c;

<.' .

' cr. <c

c*.- ..

cc

</<- <r c<
<?: a c<

c

r<7v"<ST<r <

S <^^ -
: f?'

<

.. . (

i^Sfl^^c-S

' "

>S^v> <0 SP<^ .c cc

C<SL CS::[!C !; ^XCC ( CC < <
* <sc^ j .K; Sara^c^^fe,

^^Jgs^iSiim* r

:

&1SE ^

""

'^^TcX'C'

<.'C<

tcv<crc

.:*C.

< ^

S'Xlcr

.-rc-.ee

'

fc <L .

|:

C ?

c c

ANNALS

OF THE

SOUTH AFRICAN MUSEUM

VOLUME XII

ANNALS

OP THE

SOUTH AFRICAN MUSEUM

VOLUME XII

DESCRIPTIONS OF THE PALAEONTOLOGICAL MATERIAL
COLLECTED BY THE SOUTH AFRICAN MUSEUM
AND THE GEOLOGICAL SURVEY OF SOUTH AFRICA

PRINTED FOR THE

TRUSTEES OF THE SOUTH AFRICAN MUSEUM

AND THE

GEOLOGICAL SURVEY OF THE UNION OF SOUTH AFRICA

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

1913-1924.

TRUSTEES OF THE SOUTH AFRICAN MUSEUM.

The Right Hon. JOHN XAVIEH MERRIMAN, P.O.

Sir THOMAS Mum, Kt., M.A., F.R.S., F.R.S.E., C.M.G., LL.D.

The Hon. JOHN WILLIAM JAOGER, M.L.A., F.R.Stat.S.

SCIENTIFIC STAFF OF THE SOUTH AFRICAN

MUSEUM.

Louis ALBERT PERINGUEY, D.Sc., F.Z.S., F.E.S., Director.

KEPPEL HARCOURT BARNARD, M.A., F.L.S., F.R.S.S.Afr., Assistant Director, in

Charge of Fish and Marine Invertebrate Collections.
STAR GARABEDIAN, B.A., Assistant in Charge of the Herbarium.
REGINALD FREDERICK LAWRENCE, B.A., Assistant in Charge of Aithropoda (Insects

and Crustacea excluded).

ALBERT JOHN HESSE, B.Sc., Ph.D., Assistant in Charge of Insects.
ARTHUR LEWIS HALL, M.A., Hon. Keeper of the Geological and Mineralogical

Collections.
SIDNEY HENRY HAUGHTON, B.A., D.Sc., F.G.S., F.R.S.S.Afr.. Hon. Keeper of the

Palaeontological Collections.

LIST OF CONTRIBUTORS.

R. BROOM. PAGE

On some Fishes from the Lower and Middle Karroo Beds . . 1

On a new South African Stegocephalian (Phrynosuchus irlmitsi) . . f>

On a nearly perfect Skull of a new species of the Gorgonopsia . . 8

Man contemporaneous with Extinct Animals in South Africa 13

B. BROOM and S. H. H. \UGHTON.

On the Skeleton of a new Pareiasaurian (I'/ir/ iii.tiK-lnix / r!/////i/ >/! g. et sp.

nov.) . .17

On a new species of Scymnognathus (*S'. tigrir.e i>n~) . . . .26
On two new species of Dicynodon . . 36

Some new species of Anninnilnntin (Reptilia) . .119

F. CHAPMAN.

Foraminifera and Ostracoda from the Upper Cretaceous of Need's Camp,

Buffalo River, Cape Province . .107

S. H. HAUGHTON.

On a Skull of Tapinocephalus atherstoni, Owen ..... 40

On a new species of Propappus ....... 43

Investigations in South African Fossil Reptiles and Amphibia.

1. On a new species of Trematosaurus (T. W >/i) 47

2. On a new Dinocephalian from the Gouph .... 52

3. On two new TherocephaUans from the Gouph . . . .55

4. On some new Anomodonts ....... 58

5. On the genus lilhifxiir/iti*, UK mm, with Notes on the described

species ..... 65

6. On a new type of Dinocephalian ...... 78

7. On some new Gorgonopsians ....... 82

8. On a Skull of the genus Kannemeyerla . . . . .91

9. A new Thecodont from the Stormberg Beds (HplirnnsucliHs acutus,

g. et sp. nov.) . . . . . . . .98

10. Descriptive Catalogue of the Anomodontia, with especial reference

to the examples in the South African Museum. Part 1 . . 119

11. Some new Carnivorous Therapsida, with Notes upon the Brain-

case in certain species . . . . . . .175

12. On some Gorgonopsian Skulls in the Collection of the South

African Museum ........ 499

The Fauna and Stratigraphy of the Stormberg Series .... 323

L. F. SPATH.

On Cretaceous Cephalopoda from Zululand ..... 217

vi

LIST OF NEW GENERA PROPOSED IN THIS

VOLUME.

Aelurognathus (gen. nov.)
Akidnognathus (gen. nov.)
Archaegryllodes (gen. nov.)
Caruichthys (gen. nov.)
Chelyrhynchus (gen. nov.)
Diaziceras (gen. nov.)
Galesuchus (gen. nov.)
Gorgonognathus (gen. nov.)
Lycorhinus (gen. nov.)
Macroscelesaurus (gen. nov.)
Melanorosaurus (gen. nov.)
Moschosaurus (gen. nov.)
Myosaurus (gen. nov.)
Pareiasuchus (gen. nov.)

Phrynosuchus (gen. nov.)
Prolystrosaurus (gen. nov.)
Pseudophacoceras (gen. nov.)
Pseudoschloenbachia (gen. nov.
Scylacops (gen. nov.)
Spheniscoceras (gen. nov.)

Sphenosuchus (gen. nov.)
Striatotegmen (gen. nov.)

Struthiocephalus (gen. nov.)
Subschloenbachia (gen. nov.)
Sycosaurns (gen. nov.)
Trochosaurus (gen. nov.)
Whaitsia (gen. nov.)

HAUGHTON

SPATH

HAUGHTON

PAGE

HAUOHTON 503
192
336
4

156
242
82
84
343
175
428
78
164
BROOM and 17

HAUGHTON
BROOM 6

HAUGHTON 167
281
236

BROOM 8

CRICK MSS. 242

SPATH

Pseudosuchia (Reptilia) HAUGHTON 98

Mesoblattinidac (Orthoptcra- 331

Insecta)

Dinocephalia (Reptilia) ,, 52

Dipoloceratidae (Cephalopoda) SPATH 284

Gorgonopsia (Reptilia) HAUGHTON 509

Tlicrocephalia (Reptilia) ,, 55

Therocephalia (Reptilia) 184

Gorgonopsia (Reptilia)
Therocephalia (Reptilia.) ,,

Gryllidae (Orthoptera-Iiisecta) ,,

Platysomatidae (Pisces) BROOM

Anomodontia (Reptilia)
Prionotropidae (Cephalopoda)
Gorgonopsia (Reptilia)
Gorgonopsia (Reptilia)
Cynodontia (Reptilia)
Therocephalia (Reptilia)
Plateosauridae (Reptilia)
Dinocephalia (Reptilia)
Anomodontia (Reptilia)
C< it ylosauria ( Reptilia.)

Strgorcphalin (Ani])hibia)
Anomodontia ( !!< \>\ ilia)
I >i|ioloceratidae (Cephalopoda) SPATH
I 'rionotropidae (Cephalopoda)
Gorgonopsia ( Reptilia)
Prionotropidae (Cephalopoda)

DATE OF ISSUE OF THE PARTS.

Part 1. 30th May 1913.

Part 2. 14th January 1915.

Part 3. 2nd September 1915.

Part 4. 22nd December 1916.

Part 5. 12th December 1917.

Part 6. 6th November 1918.

Part 7. 30th April 1921.

Part 8. April 1924.
via

37224

ERRATA.

(Part 8.)

Page 348, line 33, for " from " read " form."

,, 355, ,, 6, ,, " Omitosuchus " rat'! " Ornithosuchns."

355, ,, 21, ,, " Spenosuchus " ,, " Sphenosuchus."

357, 34, " doracoid " " coracoid."

357, ,, 38, ,, " Spenosuchus " " Sphenosuchus."

358, 10, "Broem" "Broom."

395, 27, "public" "pubic/'

398, 19, "to" "no."

409, 5, "width" "with."

416, in description of fig. 36, for " pelvis " read " pubis."

,, 417, line 12, delete " A figure is given herewith."

439, line 17, for " builders " read " boulders."

439, 24, ,, second " of " read " or."

444, last line, for " place " read " plane."

,, 457, last word, for " microline " read " microcline."

464, line 5, for " it," read " if."

464, 25, " tick " read " thick."

468, 24, " pruvial " read " pluvial."

475, ,, 13, " transportion " n-oil " transportation."

546, 38, "are" "is."

