Cape of Good Hope appears to be in L. Cayeux: The Phosphatic

Nodules of the Agulhas Bank. Ann. S. Afr. Mus. XXXI, p. 133. 1934. [Ed.]

Vel AEM Part IV:

SUN 4 1956

328 ANNALS OF THE SOUTH AFRICAN MUSEUM

The more diagnostic features of these Cape of Good Hope nautiloids are

elucidated by the accompanying illustrations. In general physiognomy, this

form is reminiscent of similar-sized Aturias from the Americas. Because of

the width and shape of the umbilical lobe of the sutures, it can be said to

belong in a group typified by Aturia angustata (Conrad) of the Oligocene and

Miocene of the North American Pacific coastal region. The nearest geographic

occurrence of the genus is that of A. /Jotzi Bohm at Bogenfels, some 500 miles

north. Insofar as we can tell from the published information in regard to

the type specimens of that species, they are conspecific with those we are

studying. All of them came from a sandstone which carries a fauna generally

identified as Miocene.

Each of the four specimens under consideration represents a different

portion of the phragmocone, and it is possible that all are parts of a single

large individual, which (including the body chamber) had a diameter of

250 mm., or more. In every case, the surface is smooth and polished but,

nevertheless, bears serpuloid worm tubes, even on the exposed septa.

Clearly, the pieces lay loose on the bottom of the sea and were dissociated.

It should perhaps be stated that Aturias are known from latitudes as far

north and south as 55 or 60 degrees. Specimens have been collected from

Tierra del Fuego, Tasmania, and New Zealand. Therefore, their discovery

near southernmost Africa is not surprising.

REFERENCE

Mitter, A. K.: Tertiary nautiloids of west-coastal Africa: Mus. Congo Belge, Ann.,

Sér. in 8°, Sci. Géol., vol. 8, pp. 1-88, pls. 1-31. 1951.

Fic. 2.—Aturia lotzi Bédhm. Drawing traced from a photograph to show position

of siphuncle. This excellently preserved specimen was obtained by the South African

Museum from off Cape Columbine, Saldanha Bay area, after this paper had been

received from the authors, and has not been seen by them. Natural size. [Ed.].

PLATE XXVI

Ama, So UNae, Wis. Wolle, SEIU

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portrayed by figures I & 2.

16. Phosphatic-Glauconitic Deposits off the West Coast of South Africa.

By S. H. Haueuton, D.Sc., F.G.S.

From time to time there have come into the possession of the South

African Museum in Cape Town various phosphatised invertebrate and verte-

brate remains and phosphatic nodules that have been dredged off the west

of the Cape Peninsula at depths of over 100 fathoms. Most of those which

were received after 1930 were confided to me for examination; but pressure

of other work prevented an attempt at systematic study of this very interesting.

material.

In the Annual Report of the South African Museum for 1905 (p. 10),

Dr. J. D. F. Gilchrist, who was in charge of the collections made during

the dredging and collecting expeditions of the Government research vessel

“Pieter Faure’’ stated: ‘“The working out of the fossil remains in a series.

of deep sea deposits has been undertaken by Dr. Broom.’’ There is no:

record of this work having been completed; but in a letter to Dr. Gilchrist,.

dated 2oth August, 1905, from Stellenbosch Broom referred to a ‘“fragment.

of a Nautiloid shell’’ which he had no hesitation in identifying as “‘Atuna

aturt, a Miocene Nautiloid found in Miocene of France, Italy, Malta and

Australia’’. In the same letter he referred to a “‘Terebratula’’ and to

“‘whales’’. The Atuwna specimen is not in the South African Museum

collection; the other material referred to by Broom has not been definitely

located.

Early in 1938, Capt. J. T. R. Gibson, skipper of a trawler owned by

Messrs. Irvin and Johnson, deposited at the Museum some material which

he had dredged from a depth of 160-170 fathoms between Cape Point and

Slangkop Lighthouse. This included internal casts of a Xenophora-like shell

and four fragments of an Atuna, all phosphatised. The Aturia seemed to be

indistinguishable specifically from Atura lotzi, the type of which came from

Tertiary beds near Bogenfels in South West Africa. In the latter half of

1953 these four fragments were sent for examination to Dr. A. K. Miller

of the State University of Iowa whose publication on ‘Tertiary Nautiloids of

West-Coastal Africa’’ (Ann. Mus. Congo Belge, Sci. Géol., Vol. 8, 1951) is

the authoritative work on the subject. Dr. Miller confirmed their attribution

to A. lotzi Bohm, and is publishing a separate note on them.

In the Annual Reports of the South African Museum for 1948, 1950,

1952, and 1953 Dr. K. H. Barnard made brief mention of further phosphatised

330 ANNALS OF THE SOUTH AFRICAN MUSEUM

material obtained from Capt. Gibson, the Fisheries Survey, and from a depth

of approximately 200 fathoms off Cape Columbine, the last presented by the

Department of Geography of the University of Cape Town. Cape Columbine

lies to the north of Saldanha Bay, forming the southern headline of Klein

Paternoster bay.

The petrological and mineralogical characteristics of phosphatic -material

dredged by the S.S. ‘‘Pieter Faure’’ from the Agulhas Bank, together with

an account of the probable geological history, were considered in detail by

L. Cayeux (Ann. S.Afr. Mus., XXXI, 1934, p. 108). Some of the material

collected by Capt. Gibson was studied in the Geological Survey Laboratories

by Dr. J. E. de Villiers, now Professor of Geology in the University of the

Orange Free State, Bloemfontein.

Dr. de Villers recognised two major varieties of phosphate rock. One is

a conglomeratic type, composed of irregular nodules set in a cement; the

other is more homogeneous and similar to the ‘‘cement’’ of the conglomeratic

type. He reported that “‘the nodules consist of yellow-brown, amorphous-

appearing phosphate which for the most part is crowded with calcareous organic

remains. Angular mineral fragments (quartz, albite, orthoclase, labradorite,

zircon, garnet, tourmaline and muscovite) and small, rounded grains of

glauconite are also present. X-ray examination of the phosphate showed it

to have a pattern identical with that of francolite from the Richtersveld.

In a few instances it was found that the nodules had been glauconitised at

or near the contact with the matrix; the glauconitisation took place before

cementation of the nodules — uncemented nodules occur which have a mm.

of glauconite.’’ )

“The cementing material of the matrix is collophanite. Calcareous

inclusions and tests of foraminifera are more rare than in the nodules, and

quartz and round glauconite grains are more plentiful. The fragments and

grains of these two minerals are larger in the matrix than in the nodules and

the ratio of glauconite to quartz is about 4 : 1. Tiny dust-like specks of iron.

oxide and perhaps carbon are irregularly distributed and cause the colour

to vary from light to darker yellow-brown’’ (De Villiers).

The ‘‘conglomeratic’’ material, therefore, has had a composite geological

history. The ‘‘nodules’’, which are often angular or semi-angular and of

irregular shape, may be pieces of an originally calcareous sediment containing

fragments of bone, shells, and foraminiferal tests which suffered a degree of

phosphatisation before fragmentation. The fragments which now form the

nodules became embedded in a glauconite sand containing some foraminifera

together with the grains of quartz and other terrigenous material which in its

turn became phosphatised and cemented, forming a phosphorite. The frag-

ments as now dredged are irregular in shape. In the conglomeratic samples,

the older fragments exist in juxtaposition to the newer cement at the surface

PHOSPHATIC-GLAUCONITIC DEPOSITS OFF WEST COAST OF S.A. Ban

of the specimens, sometimes protruding. Many have a veneer of glauconite;

many are partially encrusted with calcareous material, serpulid tubes, and

sponge spicules. In cavities in the irregular dredged fragments there was

sometimes found sand consisting of abundant grains of quartz and felspar,

some rounded glauconite grains, and organic material consisting of numerous

sponge fragments, calcite plates minutely perforated, a few globigerina tests,

and plates of chitin. This, and the sand found in phosphatised ear-bones of

whales, is the youngest material in the collection.

Cayeux (Ann. S. Afr. Mus., XXXI, 1934, p. 124) in his description

of a ‘‘nodule formed of two differing rocks’’ was obviously dealing with

material similar to the ‘‘conglomeratic type’’ of de Villiers, although his

specimen was of such a nature that he was unable to say that one rock

served as a matrix to the other. The lighter-coloured rock he considered to

be a phosphatised ‘““bryozoan foraminiferal limestone’, the other a

phosphatised greensand. He pointed out, however, that these two rock

types must have been formed under differing bathymetric conditions and

formulated the idea of remamnié material from a deposit formed originally at

a greater depth being incorporated in shallower water greensands.

The four specimens assigned by Miller to Aturia lotzt are composed

lithologically of the same kind of material as forms the “‘nodules’’ in the

“‘conglomeratic type’’. The Bogenfels specimens of A. Lotzi described by

Bohm were obtained from a marly sandstone of terrigenous origin which now

stands 140-160 m. above the sea, considered by him (1926) to be of Middle

and Upper Eocene age. The general nature of the succession of beds in the

Bogenfels area shows that they are near-shore deposits; uplift of the order

of 400-500 feet is thus indicated for them. The fragments studied by Miller

_must have been deposited therefore in an environment different from those

found in the Bogenfels beds, the former at a greater depth below sea-level and

farther from the shore than the latter.

More recently two more specimens of Aturia lotzi were acquired by the

South African Museum. They had been trawled from the Saldanha Bay-

Cape Columbine area of the west coast, but the depth and actual position

of the find are unknown. The discovery area is well to the north of that

from which Capt. Gibson obtained the four specimens examined by Miller,

and therefore nearer to Bogenfels, but the matrix is said by Dr. Barnard to

be like that of the others.

