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ANNALS, OF TEE,

SOUTH AFRICAN MUSEUM

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LIST OF CONTENTS

BARRY, -L..H

The cranial morphology of the Permo-Triassic anomodont Pristerodon buffaloensis

with special reference to the neural endocranium and visceral arch skeleton

(published December 1967)

Barry, T. H.

Sound conduction in the fossil anomodont Lystrosaurus (published June 1968)

BoonstRA, L. D.

The girdles and limbs of the Dicynodontia of the Tapinocephalus zone (published

November 1966)

Boonstra, L. D.

The dinocephalian manus and pes (published November 1966)

BoonstrA, L. D.

An early stage in the evolution of the mammalian quadrupedal walking gait (pub-

lished March 1967)

BoonstrRA, L. D.

The braincase, basicranial axis and median septum in the Dinocephalia (published

May 1968)

Hesse, A. J.

Additions to the Cyrtosiinae (Bombyliidae) of South Africa (published December

1967)

Joun, H.

Neue Spezies von Notiophygus Gory nebst Erganzungen (Discolomidae Col.) (pub-

lished December 1967)

KENSLEY, B. F.

Deep sea decapod Crustacea from west of Cape Point, South Africa (published

June 1968)

MiILiLarp, N. A. H.

Hydroids from the south-west Indian ocean (published December 1967)

PEenrRITH, M.-L.

Studies on the South African Clinidae. II. Two new species of Clinus from the

western Cape (published October 1967)

Voss, G. L.

Some bathypelagic cephalopods from South African waters (published November

1967)

Page

195

163

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NEW GENERIC NAMES PROPOSED IN THIS VOLUME

Pseudoglabellula Hesse, 1967 (Bombyliidae), 118

Psiloderoides Hesse, 1967 (Bombyliidae), 121

NEW SUBGENERIC NAME PROPOSED IN THIS VOLUME

Aetheoptilus Hesse, 1967 (Bombyliidae), 112

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INDEX TO GENERA AND SUBGENERA

(SYNONYMS IN ITALICS)

* Dicynodon, 1, 2, 131, 135, 137, 139, 141,

Abralia, 66 _ 198, 260, 270

Abraliopsis, 62, 66 Dimetrodon, 31, 139, 268, 271, 272

Acanthephyra, 284, 285, 310 Doliopteryx, 92, 112

Acryptolaria, 171, 172, 192 E

Aetheoptilus, 92, 108, 112

as as : Eledonella, 63, 85

Empidideicus, 92, 93, 105, 107, 116, 119

Emydochampsa, 155

Emydura, 155

Endothiodon, 151

Enoploteuthis, 70

Ephyra, 310

Eryoneicus, 284, 297

Euanthobates, 89, 93, 106, 116, 118

Alepisaurus, 61, 72, 73, 76, 79, 82, 86

Alopecognathus, 41

Amalopenaeus, 301, 302

Amphisbaena, 155

Amphitretus, 63, 85

Anniella, 140, 141, 153, 280

Anomaloptilus, 93, 108, 113

Anteosaurus, 198, 230, 260

Aphaniotes, 280 F

Aprasia, 280 ’

Austrorossia, 62, 65 Filellum, 171, 175, 192

B G

Blennius, 43 Galacantha, 284, 292

Brachyuraniscus, 1 Galiteuthis, 63, 85

Broilius, 1 Gennadas, 284, 285, 301

Brucella, 176 Glabellula, 92, 105, 107, 112, 118

Glabellula, 107

C . Glyphocrangon, 286, 318

Calliteuthis, 74

Calotes, 153 H

Ceratolaemus, 91, 96, 99, 100, 119 Halicornaria, 172, 184, 191, 192

Ceratophora, 280 Haliporus, 284, 285, 299

Chalydra, 155 Hemisepius, 62, 64

Chelone, 147 Heteroteuthis, 62, 65

Chiroteuthis, 63, 76, 87 Hincksella, 171, 176, 192, 193

Chrysemys, 278 Hipposaurus, 34

Cistecephalus, 132 Histioteuthis, 63, 74, 75

Cladocarpus, 172, 188, 192, 193 Hymenodora, 285, 309

Clinus, 43 Hymenodora, 312

Cophotis, 280 Hymenopenaeus, 299

Cordylus, 140

Cranchia, 63, 82 I

Criocephalus, 196, 197, 229

Crocodilus, 278

Cryptolaria, 172, 174

Ctenopteryx, 63, 73 J

Cyrtisiopsis, 92, 93, 98, 100, 119

Cyrtoides, 107

Cyrtosia, 91, 93, 96, 100, 104, 107, 116, 119,

Iguana, 149

Inioteuthis, 62, 65

Jonkeria, 13, 22, 197, 198, 238, 258, 260

122 K

Kannemeyeria, 155

D Keratocephalus, 196, 197, 219

Daptocephalus, 138, 142, 145, 156 Kingoria, 145, 155, 156

Dasypus, 140 Kirchenpaueria, 172, 184, 192

Desmoteuthis, 84 Koupia, I

VY?)

Labidosaurus, 155

Lacerta, 153, 278

Lacertilia, 149

Lafoea, 171, 175, 192

Lafoéa, 175

Leachia, 81

Leptodontoteuthis, 66

Locustana, 121

Lycoteuthis, 62, 66

Lyriocephalus, 153, 280

Lystrosaurus, 142, 146, 156, 158, 275

Lytocarpus, 191

Maraisaurus, 198, 264

Megalocranchia, 63, 82, 87

Meleagroteuthis, 75

Micrabralia, 70

Micranteosaurus, 13, 14, 17, 23, 25

Monopeltis, 140, 149, 280

Mormosaurus, 196, 211

Moschoides, 13, 15, 19

Moschops, 13, 14, 196, 223, 260

Munida, 284, 287

Munidopsis, 284, 288

Nematocarcinus, 283, 284, 285, 286, 317

Nemertesia, 172, 175, 185, 192

Neolithodes, 284, 286

Notiophygus, 163

Notostomus, 285, 310

Notostomus, 314.