There are a number of typographical errors scattered through the two papers,
which have largely entered in since the reading of the final proof -sheets ; but these
will be obvious to anyone acquainted with the English tongue, and have not
therefore been listed above.

The title to Dr. Haughton's paper on p. 499 is incorrect and should be replaced
by the following :
18. Investigations in South African Fossil Reptiles and Amphibia (Part 12). By

S. H. HATJGHTON, B.A., D.Sc., F.G.S., Honorary Curator of the Palaeonto-

logical Collections.

12. On Some Gorgonopsian Skulls in the Collection of the South African Mii.iemn.
(With 8 Text-figures.)

LIST OF PLATES.

f

XIV.

XV.

PLATE

II.
III.

TV.

V.

VI.

VII.

VIII.

IX.

X.

*'{

XII.
XTTT. /

Caruichthys ornatus, Broom.
Atherstonia cairncrossi, Broom.
Palaooniscus capensis, Broom.
Elonichthys whaitsi, Broom.
Pareiasuchus peringueyi, Broom and Haughton.
Pareiasuchus peringueyi, Broom and Haughtoii.
Propappus parvus, Haughton.
Pareiasuchus peringueyi, Broom and Haughton.
Scymnognathus tigriceps, Broom and Haughton.
Scylacops capensis, Broom.
Dicynodon alticeps, Broom and Haughton.
Dicynodon testudirostris, Broom and Haughton.
Trematosaurus sobeyi, Haughton.
Trematosaurus sobeyi, Haughton.
Struthiocephalus whaitsi, Haughton.
Dicynodon mustoi, Haughton.
Dicynodon breviceps, Haughton.
Lystrosaurus oviceps, Haugliinn.
Rhinesuchus senekalensis (v. Hoopon).
Bhinesuchus whaitsi, Broom.
Gorgonognathus longifrons, Haughton.
Scymnognathus serratidens, Haugliton.
Haplophragmium neocomianum, Chapman.
Spiroplecta deflata, Chapman.
Spiroplecta anceps (Reuss).

Spiroplecta anceps (Reuss) var. infracta, Chapman.
Nodosaria zippei, Reuss.
Nodosaria sulcata, Nilsson.
Cristellaria parallela, Reuss.
Cristellaria intermedia, Reuss.
Cristellaria secans, Reuss.
Trimcatulina schloenbachi (Reuss).
Discorbina pileolus (d'Orbigny).
Anomalina ammonoides (Reuss).
Pulvinulina karsteni (Reuss).
Trimcatulina ungeriana (d'Orbigny).
Mtiirdia subdeltoidea (Minister) var.
Cyllu-iv postcultrata, Chapman.
Bairdia africana, Chapman.
ix

List

PLATE

XVI.

XVII.

XVIII.

XIX.

XX.

XXI.

XXII.

XXIII.

XXIV.

XXV,

xxvi.

Dicynodon grandis, Haughton.

Eocyclops longus, Broom.

Prolystrosaurus natalensis, Haughton.

Diaziceras tissotiaeforme, Spath.

Parapuzosia sp. nov. ? ind.
C Parapuzosia sp. nov. ? ind.
I Pseudoschloenbachia umbulazi (Baily).
j Pseudoschloenbachia umbulazi (Baily) var. acuta Spath.
"- Mortoniceras soutoni (Baily).

{Mortoniceras woodsi, Spath.
Placenticeras subkaffrarium. Spath.
C Parapachydiscus sp. aff. colligatus (Binkhorst).
< Nostoceras ? natalense, Spath.
*- Nostoceras ? subangulatum, Spath.
r Peroniceras cf. czornigi (Redtenbachcr).
J Mortoniceras stangeri (Baily).

Mortoniceras vanuxemi (Morton).
" Diplomoceras ? indicum (Forbes).

Parapachydiscus cf. wittekindi (Schluter).

Bostrychoceras ? sp. ind.

Parapuzosia sp. ind.

Baculites cf. aspero-anceps, Lasswitz.

Baculites cf. brevicosta, Schluter.

Baculites capensis, Woods.

Kossmaticeras (Madrasites) bhavani (Stoliczka).

Subschloenbachia bispinosa, Spath.

Subschloenbachia prerostrata, Spath.
r Pseudophacoceras nianuanense, Spath.
J Dipoloceras cristatum (Deluc).
] Dipoloceras quadratum, Spath.

Subschloenbachia cf. trinodosa (Bose).

Cheloniceras gottschei (Kilian).

Cheloniceras delagoense (Krenkel).

C'heloniceras hambrovii (Forbes).

Aconeceras nisoides (Sarasin).

Dipoloceras sp. nov. ?

Dipoloceras cristatum (Deluc).

Anisoceras sp. ind.

INDKX OF GENERA.

^4 canthocera*
Aconeceras
Adolphia .
Aelurognathus .
Aetonyx .
Akidnognathus .
Alopccognathus .
Anisoccras
Anomalina
Archaegryllodcs
Arctognathus
Aristosaurus
Atherstonia

Baculitcs .
Bairdia
Barroisiceras
Bostrychoceras .
Bubalus

Caruichthys

Cheloniceras

Chelyrhynchus .

Cobus

Connochaetes

Cristellaria

Cymatoceras

Cynosuchus

Cythere

Cytherina .

Cyzicus

I!

C

PAGE

U

PAGE

. 312

Diaziceras .

. 242

. 311

Dictyopyye

. 340

. 312

Dicynodon

. 36, 58, 119

. 503

Diictodon .

. 160

. -4<M

Diplomoceras

. 256

. 192

Dipoloceras

. 277

180, 210

Discorbina

. 112

256, 288

I ><>/ir ill* ic< nix

. 312

108, 114

Dromicosaurus .

. 405

. 336

. 507

. 379

E

3

Elonichthys

2

Emydops .

. 124

. 257

Eocyclops .

. 160

. 114

Equus

14

237, 244

Erythrochampsa

. 367

252, 308

Esther la .

. 326

14

Eucnemesaurus .

. 428

Eiilophoceras

. 242

Euskelesaurus .

. 423

Eutrephoceras .

. 262

4

. 312

. 156

G

15

14

Galesuchus

82, 499

. 110

Gauthiericeras .

. 237

264, 290, 301

Geranosaurus

. 436

. 197

Gigantoscelus

. 427

114, 116

Gorgonognathus

84, 209, 505

. 115

Gryponyx.

. 417

. 326

Gyposaurus

. 379

XI

Xll

Index of Genera.

H

PAGE

PAGE

Handle fi .

. 256, 288

P achy discus

. 226

Haplophragmium

. 109

Pachygenelus

. 340

Hauericeras

. 237

Palaeoniscus

1

Helichthys

. 340

Paralenticeras .

. 244

Heteroceras

. 308

Parapachydiscus

. 226

Hippopotamus .

14

Parapuzosia

224

Holcodiscus

. 299

Pareiasuchus

17

Hortalotarsu-s

370, 379

Pedeticosaurus .

. 366

Peroniceras

. 295

Phacochoerus

14

K

Phrynosuchus .

(i

Phthartus .

. 331

Kannemeyeria .

1)1, 172

Phylloceras

. 273

Kossmaticeras .

. 299

Placenticeras

. 247, 300

Plateosaurus

. 406

Prohauericeras .

. 238

L ' Prolystrosaurus .

. 167

Propappus

43

Lenticeras

. 2H

Pseudophacoceras

. 281

Lepidurus .

328

Pseudoschloenbachia

. 236

Lycorhinus

. 343

Pulvinulina

108, 114

Lystrosaurus

. 01

Puzosia

. 237, 274

M

R

Macroscelesaurus

. 175

Madrasites

. 299

Rhinesuchus

65

Massospondylus .

. 370, 383

Rosalina .

113, 114

Melanorosaui'us .

. 428

Rotalia

. 114

Miliolina .

. 107

Rotalina .

. 113

Mortoniceras

232, 277, 308

Moschosaurus

78

Muiiiericeras

. 237

Myosaurus

. 164

S

Schloenbaclda

240, 281, 285, 295

N

Scymnognathus .

. 26, 88, 205, 503

Scylacops .

. 8, 210, 501

Nautilus .

262, 264, 290, 301

Seinionotus

. 338

Nodosaria .

. 110

Spheniscoceras .

. 242

Nostoceras

. 248

Sphenosuchus .

98, 344

Notoohampsa .

. 358

Spiroplecta

. 109

Stoliczkaia

. 286

Striatotegrnen .

. 331

Struthiocephalus

52

Subschloenbachia

. 284

Oppelia

. 311

Sycosaurus

. 509

Index of Genera,.

xni

U

Tapini (cephalus

Taurotragus

Textularia

Thecodontosaurus

Titanosuchus

Torneutoceras .

Trcmatosaurus .

Tritheledon

Trochosaurus

Truncatulina

Turrilites .

PAGE

PAGE

40

Uhligella . ... 275

15

108,

109

.

370

V

.