Among the material dredged off Cape Columbine were nodules containing

more or less fragmentary lamellibranchs and brachiopods. These were sent

for study to Dr. Helen Muir-Wood of the Department of Geology of the

British Museum (Natural History) who, in turn, passed the lamellibranchs on

to her colleague Dr. L. R. Cox. Both these experts have been good enough

to favour me with their comments on the material examined.

332 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dr. Muir-Wood stated : —

‘‘T have looked at the four brachiopods dredged off Cape Columbine, but

can give you no specific determinations or any definite information, about their

age. They are not comparable with anything from South Africa, and we

have no specimens from other areas that are altogether similar. They could

be Miocene in age as suggested by Dr. Cox, and the two Terebratulids are

externally rather like specimens of ‘Terebratula’ miocenica Michelotti from the

Miocene of Italy. I have no idea whether the African forms are actually

related to the Italian species, since the internal characters of both are

unknown.

‘‘The larger of the two Terebratulinids is probably new. I have compared

it with Cancellothyris which does occur in the Miocene, but do not think it

belongs to this genus. The smaller specimen will have to be compared with

the Recent species Terebratulina retusa (Linn.). This species, usually quoted as’

T. caputserpentis (Linn.) appears frequently in fossil lists of species from the

Miocene or Pliocene. ,

‘“‘The Tertiary brachiopods are in great need of revision and it is difficult

at present to give specific determinations.”’

Dr. L. R. Cox reported on the molluscan fossils as follows : —

“They have nothing in common with the assemblage described by Bohm

and Weissermel from the Bogenfels diamond fields and do not, in fact, link

up at all with any fossels hitherto described from Africa. They may be

recorded as follows :— ; !

“Ostrea (Pycnodonta) ct. brogmiarti Bronn. This is the Gryphaea-like

oyster, of which there are several specimens, some with a very spongy

texture. O. brogntarti, which is scarcely distinguishable from the Cretaceous

O. vesicularis Lamarck, is widespread in the Eocene of the Tethyan province,

persisting into the Oligocene of several Mediteranean countries, and being

last met with in the Lower Miocene (Aquitanian) of N. Italy. Of its recorded

localities, the nearest to S. Africa is the Belgian Congo, where it occurs in the

Paleocene. It could conceivably have survived to the Miocene off S.W.

Africa.

““Pecten sp.nov. This species, of which there are two or three specimens,

has the right valve strongly convex and the left valve almost flat, and there-

fore belongs to Pecten s. str. The left valve has fine, evenly spaced con-:

centric threads, crossing ribs and intervals. This species belongs to a group.

well represented in the Oligocene and Neogene of the Mediterranean and

N. W. Indian Ocean areas, but absent in the Eocene except in California.

It does not appear to be identical with any described species.

“‘Chlamys spp. indet. There are three ill-preserved specimens of equivalve

iipbed Pectinids, probably belonging to two different species. The details

PHOSPHATIC-GLAUCONITIC DEPOSITS OFF WEST COAST OF S.A. 333

of the ornament cannot be observed and the specimens throw no light on

the geological age of the deposit.

“Conus sp. indet. One ill-preserved internal mould.

“Cadulus sp. indet. One internal mould.

“The material is too limited and ill-preserved to allow the age to be

established definitely. A lower Miocene age would, however, be compatible

with the presence of oysters indistinguishable from O. brogmniarti Bronn

associated with specimens of Pecten s. str., a group which, although found in

the Oligocene, does not become abundant until the lower Miocene. The

moulds of worm tubes and of two species of corals do not assist in the

determination of age.”’

Apart from the material submitted for examination to overseas specialists,

there remains in the collections of the South African Museum a considerable

number of phosphatised specimens obtained from dredging off the west coast

of the Cape Peninsula at various times since the S.S. ‘‘Pieter Faure’’ was

used as a governmental research ship under Dr. J. D. F. Gilchrist. These

hav been examined by Dr. K. H. Barnard, who has supplied the following

identifications for purposes of record : —

MAMMALIA

Otoliths of Balaena (Right Whale), Balaenoptera (Finner Whale),

Megaptera (Humback Whale). Portion of skull of unidentified

cetacean (S.A.M. Reg. No. 19475).

Anterior portions of skulls of Mesoplodon spp.

(a) With maxillary and premaxillary foramina on same _ transverse

line (S.A.M. Reg. Nos. 19473 and 19934). Cf. M. europaeus,

M. bidens, M. layardu.

(b) One with premaxilliary foramina posterior to maxillary foramina

(S.A.M. Reg. No. 19930). Cf. M. densirostris, M. grayt, M.

australis.

Anterior portion of skull of ? Ziphius, with gibbous maxillaries

(S.A.M. Reg. No. 19942).

Hind portions of left mandible with 7 tooth-sockets of Orca sp.

PISCES

Carcharodon sp. — teeth.

Carcharias sp. — teeth.

Isurus or Lamna sp. — teeth.

Histiophorus sp. — portion of rostrum.

334 ANNALS OF THE SOUTH AFRICAN MUSEUM

GASTROPODA. (Internal casts.)

Xenophora sp.

Conus (?) sp.

Cf. Pleurotoma sp.

Cf. Pustonella sp.

Fusus (?) sp.

Natica sp.

LAMELLIBRANCHIA. (Internal casts.)

Cuspidaria sp.

Venerid (?).

SCAPHOPODA. (Internal casts.)

Dentahum sp.

Most, if not all, of this material would appear to be associated with the

later glauconitic greensand type of deposit. The impossibility of specific

identification of these various forms renders comparison with other described

faunas hazardous; and clearly further attention should be paid to the

collection of additional material and to intensive study of a fossil assemblage

that seemingly is contained in rocks of at least two differing ages. Study

of the foraminifera, which occur abundantly in the ‘‘nodules’’ and more

sparsely in the ‘‘cement’’ should be of assistance. Such evidence as has been

yielded by the macro-fossils, however, suggests that the foraminiferal

‘“‘nodular’’ part of the dredged material is of Miocene (and even of Lower

Miocene) age and that it may represent a somewhat deeper-water facies of

the Bogenfels beds which carry Aturia lotzt.

17. Fossil Carnivora from Hopefield. By R. F. Ewer, Zoology Department,

Rhodes University, Grahamstown, and R. SINGER, Anatomy

Department, University of Cape Town.

(With Plates XXVII-XXXII and 1 text-figure.)

I. INTRODUCTION

The history and general appearance of the fossil site on the farm

“‘Elandsfontein’’, I0 miles from Hopefield —a village 90 miles north of

Cape Town —has already been described in a fairly detailed manner

(Drennan, 1954; Singer, 1954). The movement of the exposed crescentic

sand-dunes situated in the scrub-covered bushveld has uncovered calcareous

floors from which a wealth of fossil material has been recovered by a research

team of the University of Cape Town. Besides the human cranial fragments

(see above publications, and also Drennan and Singer, 1955) and stone

implements of the final phases of the South African hand-axe culture and of

the Still Bay culture, many species of extant and extinct mammals have

already been identified — Equidae, Homotoceras (?Bubalus) sp., Palaeo-

loxodon cf. antiquus recki, Ceratotherum and _ Diceros, Giraffidae

_(2Stvatherium), Hippopotamus amphibius and Mesochoerus lategam (Singer

and Keen, 1955), and numerous types of antelopes (eland, wildebeest, lechwe,

etc.). A small number of fossilized tortoise carapace fragments, shells of

Succinea, land snails (Trigonephrus globulus), and a cranial fragment of a

scaly anteater, have also been recognized. Fossil Carnivora are not abundant

at Hopefield, but the remains which have been recovered to date show that

the fauna included a viverrid, two hyaenids, a small felid, a Lycaon, a jackal

and a mustelid. The material is described in detail below.

Il. DESCRIPTIONS

1. Family Viverridae

Specimen E.C.7: the top of a braincase with the greater part of the

occiput.

In size and general conformation the specimen closely resembles Herpestes

tchneumon (Linn.), but in view of its incompleteness no detailed comparisons

are possible.

336 ANNALS OF THE SOUTH AFRICAN MUSEUM

2. Family Hyaenidae

The Hyaenid remains represent two species, one of which is a Hyaena,

Closely resesmbling the living H. brunnea, and the other a large Crocuta.

(1) Hyaena brunnea Thunb.

Specimen E.C.2: right mandibular ramus, lacking the coronoid process

and with I, and I. broken off; all the other teeth are present in a good state

of preservation and showing only slight wear.

Specimen E.C.3: portion of left mandible bearing the canine, P2, P:

lacking its anterior end, P. and slightly damaged M,. This almost certainly

belongs to the same individual as E.C.z2.

Specimen E.C.4: maxillary fragment with P*, P* and M’. These teeth

occlude perfectly with those of E.C.3, and there is no doubt that they belong

to the same individual.

Specimen E.C.6: maxillary fragment with P* and the anterior roots

of Pr;

These specimens do not show any important differences from the living

Hyaena brunnea Thunb. The measurements of the teeth are given below

and compared with the corresponding values found for a sample of 15

specimens (13 measurements for mandibles) of the living brown hyaena

(table 1). For each measurement of the latter the mean and standard

deviation are given. From the ratio of the difference between the fossil

measurement and the mean to the standard deviation can be found P, the

probability of a specimen from the same population as the living specimens

having a measurement as divergent as that of the fossil.