Octopodoteuthopsis, 63, 73

Octopus, 63, 86

Ocythoe, 63, 86

Onchopelma, 91, 104, 121, 123

Onychoteuthis, 62, 72

Onykia, 62, 72

Ophioceps, 153, 155

Ophthalmolophus, 43

Ornithoteuthis, 63, 76

Otsheria, 272

Oudenodon, 138, 146, 156

Palaeogyrinus, 267

Parascapanodon, 15, 23

Pelomedusa, 144

Peltosaurus, 155

Pentacheles, 293

Phrynops, 145, 156

Placerias, 145

Platypygus, 91, 93, 96, 98, 100, 119

Plesiopenaeus, 284, 285, 298

Plumularia, 172, 175, 185, 192, 193

Plumularia, 184

Polycheles, 284, 292

Polycheles, 293

Pontophilus, 283, 284, 286, 319

Pristerodon, I, 131

Pristerognathoides, 198

Pseudoglabellula, 92, 106, 112, 118

Psilodera, go, 121

Psiloderoides, 90, 121

Pterygioteuthis, 62, 71

Pyrgopsis, 63, 79

Pyroteuthis, 62, 70

Reticularia, 175

Rhachocaris, 318

Rhineura, 280

Rhombosepion, 64.

Robertia, 1, 3

Rossia, 62, 65

Salacia, 171, 179, 192

Scelotes, 141, 280

Scleyocrangon, 284, 286, 318

Scymnosaurus, 40

Semirossia, 65

Sepia, 62, 64

Sergestes, 284, 285, 302

Sergia, 303, 308

Sertularella, 171, 180, 192

Sertularella, 182

Sertularia, 176, 179

Sphenodon, 140, 146, 152, 156, 278, 279

Spirula, 62, 63

Stahleckeria, 155

Stegopoma, 171, 172, 192

Stephanoteuthis, 65

Stereomastis, 284, 293

Stigmatoteuthis, 74.

Struthiocephalus, 14, 15, 195, 196, 197, 199,

256, 260

Symplectoscyphus, 171, 182, 192

Synostocephalus, 158

Systellaspis, 284, 285, 309

Systoechus, 124

Tapinocephalus, 1, 13, 22, 27, 255, 264, 266

270, 271, 272, 273

Terphis, 121

Testudo, 278

Tetronychoteuthis, 62, 72

Teuthowenia, 84

Thyllis, 121

Titanophoneus, 13, 18, 24, 25

Todaropsis, 63, 76

Tremoctopus, 63, 86

Tripteris, 98

Trogonophis, 153

Tropiocolotes, 152

Tympanocryptis, 280

Typhlosaurus, 280

Venjukovia, 137

Willemoesia, 284, 294

Zygophylax, 171, 172, 176, 192

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4 :

LIEUWE DIRK BOONSTRA

THE GIRDLES AND LIMBS

OF THE DICYNODONTIA OF THE

TAPINOCEPHALUS ZONE

November 1966 November

Volume 50 Band

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THE GIRDLES AND LIMBS

OF THE

DICYNODONTIA OF THE TAPINOCEPHALUS ZONE

LIEUWE Dirk BOONSTRA

(With 6 Text-figures)

CONTENTS

PAGE

Introduction I

Material 2

Pectoral Girdle ‘ , . i Sinead

Pelvic Girdle : ; : : F ies

The Humerus : : : wo

The Femur . PRUE NG

The Anterior Epipodial, Carpus ang Maris a >,

The Pes : F a se akO

Discussion . ; : : , : fy Ed

Summary . é : : ; a

Aguneny seamen : : é : mah

INTRODUCTION

The oldest specimens of the Dicynodontia as yet recovered from the Karroo

are from the Tapinocephalus Zone of the Lower Beaufort Beds. These are of both

the families Endothiodontidae and Dicynodontidae. Hitherto six genera and

sixteen species have been described—all based on cranial material alone, viz.

Endothiodontidae

Brachyuraniscus broomi

Brachyuraniscus merwevillensis

Brachyuraniscus reuningt

Broilius antjtesfonteinensis

Koupia koupensis

Pristerodon brachyops

Robertia broomiana

Dicynodontidae

Dicynodon antjiesfonternensis

Dicynodon gamkaensis

Ann. S. Afr. Mus. 50 (1), 1966: 1-11, 6 figs.

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dicynodon haughtoniscus

Dicynodon huener

Dicynodon jouberti

Dicynodon megalorhinus

Dicynodon pseudojouberti

Dicynodon schroedert

Dicynodon vanderhorstt.

The Dicynodont fauna at this time consisted of an assemblage of small

reptiles, with skull lengths varying from 40 to 130 mm. These small reptiles

occur abundantly in the Tapinocephalus Zone. The South African Museum

Collection alone contains 705 specimens. By far the greater number of specimens

have been found as eroded fossils lying loose on the surface. Although a number .

have been found as isolated specimens—some still in sztu—they mostly occur

concentrated in small patches of loose lying rubble. These have been weathered

out of thin layers of about 3-5 inches thickness in the mudstone. In the upper

part of the zone these layers tend to be arenaceous. In a number of places in

the Koup I have collected over a hundred specimens still zm situ in such thin

layers. These specimens are mostly of skulls and parts of skulls with postcranial

parts very rare. This material has obviously been washed into shallow pans or

depressions, where they were covered up by the inflowing silt. The paucity of

postcranial material is in all probability due to the activities of the contemporary

carnivores and carrion eaters.

In one locality only—on the farm Michau’s Request—have I found a

number of skeletons entombed together in a fairly complete state in a purplish

layer of fine mudstone of about 12 inches thickness. This find is of a number of

these reptiles which had been overwhelmed by some catastrophe and rapidly

imbedded.

These small Dicynodontia must have been present in large numbers

(flocks?) and formed the diet of the numerous contemporary Therocephalia.

Although occurring throughout the zone the patches where concentrated

assemblages have been encountered are mostly in the upper part of the zone.

Often these patches include remains—also mostly skulls—of medium-sized

Therocephalia.

Hitherto no part of the postcranial skeleton of any Dicynodont from the

LT apinocephalus Zone has been described.

MATERIAL

In this contribution an account will be given of the girdles and limbs of

these specimens from the Tapinocephalus Zone in the collection of the South

African Museum in which parts of these structures are preserved. From the

following table of the available material it is evident that much more collecting

will have to be done before a really adequate picture of this early stage in the

development of the girdles and limbs of the Dicynodontia can be given.

DICYNODONTIA OF THE TAPINOCEPHALUS ZONE 3

SAM 11588 Endothiodontid. Skull, partial pectoral girdle and humerus.

Cypher, Beaufort West, Low Tapinocephalus Zone. Collected

Boonstra 1940.