57

.

288

Valfulinu . .112

.

47

.

341

.

55

W

.

113

28<J

Whaitsia . 184

ANNALS

OF THE

SOUTH AFRICAN MUSEUM.

(VOL. XII.)

1. On some Fishes from the Lower and Middle Karroo Beds. By

E. BROOM, M.D., D.Sc.

WITH the exception of a couple of species of Atherstonia, described
by Smith Woodward, almost nothing has been known of the fishes
of the Lower and Middle Karroo. The fishes of the Burghersdorp
beds and of the Cave sandstone have been dealt with by Smith
Woodward and by myself, and a good many forms are now known.
The fishes of the older rocks, though less varied than those of the
upper, are nevertheless interesting. The types of the following
specimens are in the collection of the South African Museum.

PAL^ONISCIDAE.

PAL^ONISCUS CAPENSIS, sp. nov.

This new species is founded on three specimens from the Hantam
Mountains, 12 miles west of Calvinia, and from beds which are
probably Upper Dwyka, and thus of Lower Permian age. The
specimens unfortunately are all of the posterior part of the fishes-
one shows almost all except the head, but the other two only the
tail halves. The preservation of the specimens is marvellously
perfect.

The total length of the specimen was probably 300 mm., and the
greatest depth of the body about 75 mm.

1

2 Annals of the South African Museum.

The dorsal fin is placed more anteriorly than in Palaoniscus
macropomns, and the distance between the pectoral and pelvic fins is
also less. The front of the dorsal fin is opposite the point midway
between the front of the pectoral and pelvic fins. The dorsal fin has
38 rays, of which the 9th is the longest. Distinct fulcra are
present. All the rays are jointed. The anterior short rays and
the first three of the long ones are undivided, but the later rays
are all bifurcated at their tips.

The structure of the pectoral arid pelvic fins cannot be very clearly
made out. The rays are jointed and apparently bifurcated.

The anal fin is of moderate size. It consists of 52 rays, all jointed,
and the posterior ones bifurcated distally. There are well-developed
fulcra.

The caudal fin is large and deeply bifurcated. The lower lobe is
smaller than the upper.

The scales on the anterior part of the body are ornamented by a
series of obtuse ridges. On the upper part of the scale the ridges
run backwards a short distance, then curve downwards and run
parallel to the posterior border of the scale. From the anterior
border of the scale other ridges run backwards, stopping at the
point where the descending ridges would meet them. On the
posterior scales there are only a few transverse blunt ridges on
the anterior part of the scale. The scales on the ventral surface
between the pectoral and pelvic fins are considerably enlarged.
Between the pelvic and anal fins are a number of enlarged scales.
Two moderately large scales are placed on either side just in front
of the anal fin, and in front of these two is a much larger median
scale. Enlarged median scales are also placed over the upper lobe
of the tail largest at its base.

ELONICHTHYS WHAITSI, sp. nov.

This very fine fish was discovered by the Rev. J. H. Whaits on
the farm Droogvoets, Praserburg District. The specimen is nearly
complete, only a portion of the upper border being lost. The head
is crushed and not well preserved.

The total length of the type is 280 mm., and the greatest depth of
the body is probably 80 mm.

The fins are large and powerful. The dorsal is situated about the
middle of the body, and the anal begins a little in front of the
posterior part of the dorsal.

The most of the front of the dorsal fin is missing, but there are

On some Fishes from the Lower and Middle Karroo Beds. 3

30 rays behind the 1st long ray. There are well-developed fulcra.
All the rays are jointed, and all except perhaps the first two long
ones are branched.

The pelvic fin has 27 rays, all jointed, and the posterior rays
branched.

The anal fin is very large. It has 48 rays, of which the first
nine are short. All the rays are jointed, and the short anterior rays
and the first six of the long rays are unbranched. All the posterior
rays are bifurcated.

The caudal fin is very powerful and deeply bifurcated. The lower
lobe is quite as large as the upper. All the rays are branched, but
most only bifurcated.

The scales of the anterior part of the body are rhombic, with
feeble transverse irregular ridges and denticulate posterior margins.
The posterior scales are unornamented.

Some very large scales are situated in front of the anal fin, and
large ridge scales are also above the root of the tail.

ATHERSTONIA CAIRNCBOSSI, sp. nov.

This new species of Atherstonia is founded on the greater part of
a fish found by Mr. J. L. Cairncross at Coleskop in 1906, and
presented by him to the South African Museum. It was regarded
at the time as a small specimen of Atherstonia scutata, S.-W.,
but on comparing it with undoubted specimens of the type species
in the Albany Museum it is seen to be a different species.

It differs from A. scutata in being relatively shorter and deeper.
In A. scutata the length from the clavicle to the base of the tail is
about 200 mm., and the greatest depth of the body about 75 mm. ;
in this new species the length from the clavicle to the base of
the tail is 117 mm., and the greatest depth 57 mm. In other
words, the depth of the body in the new species is half the length
without the head and tail; in A. scutata the depth is only a little
more than ^ the length.

The dorsal fin has a long base and is powerful. It begins
nearer to the clavicle than to the base of the tail, and not as in
A. scutata nearer to the tail. The enlarged ridge scales do not
strengthen the front of the fin so markedly as in A. scutata, and
the fulcra are very rudimentary. There are about 43 rays, which
are jointed but not bifurcated.

The pelvic fin is in front of the dorsal, ending opposite the
point where the dorsal begins. It has about 25 jointed rays.

4 Annals of the South African Museum.

The anal fin is about the same size as the dorsal, and begins
opposite the point of union of the latter and middle thirds of the
dorsal. It has about 36 jointed but unbranched rays. The fulcra
are rudimentary.

The caudal fin is very imperfectly preserved.

There is, as in A. scutata, a row of ridge scales both in front
of and behind the dorsal fin. Those between the dorsal and tail
are very large and only feebly ornamented. Those in front of the
dorsal are smaller, thinner, and more distinctly striated. The body
scales are obliquely striated and posteriorily feebly but distinctly
denticulated.

FAMILY PLATYSOMATIDAE.

CAKUICHTHYS ORNATUS, g. et sp. n.

The type of this species is a single specimen found at Doom
Eiver, in the Cradock district, and most probably from beds of the
Lystrosaurus zone.

A small portion of the front of the head is missing, and the whole
of the tail from behind the dorsal fin.

The trunk is deeper than in Eurynotus, but not so deep as in
Platysotnus. The head is large and deep, somewhat resembling
the head of Platysomus, but with relatively much more power-
ful jaws and with the branchiostegal rays large and extending
much below the mandibles. The teeth are small and styliform.
The dorsal margin curves up and back above the head and then
passes nearly directly backwards to the dorsal fin. Along this
dorsal margin are a series of enlarged much ornamented scales.
At the commencement of the dorsal fin the line of the body
margin abruptly passes downwards and backwards as in
Platysomus.

The pectoral fin is large and possibly not unlike that in Eury-
notus. It is too imperfectly preserved to show its structure. Only
the anterior part of the dorsal fin is preserved. There are a
series of 9 or 10 short but gradually lengthening rays in front of
what is probably the longest ray of the fin. These short rays are
jointed but unbranched. In front of the first long ray are feeble
fulcra. The later rays of the fin are long and much feebler than
the anterior. They are jointed and unbranched. The pelvic fin
is much larger than that in Eurynotus. It has about 40 rays all
jointed and unbranched. The posterior part of the fin is distinctly

On some Fishes from the Lower and Middle Karroo Beds. 5

in front of the beginning of the dorsal. No parts of the anal or
caudal fins are preserved.

The scales are very narrow and deep and much imbricated.
Each is ornamented by numerous irregular prominent transverse
ridges. The rows of scales below the dorsal fin are smaller, and
the scales immediately above the pelvic fin are small.

The length from the snout to the front of the dorsal is 140 mm.
The whole length of the fish is probably about 210 mm. The
greatest depth of the body is 100 mm.

(6)

2. On a New South African StegocepJialian (Phrynosuchus whaitsi).

-By E. BROOM, M.D., D.Sc.

THE little Stegocephalian about to be described was found by the
Rev. J. H. Whaits on the farm Droogvoets, in the Fraserburg
district. The exact horizon of the locality is not certain, as the
other fossil remains are either dissimilar from those known in the
South or are too fragmentary to be of service. Most probably
the zone corresponds to the upper part of Endothiodont zone of the
southern Karroo.

The specimen consists of much of the skull in a very weathered
condition, remains of most of the vertebrae and short ribs as far as
the pelvic region, the most of the right front and hind limbs, and
considerable indications of the dermal covering. Allowing for a
moderate tail, the whole animal probably measured about 350 mm.

The skull shows the impressions of most of the bones of the
cranium, and is so broken across as to reveal the structure of at
least the front half of the palate. The whole head is broad and
very flat. The nostrils are wide apart, and the orbits, which are
also far apart, are entirely in the anterior half of the skull.