In the dimensions of the upper teeth none of the fossils differs significantly

from the living species. In the case of teeth from the mandibles some slight

differences are apparent: the carnassials and P; are slightly smaller, and

P, is slightly narrower both absolutely and relatively. The smaller lower

carnassial and slightly lesser development of the crushing specialisations of

P, and P,, as reflected in their slightly smaller widths, are points in which

the fossil specimens appear to be a little more primitive than their living

counterparts. The differences are so slight, however, that it has not been

considered necessary to place the fossil specimens in a distinct subspecies.

A subspecies of Hyaena brunnea, H. brunnea dispar, has been described

by Ewer (1955) from Swartkrans. In this subspecies P* is slightly shorter

than in the living form and P* is more primitive, the posterior cusp being

rather large and the anterior ridge situated less internally. The Hopefield

specimens do not show these characteristics. The only point of resemblance

between the two is that P,; in each case is shorter than in the hving form, but

FOSSIL CARNIVORA FROM HOPEFIELD 337

in the Hopefield specimen the tooth is smaller in both its linear dimensions,

whereas in the dispar subspecies the width is not significantly different from

that of the living form. There is thus little justification for including the

Hopefield specimens in the dispar subspecies.

TABLE 1.

Measurements of fossil Hyaena brunnea compared with those of a sample of 13 mandibles

and 15 skulls of living H. brunnea. For the latter the mean and its standard deviation are

given. = is the difference between the fossil measurement and the mean divided by the

standard deviation of the latter. P gives the probability of a measurement as divergent as that

of the fossil specimen being found in the population from which the sample of living H. brunnea

‘was drawn. Values marked with an asterisk are statistically significant.

All measurements, in this and the succeeding tables, are given in millimetres.

Living

Specimen No. Hi. brunnea da

| —_—-—___— -— ' acl oy

| | E.C.2| B.C. 3| Mean | S.D. P

I, Jength bas a is pl ae | 6°60 -56 | -°89 | >-05

breadth T°2 7:56 | -41 -88 | >-05

breadth W262) ie U9 yA 3235 -84 | 1-05 | >-05

P, length 14-8 | 15-1 15:55 -50 | 1-50 | >-05

breadth .. ne 10-0 | 10-5 | 11-03 -50 | 2:03 | <-05>-04

length : breadth ratio 1-48 1-44) 1-41 06 | 1-16 | >-05

P, length 19-6 21-17 ‘40 | 3:70 | <-0004*

breadth .. ay 12-7 |.12°7 114-40 -60 | 2°83 | =-005*

length : breadth ratio 1-54 1-47 | -06 | 1-20 | >:05

P, length D322 4 | 23e lyk lean 52 -56 88 | >-05

breadth .. te: 12-4 | 12-4 | 13-80; -51 | 2:75 | =<-01>-005*

length : breadth ratio 1-87 1-86 1-71 -06 | 2:67 | <-01>-005*

M, length .. dis et ey 224 | 2204. 2a D6 -89 | 2-43 | <:02>-012*

breadth .. i, We See IAS IEA «| 12761 -43 | 3-05 | <-:003>-001*

| E.C. 4 | E.C. 6 | | | |

P? length .. oh BN Dp PZB AL GIO PAIN 9.220105) °63 | 1:08 | >-05

breadth .. he Ps «. | 15°9 | 15:9 | 15-98 -70 -11 | >-05

length : breadth ratio .. a 1-45 1-40 1-43 -05 ‘06 | >-05

P'dength .. sia si Le ors 35-31 | 1-08 -45 | >-05

breadth .. oy, ao Cea ae) ite | 21-48 68 -56 | >-05

M' length .. lid ie ie 6-0 5-62 -38 | 1:00 | >-05

=> -05

breadth .. 2 hs .. | 14-0 13-07 enh | LeO7,

‘ i A

(11) Crocuta spelaea (Goldf.).

Specimen E.C.1: posterior half of skull, broken off at about the level

of the post-orbital constriction. The zygomata are missing and the auditory

bullae broken, but otherwise the specimen is almost perfect (Pls. XXVII-

XXX).

338 ANNALS OF THE SOUTH AFRICAN MUSEUM

Specimen E.C.g: mandibular fragment with much damaged P., Ps, P.

and the alveoli of M,.

Specimen E.C.8: mandibular fragment with the anterior two thirds of

M, (Pls. XXX, XXXI). |

Specimen E.C.5: maxillary fragment with P* and damaged P*

(Pls. XXX, XXX).

The skull is large and heavily built, the occiput high and narrow, and

very distinctly ‘‘shouldered’’ (see Pl. XXVIII). The lambdoid and sagittal

crests are well developed, and the latter is very high posteriorly. The con-

formation of this region is very similar to that of a skull of Crocuta spelaea

from Sundwig in the British Museum of Natural History (No. 28558) and an

occipital fragment (M. 4570) from Torbryan Caves also shows similar

development of the crests.

Goldfuss (1821) notes as a characteristic of Crocuta spelaea the large

development of the post-glenoid process. This is not at all an easy character

to assess or to measure, particularly since in fossil material the tip of the

process is commonly slightly damaged. From an examination of the

C. spelaea material in the British Museum of Natural History it appears that,

apart from its size, the post-glenoid process of C. spelaea differs from that of

C. crocuta (Erxl.) in its orientation. In C. spelaea the process lies almost

in the transverse plane of the skull, whereas in C. crocuta it slopes distinctly

forward from its outer to its inner end. In specimen E.C. 1 the process is.

orientated as in C. spelaea (Pl. XXIX). |

Unfortunately both the carnassials are damaged posteriorly, but the

presence of posterior roots makes it possible to restore the missing portions.

with a fair degree of accuracy, and at least to determine a minimum length.

In specimen E.C. 5 the palate is somewhat damaged at the posterior end

of P*, and it is therefore impossible to be quite certain that M’ was absent,

but clearly if present at all it can only have been very small. The upper

carnassial possesses the long metacone and relatively short parastyle

characteristic of advanced Crocutas, but the protocone does not slope forward

as much as is usual in C. spelaea. The lower carnassial, apart from its large

size, 1s remarkable only for the structure of the anterior cingulum. This

forms a sharp shelf-like excrescence round the antero-external margin of the

tooth extending for just over 2 mm. and then ceasing abruptly at either end.

Since the posterior portion of the tooth is missing the characters of the

talonid are unknown.

The premolars do not provide any distinctive characters but closely

resemble those of Crocuta crocuta, except for their larger size. The posterior

part of P. is missing, but the posterior root is very broad: P. in C. spelaea

is commonly very wide at the posterior end.

FOSSIL CARNIVORA FROM HOPEFIELD 339

The measurements of the specimens are given below, compared, in the

same way as before, with the corresponding figures for C. spelaea (table 2).

The only points of difference are that the premolars are a trifle narrower in

the Hopefield than in the European specimens. In this character the Hope-

field specimens resemble the capensis subspecies of C. spelaea described by

Broom (1939) (see also Ewer 1954) from Kromdraai. C. spelaea capensis,

however, differs from typical C. spelaea also in the large and almost square

M’ and in the very long P’, neither of which characters is shown by the

Hopefield specimens. In addition the protocone of P* in C. spelaea capensis is

large and slopes forwards very considerably. The general narrowness of the

premolars is a primitive character, and cannot, by itself, be taken to indicate

any particular close relationship of the Hopefield specimens to C. spelaea

capensis. The former are best regarded as closely resembling the typical

European C. sfelaea, differing only in the slightly more primitive character

of the premolars. This difference does not seem of sufficient importance to

TABLE 2.

Measurements of Hopefield specimens of Crocuta spelaea. The dimensions of the teeth

are compared with those of a sample of C. spelaea from various European localities: the

figure in brackets after the mean for the latter gives the number of specimens on which the

mean is based. Other symbols as before.

Specimen E.C. 1.

Maximum vertical height of sagittal crest above upper edge of foramen magnum 81:6 mm.

Maximum vertical height of external occipital protuberance above foramen magnum 55:0 mm.

Maximum width across occipital condyles 47-0 mm.

Maximum width of skull (intersquamosal) 95-0 mm.

Maximum width across mandibular fossae ca. 124-0 mm.

| European sample 4

| Mean S.D. | oO P

Specimen E.C. 9

Pyiength .. Ne ie Sg Meee 7/3) 16-9 (31) 85 TPA SPOS

Peneneth |. a BY. ee | 22D 227) (35) -78 -64 | >-05

breadth .. 8 ih abt 14-6 | 16:5 -93 | 2:04 | =-04

length : breadth ratio .. Weel 2 1-38 -054 | 2-59 | =-01*

Pare er rs A oaes 193.5) (32). 1131) 89, 1s s8205

breadth .. ie Me: MY 13-5 14-9 87 1-61 | >-05

Jength : breadth ratio .. an 1-8] 1-58 -088 | 2-61 | =-O1*

Specimen E.C. 8 |

M, length .. a i re Caz 30°'S.) id 24i (26) +2 1,233 1:43 | >-05

breadth .. ae ie Ayal 12-8 Sp) 67/5) 1-23 | >-05

Specimen E.C. 5 |

PoOMenethy ..),. me i! bien 24:3 | 24-1 (19) | 1-34 “15, |=="05

‘breadth .. ie Ne chee 16:6 | 17-9 a7 1-11 | >-05

length : breadth ratio .. 5 1-46 1s -§2 | 2:11 | =<:04—:03

Pedenpty ii)5 . ‘. hs vol ca 40 40:9 (27) “92 -98 | >-05

225) L222 es L238) e205

breadth .. a ie ABA ce OA

340 ANNALS OF THE SOUTH AFRICAN MUSEUM

warrant the erection of a new subspecies for the Hopefield specimens, but

should be borne in mind in considering the probable age of the deposit.