SAM 11760 Robertia broomiana. Skull, humerus and epipodial.

Klein-Koedoeskop, Beaufort West, Low Tapinocephalus Zone.

Collected Boonstra 1929.

SAM 11825 ? Partial pelvis.

Dubbelefontein, Beaufort West, High Tapinocephalus Zone.

Collected Boonstra 1947.

SAM 11883 ? Vertebral column and ribs with partial girdles and a femur.

Steenboksfontein, Laingsburg, Middle Tapinocephalus Zone.

Collected Boonstra 1948.

SAM 11885 Endothiodontid. A number of fairly complete skeletons.

Michau’s Request, Beaufort West. Low Tapinocephalus Zone.

Collected Boonstra and Jooste 1948.

SAM 12255 ? Part of pectoral girdle, humerus and epipodial.

Beukesplaas, Fraserburg. Low? Tapinocephalus Zone. Collected

Boonstra and Zinn 1959.

SAM K259 ? Partial pectoral girdle.

Plaatdorings, Beaufort West. High Tapinocephalus Zone. Collected

Boonstra and Zinn 1960.

SAM K1i134 ? Skull and pectoral girdle.

Lammerkraal, Prince Albert. High Tapinocephalus Zone. Collected

Boonstra and Zinn 1959.

PECTORAL GIRDLE

(Fig. 1)

The following description of the pectoral girdle is based on SAM 11588

consisting of a left scapulo-coracoid and cleithrum, SAM 11885 in which there

are a number of girdles partially preserved, SAM 12255, with most of the left

half of a girdle, SAM K259, with an incomplete left scapula and an interclavicle

and SAM K1134 with a complete girdle preserved.

Of the last specimen I have made a plaster model enlarged three times and

checked with the others, and this forms the basis of the accompanying figures

(ier):

The coraco-scapula is well developed with the scapular blade lying at

right angles to the vertebral axis, with only a slight curving around the thorax;

the coracoidal plate is fairly large, moderately long but low, with the greater

part of its ventral edge resting on the stem of the interclavicle. There is no

supra-glenoid buttress or foramen. The scapular facet of the glenoid faces

mostly backwards and the coracoidal facet upwards and outwards. The

precoracoid does not enter the glenoid.

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

The scapular blade is high with its dorsal part expanded and its outer

face flattened. Along the inside of its anterior face it receives the splint-like

cleithrum. Lower down the anterior edge of the scapula is everted and forms a

well-developed and prominent antero-laterally directed acromion process. The

Dicynodontia are thus the earliest Therapsids in which the acromion process is

developed and when encountered as low down as the Tapinocephalus Zone it is

already as well developed as in any later Dicynodont. None of the other

contemporary Therapsids have even an incipient acromion process. The upper

end of the clavicle is applied to the inner face of the acromion process. No scar

for the scapular head of the triceps can be seen on the posterior face of the

scapula above the glenoid.

Fic. 1. Dicynodon jouberti SAM K1134. Pectoral Girdle x 1.

a. Ventral. b. Anterior. c. Lateral. All figures are orthoprojections.

ac—acromion process PC—precoracoid (anterior coracoid)

C—coracoid (posterior) SC—scapula

CL—clavicle scf—supracoracoid foramen

CM-—cleithrum ST—sternum

ICL—interclavicle

The precoracoid is relatively small; this is mainly due to the lack of

development in its anterior part. Its anterior edge does not reach the plane

of the clavicle as it does in all the other contemporary Therapsids. The pre-

coracoid is pierced by a moderate foramen supracoracoideum lying in the acute

angle formed by the precoracoid anterior to and below the lower edge of the

glenoid.

The coracoid is a robust bone with a fairly long upturned posterior process,

but there is no special protuberance or scar for the coracoidal head of the

triceps.

The dermal clavicular girdle is moderately well developed but the

cleithrum is a weak splint-like element lying applied to the inner anterior edge

of the scapular blade.

DICYNODONTIA OF THE TAPINOCEPHALUS ZONE 5

The interclavicle is fairly short; its anterior end, which is expanded,

curves slightly upwards and its lateral corners are underlain by the truncated

lower ends of the clavicles; the median stem is broad but short without a

definite waist or expanded posterior end. The function of an expanded posterior

end is apparently exercised by the well-developed ossified sternum.

The clavicle is a well-developed bone, with its expanded ventral end

truncated and underlying the antero-lateral corner of the spatulate anterior

end of the interclavicle. It ends well away from the median line and does not

extend posteriorly under the interclavicle. This relation is thus much as in the

contemporary Gorgonopsians and Dinocephalians and quite different from

that obtaining in the pristerognathid Therocephalians. From its ventral

expanded end the clavicle narrows and curving upwards has its upper end

applied to the inner face of the acromion process. The lower part of the scapula

and the precoracoid, having little anterior extent, do not lie applied to the

inner face of the clavicle, but lie free of it with their anterior edges in a more

posterior plane.

The sternum is ossified as a large broad and squarish plate, with its

anterior end overlying the posterior end of the stem of the interclavicle, and its

antero-lateral edges are underlain by the inner edge of the coracoids. There

are no ossified ribs articulating with the sternum.

PELvic GIRDLE

(Fig. 2)

Little and poorly preserved pelvic material is available for study. The

accompanying figure is a reconstruction based mainly on the specimens

SAM 11825 and 11885.

Notwithstanding the inadequate material it is possible to give, in general

terms, a statement on the overall nature of the pelvis in these oldest known

Dicynodontia, viz. the pelvic girdle is high, short, the pubo-ischium not plate-

like, but V-shaped, symphysis absent, pubo-ischiadic fenestra developed; the

acetabulum situated above the pubis with its anterior rim on the anterior

border of the pelvis. The Dicynodonts of the Tapinocephalus Zone are thus the

first Therapsids to accomplish the rotation of the pelvis in relation to the

acetabulum. This is a fatt accompli and we know no older forms showing the

steps by which this was achieved.

The ilium is high; the iliac blade is large and lies diagonally with the

large anterior process directed upwards; the posterior process is weaker, but

well developed; the supra-acetabular buttress is strong; about half the

acetabulum is formed by the ilium.