The nasal is probably very large, the prefrontal and lachrymal
very small. Each frontal is broad, and shut out from the orbital
margin by the meeting of the postfrontal with the prefrontal.

The postfrontal is large and forms the greater part of the upper
orbital margin. The postorbital is much smaller.

The parietal is a large bone, with a small pineal foramen between
the pair of bones. The squarnosal is rather smaller than the parietal.
Outside of the squamosal is a large quadrato-jugal.

Behind the parietal is a large dermo-occipital.

The parasphenoid (or vomer) is of large size and extends far
forward as a broad, flat plate. The prevomers are large and the
internal nares far apart. The palatines are also large.

The teeth are not well shown but are manifestly labyrinthodont.

The length of the skull is about 68 mm., and the width almost
exactly the same.

On a Neiv South African Stegocephalian (Phrynosuchus whaitsi). 7

Behind the occiput are some crushed bones which may be parts
of the clavicles and interclavicles, and by the sides of the neck are
some small ossicles which may be the remains of gill arches.

The body is long and salamander-like. The vertebrae are very
imperfectly ossified and represented by paired ossifications, probably
representing pleurocentra. The ribs are short as in the Branchio-
sauridae and almost straight.

FIG. 1. Skull of Phrynosuchus whaitsi. Very slightly reduced.

The shoulder girdle and clavicular apparatus are not well pre-
served, but the humerus, radius, and ulna of the right side are seen.
The humerus is a short bone with a moderately wide distal end.
The length is 16 mm. and the distal end is 9 mm. wide. The radius
is a slender rod 10 mm. in length and the ulna is rather stouter.

Of the hind limb the only parts preserved are a portion of the
tibia, five metatarsals, and two phalanges. There are certainly five
digits, and the tarsus is unossified.

There is a complete exoskeleton of thin, ossified scales.

The type, which has been named after the Eev. J. H. Whaits, has
been placed in the South African Museum.

(8)

3. On a Nearly Perfect Skull of a New Species of the Gorgonopsia.

-By E. BROOM, M.D., D.Sc.

THE skull here described was found by Mr. S. H. Haughton on
the farm Dunedin, Beaufort West, about 3 miles WSW. of the
homestead. The exact horizon of the- beds is still uncertain, but
they must be at least 2,000 feet above the horizon of Beaufort West.
Whether they correspond to the Cistecephalus beds or may be as
high as the Lystrosaurus zone is at present in doubt. We know
that Cistecephalus occurs at Kuilspoort probably 1,500 feet above the
horizon of Beaufort West, and about 500 feet higher up there are
obscure indications which incline me to believe we are into the
Lystrosaurus zone, but as Dr. Eogers, who has gone over the ground,
is rather opposed to this idea, we must at present leave the matter
in doubt. In any case it is quite certain that the specimen comes
from a very much higher zone than does Gorgonops tor mis, Owen.
Owen's type comes from Fort Beaufort, and it has also been obtained
at Beaufort West, so that there is no doubt it belongs to the lower
part of the Endothiodon zone, not more than a few hundred feet
above the Pareiasaurus zone. Another Gorgonopsid occurs in the
same zone, viz. Scymnognathus whaitsi recently described by me.
The occurrence of a new Gorgonopsid at a very much higher horizon
is thus particularly interesting, and the fact that the skull is the
most perfect known and reveals most of the details of the structure
makes the find a most important one.

The skull measures 190 mm. in length, and the greatest width
across the temporal region is about 128 mm. The snout is rounded
and broader than deep. The nostrils are directed forwards and are
much flattened as in Gorgonops torvus. The frontal region is broad
and the orbits look more outwards than upwards. The parietal
region is as broad as the frontal and the temporal fossae of fair size.
The occiput is nearly vertical, sloping only very slightly backwards
from the parietal.

The premaxillary is similar to the Therocephalian type. It bears
5 rounded and pointed incisors. I do not find any evidence of

On a Nearly Perfect Skull of a New Species of the Gorgonopsia. 9

serration, though the evidence is not quite conclusive. The 5
incisors measure 23 mm.

The septomaxilla is also as in Therocephalians. The posterior
process is long and separates the anterior part of the nasal from the
maxilla.

The maxilla is large and deep. In front it overlaps the premaxilla.
There is a single fair- sized canine situated 8 mm. behind the last
incisor, and measuring in section 8*5 mm. by 6 mm. There are a
few small rounded molars, but the exact number cannot be made out
with certainty. Three are preserved. Possibly the original number
was 5. These molars are only about ^ of the size of the incisors.

The nasal is large and fairly wide. It is only slightly wider at
the front and behind than it is in the middle. The internasal
pi'ocess is evidently feeble.

The prefrontal is large and forms the upper and anterior quarter of
the orbital margin. It unites behind with the postfrontal, shutting
out the frontal from the orbit. In this it agrees with the conditions
in the higher Cynodonts and differs from all known Therocephalians,
and also from the lower Cynodonts and from the Anomodonts.

The lachrymal is of moderate size, and the canal seems to be
inside the orbit.

The jugal is fairly large but much hidden by the overlapping
squamosal and postorbital.

The frontal is large, the two forming most of the interorbital
region. On its outer side it articulates with the prefrontal and post-
frontal. Behind it meets the parietal and preparietal.

In the middle line between the two frontals and the two parietals
is a small median bone, manifestly the bone which in the Anomo-
donts has been called the preparietal. This character is perhaps the
most interesting in the skull. For long the little median bone which
lies mainly in front of the parietal foramen has been known in the
Anomodonts. It has been variously named parietal, interparietal
and preparietal. Quite certainly it is neither the parietal nor the
interparietal, and the name preparietal proposed by Seeley and
adopted by myself and others seems the best name for it. It is
known in all Anomodonts except Cistccephaius, but hitherto it has
not been recognised in any other group. The discovery of a pre-
parietal in another group is thus of great interest. It pretty
certainly does not occur in most Therocephalians. That it should
occur in the Gorgonopsidae, the group which in some other respects
most approaches the Anomodonts adds further confirmation to the
view I expressed many years ago that the Anomodonts are descended

10 Annals of the Sontli African Museum.

from a Therocephalian ancestor. In some Anomodonts the foramen
is in the middle of the bone ; in others the bone lies in front of the
foramen though forming its anterior wall. In no known Anomodont
is the foramen as here between the parietals and the preparietal
away from the foramen.

The postfrontal is unusually large. In Anomodonts it is always
small, and in Therocephalians it was also small where it could be
made out distinctly. Here it forms much of the orbital margin and
quite a large part of the upper cranial wall. For the most part it
lies between the frontal and the postorbital. Posteriorly it meets the
parietal.

The parietal is about as large as the frontal. In front it meets the
preparietal, the frontal, the postfrontal; while laterally it is sup-
ported by the postorbital, which completely shuts it out from the
temporal fossa. Posteriorly it meets the large median interparietal,
and posterior-laterally the squamosal.

The postorbital is large. It forms most of the postorbital arch,
and its posterior extension forms the whole of the upper margin of
the temporal fossa.

The squamosal is very large and forms practically the whole of the
back of the temporal fossa. Its inner side articulates with the
interparietal and the exoccipital, and forms about 1 of the occiput.
The ends of the parietal and the postorbital are clasped by the upper
and inner part of the bone. Immediately outside the lateral
occipital foramen the squamosal has a vertical posterior ridge which
apparently delimits the occiput proper from the groove which is
probably the auditory groove. The lower and outer part of the
squamosal passes down and almost entirely conceals the quadrate.
The outer part of the bone curves outwards and then forwards,
forming most of the zygomatic arch. Much of the inner side of the
arch is, however, formed by the jugal.

The interparietal is a large median bone a little broader than deep
It is bounded above by the parietals, laterally by the squarnosals and
inferioriy by the exoccipitals.

The exoccipitals appear to form the whole of the occipital region
above and to the sides of the foramen magnum. There is a well-
developed lateral process which apparently meets the quadrate.

The quadrate lies almost entirely in front of the descending part
of the squamosal, and is thus scarcely seen when viewed from
behind. Towards the quadrate there runs out a long process of the
pterygoid, but whether it reaches the quadrate is not quite clear.
The quadrate certainly has a short process which runs forward and

On a Nearly Perfect Skull of a Neu- Species of the Gorgonopsia. 11

inwards towards the pterygoid, but between the two there seems to
be another element. If this is so it will probably prove to be the
alisphenoid. As the specimen is slightly crushed one cannot be
quite positive on this point.

There is an elongated stapes not unlike that of Cynoqnathus, which
stretches from the quadrate to the foramen ovale, which lies as in
the Anomodonts in the process formed by the basioccipital, basi-
sphenoid, and doubtless the pro-otic and opisthotic.

The basioccipital is a small bone which appears to form most of
the single occipital condyle. The foramen for the IXth, Xth, Xlth,
and XTIth nerves is as in the Anomodonts far back by the side of the
condyle.