3. Family Felidae

(°?) Leptatlurus serval (Schreber).

Specimens E.C.15, E.C.17: mandibular rami, incomplete anteriorly and

posteriorly, bearing P; Ps and M,.

Specimen E.C.16: portion of mandibular ramus with P. and Mh.

The specimens all belong to a small felid, and the teeth do not show any |

peculiar characters. A set of skulls is not available to us for detailed

comparison, but, judging from the measurements given by Roberts (1951),

the specimens are a trifle too large to belong to the Cape wild cat, Fels cafra

Desmarest, and fall within the range of Leptailurus serval (Schreber), to which

species they are tentatively referred.

TABLE 3.

Measurements of fossil specimens of Leptailurus serval.

Specimen No.

| E.C. 15 | E.C. 16 | E.C.17

Depth of mandible between P, and M,.. ie Ag Ne 13°5 1562), eae +S

P, length .. vy oy uth i Wy uk re 8°7 7°5

breadth .. uM et iG i tei) an 4°8 3°9

P, length .. Me ws ae i, A 4) in 10:5 10-7 9-7

breadth .. a Hs Mi aD Li Hs gi 5-8 5-1 5-1

M, length .. Mi eit she ir i, a AN 12-3 11°4 11-5

breadth. . bs uh ae a i ua 6:2 4-9 5°5

4. Family Canidae

(i) Lycaon pictus (Temm.) n. subsp. magnus.

DIAGNOSIS:

A subspecies of Lycaon differing from the extant form in the greater

length of the mandible, larger lower canines and incisors and longer but

relatively narrower lower premolars.

Specimen E.C.13: portion of left mandibular ramus, incomplete below

and bearing P, to P, and the anterior root of M,. (Pls. XXX, XXXI.)

Specimen E.C.121: right mandibular fragment bearing I., I, and C.

Specimen E.C.30: isolated damaged right M,.

All three specimens show moderate wear and are in a very similar state

of preservation: it is probable that they belong to a single individual. Apart

FOSSIL CARNIVORA FROM HOPEFIELD 341

from their distinctly greater length the teeth differ in no way from those of the

living Lycaon pictus (Temminck). The measurements are given below,

compared as before with those of the extant species. It will be seen that the

fossil jaw is considerably longer than that of the extant species and that the

teeth differ significantly from those of the latter in the following points: the

premolars are considerably longer, but not much broader; the canine and I.

are significantly larger in both dimensions while the significance of the slightly

greater breadth of I. is less certain. It is considered that these differences

are sufficiently clear cut to warrant placing the fossil specimens in a distant

subspecies.

Wells & Cooke (1942) record from Vlakkraal a damaged M, and a worn

I,; the latter is said to be “‘a little larger than any specimen of this genus

actually axamined.’’ It seems not improbable that this belongs to the same

subspecies as the Hopefield material.

TABLE 4.

Dimensions of teeth of fossil Lycaon compared with those of a sample of 1 extant

Lycaon pictus. Symbols as before.

Extant |

Specimen No. L. pictus | F

)EC. 12 EC13/ Mean | SD. | o P

Length from back of P, to front

of P, .. ws ee i 52-8), |40-7 1:79 | 6-76 | <-0004*

I, length 555) 4-48 “57 04 | >-05

breadth 5-6 4-71 -39 | 2-29 | <-:03>-02

_ I, length re mt te 7:6 6:03 -47 | 3-34 | <:-001> -0004*

breadth ct sf ia 6°5 5-66 -30 | 2-80 | =-005*

C length a a .. | 13-0 10-61 ‘72, | 3-32 | <:001> -0004*

breadth a! a ib 9-1 7-80 53 | 2:45 | <:02>:012*

P, length 8-1 6:45 -50 | 3:30 | <-001> -0004*

breadth if ae 5199) 4:71 350 WV Ned ipl 205

length : breadth ratio 1-560) 373 *119 | 1-57 | >-05

P, length 12-4 10-05 -62 3:79 | <-0004*

breadth nas ap Bop) 5:35 “25 60 | >-05

length : breadth ratio 225 1-879 -076 | 4-88 | <-0004*

P, length 14-3 11-95 54 | 4-37 | <-0004*

breadth (01) |. 6-3 | 6-08 | -39 56 | >-05

length : breadth ratio 224 1-970 | -130 | 2:30 ; <-03>-02

P, length 16:5 13:75 | -79 3°48 | <-0004*

breadth om My 7:3 7-11 | 47 40 | >-05

length : breadth ratio 2:26 | 1-938 : 100 | 3:22 | <-003>-001*

M, breadth... ye is 10-4 992

342 ANNALS OF THE SOUTH AFRICAN MUSEUM

(ii) Canis mesomelas Schreber.

Specimens E.C.18-20, 22-27 and 29; various incomplete mandibular

fragments. :

Specimen E.C.31: isolated Mi.

Specimen E.C.32: isolated Py.

Specimen E.C.21: maxillary fragment bearing M’ and M’.

In another paper (Ewer, in the press) only a single character was found

in which there was no overlap between the two living species of jackal, Cams

mesomelas Schreber and C. adustus Sundevall, and which therefore allowed

of a certain assignment of a single specimen to one or the other species. This

character is the length of the carnassial tooth relative to that of the succeeding

molar. This is referred to as the ‘‘carnassial: molar ratio’’ and either upper

or lower ratios serve to separate the two species.

In the case of the material under discussion at present four specimens,

E.C.18, 22, 25 and 26, include both M, and M: and therefore allow of the

calculation of the lower carnassial: molar ratio. The values found are 2.03,

1.98, 2.10 and 2.30 respectively. For the samples of 14 C. mesomelas and

10 C. adustus used in the previous study the values range from 1.99 to 2.57

for the former and 1.60 to 1.81 for the latter. The Hopefield specimens

clearly belong to mesomelas, and in the table of measurements the mean and

range for the sample of extant mesomelas are given for comparison (table 5).

It will be seen that although the means for the Hopefield specimens lie within

the normal range for mesomelas a number of individual measurements lie

beyond the upper size limit found for the living species.

A fossil subspecies of C. mesomelas has been described (Ewer, in the

press) from the Transvaal deposits, differing from the living form principally

in the fact that relative to the length of the carnassial M, is less reduced and

the premolars are longer. It is therefore of some interest to compare the

lengths of M. and of the premolars in the Hopefield specimens with the

corresponding values for the Transvaal fossils. Figure 1 shows the mean

values for the relevant dimensions, with those of the extant form for

comparison. It will be seen that in each case the Hopefield value is inter-

mediate between those for the Transvaal subspecies and for the living form.

There seems to be little doubt that we are here dealing with a phylogenetic

series in which a reduction of the molars and a shortening of the premolars is

taking place; the Hopefield specimens representing a form ancestral to the

living black-backed jackal and the Transvaal form in turn ancestral to the

Hopefield. The functional significance of the changes is not at once apparent,

and it may be that they would be better expressed by saying that a general

reduction in overall size is taking place, but without any reduction in size

of the carnassials, which therefore show an increase in relative size. Without

FOSSIL CARNIVORA FROM HOPEFIELD 345.

tig. 1

P, AXA !

GI fransvaal subspecies.

x Hopefield specimens.

A Living GC. mesomelas.

Fe Aye!

2 P.

R= "3 AX———_f]

@ ;

re) PR

sa 4: A—xX—)

M, A a

4. Sar “ta 3 10 7 12

Tooth length (mm)

VALUES.

Length(mm.) Transvaal Hopefied. Living.

Pen 4.5 4.2 4.0

P, 9.0 8.3 8.0

Sela 9.8 8.9 8.8

Pp, Th. 10.8 10.3

M, 9.6 9-4 8.5

whole skulls and skeletal material available it is not possible to decide whether

this latter formulation is the more correct, It may be noted in passing that

the Hopetield Lycaon also differs from the living species in possessing a longer

premolar row. | :

A second fossil subspecies of C. mesomelas is known from African deposits.

This is Canis mesomelas latirostris (Pohle) from Olduvai and Serengeti. This

subspecies is distinguished from the living form by its very broad snout and

shorter and less inflated auditory bulla. In the absence of a complete skull

no adequate comparison of our material with this subspecies is possible, but

Pohle (1928) gives measurements for the lengths of the teeth of his three

specimens. The lower premolars resemble those of the Hopefield specimens in

being rather longer than is usual in living C. mesomelas, but M. (present in

Annals—2

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SS | TS | LT | | | RT | RS | ES | TS | | TTS | A TTS | RES | RNAS

ANNALS OF THE SOUTH AFRICAN MUSEUM

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344

FOSSIL CARNIVORA FROM HOPEFIELD 345

2 specimens) is shorter than in the Hopefield material. In the absence of a

skull it seems unwise to assign our material to Pohle’s subspecies, particularly

since the Transvaal subspecies does not show the characters of snout and

bulla diagnostic for C. mesomelas latirostnis.

The Hopefield specimens thus appear to be the middle term of a series

leading from the Transvaal subspecies to the extant black-backed jackal.

As must always be the case when a fossil record approaches completeness,

this situation raises difficulties in nomenclature. The exact point as which a

progressive change is taken to be of sufficient importance to warrant

nomenclatural recognition must be largely a matter of individual judgment.