The pubis is very short, with its anterior border in a plane posterior to the

anterior rim of the acetabulum; its anterior edge carried a well developed tuber

which is strongly everted; there is no pubic foramen, but the pubis forms the

anterior border of the large pubo-ischiadic fenestra. Ventrally the two pubes

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 2. Dicynodont. Pelvic Girdle x 1. Lateral view.

ap—anterior process of the iliac blade IS—ischium

b—supra-acetabular buttress P— pubis

f—pubo-ischiadic fenestra pt—everted pubic tuber

IL—ilium

meet at an acute angle, but there is no symphysis and no median keel is

developed.

The ischium is long; the pair lie at an acute angle, with each other and no

median keel is developed. The ischiadic tubera are moderately well-developed.

Anteriorly the ischium is pierced by a well-developed pubo-ischiadic fenestra.

Here again these early Dicynodonts are the first known Therapsids where the

old pubic foramen is superseded by a pubo-ischiadic fenestra.

THe HumERuS

(Fig. 3)

An indifferently preserved humerus is present in SAM 11760 and in

SAM 11885 there are seven humeri and SAM 11588 and 12255 have each a

well-preserved humerus.

The humerus is a robust bone with greatly expanded ends, a very strong

delto-pectoral crest and a short stout shaft. The rotation of the ends on the

shaft varies from 30°—56°.

The proximal surface has the processus medialis and processus lateralis

indistinctly demarcated from the caput, which is flattened and narrowly oval

in outline.

The delto-pectoral crest is very strongly developed and extends far distally

with a thickened bulbous corner for the insertion of the strong m. pectoralis;

from here it subsides abruptly into the shaft from where a rounded ridge extends

obliquely on to the entepicondyle.

DICYNODONTIA OF THE TAPINOCEPHALUS ZONE 7

The bicipital fossa is large and fairly deep with a rounded posterior rim,

whose posterior face forms a strong and long rectangular area for the origin

of the medial head of the m. triceps.

The proximo-dorsal face of the humerus is divided by the anterior dorso-

ventral line (ADVL) into two parts. Preaxially to this line lies a triangular area

for the insertion of the strong m. deltoideus. Posterior of this line lies the area of

insertion for the strong m. latissimus dorsi, whose main insertion is into a hollow

extending up to the processus medialis.

Af =

qd b @ d

Fic. 3. Dicynodont. SAM 11588. Humerus x 2

a. Proximal. b. Posterior. c. Dorsal. d. Ventral.

advl—anterior dorso-ventral line f—entepicondylar foramen

bf—bicipital fossa ld—insertion of m. latissimus dorsi

c—caput humeralis Iml—latero-median line

dpc—delto-pectoral crest pl— processus lateralis

ec— ectepicondyle pm—processus medialis

en—entepicondyle rc—radial condyle (capitellum)

Distally the epicondyles are both well developed and robust indicating

strong flexors and extensors. The entepicondylar foramen is large and oval in

outline. There is no ectepicondylar foramen.

The distal condyles are not strongly developed or well moulded. The

radial condyle is directed much ventrally and the epipodial capable of full

extension. On the dorso-distal surface the trochlear fossa is very shallow, which

feature is related to the absence of an olecranon process to the ulna.

THE FEMUR

(Fig. 4)

In SAM 11855 there are six femora, all small and with the various features

not very well shown. The accompanying figure is composite with the features

shown only diagrammatically correct.

8 . ANNALS OF THE SOUTH AFRICAN MUSEUM

pifi

a D

Fic. 4. Dicynodont. SAM 11885. Femur x 1.

a. Dorsal. b. Ventral.

c—caput femoris in—internal trochanter (minor)

ec—ectocondyle pifi—insertion of m. pubo-ischio-femoralis

en—entocondyle internus

ex—external trochanter (major) 3— ? third trochanter

The femur, longer than the humerus, is a fairly light bone with only

slightly expanded ends and a long fairly slender shaft; the preaxial border is

concave longitudinally.

The distal condyles lie in the same plane, not well moulded, terminal but

facing somewhat ventrally.

Proximally the caput femoris is terminal, but directed appreciably pre-

axially and dorsally; widely oval and curving towards the external trochanter

into which it flows. From the not prominent external trochanter the postaxial

edge is thickened rugosely and is turned slightly ventrally; the distal end of this

edge forms a slight protuberance and this muscular scar apparently represents

a third trochanter.

On the dorso-proximal surface, near the caput femoris, there is an

indication of a muscle-scar for the m. pubo-ischio femoralis internus.

On the ventro-proximal surface there is a slight, low mound near the

preaxial border of the bone and this represents an internal trochanter for the

insertion of the m. pubo-ischio femoralis externus.

THE ANTERIOR EPIPODIAL, CARPUS AND MANus

(Fig. 5)

A good epipodial is preserved in SAM 12255 and in SAM 11760 and in

SAM 11885 there are six fairly complete radii and ulnae preserved in natural

relation. Under this number there are also eight more or less complete carpi

and manus. All are small and not very well preserved so that few details of the

structure of the individual bones can be given. The figure given is composite

incorporating features from the various specimens.

The olecranon process of the ulna is feebly developed. Proximally the

radius abuts against a facet on the ulna and thus forms a continuous articulatory

DICYNODONTIA OF THE TAPINOCEPHALUS ZONE 9

face sliding round the distal humeral end in extension and flexion. Both the

ulna and radius have expanded distal ends.

The proximal row of carpals consists of three elements—a radiale of

moderate size, rounded in outline, an ulnare, well developed and elongated,

and a laterally compressed intermedium. A disc-like pisiforme lies laterally

of the ulna-ulnare articulation. :

In the middle row there are two centrals—one, elongated lies between the

radiale and the last three distals; the central one, squarish in outline, lies

flanked by the ulnare and the first central, with the intermedium proximal to it

and the enlarged fourth distal distal to it.

There are four distals; the first three small and the fourth, with the fifth

incorporated, large.

The metacarpals are moderately well developed —all elongated and fairly

slender; they increase in robustness from one to four and the fifth about the

size of the third.

The phalangeal formula is 2, 3, 3, 3, 3.

The proximal phalanges are all elongated, with a long shaft or waist. The

first is the smallest and is lightly built and fairly short; the second is longer

and the third is quite a long and robust bone; the fourth of the same length

_ but of lighter build and the fifth much shorter.

The middle phalanges, smaller than the proximal ones, follow the same

pattern of size as the proximal phalanges.