The basisphenoid meets the basioccipital very much as in the
Anomodonts, but in front the bone differs in having a strong deep
median keel as in Bauria.

The pterygoid is rather peculiar. It has a large posterior process
towards the quadrate and also a broad support along the basi-
sphenoid. It has, as in most Therapsida, a powerful lateral process
by the side of the mandible, but the anterior development is smaller
than in any other of the Therapsida except the Anomodonts. I fail
to find any teeth on the pterygoid.

The transpalatine is well developed but also of an unusual
character and quite unlike that of the Therocephalia. It forms
quite a large part of the bony palate and the anterior part of the
lateral descending process.

The palatine is a large bone which takes the place usually occupied
by the anterior part of the pterygoid. It also forms most of the
bony palate. On the inner part are a few fairly large teeth.

The vomer is not displayed, but probably is as in Gorgonops and
Scymnognatlius a true median vomer.

The greater part of each mandible is preserved. The dentary is
powerful but the coronoid process probably short. The splenial
is also well developed but does not extend far back. The angular
is apparently pretty similar to that of Scymuognathus. It passes well
forwards between the dentary and splenial. The exact nature of
the articular region cannot be made out with certainty. Besides
an articular it seems there may be a prearticular.

The skull is particularly interesting as showing that the Gorgono-
psidae differ in many important characters from the Therocephalian
type seen in Scylacosaurus, and also by showing some striking resem-
blances to the skulls of Anomodonts and Cynodonts. In the structure
of both the upper cranial region and the palate there is a fore-

]2 Annals of the South African Museum.

shadowing of the peculiar Anomodont type. In the arrangement of
the frontals, postfrontals, postorbitals, preparietal, and parietals we
have almost the Anomodont condition. The palate of course differs
markedly. Still the pterygoid with its very short anterior part meet-
ing the palatine instead of the prevomer and the peculiar condition of
the transpalatine are just what we should expect in the Anomodont
ancestor. The relations of the stapes, basioccipital, and basi-
sphenoid are almost exactly as in the Anomodonts.

Though the relationships to the Cynodonts are not quite so evident,
still this type comes nearer to the Cynodont than does Scylacosaurus.
If the palatine bones curved over and made a secondary palate and
the maxillae also joined over, the whole palate would be not unlike
that of the Cynodont. The occiput would resemble that of the
Cynodont if the opisthotic were shown on the posterior surface, but
this is not a very important character when we consider how variable
it is in mammals.

It seems necessary to revive Seeley's Gorgonopsia as a distinct
sub-order of the Therapsida for reptiles of the Gorgonops type.

The relationships of the various sub-orders of the Therapsida may
be expressed as follows :

For the new skull I suggest the name Scylacops capensis, g. et
sp. nov.

(13)

4. Man contemporaneous with Extinct Animals in South Africa.

By E. BROOM, M.D., D.Sc.

FOR a good many years there has been some evidence that man
was contemporaneous with extinct animals in South Africa, Human
implements have been found among the gravels at the Vaal River
diamond diggings, and from the gravels have also been found teeth
of Equus capensis and a species of Mastodon. But it has been quite
impossible to say that man was contemporaneous from the uncer-
tainty of the gravel deposits being of the same age.

At Bloembosch, near Darling, Cape Province, the sand-hills on
shifting have revealed remains of Bubalus baini and Equus capensis
in association with human implements. But though the evidence
of contemporaneity was quite sufficient for me, it was not as
convincing as could be desired.

At Haagenstad saltpan, about 30 miles north of Bloemfontein,
a discovery has just been made which removes all possibility of
doubt of a human race having inhabited South Africa at a time
when there lived many large mammals now quite extinct.

Near the saltpan there is a hot spring which has issued from a
large sand-hill. Baths have been erected to utilise the hot water,
and recently excavations have been made into the hill to get nearer
to the source of the spring. In cutting into the hill a thick deposit
of beautiful peat was met with. The peat has evidently been formed
by the roots of bushes and trees, and as the deposit is 8 or 10 feet in
thickness, it must represent the growth of many years. Below the
purer peat are another 8 or 10 feet of peaty sand, and underneath
this a layer of broken bones, burnt wood, and human implements.

The Bloemfontein Museum authorities heard of the find, and Mr.
Leviseur authorised Professor Potts and myself to visit the spot.
That the more important finds have been preserved we owe to Mrs.
Martha Johanna Venter, the very intelligent wife of the farmer who
lives at the spot. She has made a large collection of the bones and
other remains, and though unfortunately she has allowed many of
the human implements to pass into the hands of unscientific curiosity

14 Annals of the South African Museum.

hunters, her collection reveals all that is required for at least a
preliminary note.

The animals whose remains have been found form a most
interesting and varied fauna.

Equus capensis is represented only by a single tooth so far as I
observed in looking over the collection.

Equus sp. There are teeth of a small Equus probably one of
the small Zebras.

Hippopotamus amphibius. The remains of Hippopotamus are
abundant. Very fine tusks, teeth, and jaws are represented in the
collection besides limb bones, etc. I do not think the Hippopotamus
differs from the present-day species, and some of the specimens
must have been of great size.

* ...

FIG. 2. Connochaetes antiquits. Broom, x j.

Phacochoerus aetliiopicus. There are many fragmentary jaws,
tusks, and other bones of the Wart Hog.

Bubalus baini is represented by numerous specimens of horn
cores, skulls, jaws, teeth, limb bones, and vertebrae. Most of the
bones have been broken by human agency.

Connochaetes antiquus, n. sp. The animal of which the remains
seem most abundant is a new species of Wildebeest. In the col-
lection are many horn cores. In type the horn is intermediate
between that of the two surviving South African species. Like that
of the white-tailed Wildebeest, there is a very prominent flattened
backward process at the base of the horn. From the base the horn
curves much more outwards in this new species, and then less
abruptly upwards and backwards. The curving will be better
understood from the illustration.

Man contemporaneous with Extinct Animals in South Africa. 15

The greatest length of the core from the base to the outer curve
in a straight line is 343 mm., and from the posterior knob at the
base to the tip following the posterior curvature is 465 mm.

C. antiqnus is so nearly intermediate between the two South
African living forms that one wonders if it may perhaps be the
ancestor of both.

Taurotragus oryx. There are a number of bones of the Eland,
including the frontal region with horn cores.

Cobus venterae, sp. nov. There are many horn cores of a species
of Cobus. It is somewhat intermediate between the Waterbuck
(C. ellipsipyrmnus) and the Lechwe (C. leche), but much nearer to

FIG. 3. Colitis rcnterae. Broom, x |.

the latter. From the existing Lechwe it differs in the curve of the
horn and in the mode of implantation on the frontal bone.

Besides these readily recognisable forms there are remains of a
number of small Carnivores and Ungulates, represented for the most
part by teeth. Much work yet remains to be done in identifying
these small forms. I failed to observe anything strikingly new, and
probably most belong to existing species.

It is much to be regretted that the most interesting human
implements have been scattered. The only two stone implements
still left in the collection are a spear-head and a knife. The spear-
head is 8 inches in length, about 2^ inches in greatest width, and
about f inch thick at its thickest part. In section it is triangular,
the flat side having a bulb of concussion, and the other side formed
by the removal of two long flakes. The base has been trimmed by

16 Annals of the South African Museum.

the removal of another oblique flake, and by the formation of some
small notches for the attachment of the spear-head to a wooden
shaft. The other stone implement is evidently a knife. It is 5 inches
in length. One side is flat and the other formed by the removal of
three longitudinal flakes. Owing to incrustation from the hot spring
the material of the implements could not be made out without
chipping them.

Besides the stone implements there are a number of small wooden
articles pins, etc. clearly shaped by human agency. And there are
great quantities of burnt wood. Some of the logs are of large size,
and show that there must have been much larger trees growing in
the district than is now the case. I failed to detect any human
bones in the collection. Possibly further exploration may reveal
some. Till then some doubt must remain as to what human tribe
lived in South Africa at the time of the Cape horse, the Bubalus
baini, and the extinct Wildebeest.

(17)

5. On the Skeleton of a New Pareiasaurian (Pareiasuchus
peringueyi, g. et sp. nov.). By E. BROOM, M.D., D.Sc., and
S. H. HAUGHTON, B.A.

THE almost complete skeleton described in this paper was found
early in the year 1912 at the drift across the Zak Eiver on the
main Beaufort road on the farm Dunedin, in the division of Beaufort
West. The skeleton was found in sitil on the side of a kopje,
embedded in a comparatively hard mudstone. On extraction it
was found that the limb bones of the left side were not present, and
some of these were discovered at the foot of the kopje in a slightly
weathered condition. Further, it was discovered that the whole
of the right hind-limb was missing. On development, the animal
showed signs of crushing, especially in the anterior region.