In the present case it is felt that since the majority of measurements for

the Hopefield specimens fall within the range of values found for the living

specimens, the erection of a separate subspecies is not likely to be useful.

Nevertheless, when questions of dating are being considered the intermediate

character of the Hopefield specimens between the extant C. mesomelas and

the Transvaal fossil subspecies must be borne in mind.

5. Family Mustelidae

Mellivora capensis (Schreber).

Specimen E.C.14: an almost perfect skull, with the zygomata missing

(Pl. XXXII).

Specimen E.C.14 very closely resembles the living honey-badger,

Mellivora capensis (Schreber), and differs only in its rather smaller size and

the fact that the palate is considerably less prolonged posteriorly. The

measurements of the fossil specimen are given below compared with those

of the living form (table 6). It will be seen that although the fossil skull is

considerably shorter than the mean for the sample of 15 living Mellivora

capensis the variability of the latter is so high that the difference is not

significant. The post-orbital width is significantly less in the fossil than in the

living specimens; but when the smaller size of the fossil is taken into account,

by expressing the post-orbital width as a percentage of the basilar length,

the difference is no longer significant. The palate is, hower, significantly

shorter in the fossil than in the living species, both absolutely and when

expressed as a percentage of the basilar length.

The fossil specimen thus differs from extant Mellivora capensts only in

having the palate less prolonged posteriorly. On the basis of a single specimen,

and bearing in mind that the difference is significant only at the 2% level,

it does not seem desirable to erect a separate subspecies for the Hopefield

Melhvora; but, as has been emphasised before, the fact that the Hopefield

specimen is distinguishable from the living species must not be lost sight of

in considering problems of dating.

346 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 6.

Measurements of Hopefield Mellivora compared with the corresponding measurements

for a sample of 15 extant M. capensis. Symbols as before.

Extant

M. capensis

Specimen fu et el

E.C. 14 Mean S.D. oO P

Basilar length ... at) e Sh 113-0 123-43 | 6-636 | 1-57 | >-05

Palate length ; ae $e Ris 50-4 60-65. | 4-389 | 2:34 | ==-02*

Posterior palate width? dT a Me 16-6 19-45 1-354 | 2-10) —-04=>-03

Postorbital width é “a LY. 28'-2 34-47 1-533 | 4:09 | <-0004*

Maximum cranial width Ne een ere (CIS aN 65:81 3-540 712 i = 0S

Width of nasal aperture. ¥ Tee aor £3709 1-579 | 1°48: |) "05

Length of P4 & tt 2048 12:67 1-133 |: 1-65) 3-05

Width of palate at level of P! NU oS Getta 45-33 3-114 | 1-84 | >-05

Palate length as % of basilar lengih .. 44-60 49-226 |" 1-990") 2432 | 202"

Posterior palate width as % of basilar

length .. 14-69 15-780 -968 | £13 |S =05

Postorbital width as H4 of basilar length 24-96 28:026 | 1-704 | 1-80 | =>-05

1 This is the width acros the posterior end of the hard plate.

III. DzIscussIon.

The faunal remains described above may be considered in terms of the

light they may throw on two problems: the date at which the Hopefield

deposit was laid down, and the ecological nature of the locality at that time.

As regards the latter point the carnivore fauna adds little to our

knowledge. All the species described are very similar to species which have

existed within the area during historic times. One curious point is that

although hyaenas and jackals are present, no remains of large carnivores,

such as leopard and lion, have to date been discovered. Artiodactyl and

horse remains are abundant, and it seems very improbable that in fact no

large carnivore predators were present. It seems likely that further work °

may bring lion or leopard remains to light, and their absence cannot at this

stage be taken as established.

As regards the date of the deposit it is quite clear on the one hand that it

is much more recent than the Transvaal deposits. The species represented

are all closely related to living forms, and no trace has been found of the

archaic hyaenids, Lycyaena and Leecyaena, and the sabre-tooths which exist

in the latter deposits. At the same time the fact that most of the specimens

show slight differences from their living counterparts indicates that the deposit

cannot be of very recent origin, but is likely to be at least as old as the upper

Pleistocene. The Vlakkraal deposit in which was found the large Lycaon

incisor previously mentioned is estimated to belong to the upper Pleistocene

(Wells & Cooke 1942). Chemical analyses, carried out through the courtesy °

of Dr. K. Oakley of the British Museum, on a fragment of Hyaena brunnea

mandible indicate that the fluorine content does not differ from that of the

FOSSIL CARNIVORA FROM HOPEFIELD 347

human skull and of Mesochoerus lategani. The carnivore fossils per se do not

provide any grounds on which it would be possible to decide whether the

deposit belongs to the upper Pleistocene, as has been suggested (Singer &

Keen, 1955), or is as old as the top of the middle Pleistocene.

SUMMARY

All the carnivore remains so far recovered from the Hopefield site are described.

These include a new subspecies of Lycaon, Lycaon pictus magnus, a Crocuta spelaea

and a jackal showing characters intermediate between the fossil subspecies from the

Transvaal cave deposits and the living form, together with a Hyaena brunnea and a

Mellivora capensis, each differing very slightly from the corresponding living form.

In addition there are fragmentary remains of a viverrid resembling Herpestes ichneumon

and a felid, probably Leptailurus serval.

It is concluded that the carnivore fauna is consistent with the upper Pleistocene

dating which has previously been suggested for the deposit.

ACKNOWLEDGEMENTS

We are indebted to Dr. Hewitt of the Albany Museum, to Dr. K. H. Barnard

of the South African Museum and to Mr. Skead of the Kaffrarian Museum for their

allowing us every facility for studying skulls in their collections and for the loan of

material. One of us (R.F.E.) is also indebted to the authorities of the British Museum.

(Natural History) for facilities for examining and measuring material in their collections,

and to the South African Council for Scientific and Industrial Research for a research

grant.

Mr. G. McManus, Department of Surgery, University of Cape Town, kindly

photographed the specimens. :

Part of a research grant from the Dr. C. L. Herman Research Fund of the

University of Cape Town allocated to Drs. Singer and Keen was utilized for studying

the carnivora.

One of us (R.S.) is indebted to the Wenner-Gren Foundation for Anthropological

Research, New York, whose motor vehicle, donated to the University of Cape Town,

is being extensively used for collection of material from the site.

Publication of this paper was assisted by a grant-in-aid made by the Council of

the University of Cape Town.

REFERENCES

Broom, R. 1939. A preliminary account of the Pleistocene carnivores of the Trans-

vaal caves. Ann. Transv. Mus. r9, 331-338.

Drennan, M. R. 1954. Saldanha Man and his Associations. Amer. Anthropologist

56, no. 5, 879-884.

DRENNAN, M. R. and Sincer, R. 1955. A Mandibular Fragment, probably of the

Saldanha Skull. Nature 175, 364.

Ewer, R. F. 1954. The fossil carnivores of the Transvaal caves. The Hyaenidae of

Kromdraai. Proc. Zool. Soc. Lond. 124, 565-585.

Ewer, R. F. 1955. The fossil carnivores of the Transvaal caves. The Hyaenidae,

other than Lycyaena, of Swartkrans and Sterkfontein. Ibid. 124, 815-837.

Ewer, R. F. (in press). The fossil carnivores of the Transvaal caves. Canidae. Ibid.

Gotpruss, G. A. 1821. Osteologische Beitrage zur Kenntniss verschiedener Satige-

thiere der Vorwelt. Uber die Héhlen-Hyane (Hydne spelaea). Nova. Acta Akad.

Caes. Leop. 456-462.

Poute, H. 1928. Die Raubtiere von Oldoway. Wiss. Ergeb. Oldoway-Expedit 1913.

N.f. Hfr. 3. 45-54.

Roperts, A. 1951. The Mammals of South Africa. Published by the Trustees of

the ‘‘Mammals of South Africa’’ Book Fund.

SINGER, R. 1954. The Saldanha Skull from Hopefield, South Africa. Amer. J. Phys.

Anthrop. n.s. 12, no. 3, 345-362.

SmncER, R. and Keen, E. N. 1955. Fossil Suiformes from Hopefield. Ann. S. Afr.

Mus. 42, 169-179.

WELts, L. H. and Cooke, H. B. S. 1942. The associated fauna and culture of the

Vlakkraal thermal springs, O.F.S. Trans. Roy. Soc. S. Afr. 29, 203-233.

‘

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Ai |

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Mas Nth {

} ASTI ‘ i x WP een }

a * ih ‘ ¥ ; %

8 q i 4H

ANCE

V AN,

Prate XXVII

Name oe arr. Mus. Vol. XLII

Pirate XXVIII. Crocuta spelaea Goldf, Side view of specimen E.C.1,

ANG, S\, Aver, WIE, Woll, SGU

PLATE XXVIII. Crocuta spelaea Goldf.

PEATE SOGviial

Posterior view of specimen E.C.r.

PLATE XXIX

Nii SmeAdr. Mus), Volk XETI

scale cm

1 2 3

Prare XXIX. Crocuta spelaea Goldf, Ventral view of specimen E.C.1.

ANG, S), Boe, Whos, Woll, SLICE

PLATE XXX. Lingual view of:

Top: Lycaon pictus magnus n.subsp. Specimen

Centre: Crocuta spelaea. Specimen E.C. 5.

Bottom: Crocuta spelaea. Specimen E.C.8.

Nimes Air. Mus Vol. XII PLaTE XXXI

cee

PATE Oe Labial views of the specimens illustrated on Plate XXX.