Fic. 5. Dicynodont. SAM 11885. Epipodial and forefoot x .

c—centrals R—radius ~ u—ulnare

i—intermediam ~ r—radiale ~ -1-4—distal carpals

p—pisiforme U—ulna m—fifth metacarpal

IO ANNALS OF THE SOUTH AFRICAN MUSEUM

The ungual phalanges are quite broad and apparently carried broad and

long nails adapted to digging.

The third digit is the longest and the purchase of the foot mesaxonic.

Tue PEs

(Fig. 6)

In SAM 11885 I have a nearly complete hindfoot. The preserved

mineralized bone has been dissolved with dilute formic acid to leave a good

impression of the dorsal surface. ,

In the proximal row of the tarsus there is a robust astragalus broader than

long and a large disc-like calcaneum. The tibia and the fibula articulate with

the distal surfaces of the astragalus and calcaneum respectively. |

In the middle row there are two centrals. The preaxial one is the larger

and lies distally of the astragalus; the smaller central lies lateral of the first

and it articulates with the inner edge of the calcaneum.

There are five distal tarsals, all pebble-like except the first, of which the

impression in the mould shows only a crescent-shaped ridge. The fifth distal

lies far proximally to abut against the outer edge of the calcaneum.

The five metatarsals are of moderate length, but the first is squat.

The phalangeal formula is 2, 3, 3, 3, 3.

Fic. 6. Dicynodont. SAM 11885. Epipodial and hindfoot x 3.

a—astragalus F—fibula 1—5—distal tarsals

ca—calcaneum T—tibia m—first metatarsal

c—centrals

DICYNODONTIA OF THE TAPINOCEPHALUS ZONE i

The phalanges are all fairly long with a good shaft. The proximal one of

the third digit is the longest. The ungual phalanges are broad as in the forefoot

and probably also carried broad and long nails.

The third digit is the longest and the purchase of the foot was mesaxonic.

DISCUSSION

As I have now completed my study of all the available material of the

girdles and limbs of all the Therapsids of the Tapinocephalus Zone, I am engaged

on a comparative account which will form the concluding paper of this series.

SUMMARY

Descriptions are given of the girdles and limbs of the Dicynodontia of the

Tapinocephalus Zone in South Africa. Postcranial parts are rarely found and of

the 705 specimens in the South African Museum eight have parts of the girdles

and limbs present and this account is based on these specimens.

ACKNOWLEDGEMENT

The Trustees of the South African Museum are grateful to the Council

for Scientific and Industrial Research for a grant to publish this paper.

INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,

including TABLE OF CONTENTs and SumMArRy. Position of text-figures and tables must be

indicated. ;

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 43 in. = 7 in. (74 in.

including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in

alphabetical order of authors’ names (Harvard system). References at end of paper must be

given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,

not ‘and’. Year of publication; several papers by the same author in one year designated by

suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper

names (except in German). Title of journal, abbreviated according to World list of scientific

periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B.).

Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (bold type).

Part number, only if separate parts of one volume are independently numbered. Page numbers,

first and last, preceded by a colon (without prefix ‘p’). Thus:

SmitH, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;

date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,

with wavy underlining; place of publication; name of publisher. Thus:

BRown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of

author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of

author of book; his initials; title of book, underlined; edition, if any; volume number, if any,

in arabic numerals, with wavy underlining; pagination of paper; place of publication; name

of publisher. Thus:

SmiTH, C. D. 1954. South African Plonias. Jn Brown, X. Y. Marine faunas. and ed. 3: 63-95.

London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species

has been previously designated (subsequent to 1758) are listed in chronological order, with

abbreviated bibliographic references to descriptions or citations following in chronological

order after each name. Full references must be given at the end of the paper. Articles and

recommendations of the Iniernational code of zoological nomenclature adopted by the XV International

congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:

Euplonia capensis (Smith) Brown, 1955: 259.

When misidentified as another species:

Plonia natalensis (non West), Jones, 1956: 18.

When another species has been called by the same name:

[non] Plonia capensis: Jones, 1957: 27 ( = natalensis West).

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LIEUWE DIRK BOONSTRA

THE DINOCEPHALIAN MANUS AND PES

November 1966 November

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THE DINOCEPHALIAN MANUS AND PES

LIEUWE Dirk BOONSTRA

(With to Text-figures)

CONTENTS

PAGE

Introduction . : 5 shag

Material . ‘ ‘ ; A), 5b8

Forefoot

Tapinocephalia . ; Mens

Titanosuchia ‘ , 706

Anteosauria : ; AN ity

Hindfoot

Tapinocephalia . ; Pritt

Titanosuchia f ‘ DD

Anteosauria : : itn

Discussion d : : » 25

Summary : : ; 35 20

Acknowledgement . j 526

References ‘ : ‘ Hi 26

INTRODUCTION

Very little is known of the feet of the South African Dinocephalia.

From a mixed lot of disarticulated elements Gregory produced the mount

of Moschops, which includes the reconstructed feet, but in his description of

1926 admits his inability to assemble either foot.

In 1929 Broom described the two proximal tarsal elements of Jonkeria.

In 1940 Byrne very briefly described both the fore- and hindfeet of

Moschoides.

In 1954 I described a partial Moschopid carpus and gave descriptions and

figures of the fore- and hindfoot of Micranteosaurus, which I at the time completely

misinterpreted.

From the Russian deposits Orlov in 1958 described and gave beautiful

figures of both fore- and hindfeet of Titanophoneus.

MATERIAL

After 35 years of collecting in the Tapinocephalus zone, during which period

I have excavated two hundred and fifty-one specimens of the Dinocephalia, I

45)

Ann. S. Afr. Mus., 50 (2), 1966: 13-26, 10 figs.

ANNALS OF THE SOUTH AFRICAN MUSEUM

have a poor collection of foot material. This is due to the nature of the preserva-

tion of the Dinocephalian material in this zone. I know of only one case where

a skeleton more or less articulated has been found. In all other cases the bones

of the skeleton are disarticulated and scattered with the loss of most of the

smaller elements composing the feet.

SAM 4323

SAM 9157

SAM 12011

SAM 12017

SAM 12033

SAM 12065

SAM 12104

SAM 12105

SAM 12109

SAM 12110

SAM 12210

Micranteosaurus parvus. An incomplete fore- and hindfoot associated

with a good snout.

Merweville Commonage, Low Tapinocephalus zone. Coll.

Haughton 1917.