The skull is almost complete, and, with the exception of the atlas,
there is a complete vertebral column from the axis to the llth caudal
vertebra. The shoulder girdle is complete, and only portions of the
right innominate are missing from the pelvis. The right fore-limb
is almost complete, the carpus and manus being fragmentary, and
the left ulna was found weathered out together with the left femur
and tibia. A large number of ribs were preserved, and the skeleton
was covered along the back, ribs, and pelvis, by dermal ossicles,
ranging in length from 15 mm. to 70 mm.

SKULL.

The skull, though practically complete, is much crushed, and the
bone so friable that perfect development is impracticable. And thus
though it is possible to describe the general features we must wait
for other specimens before we can give a full description of the
Pareiasaurian skull in detail.

Though typically Pareiasaurian, this skull differs considerably
from any of the known South African Pareiasaurus skulls, and it
also differs from the very imperfectly known skull of Propappus
omocratus.

The snout is broad and flat, and though the flattening is partly
due to crushing, one need have no hesitation in saying that the snout

2

18 Annals of the South African Museum.

is twice as broad as deep. The nostrils are large and separated by a
very feeble internasal process of tbe premaxillaries. Immediately
behind each nostril is a distinct but small boss. With these excep-
tions the whole of the upper surface of the skull is characterised
only by the typical Pareiasaurian sculpture. Even above the orbit
there is only a rudimentary indication of a boss. The pineal
foramen is large. There are no indications of the positions of the
sutures of the upper cranial bones, and these could only be made
out by seriously damaging the specimen. The total length of the
skull from the snout to the middle of the occipital region is about
358 mm. From the middle of the snout to the back of the orbit
is 230 mm.

The most noteworthy feature of the skull is the great size of

the temporal roof. In the Pareiasaurs from the Pareiasaurus

zone the quadrato-jugal region does not descend very greatly below

the quadrate, and the angle of the region is almost as far back as the

occiput. In this specimen the large quadrato-jugal portion passes

far outwards and is turned much forwards, so that what is really its

lower border makes an angle of about 120 to the maxillary edge.

In this it agrees with the Russian Pareiasaurs obtained by

Amalitsky. These Russian Pareiasaurs are probably from a higher

zone than the South African species. They are associated with

a large species of Dicynodon and with a number of large Thero-

cephalians. Were such an associated fauna found in South Africa

without Pareiasaurus, it would be referred to a zone very high in

the Permian possibly corresponding to our Cistcceplialus zone.

The Russian Pareiasaurs differ so much from our South African

forms that it seems not improbable they will have to be placed

in a different genus. In any case, the skull certainly resembles

much more closely this South African Pareiasuclius, which is also

probably as high as the Gistecephalus zone.

Almost the whole of the lower half of this great lateral bony plate
is formed by the quadrato-jugal. At the lower angle is a huge
rounded boss about 45 mm. in diameter. Immediately in front
of this is a boss about half the size of the large one. Between
this and the point where the quadrato-jugal meets the jugal the
edge of the bone is slightly thickened and forms a prominent curve,
which when viewed from' the side looks like another boss. On the
posterior border of the quadrato-jugal are three small bosses. The
jugal is relatively a small bone which forms the lower border of
the orbit and passes backwards some distance between the quadrato-
jugal and the bone which is probably the postorbital.

On the Skeleton of a Neio Pareiasaurian. 19

Above the quadrato-jugal is a moderately large bone, which
is pretty certainly the squamosal. If this identification be correct
the arrangement of bones in the temporal region would agree
essentially with the arrangement in Procoloplion, Diadectes,
Captorhinus, and most Cotylosaurs, and differ from that of Pantylus
in having no prosquamosal. The arrangement of the bones in the
occipital and postparietal region cannot be satisfactorily made out.
There are two low bosses in the region corresponding to that
formed by the post-temporal in Procoloplion, and a third boss
near what is apparently the lower end of the squamosal. Immedi-
ately above this last boss is a marked groove which curves round the
border of the bone. There can be little doubt that this is the groove
for the auditory canal.

The teeth are not very well preserved. In the upper jaw there
seem to be 13 in each side. The teeth are relatively larger and
flatter than in Pareiasaurus, and the cusps are either smaller or
more rapidly worn down. In one tooth, probably the 12th, in which
the crown is completely preserved, there are 13 cusps, of which the
middle one is the largest, and they are arranged in a semicircle
around the anterior half of the tooth. In the anterior teeth there
are possibly 15 cusps.

The palate is not in good condition for showing the detailed
structure. So far as can be seen it agrees closely with the
Pareiasaiirus palate photographed by Amalitsky. There is appar-
ently a distinct transpalatine separated from the palatine by a
large oval foramen. The large pterygoid is firmly fixed to the
basisphenoid, and there is no possibility of movement.

The structure of the basicraniurn could not easily be made out
without damaging the specimen.

The lower jaw is pretty well preserved. The anterior part is
broad and massive. The splenial is strongly developed and forms
most of the inner and lower half of the front of the jaw. The
splenial appears to pass back as far as the plane of the middle
of the large boss in the angular. The angular is massive, but is
not of great antero-posterior length. On the lower border is a
single, very powerful boss, which differs from the corresponding boss
in Pareiasaurus in being, not hornlike, but slightly expanded at its
end, which is flattened. In Propappus omocratus there are two
hornlike processes. The articular is broad, but short. There
appears to be a distinct but feeble prearticular or goniale. There
is also apparently a distinct coronoid bone, but it and the surangular
are not very well shown.

20 Annals of the South African Museum.

Bemains of the hyoid apparatus are preserved, but not sufficiently
satisfactorily to enable one to identify the elements with certainty.

VERTEBRAL COLUMN.

Of the vertebral column there is a continuous series of 33 vertebrae,
ranging from the axis to the llth caudal, and containing
18 praesacral, 4 sacral, and 11 caudal vertebrae.

The axis is complete, save for the neural spine, which is represented
by a base 35 mm. long. The body of the axis is much shorter than
those of the succeeding vertebrae, being 35 mm. in length. It
is 40 mm. broad, and the total height of the vertebra to the base
of the spine is 60 mm. The distance between the posterior
zygapophyses is 50 mm. There is a short transverse process in
the anterior half of the vertebra for the head of a short, apparently
single-headed rib, which is almost on a level with the top of the
body. Intercentra were probably present in all the cervical
vertebrae, but \vere certainly small.

The 3rd and 4th cervicals agree in possessing bodies whose
length is equal to the greatest width, neural spines strongly
directed forward, and articular surfaces for double-headed ribs,
one surface being a process in the anterior part of the middle of
the centrum, and the other the transverse process just above
the level of the body. The bodies of these vertebrae, and of the
5th cervical, are provided with a strong longitudinal ventral ridge,
with slight excavations on either side. The zygapophyses are
not so wide apart as in the dorsals.

The 5th and 6th vertebrae are displaced, but are provided with
complete neural spines, which increase in thickness upward, and at
the top contain a pronounced depression. Each spine is 47 mm.
high. The 5th still possesses two articular surfaces for the ribs ;
but the 6th has the two surfaces fused for the articulation of a
single-headed rib. The single surface projects as a strong plate
beneath the prezygapophysis. The cervical ribs increase rapidly
in size and strength up to the 7th, which is comparatively massive.
Similarly the zygapophyses become wider apart and more strongly
developed, the following being the width between the points of
the postzygapophyses of the vertebrae up to the 8th : 2nd, 50 mrn. ;
3rd, 62 mm. ; 4th, 63 mm. ; 5th, 74 mm. ; 6th, 94 mm. ; 7th,
102 mm. ; 8th, 120 mm. The 5th vertebra, which is the most
complete of the cervicals (although somewhat crushed), gives the
following measurements : Length of centrum, 51 mm. ; whole

On the Skeleton of a New Pareiasaurian. 21

height, 119 mm.; height of spine, 48 mm.; width between points
of postzygapophyses, 74 mm.

The vertebrae from the 10th to the 19th are roughly similar,
increasing gradually in size and strength backwards. The neural
spines become longer and more regularly diamond shaped in section.
The 16th to 19th inclusive differ from the others in that the centra
in these are considerably wider than long. The following give the
measurements of one of each of the typical vertebrae :

12th (anterior dorsal) : length of centrum, 59 mm. ; width of
centrum, 60 mm. ; distance between points of transverse processes,
154 mm. ; distance between points of postzygapophyses, 128 mm.

18th (posterior dorsal) : length of centrum, 52 mm. ; width of
centrum, 65 mm. ; whole height, 163 mm. ; height of spine,
38 mm. ; distance between points of transverse processes, 174 mm. ;
distance between points of postzygapophyses, 165 mm.

The articular surface for the 13th rib is 62 mm. long, while that
for the 18th rib is 34 mm. long. The surfaces decrease in size on
passing backwards.