AWW, S, Ase, Wins, Woll, SILI

PrATE DOxOxXi

—

PLATE XXXII. Mellivora capensis (Schreber).

IS (Caikal.

Dorsal and palatal views of specimen i

18. Further Fossil Suidae from Hopefield. By E. N. KEEN and R. SINGER,

Anatomy Department, University of Cape Town.

(With Plates XXXIII-XXXV.)

INTRODUCTION

In a previous publication (Singer and Keen, 1955) the available suid

material from the fossil site on the farm ‘‘Elandsfontein’’ was ascribed to a

single new species Mesochoerus lategant. Subsequently, on a number of field

trips, one of us (R.S.) and a medical student, A. J. van Niekerk, recovered

not only further specimens of the above genus, but also a fragment of a

mandible containing deciduous third and fourth molars and a permanent

unworn first molar (specimen S. 22) of this genus, as well as a second and

third molar (probably upper) of a different genus, namely Tapinochoerus.

It is the purpose of this paper to describe the new material in the

Anatomy Department : —

A. The additional Mesochoerus specimens so as to increase the range of

variation in Mesochoerus lategani and to evaluate the status of this

species better, and to describe a specimen (S.28) of Mesochoerus

paiceae.

B. The milk dentition in the fragment of mandible numbered $.22.

C. The Tapinochoerus specimen (S.26) and to establish its specific rank.

DESCRIPTION AND DISCUSSION

A. Genus MESOCHOERUS Shaw and Cooke

DESCRIPTION

1. Mesochoerus lategant Singer and Keen.

The general description of these four specimens (S.23, 24, 25, 27) is the

same as in our previously published material (1955), and it is only necessary

to note the specimens and their dimensions, and to outline any additional

distinctive features.

- ae ame et we

FURTHER FOSSIL SUIDAE FROM HOPEFIELD So

Specimen S.23: A left upper third molar in which the posterior part of the

talon consists of only one pillar behind the third lateral pair. This single

pillar is vertically grooved as if in partial separation. The anterior median

pillar and its outriders project beyond the root margin giving the tooth a

pronounced rounded bulge anteriorly, well seen in side view (Pl. XXXIII A).

On reviewing our specimens, this feature is seen in all the third molars

particularly when in early wear. It is also observed in Metridiochoerus as

well as in Mesochoerus olduvaiensis and paiceae, while in Sus limnetes from

Omo (now classified as Kowropotamus = Potamochoerus by Hopwood and

Hollyfield, 1954) the anterior median pillar and its outriders form a separate

plate. In Notochoerus and Phacochoerus species the bulge is absent.

Specimen §.24: A left upper third molar in which the talon behind the

third lateral pair consists of 5 small pillars. The buccal pillar of the second

lateral pair also has an accessory nodule anterior to and confluent with it.

Specimens S.23 and S.24 are in approximately the same stage of late wear,

yet in S.23 the posterior pillars are fused into one massive posterior median

pular. This variation in number of pillars in the talon has already been

commented on in our previous publication.

--Specimen $.25: A right upper third molar whose front portion is missing —

as it is broken through the anterior pair of lateral pillars. The lingual surface

of the 2nd and 3rd lateral pillars shows two deep vertical grooves, giving

an exaggerated appearance not yet encountered in this species and suggestive

of incomplete fusion of the cones which form the complex occlusal pattern

of the teeth (Pl. XXXIII B). Viewed from the occlusal aspect the tooth

presents an unusual concave curvature on the labial side of the posterior part

of the tooth (Pl. XXXIII C) which is produced by a bulge at the crown-root

junction of the 2nd lateral pillar. The roots are missing.

Specimen $.27: An almost complete right upper third molar. The anterior

pair of roots are broken off, and a small piece of the central portion of the

first lateral pillar on the lingual side is missing. The part of the talon behind

the 3rd lateral pair in this massive tooth also consists of 5 pillars. The anterior

median pillar and its outriders are so large as to almost form a separate plate,

while the 2nd lateral pillar on the lingual side has an accessory cone in front

of it (compare S.24) while the pillar on the buccal side has a small nodule

projecting up in front of it at the cingulum. The buccal pillar of the 3rd

lateral pair has 3 small accessory nodules on its outer side. The outer root of

the 2nd pair of lateral pillars, though broken near its tip, is projecting back

acutely to overlap and cover the plate-like roots of the 3rd pillar with which

it is partially fused just below the crown-root junction (Pl. XX XIII D).

352 ANNALS OF THE SOUTH AFRICAN MUSEUM

Maximum Maximum Occlusal Height of unworn

Specimen. Length. Breadth. Breadth. pillars above root.

Se 7s Ot Re MM RD TORY De URN o's LORD wh NT OEE 25-4 212 —

SOSA Ty atin aces WVelee te Mie ete Neen oa yy CO) 25-9 21.4 26.0 (posterior)

SS HTS EES AC CE PE: — C27 C.25 30.0 (posterior)

SA Ant ee) Gent as MTN Cah oe, MI oS Bs 29.7 24.2 28.0 (posterior)

Range of previous specimens (7) 63-70 24-28 20-23 22-35

TaBLE I: Dimensions of the crowns of the upper third molars (mm.).

The length of S.23 and S.24 might appear to exclude them from the range

of variation revealed by the previous series. However, in the previously

described series of lower third molars, of which 6 could be measured, the

smallest was 67 mm. long, while the other 5 ranged from 73 to 77 mm. (mean

75 mm.), but there seemed to be no other reasons for excluding the short tooth

from the series. In the same way, the shortness of S.23 and S. 24 will not

exclude these two upper molars, identical in all other respects, from the

species. The range of variation in length is now increased to 59-70 mm.

Bearing in mind the following description of a different species of Mesochoerus

from our site, and also that no upper teeth of Mesochoerus paticeae have

previously been described for comparison, there is a possibility that the

smaller upper teeth which we have included in our range for Mesochoerus

lategani (e.g., S.24) may yet prove to be the uppers of Mesochoerus paiceae.

From table 1 it is also seen that $.25 and S. 27 are at the upper end of

the range of variation for maximum breadth, and actually $.27 outstrips the

other specimens and increases the upper end of the range to 29.7 mm.

2. Mesochoerus paiceae Broom.

Specimen S.28: A mandibular fragment containing a right third molar

tooth in early wear. There are only 4 pairs of lateral pillars and these are

more widely separted from each other than in Mesochoerus lategani specimens.

The pillars taper to a pointed occlusal end, their bases being wide and partially

fused at the cingulum. The 2nd lateral pillar on the labial side has an

accessory cone anterior to it (Pl. XXXIII E). The anterior median pillar

is pushed somewhat to the lingual side by a large outrider which is continuous

with 6 low nodules fused together to form a plate anterior to the anterior

median pillar.

Dimensions of S.28 (mm.):—

Maximum length 9) 0 OS ty A

Maximum’ breadth... 0° S0 oe" Ve Re eke

Occlusal breadth EUV AMER ec ha 179

Height of unworn 3rd pillar above root .. .. .. .. 31.5

FURTHER FOSSIL SUIDAE FROM HOPEFIELD 353

The individual dimensions of this tooth fall within the previously reported

range of Mesochoerus lategani, with the exception of the maximum length

which is 2 mm. short of the lower end of the range. In itself this would

have been of minor importance. Hower, there are only 4 pairs of well-

separated lateral pillars, as compared with 5 pairs in Jategani, and behind the

fourth pair is a single undivided median pillar. In addition, despite its

shortness, the breadth is at the upper limit of our previous range. Taking

these 3 points into consideration, it cannot be distinguished from the 2

specimens (Broom’s type and Shaw and Cooke’s neotype) on which

Mesochoerus patceae is based.

DISCUSSION

Since our previous publication on the Hopefield Suidae, Hopwood and

Hollyfield’s monograph (1954) has been received. In this they classify

Mesochoerus with Hylochoerus, hereby following Arambourg (1947), although

the latter was not quite certain of this attribution. We have weighed up this

opinion in the light of the reasons which led Shaw and Cooke (1941) to decide

that their genus should be separated from Hylochoerus. Briefly, these were

that the third molars were elongated and not so brachyodont as in Hylochoerus,

and that the premolars were not reduced as in the modern animal. These

points remain, in our opinion, ample justification for maintaining the genus

Mesochoerus which now comprises three species, namely, patceae,

olduvaiensts and lategani. Leakey’s Mesochoerus heselont has been attributed

by Arambourg (1947) to Omochoerus. However, Leakey (personal commu-

nication, 1955) still maintains that Omochoerus should be included under

Mesochoerus. Consequently the position of heselont is held in abeyance.

On a phylogenetic sequence, Mesochoerus would seem a logical intermediate

stage between Hylochoerus and Metridiochoerus. This problem may finally

be resolved by further discoveries, particularly of tusks and other skeletal

remains. |

The specimen S.28, as described above, cannot be grouped with the

previously described M. specimens of Mesochoerus lategani. Its dimensions

and general appearance are such that it must be identified as Mesochoerus

paiceae. In our previous series S.21 was included on the assumption that

a 5th pair of lateral pillars was broken away posteriorly. Re-examination

of this specimen, in the light of the fresh discovery, makes this seem less

likely, and S.2t may represent a second example of a tooth which must be

referred to the genotype patceae.