Moschops? An incomplete carpus associated with a humerus, ulna

and radius.

Wolwefontein, Prince Albert, Low Tapinocephalus zone. Coll.

Boonstra 1929.

Struthiocephalus ? ‘Tibia and astragalus.

Rietfontein, Beaufort West. Low Tapinocephalus zone. Coll.

Boonstra and Zinn 1956.

Tapinocephalian. A calcaneum and fibula with a piece of skull.

Spitskop, Laingsburg. Low Tapinocephalus zone. Coll. Boonstra

and Zinn 1956.

Tapinocephalian. Radiale.

Worsteling, Laingsburg. Low? Tapinocephalus zone. Coll. Boonstra

and Zinn 1956.

Struthiocephalus ? Two calcanei found on a small slope which yielded

a lot of bones together with some cranial material of Struthioce-

phalus.

Skoenmaker, Beaufort West. Low Tapinocephalus zone. Coll.

Boonstra 1957.

Titanosuchian. An isolated astragalus.

Kalkkraal, Prince Albert. Low Tapinocephalus zone. Coll. Boonstra

and Zinn 1957.

Titanosuchian. An isolated astragalus.

Kalkkraal, Prince Albert. Low Tapinocephalus zone. Coll. Boonstra

and Zinn 1957.

Titanosuchian. An isolated radiale.

Kalkkraal, Prince Albert. Low Tapinocephalus zone. Coll. Boonstra

and Zinn 1957.

Titanosuchian ? Radius and intermedium.

Kalkkraal, Prince Albert, Low Tapinocephalus zone. Coll. Boonstra

and Zinn 1957.

Titanosuchian. An isolated astragalus.

Kroonplaas, Beaufort West. High? Tapinocephalus zone. Coll.

Boonstra and Zinn 1959.

THE DINOCEPHALIAN MANUS AND PES 15

SAM 12226 Struthiocephalus sp. Hindfoot and manus associated with some

teeth.

Knoffelfontein, Beaufort West. Coll. Boonstra and Zinn 1959.

SAM Keo1 Struthiocephalus sp. Isolated astragalus.

Paradys of Rietfontein, Prince Albert. Middle Tapinocephalus

zone. Coll. Boonstra 1959.

SAM K249 Parascapanodon sp. Disarticulated skeleton without skull, including

two calcanei, one astragalus and other carpal, tarsal and digital

bones.

Steynskraal, Beaufort West. Middle Tapinocephalus zone. Coll.

Boonstra and Zinn 1959.

SAM Ka71 Tapinocephalian. Isolated calcaneum and astragalus.

Wonderboom of Plaatdorings, Beaufort West. Low Tapinocephalus

zone. Coll. Boonstra, Zinn and Boonstra, 1960.

SAM K323 Tapinocephalian. Scattered foot bones associated with teeth.

Die Bad, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra,

Zinn and Gow, 1960.

SAM K362 Tapinocephalian. Disarticulated carpals, tarsals and phalanges

associated with cranial and dental material.

Twee Susters of Grootfontein, Fraserburg. Low? Tapinocephalus

zone. Coll. Boonstra and Zinn 1962.

SAM K366 Moschopid? Isolated Calcaneum.

Moutonsfontein, Fraserburg. Low? Tapinocephalus zone. Coll.

Boonstra and Zinn 1962.

FOREFOOT

LT apinocephalia (Fig. 1)

With so little and such poor material available only a tentative description

can be given of the Tapinocephalian manus. Of the seven specimens in which

elements of the forefoot are preserved four have only disarticulated carpal and

digital bones preserved. In SAM 9157 parts of four proximal carpals are

present in articulation; in Romer’s Chicago specimen of Moschoides (which I

have not seen) an articulated manus is preserved and in SAM 12226 an incom-

plete and partially disarticulated manus of a species of Struthiocephalus is

available for study.

The Tapinocephalian manus is broad and very short with little difference

in the length of the toes, but the fourth is the longest digit. The purchase of the

foot is thus mainly mesaxonic but somewhat more post- than preaxial. The

body weight is mainly transmitted through the radius on to the robust ovoid

radiale. The extension and twist of the foot during the stride is executed through

the ulna articulating with a flattish plate-like ulnare supported postaxially by

the pisiforme and preaxially by a laterally uncompressed intermedium.

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 1. Struthiocephalus sp. SAM 12226 x 4

Dorsal view of left manus as restored.

In the middle of the foot there are two centralia, of which the proximal one

is the larger, circular in outline; the distal or inner centrale is oval in outline.

There are five distalia of which the fourth is the largest; the first three are

broader than long and the other two approximately as long as broad.

The first four metacarpals are very short, but the fifth is quite a large bone.

The digital formula is 2, 3, 3, 3, 3. In each digit the proximal phalanx

is very short; in the last four digits the second phalanx is slightly longer.

The terminal or ungual phalanges are broad and carried broad, slightly

convex nails.

Titanosuchia (Fig. 2)

In the Titanosuchia even less material of the manus is available. In

SAM Ke249 a number of disarticulated foot-bones are preserved in very good

condition but reassembly as in the figure is an act of faith. As reassembled there

are three proximal, two central and five distals in the carpus and the digital

formula 2, 3, 3, 3, 3.

The radiale is a strong bone oval in outline and both the intermedium and

proximal central are laterally compressed elements. A pair of beautifully

preserved ulnaria are preserved. The ulnare is a robust bone with large, well-

THE DINOCEPHALIAN MANUS AND PES 37

Fic. 2. Parascapanodon sp. SAM K24g9 x 4

Dorsal view of right manus as restored.

developed convex distal as well as proximal articulatory faces; dorsally the

surface is shallowly concave and ventrally deeply concave; medially the face

is deeply excavated and with a similarly excavated lateral face of the contiguous

centrale a long tube is formed to house the penetrating carpal nutritive and

innervating vessels.

The fourth distal is the largest of the distalia. The metacarpals are very

short and have well modelled convex faces both proximally and distally. The

fifth metacarpal is large and broad.

Anteosauria (Fig. 3)

From the Tapinocephalus zone I have only one specimen of the Anteosaurs—

the type of Micranteosaurus parvus—in which the forefoot is preserved. With

better technical equipment now available I have prepared the specimen

further and with increased knowledge of the structure in related forms I wish

to correct the misinterpretation I made in my account of 1954.