The sacrum consists of four anchylosed vertebrae, differing thus
from that of Pareiasaunis serridens, in which the sacrum is formed
of two vertebrae only. The sacral vertebrae diminish in length and
width from the first to the last, while the following give the
heights : Whole height : 20th, 138 mm. ; 21st, 107 mm. ; 22nd,
103 mm. ; 23rd, 120 mm. ; height of dorsal spine, 43 mm., 40 mm.,
40 mm., 48 mm. respectively. The first sacral pleurapophysis is very
massive. From the body of the vertebra to the ilium it measures
158 mm., while its articulation with the ilium measures 127 mm. in
length. The articular surface is hollowed out into a deep groove,
which accommodates the inwardly bent lower portion of the anterior
part of the ilium. The second and third sacral ribs are compara-
tively small and thin, and apparently coalesce into a single thin
plate above the posterior part of the articulation of the ilium and
first sacral rib. The fourth sacral rib is stouter, being 82 mm. long
and 50 mm. broad. The last three ribs seem to have been attached
to the ilium partly by ligament.

The caudal vertebrae present are 11 in number, and decrease
regularly in size : the 1st is 110 mm. high, the llth 50 mm. The
first six resemble the praesacrals in form, having similar dorsal
spines and bodies. From each side of the centra of these six lateral
processes are formed, the longest being 72 mm. long. Chevrons
begin on the 7th caudal. The series is incomplete.

A fairly complete set of ribs is preserved. The largest measures

22 Annals of the South African Museum.

430 mm. in length, has an average height of 20 mm., and is about
15 mm. thick. The cervical ribs are displaced and thrust forward.

SHOULDER GIRDLE.

Both sides of the girdle are well preserved, the left one being
complete save for a small part of the precoracoid, and the right one
lacking rather more of the coracoid and precoracoid. In general
appearance the girdle resembles the known parts of Propappus rogers i,
the chief differences lying in the smaller size of the glenoid cavity
and the greater width of the scapula.

The scapula is anchylosed to both the coracoid and precoracoid,
and with the former forms the whole of the glenoid cavity. This
consists of two unequal parts, that formed by the scapula being the
larger. At the anterior end of the cavity on the suture line there is
an appreciable pit. The cavity is much smaller than in Propappus
rogersi. The large precoracoid foramen is present, and the groove
continuous with it in the scapula is very deep.

The coracoid is much thicker than the precoracoid, and is
truncated at its posterior border.

The precoracoid is a thin, expanded bone which almost certainly
forms no part of the glenoid cavity. The anterior border is thickened,
but the whole of the lower border is missing.

The following are the principal measurements :

Posterior end of coracoid to top of acromion 272 mm.

Greatest width of glenoid 82 , ,

Height of scapula above acromion 237 , ,

Width of scapula immediately above acromion 79 ,,

Thickness of scapula immediately above acromion 38 ,,

Width across top of scapula 124 ,,

Length of acromion 73 , ,

Maximum height of acromion 52 ,,

Distance between anterior and posterior borders of cora-
coid and precoracoid 197 ,,

The clavicles and interclavicle are almost complete. As in
Pareiasanrus serridens, the clavicles fit into an irregular groove in
the front of the lateral process of the interclavicle. They meet just
in the middle line without fusion. The interclavicular process
measures 245 mm. from side to side. The ventral process was
probably about 200 mm. long, possessing an anterior median ridge
proximally which dies away about half-way along the length. The

On the Skeleton of a New Pareiasaurian. 23

distal end is bi-lobed, possesses two main, low ridges, and is
longitudinally striated.

There is no trace of a clei thrum.

HUMEEUS.

The right humerus is almost complete, and well preserved, but
somewhat crushed, while portions of the distal and proximal ends of
the left humerus were found weathered out.

The greatest length of the right bone is 345 mm. The bone is
greatly constricted in the middle, at its narrowest measuring
45 mm. x 40 mm. The shaft is rotated to a greater degree than in
Propappus rogersi, but less than in Pareiasaurus serridens, the angle
which the head makes with the distal end being about 45 much
the same as in Propappus omocratus (S.A.M. specimen).

The proximal end measures 184 mm. in breadth, and is compara-
tively flat. The delto-pectoral ridge is well developed, but shows
none of the twisting so prominent in Propappus rogersi. The
articular surface is long and narrow, and there is a well-marked
ridge running on the outer surface of the bone from the anterior end
of the articular surface to the constricted shaft.

The distal end differs both from that of Propappus omocratus and
from that of P. rogersi in being much narrower, and in having the
condyles much nearer together. The inner condyle appears to have
been crushed, and is rather incomplete, but it forms a prominent
narrow boss with a central ridge. The width across the distal end
is 140 mm. The large rounded prominence for the articulation of
the radius and ulna agrees in shape and position with that of
P. omocratus. There is a well-developed entepicondylar foramen.

EADIUS AND ULNA.

The right radius is perfect, and the proximal ends of both ulnae
are present.

The radius consists of a straight columnar shaft with dilated ends,
its length being 188 mm. The proximal end has an oval concavity,
79 mm. by 52 mm., about 16 mm. deep, and must have been sup-
plied with a thick pad of cartilage. The inner end of the concavity
is slightly prolonged. The distal end is irregularly triangular in
shape with the front outer angle rounded. It measures 80 mm. by
55 mm., and altogether is more massive than the proximal expan-
sion. At its narrowest the shaft measures 38 mm. by 30 mm.

The proximal part of the right ulna preserved measures 140 mm.

24 Annals of the South African Museum.

in length and 105 mm. at its greatest breadth. The olecranon
process, so prominent in Pareiasaurus baini, is not developed to
quite so great a degree, although it is much more pronounced than
in Pareiasaurus serridens. The process which lies to the outer side
of the head of the radius is triangular and has a base of 40 mm.

A number of isolated metacarpals and phalanges were found, but
their positions had been considerably disturbed.

PELVIS.

The left innominate is practically entire, and there is a large
portion of the right side. The pelvis shows differences from those
of Pareiasaurus and Propappus rogersi and from that provisionally
assigned by Lydekker to Propappus omocratus.

The axis of the ilium is directed more forwards and outwards than
in Propappus rogersi, and the whole ilium is longer and of a more
slender appearance. The crest is almost straight, 225 mm. in
length, posteriorly being turned over a little inwards. Anteriorly
the ilium is twisted, the anterior edge being turned outwards below,
so that, from the front, the ilium appears to be almost horizontally
flat. Internally the ilium is concave for the reception of the sacral
ribs. Transversely it measures 90 mm., while in its narrowest part
it measures 67 mm.

The acetabulum is shallow, and looks much more downwards than
in Pareiasaurus or Propappus rogersi. It is smaller than that of the
latter, its greatest width being only 97 mm., and its transverse
measurement 86 mm. The greatest part of the acetabulum is
formed by the ilium, which is anchylosed to the ischium. The
sutures between the pubis and the other bones cannot be
distinguished.

The distance between the anterior part of the symphysis and the
point where the pubis probably joins the ilium is 165 mm. Down-
wards from the acetabulum the pubis is thickened and bent, as in
Propappus rogersi and Pareiasaurus, forming a surface for the
supposed cartilaginous prepubis. This surface is 85 mm. long and
up to 24 mm. broad. The pubic foramen lies behind, and at the
base of, this thickening, and is much narrower and deeper than the
foramen in Propappus rogersi .

The ischium measures 165 mm. by 155 mm., and is quadrangular
and plate-like in form. Posteriorly the corner is furnished with
a short, outwardly directed process. The two sides of the pelvis
are anchylosed along the symphysis, which is considerably thickened,

On the Skeleton of a New Pareiasaurian. 25

being about 60 mm. thick in the pubic region and thinning down to
20 mm. at the posterior end of the ischium.

FEMUR.

The left femur is beautifully preserved, and differs little in general
appearance from the femur of the S.A.M. specimen of Propappus
omocratus.

The chief measurements are as follows :

Pareiasuchus Propappus Propappus
pi'riiKjiiei/i. omocratus. roye-rsi.

Length of femur 320 mm. 335mm. 290mm.

Width of head 54 75 75 ,,

Length of head 126 ,, 117 ,, 112

Width of shaft at narrowest part 70 ,, 60 ,, 60 ,,

Width of distal end of bone ... 152 ,, 137 147

The chief differences, therefore, lie in the greater stoutness as
compared with the femur of Propappus omocratus, and in the
narrower, longer, and flatter head compared with both species of
Propappus. The concavity between the lesser and greater
trochanters is not so deep as in P. omocratus, and the lesser
trochanter is less twisted. The face of the bone is of the usual
Pareiasaurian type.

TIBIA.

The left tibia is present, but it is so deformed by crushing that it
is better to give no description of it.

DERMAL ARMOUR.

Dermal ossicles occur all along the back, between the proximal
parts of the ribs, especially in the neck, and around the pelvis. The
scutes are smaller than those of Propappus.