354 ANNALS OF THE SOUTH AFRICAN MUSEUM

B. MESOCHOERUS. Milk Dentition

Specimen $.22: A left mandibular fragment bearing an unworn M:, Dy

and a root fragment of D, (Pl. XXXIV). |

Dimensions of the fragment :—

Potal engsth (2 Pay Os” ED: ae

Maxamiunt breadth’), 2") 2°05), REI Qo) Oe ht

Height “or mandible at IMG) 08 ey er eae

The first molar is unworn and is composed of 2 pairs of lateral pillars

with a double median pillar between and separating them, small anterior

outriders, and a large complex posterior median pillar. The posterior lobe

of the tooth is broader than the anterior, and its 2 pillars are more separated

from each other. The enamel is coarsely rugose and its irregularities form

a pattern which is best seen on the labial side of the posterior lobe. The

anterior and posterior lateral pillars fuse with each other about 4 mm. above

the cingulum, and the posterior median pillar fuses with the posterior lateral

pillar on both sides. The cingulum is only slightly developed. There are

4 roots which cannot be directly inspected as they are embedded in the

mandible, but they were examined on a skiagram. The posterior roots are

larger than the anterior.

-Dimensions of M, (mm.):—

Maximum length Rian wate uniac mre Ol ieee. 23

Maximum breadth (posteriorly) )0s" 2...) toe 13

Height "ortrown (amterionhyy: Ve yey vee 16

Height" of ‘crown’ (postenorly)) "2 Ve

Root length er ee So ee rr

Behind the tooth is a broad incomplete cavity which presumably contained

the developing M.. The breadth of the mandible diminishes noticeably behind

The last (fourth) deciduous molar is a trilobed tooth with 3 roots on the

labial side and 2 on the lingual side. It is in moderate wear. The 3 pairs

of lateral pillars increase in size and decrease in wear from front to back.

The 2nd lingual pillar is broken. The general appearance of the tooth is

seen in Plate XXXIV. Examination of the fragment and of the skiagram

reveals that the roots of this tooth straddle a cavity which one would have

expected to contain the developing tooth germ of P.. This cavity commu-

nicates widely with the mandibular canal and it seems logical to assume

escape of the tooth germ through this opening. A skiagram of the mandible

of a modern domestic pig at about the same stage of development shows an

unerupted tooth resting in a cavity at exactly the same position in relation

to D, as in the fossil specimen.

FURTHER FOSSIL SUIDAE FROM HOPEFIELD 355

Dimensions of D, (mm.) :—

Mermum tenethe 2.) OF... Mie TAS ae 24

Maximum breadth at bier th (hdsterioniyy) ce) ee Te

Mersin lensth: of roots i(irem! Xray) NP Ae err 8

The third deciduous molar is only represented by a root fragment resting

in one of the 2 alveolar root canals in front of D,.

The bulk of the suid remains collected from Elandsfontein have proved to

belong to the genus Mesochoerus. The dimensions of M. of $.22 compare

very favourably with those of the 2 worn M, teeth of Mesochoerus lategan

previously described, especially if the measurements are taken close to the

cingulum. The unworn height of the crown (16 mm.) compares reasonably

with the unworn height of M; (3rd pillar), which averaged 29 mm. in 4

specimens in which it could be accurately measured. The ratio of the unworn

height of M,/M; is thus 1:1.8. There seems good reason to conclude that

this specimen is derived from a young Mesochoerus.

C. Genus TAPINCCHOERUS van Hoepen and van Hoepen

Tapinochoerus meadowsi Broom

DESCRIPTION

Specimen S.26: A third and a second molar, found lying in apposition,

were subsequently joined together with plaster-of-paris. The second molar is

partially embedded in bone. The third molar resembles the tooth on

which Broom (1928) originally based the species Notochoerus meadows?

( = Tapinochoerus meadowsi, Cooke, 1949). One of us (R.S.) examined

specimens of Tapinochoerus meadowsi from the Transvaal Museum

ee Ske) 286) bE 1), now, being ‘studied by Dr" KR. FF. Ewer,

Rhodes University, Grahamstown. The former two specimens are lower

M. probably from the same jaw, and B.F.1 is a part of a skull, briefly

described by Broom, 1948, containing unerupted M° teeth and partially worn

M’ teeth. As a result of careful comparison, S.26 is considered to consist of

left upper molars.

Left upper third molar: The general structure of the tooth is phacochcoeroid

consisting of two rows of lateral pillars separated by intermediate pillars. The

anterior pair of lateral pillars is in full wear and constitutes the first lobe.

The pair behind this is just coming into wear (Pl. XXXV). _ Posterior to.

this the unerupted portion of the tooth falls away sharply at an angle of |

about 45°. This part of the tooth consists of a large number of pillars

(probably constituting 3 lobes) on each side with an intermediate pillar between .

356 ANNALS OF THE SOUTH AFRICAN MUSEUM

each pair. These lobes are less separated than the type specimen, but

exhibit a similar appearance to S.K.? 387. The extreme posterior end of the

tooth is broken off. The anterior lobe is broader than the second, and is

rounded anteriorly in the occlusal view, with a slight constriction separating

it from the 2nd lobe. The anterior extremity of the lobe consists of a number

of circular and irregularly shaped nodules which cause an anterior bulging

obvious in the lateral view (Pl. XXXV). The worn surface of the lateral

pillars of this lobe are H-shaped with the inner pair of lobules equal in size

and parallel to the outer pair. The bases of all the pillars are flattened from

side to side, and are open; no roots are apparent. This is a typical phaco-

choeroid appearance. The cement covering the tooth is scanty.

Three third molar teeth from the Olduvai Gorge (Coryndon Museum, Old.

B.K.II, Ex. 1953, nos. 109, 160, 448), studied by R. S., are almost identical

in general appearance, with the posterior unworn part of the tooth falling

away sharply at an angle of about 45° from the anterior plane of wear. The

degree of separation of the pillars varies in the 3 specimens, being more marked

in 109 and 448, but the same as S.26 in 160. However, in the latter the outer

enamel is thicker and the tooth as a whole is broader.

Maximum Maximum Occlusal Height of worn Maximum

Specimen. Length. Breadth. Breadth. anterior pillar. Height.

S20 NE eee eae Ran eae et 19.2 63.8 70.0 (2nd lobe)

Type specimen .. .. 76 19 16 55 64

poo Mine | oly Lin Mn od 19.1 15.7 53.1 68.3 (3rd lobe)

Sue ABS ay wie eek Sse IQ.1 15.9 53-0 c.67 (3rd lobe)

Old <BLK ED 10916 i. = 19.5 16.8 — —

Old? BAK E166 ees 5 20.8 19.1 78.0 99.6 (3rd lobe}

Old BUR TLAgS sani a 17-4 17.0 — —

TaBLE II: Dimensions of S.26 compared with those of other third molars

(mm.). The height of 109 and 448 could not be determined precisely,

but the maximum height was considerably in excess of 70 mm. The

lengths of S.K.? 387 and 388 and Old. 109 were more than 65 mm.

The Swartkrans specimens are not fully erupted; therefore less than

full-grown length.

Left upper second molar: This closely resembles the M* of B.F.1 which

has not yet been described in detail (Broom, 1948). Our specimen is phaco-

choeroid in appearance and consists of 2 lobes which are less separated from

each other than in B.F.1. Each lobe has a separate pair of roots. The anterior

roots are smaller than the posterior ones, and the lingual pair smaller than the

labial pair. There is a fairly well-marked cingulum anteriorly and the posterior

aspect of the posterior lobe bulges back to meet the forward projection of

the third molar. The bulge posteriorly of the heel is even more accentuated

in B.F.r. The crown of the tooth is worn in the same plane as the anterior

lobe of M’ (Pl; XXXV).

a dn ee

FURTHER FOSSIL SUIDAE FROM HOPEFIELD 257,

The occlusal pattern of the anterior lobe appears to consist of 2 lateral

pillars fused with an anterior median pillar, and the advanced stage of wear

has left 3 enamel islands within an enamel outline forming two-thirds of a

circle anteriorly and an irregular border posteriorly. The posterior lobe appears

to consist of 2 lateral pillars which have fused in a most irregular manner

with an anterior median, a middle and 2 posterior pillars (Pl. XXXV).

Maximum Maximum Occlusal Height above

Specimen. Length. Breadth. Breadth. cingulum.

5:26 Ee a ee ea 31.8 19.0 18.2 19.1 (anterior lobe)

22.3 (posterior lobe)

SPREE AGN yb 3522 18.6 17.5 —

TaBLE III: Dimensions of M’? (mm.), In S.26 the maximum length is the same

as the occlusal length. Height not measured in B.F.1 because

junction between crown and root not clear.

There is no doubt that S.26 must be identified with the species originally

named Notochoerus meadows: by Broom (1928).

DISCUSSION

In efforts to reduce the number of genera of African fossil Suidae, Hopwood

and Hollyfield (1945) and Arambourg (1947) found Broom’s species difficult

to place. Arambourg (1947) discussed the evolution of the phacochoeres in

some detail and erected a phylogenetic series based on increasing height

of the teeth, increasing length of the third molars, and decreasing separation

of the constituent columns of the third molars. Assuming an origin in some

ancestor resembling Sus or Potamochoerus, his series runs_ through

Hylochoerus to Metridiochoerus Hopwood, and then Notochoerus Broom as

a logical step to Phacochoerus. Without discussing the height or the poor

separation of the columns of the tooth on which it was based, Arambourg

agreed with Broom’s original attribution of the species to Notochoerus.

Hopwood and Hollyfield (1954), evidently unhappy with this opinion, included

the species in Metridiochoerus Hopwood 1926. The height and length of

the specimen, compared with those of the genotype Metridiochoerus andrewsi,

make them uneasy companions.