In the proximal row of the carpus there were three bones. The ulnare is a

flattened element with oval proximal and distal articulatory facets; the outer

edge is thin but the inner thickened and rounded, lying preaxially of the ulnare

is the intermedium; as preserved it is a thin bone showing a large, flat upper

face, but it is possible that it has fallen out of position and would then actually

be a laterally compressed bone. Preaxially lies what appears to be a pear-

shaped element. I interpret this as a proximal rounded radiale with the distal

part actually a central. The lateral central is not preserved.

There are five distals of which the fourth is a large bone with a roughly

rectangular upper face. The other distals are pebble-like,

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

All five metacarpals are preserved; they are all elongated bones with

expanded ends and a constricted shaft. They increase in size from I to 5, with

the fifth a robust bone and the first quite feeble.

The phalanges of only four digits are preserved. The phalangeal formula

is 2, 3?, 4, 3, 3, with the third digit the longest and the fourth and fifth only

slightly shorter. The first digit is short and feeble. The purchase of the foot thus

lies mostly in the postaxial part of the foot.

The proximal phalanges of the last three digits have greatly expanded

proximal ends.

The third phalanx of the third digit, although smaller than the first and

second phalanges, is not much reduced and apparently not in the process of

being lost.

The terminal phalanges are narrow and pointed and would have carried

sharp curved claws.

Fic. 3. Micranteosaurus parvus. Type. SAM 4323 x 4

Dorsal view of left manus as restored.

In Titanophoneus, Orlov found the carpal formula to be 3, 2, 5, with the

intermedium laterally compressed and the fourth distal enlarged. The first

metacarpal is small and short and from the second to the fifth became progres-

sively longer with the fifth a quite stout bone with expanded ends. Orlov gives

the digital formula as 2, 3, 3, 3, 3, with the second phalanx of the fourth digit

showing proximally what looks like an epiphysis, which may represent an addi-

tional phalanx fused to it. No such structure is shown in the third digit.

The reduction in the number of phalanges in the third and fourth digits

has thus followed a different course in Titanophoneus and Micranteosaurus, with

THE DINOCEPHALIAN MANUS AND PES 19

the condition in the latter more primitive than in the former.

The purchase of the foot lies more postaxially in Titanophoneus than in

Maicranteosaurus.

HINDFOOT

Tapinocephalia (Figs. 4-6)

I have 5 calcanei, 3 astraguli, a number of disarticulated metatarsals and

phalanges and one nearly complete pes available for study. In Romer’s Chicago

specimen of Moschoides there is a complete left pes.

In SAM 12226 (fig. 4) it is seen that the proximal bones of the tarsus are

very well developed, but the central and distal tarsals fairly weak. The meta-

carpals and digits are short with the fourth digit slightly the longest and the

purchase of the foot mostly postaxial. The digits of the hindfoot are weaker

than those of the forefoot.

The proximal row of the tarsus is formed by a robust astragalus and a large

flattened disc-like calcaneum.

The astragalus is a strong element of peculiar shape; it is thick, a little

longer than broad. In its anterior part the dorsal surface is convex postaxially

Fic. 4. Struthiocephalus sp. SAM 12226 x 4

Dorsal view of left pes as restored.

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

and hollowed out towards its preaxial edge. The convex surface covered with

a thick cartilage is for the reception of the tibia. Proximal of the tibial facet

lies an obliquely directed deep groove, which forms a deep proximal incisure

extending round the bone and continuing across the ventral surface as a deep

groove connecting with the groove on the postaxial surface of the bone. Proximal

to this groove the astragalus carries a strong facet for the fibula facing

proximo-postaxially.

The concave postaxial face of the astragalus faces an incisure on the cal-

caneum thus forming a passage for the vessels passing through the tarsus.

Distally the surface of the astragalus is convex and this facet for the central

tarsal carried a thick cartilage.

The calcaneum is a large flattened bone oval in outline. Its outer and

distal edges are rounded and fairly thin. Proximo-preaxially this disc-like

element is thickened and here carries an oval facet which receives a part of the

distal facet of the fibula. Distally of this thickening there is a shallow groove,

which, extending to the preaxial face, lies opposite to the concavity on the

astragalar opposing face.

Fic. 5. Tapinocephalid proximal row of tarsals x }

a—dorsal view b—ventral view

As—astragalus of SAM Kaor ff—facet for the fibula

Ca—calcaneum of SAM 12065 ft—facet for the tibia

THE DINOCEPHALIAN MANUS AND PES 21

The single centrale is a fairly large rounded bone lying between the

astragalus and the second and third distalia.

There are five distal tarsals of which the fourth is the largest.

The first three metatarsals are very short, but the fourth and especially

the fifth are larger and look more like normal metacarpals.

The first phalanges are small bones roughly triangular in shape with the

apices distally articulated with the second phalanges. The second phalanges

are somewhat larger and have expanded ends and a waist.

The terminal phalanges are broad, slightly curving bones carrying a flat

nail.

The digital formula is 2, 3, 3, 3, 3, with the toes of nearly equal length, but

the first is the shortest with the fourth only slightly longer than the other three.

In figure 5 the system of grooves in both astragalus and calcaneum are

well shown in ventral view. These grooves probably housed tendons associated

with the tarsal joint which apparently functioned in a most peculiar manner

and difficult to understand. I have attempted in figure 6 to show the peculiar

action of the ankle joint.

In a. the leg is shown at the completion of the swing forwards with the toes

just about to make contact with the ground. In this position it is evident that

the fibula has its bipartite distal facet in contact with the facet on the astragalus

Fic. 6. Tapinocephalid lower hind-limb showing ankle joint in three positions.

a—at completion of the forward swing c—at completion of the stride

b—standing and carrying body weight

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

and the facet on the calcaneum, whereas the distal tibial facet is not making

contact with the facet on the dorsal face of the astragalus and is in fact out of

articulation but held by tendons.

In b. the foot is in the standing position with the body weight transmitted

along the long axis of the tibia on to the dorsal facet of the astragalus. In this

position it is evident that the distal facets of the fibula are not making contact

with the facet on the astragalus nor with the facet on the calcaneum and are

in fact out of articulation, but held by tendons only.

In c. the foot is lifted off the ground at the completion of the backward

swing of the foot. The body weight is taken off the foot and the astragalus

moved away from its contact with the distal facet of the tibia. But the fibula

is in contact with the facet on the astragalus and the facet on the calcaneum.