A similar difference of opinion, in the opposite direction, shows that these

authorities are not wholly in agreement over the definition of the two genera.

Arambourg (1947) considered that Notochoerus dtetrichti Hopwood 1934 should

be included in Metridtochoerus; Hopwood and Hollyfield persist in the opinion

that this is a Notochoerus.

The mean dimensions and characters of the upper and lower third molars

in previously described Metridiochoerus andrewsi specimens may be sum-

marised as follows: maximum length c.60 mm., length of talon (i.e. portion

358 ANNALS OF THE SOUTH AFRICAN MUSEUM

behind the first 2 lobes) c 25 mm., maximum breadth 20-28 mm., unworn

height of crown 50 mm. or less; roots fairly well developed. Notochoerus

capensis third molars have the following characteristics: the tooth is greatly

elongated (max. length 80-100 mm.) by development of the talon (length

50-70 mm.), maximum breadth is 22-30 mm., and unworn height is c 50-60

mm.; roots are well developed and lateral columns are fairly well separated

from each other. The original Notochoeérus meadows: and those third molars

which have been classified with it under the generic name Tapinochoerus may

similarly be summarised: maximum length 70-75 mm., length of talon 35-40

mm., maximum breadth 15-20 mm., unworn height of crown 65-80 mm.

Here the columns are closely packed and there is very little root formation,

the tooth as a whole having a marked phacochoeroid appearance. The

dimensions of the specimens noted and described in this paper increase these

ranges. The degree of separation of the columns (pillars) in these specimens

varies, but none exhibit the fairly marked separation of Notochoerus capensis.

These measurements, together with the general appearance of the teeth

in question, make it in our view unjustifiable to classify meadowsi

specimens as either Notochoerus (after Arambourg, 1947) or as Metnidio-

choerus (after Hopwood and Hollyfield, 1954). In the phylogenetic sequence

Metndiochoerus-Notochoerus-Phacochoerus, the meadows: specimens form a

natural link between Notochoerus and Phacochoerus. We therefore prefer

to retain the genus Tapinochoerus. with two species, meadows: Broom 1928

and modestus van Hoepen and van Hoepen, 1932. The latter, which is the:

genotype, is based on a considerably smaller third molar recovered from

Cornelia, Orange Free State. :

It might be argued that Tapinochoerus.in this sense does not deserve

separation from Phacochoerus. Cooke (1949) wrote: ‘‘Very possibly this may

be regarded as only a sub-genus of Phacochoerus.’’ Arambourg (1947)

stressed the existence in early Quarternary times of a primitive form of

Phacochoerus africanus, larger than the modern types and showing slightly.

greater complexity of enamel pattern. To this he gave the subspecific name,

first used by van Hoepen and van Hoepen (1932), of Phacochoerus africanus

fossilts. Whether this concept should be broadened to include the teeth under

discussion is, in our opinion, doubtful. We prefer to regard the differences

between these teeth and Phacochoerus as of generic rank. .

CONCLUSION

The presence of a second genus Taptnochoerus, which at Olduvai is found:

in Beds II and IV and at Olorgesailie in deposits contemporary with the upper

part of. Bed IV, as well as two. Mesochoerus species, one of which. is:

similar to olduvaiensis found in Beds II and IV, tends to refute the previous

tah me ie i — a a

FURTHER FOSSIL SUIDAE FROM HOPEFIELD 359

hypothesis of the persistence of an isolated species at ‘‘Elandsfontein’’ (Singer

and Keen, 1955). Because Tapinochoerus and Mesochoerus have been

recovered from various parts of South Africa (Vaal River, Swartkrans,

Cornelia) as well as from East and Central Africa, it would now appear

unnecessary to postulate isolation and subsequent evolution of a new species

in the Cape. ‘‘Elandsfontein’’ at Hopefield therefore reflects the widespread

distribution of at least these two suid genera.

The large number of species and genera on record at present may indicate

that if a sufficient number of specimens were recovered the true intraspecific

and intrageneric ranges of variation would appear, and would allow con-

siderable merging of the closely overlapping types, so diminishing the number

of genera and species recognized. On the other hand, it may reflect rapid

evolutionary development in several directions produced by habitat and other

selective factors.

The discovery of Tapinochoerus, which at Olduvai is relatively archaic,

and, in the Vaal River sites is found in the younger gravels, strengthens the

conclusion previously suggested (Singer and Keen, 1955; Singer, 1955) that

the Hopefield deposits should be assigned conservatively to the early Upper

Pleistocene period, equivalent to the interpluvial between the Kanjeran

pluvial and the first Gamblian pluvial of East African chronology, a period

now recognized as being the final phase of the Kanjeran pluvial.

SUMMARY

t. Four additional upper third molars of Mesochoerus lategani are described, thus

increasing the range of variation of the species and improving the previously described

diagnostic characteristics. Another specimen, identical to Mesochoerus paiceae, is

described, and, a specimen previously included with Mesochoerus lategani is now

referred to paiceae. There is also a short discussion on maintaining the rank of

Mesochoerus.

2. The fourth deciduous molar and a first molar in a young Mesochoerus mandible

are described.

3. -Fossil M°® and M? are identified with the species originally described as

Notochoerus meadowsi by Broom (1928). Reasons are given for disagreeing with

opinion expressed that this species belongs to Metridiochoerus or Notochoerus and for

maintaining the use of the name 7apinochoerus to indicate a genus intermediate between

Notochoerus and Phacochoerus.

4. The presence of Tapinochoerus meadowsi and 2 species of Mesochoerus, con-

sidered in conjunction with the other extinct and extant forms found on the farm

‘‘Elandsfontein’’ at Hopefield, strengtherns our view that the deposit may be referred

conservatively to an early Upper Pleistocene period.

ACKNOWLEDGEMENTS

Mr. G. McManus of the Surgery Department kindly photographed the specimens.

Part of a grant to us from the Dr. C. L. Herman Research Fund of the University

of Cape Town was utilized for field trips and technical] assistance.

The Land Rover donated by the Wenner-Gren Foundation for Anthropological

Research, New York, was used for the field trips.

-Annals—3

360 ANNALS OF THE SOUTH AFRICAN MUSEUM

One of us (R.S.) is indebted to the following for their permission to study their

material and to include in this paper observations and measurements made thereon:

Dr. L. S. B. Leakey, Curator, Coryndon Museum, Nairobi;

Dr. A. C. Hoffman, Director, Nasionale Museum, Bloemfontein;

Dr. R. F. Ewer, Zoology Dept., Rhodes University, Grahamstown; and Dr.

J. fT. Robinson, Dept. of Vertebrate Palaeontology, Transvaal Museum.

Grants from the Council for Scientific and Industrial Research, the Wenner-Gren

Foundation and the Dr. C. L. Herman Research Fund made possible visits by R.S.

to these institutions for studying suid and other material.

Dr. K. H. Barnard, Director of the South African Museum, Cape Town has at

all times given us access to extant comparative material, and we are also grateful to

him for his advice and co-operation in the publication of this paper.

REFERENCES

ARAMBOURG, C. 1947. Mission Scientifique de L’Omo, Tome I, Fascicule III}

Paléontologie.

Broom, R. 1928. On some New Mammals from the Diamond Gravels of the Kimberley

District, Ann. S. Afr. Mus., ‘22, 439-444.

Broom, R. 1948. Some South African Pliocene and Pleistocene Mammals, Ann.

Transv. Mus., 21, 1-38.

Cooke, H. B. S. 1949. The Fossil Suina of South Africa, Trans. Roy. Soc., 5S.

Africa, 32, I-44.

Hopwoop, A. T. and HoLtyrietp, J. P. 1954. Fossil Mammals of Africa No. 8.

An Annotated Bibliography of the Fossil Mammals of Africa (1742-1950). London

British Museum (Natural History). ;

MIDDLETON-SHAW, J. C. and Cooke, H. B. S. 1941. New Fossil Pig remains from

the Vaal River Gravels, Trans. Roy. Soc., S. Africa, 28, 293-299.

SINGER, R. 1955. Investigations at Hopefield. Paper presented at Third Pan-African

Congress on Prehistory, Livingstone. Published in summary in Congress

. Programme, and to be published in the “‘Proceedings’’. ‘ doe

StnGER, R, and Kren, E. N. 1955. Fossil Suiformes from Hopefield. Ann. S, Afr.

Mus., 42, part 3, 169-179. ( te

Van. Hoepen, E. C. N. and Van Hoepen, H. E. 1932. Vrystaatse Wilde Varke, Pal.

Nav. Nas. Mus. Bloemfontein, 2, 39-62. ;

nip ing SEE ES il eS AE

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neon Ati Mus. ) Volk Xacil PLATE XXXIII

scale em

DRE:

PLATE XXXIII. Mesochoerus lategani Singer & Keen.

A. Buccal (lateral) view of left upper molar (S 23).

B. C. Lingual (medial) and occlusal views of right upper third

molar (S 25).

D. Buccal view of right upper third molar (S 27).

Mesochoerus paiceae Broom,

E. Lingual view of right lower third molar (S 28).

Ann. S. Afr. Mus., Vol. XLII Pirate XXXIV

PrateE XXXIV. Occlusal, buccal (lateral) and lingual (medial) views of mandibular

fragment of young Mesochoerus (S 22).

om, Gy YM, Wilms. Woll, 2S!bii PLATE XXXV

scale em

1 2 5

PrateE XXXV. Tapinochoerus meadowsi (Broom). Occlusal, lingual (medial) and

buccal (lateral) views of left upper second and third molars,

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