Everything is now ready for the forward stride with the foot in position to be

swung forwards and twisted by rotation of the fibula on its long axis to assume

its contact with the ground position.

If the above representation of the action of the ankle joint is anywhere near

being correct, the presence of strong tendons is necessary and these could be

housed in the grooves present on both astragalus and calcaneum. I am not so

rash as to attempt any description of the mechanism involved.

Titanosuchia (Figs. 7-9)

In addition to the proximal tarsal elements of Jonkeria described by Broom

and the calcaneum erroneously labelled Tapinocephalus by Gregory, I have half

Fic. 7. Parascapanodon sp. K249 x +

Dorsal view with tibia in and fibula out of articulation.

THE DINOCEPHALIAN MANUS AND PES 23

a dozen good astragali and a couple of good calcanei of Parascapanodon.

Both the astragalus and calcaneum in Titanosuchians are very similar

in essential structure to the corresponding elements as described above in the

case of the Tapinocephalians. They can be distinguished from the latter in

that the astragalus is a large and heavier bone and the calcaneum is larger,

and in the details of the ventral and penetrating grooves.

In figure 7 the tibia is shown in articulation, in figure 8 the fibula is

articulated to both astragalus and calcaneum and in figure g the bones are

shown in ventral view with the fibula articulated.

Fic. 8. Parascapanodon sp. K249 X %

Dorsal view with fibula in articulation.

Anteosauria (Fig. 10)

In the type specimen of Micranteosaurus parvus there is an incomplete foot

preserved.

The calcaneum is a typical Therapsid flattened disc-like bone—thin in its

middle part and with a thin postaxial edge. Both the dorsal and ventral surfaces

are concave centrally with a thickened proximal and distal end where good

convex facets are developed for the fibula and centrale and second distal. Its

preaxial edge facing the astragalus is excavated to form a passage for the

penetrating vessels. The system of grooves so typical of the Tapinocephalia

and Titanosuchia are not developed.

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 9. Parasapanodon sp. K249 X 4

Ventral view with fibula in articulation.

The astragalus is a stout bone and is quite different to the specialized bone

seen in the Tapinocephalia and Titanosuchia. The major part of its dorsal

surface is formed by a well-developed rounded articular facet for the tibia.

Anterior to this facet lies a groove which flows into the incisure on the postaxial

face lying opposite to that on the calcaneum. Anterior to this groove an oval

knoblike thickening apparently articulates with the central and the second

distal. The ventral surface of the astragalus appears to be convex without

grooves.

There probably were five distals and also probably a central, but none are

preserved, in their position lie two displaced phalangeal elements.

The five metatarsals are elongated bones with a waist and expanded ends.

The first is quite small and the others progressively increase in size, with the

fifth a strong element.

The first digit is complete and the first phalanx short with expanded ends

and a waist; the terminal phalanx is sharp and narrow and carried a claw.

Only the proximal ends of the first phalanx of the second and third digits are

preserved. I presume that the digital formula was 2, 3?; 4?; 3?; 3?, as in the

forefoot.

The foot was apparently weak preaxially and strong postaxially.

The pes of Micranteosaurus is very similar to that of the Russian Titano-

phoneus, but there Orlov found no centrale and a digital formula 2, 3, 3, 3, 3.

THE DINOCEPHALIAN MANUS AND PES 25

\ '

‘

)

s---nc.

~ O = Wn

ee tl

Sel

“2

Fic. 10. Micranteosaurus parvus. Type. SAM 4323 X $

Dorsal view of left pes as restored.

DISCUSSION

The Dinocephalian feet are more primitive than those of the Sphenacodonts

in that the first carpal is not elongated, but otherwise they are definitely more

advanced.

The digital formula in Sphenacodonts is 2, 3, 4, 5, 3 —with a well-developed

first digit, a great increase in length from second to fourth digit and a

comparatively short fifth digit.

In the Dinocephalia the phalanges of the three middle toes are reduced in

number, the first digit is weaker, the three middle digits are shortened and

tend to become equal in length and the fifth digit becomes relatively stronger.

Micranteosaurus, with a digital formula 2, 3, 4, 3, 3, has the most primitive

feet; then comes Titanophoneus in which the fourth digit in the forefoot shows a

fusion of two phalanges to produce the formula 2, 3, 3, 3, 3, which also obtains

in the Titanosuchia and Tapinocephalia.

Proximally the tarsus is still fairly primitive in the Anteosaurs and becomes

more specialized in the Titanosuchians and Tapinocephalians.

The distal phalanges in Anteosaurs still carried a claw as one would expect

in carnivores, whereas in the Titanosuchians and Tapinocephalians, which were

herbivores, the nails are flattened.

The direction of development in the Dinocephalia seems to have been

away from a crawling habit with sprawling feet to a more walking habit with

the feet drawn in closer to the median line of the body.

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

SUMMARY

Descriptions and figures are given of the manus and pes of the Dinocephalia

of the Tapinocephalus zone in South Africa. Owing to the nature of the preserva-

tion of the dinocephalian material in this zone only eighteen of more than two

hundred specimens included bones of the feet, and this account is based on

these specimens. There are indications that the Dinocephalia show a

development from a crawling to a walking habit.

ACKNOWLEDGEMENT

The Trustees of the South African Museum are grateful to the Council

for Scientific and Industrial Research for a grant to publish this paper.

REFERENCES

Boonstra, L. D. 1954. The smallest Titanosuchid yet recovered from the Karoo. Ann. S. Afr.

Mus. 42: 149-156.

Broom, R. 1929. On the carnivorous mammal-like reptiles of the family Titanosuchidae.

Ann. Transv. Mus. 13: 9-36.

Byrne, F. 1937. A preliminary report on a new mammal-like reptile from the Permian of South

Africa. Trans. Kans. Acad. Sci. 40: 221-229.

Grecory, W. K. 1926. The skeleton of Moschops capensis Broom, a Dinocephalian from the

Permian of South Africa. Bull. Amer. Mus. nat. Hist. 56: 179-251.

Ortov, J. A. 1958. The carnivorous Deinocephalia (Titanosuchia) from the Upper Permian

deposits of the Middle Volga. Trud. paleont. Inst. Acad. Sci. 72: 1-114 (in Russian).

Romer, A. S. & Price, L. W. 1940. Review of the Pelycosauria. Spec. Pap. Geol. Soc. Amer. 28:

1-538.

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