(26)

6. On a New Species of Scymnognathus (S. tigriceps). By
R. BROOM, M.D., D.Sc., and S. H. HAUGHTON, B.A.

THE magnificent specimen which forms the type of this new species
was discovered about 3 miles WSW. of the homestead on the farm
Dunedin, Beaufort West. It consists of the nearly perfect skull, a
number of cervical vertebrae, the almost complete left fore-limb with
the shoulder girdle and mterclavicle. The discovery is one of much
importance, as though hitherto we have known a good deal of
the Carnivorous Therapsida of the Pareiasaurus and Endothiodon
zones, and even more of the carnivorous types of the Upper Beau-
fort zones, we have known almost nothing of the carnivorous forms
of the Middle Beaufort zones. Many types have been described by
Owen, but they are nearly all imperfect snouts which give little
evidence of their affinities. And for long we have been anxious to
know whether carnivores of the Cistecephalus and Lystrosaurus
zones were Therocephalians or primitive Cynodonts, or something
distinct from either. The evidence we now have shows that
they were neither typical Therocephalians nor Cynodonts, but
Gorgonopsians.

When Gorgonops was first described by Owen in 1876 it was
believed to be distinct from the other forms with mammal-like
dentition, partly owing to an apparent peculiarity of the nose, and
partly because the temporal region was believed to be roofed. Owen
formed for it a special group the Tectinarialia. Seeley and
Lydekker have both agreed as to the roofing of the temporal
region, and Seeley in 1895 made it the type of a distinct order
the Gorgonopsia. Three years ago it was shown by one of us
(Broom) that a careful examination of the British Museum type
shows that undoubtedly there is quite a large temporal opening^
though the parietal region is broad. Still, Goryonops one has always
felt differed considerably from the typical Therocephalians. For a
time it was suggested that it perhaps came nearer the Dinocepha-
lians ; latterly it has been placed with the Therocephalians for con-
venience. Now it seems more convenient to revive Seeley's order

On a New Species of Scymnognathus (S. tigriceps). 27

or sub-order Gorgonopsia for Gorgonops and a number of other allied
forms now known.

Gorgonops torvus is met with in the Endothiocloii zone, and with it
is a larger allied form, Scymnognathus whaitsi. In the Pareiasaurus
zone a small Gorgonopsian has been found by the Kev. J. H. Whaits,
and named Scylacognathus parvus. And it is thus interesting to
note that while almost all the carnivores of the Pareiasaurus zone
are Therocephalians, when we come to the Endothiodon zone
Gorgonopsians are nearly as frequent as Therocephalians, and when
we get up 2,500 feet higher Therocephalians appear to be absent, or
at least much rarer than Gorgonopsians.

As has been shown in the paper describing Sci/lacops, the
Gorgonopsians are probably the ancestors of both Anomodonts and
Cynodonts, and being thus on the direct mammalian line all new
facts concerning their morphology are extremely important.

SKULL.

The skull is nearly perfect, though slightly crushed. It agrees
closely with Scymnognathtis wlmitsi, though more powerful. From
the snout to the back of the squamosal measures 310 mm., and the
width across the zygomatic arches is 180 mm. The depth of the
snout from the nasal to the mentum is 175 mm. The interorbital
width is 80 mm., and the interternporal 93 mm.

There is nothing of special note about the premaxilla or septomaxilla
except that the latter is very large, and partly divides the anterior
nares into upper and lower passages by a inwardly directed process
or turbinal. There is also the usual outer foramen between the
septomaxilla and the maxilla.

The maxilla is very similar to that in Scymnognathus whaitsi. It
has a long posterior process which strengthens the suborbital arch
and extends to opposite the middle of the base of the postorbital arch.

The teeth are remarkable. The dental formula is i 5, c 1, m 4
for the upper jaw. The incisors are all large except the last. They
occupy a space of 50 mm. Then follows a diastema of 23 mm. in
front of the canine, the antero-posterior diameter of which is 21 mm.
Behind the canine is a diastema of 20 mm., and this is followed by
i molars which occupy a space of 30 mm. The first three molars
are larger than the 4th. All are simple pointed teeth, and we
fail to observe any serrations. All the incisors and the canines have
their points worn down, and this must have been done during the
life of the animal. Further, this wearing could only have taken
place when the jaws were widely opened, and whatever was the

28 Annals of the Soutli African Museum.

cause of the blunting of the incisors the same was the cause of the
wearing down of the canines. Probably Scymnognathus tigriceps
preyed mainly on the contemporaneous Pareiasuchus p6ringueyi and
the prehensile teeth became ground down by friction against the
mud- and sand-crusted bony dermal plates during the struggles of
the powerful Pareiasaurian with its enemy. The fact that the
incisors and canines of the lower jaw are similarly ground down
renders this theory very probable.

The sutures of the lachrymal and prefrontal are not very clear, but
both bones are evidently fairly large. The jugal forms a very deep
suborbital bar, and on passing back clasps the zygomatic portion of
the squamosal.

The bones of the top of the skull resemble those of the more per-
fectly preserved Gorgonopsian Scylacops capensis. The preparietal
is in front of the pineal foramen, and the postf rental is very well
developed. The postorbitals and parietals are large, the former
forming the upper borders of the temporal fossae.

The squamosal is very large, and remarkable chiefly by having the
zygomatic portion clasped by an outer and inner plate of the jugal.

The interparietal is well developed and deep. It differs from that
of Scylacops in having a very prominent median ridge. At the sides
it meets the squamosal, and there is evidently no tabulare or other
bone at the side of the occiput as in some Therapsidans. In many
mammals there is a bone appearing on the lateral part of the
occiput, which is apparently rightly identified as an opisthotic.
What is possibly the same bone appears on the occiput of Cyno-
donts, and in the recent paper by one of us on the Cynodont skull it
was referred to as the opisthotic. Whether it is in the Cynodont an
opisthotic may be regarded as not yet proven, but that there is a
distinct element here is beyond question. In Dinocephalians there
is pretty certainly an element on the occiput which is neither squa-
mosal, parietal, or interparietal. In Anomodonts, Therocephalians,
and Gorgonopsians there does not appear to be usually at least any
extra element, but in Lystrosaurus there is a small lateral bone which
is evidently quite distinct from the parietal or squamosal.

The palate is well preserved and very interesting. The basi-
sphenoid is somewhat Anomodont in character. It passes down
considerably below the level of the basioccipital, and forms a
deep median ridge which in front meets the median ridge formed
by the two pterygoids. Above, the basisphenoid forms a median
septum which passes upwards and forwards to near the front of the
orbital region.

On a New /Species of Scymnognathus (S. tigriceps). 29

The pterygoids are large bones. They form behind part of the
median descending ridge. Postero-laterally a process is sent to
meet the quadrate and also to support the columella cranii. Each
pterygoid also forms a huge descending pterygoid process. In front
the development of the pterygoid is slight. It meets the trans-
palatine and the palatine. Superiorly the pterygoid forms a large
thin paramedian plate not unlike that seen in the pterygoid of the
Pelycosaurs. The huge median bony septum is probably formed by
the two pterygoid plates anchylosing with a median plate formed by
the basisphenoid or ethmoid. In section no sutures can be made out,
but the outer wall of the septum is unquestionably pterygoid. In
the allied Scylacognathus parvus the pterygoids have probably had
cartilage between them, as they are not anchylosed to form a septum.

The palatines form a large part of the palate, and send up para-
septal plates, as do the pterygoids.

The vomer is a true median vomer, as in the Anomodonts, the
Cynodonts, and Mammals. It is a large thin plate which divides
the nasal space into two. The greater part of the bone is thin, but
the lower border which formed part of the roof of the mouth is
fairly strong and bmid.

There appears to be no trace of paired prevomers.

The quadrate is fairly large, and in its relations to the squamosal
not unlike that of the Cynodonts, but relatively very much larger.
There appears to be no trace of a quadrato-jugal.

The stapes is long and fairly strong.

The lower jaw is in beautiful condition, and all the elements can
be satisfactorily made out.

The dentary is as usual the largest bone. It has a very powerful,
deep symphysis. It is not yet known for certain, but it seems not
improbable, that all those supposed Therocephalians with a powerful,
deep symphysis will prove to be Gorgonopsians. It is pretty certain
at any rate that those carnivorous Therapsida with a feeble, loose
symphysis are true Therocephalians. The coronoid process is
fairly well developed, but shorter than in typical Therocephalians.

The splenial is large. It forms much of the lower part of the
symphysis and most of the inner side of the front half of the jaw.

There is a distinct coronoid bone. It is a thin triangular splint
that supports and holds together parts of the dentary, the surangu-
lar, the angular, and the prearticular.

The angular is the second largest bone of the jaw. In front it
passes between the splenial and the dentary, and extends nearly to
the symphysis. Posteriorly it forms most of the lower half of the

30 Annals of the South African Museum.

jaw. A considerable part of its inner surface is covered by the sur-
angular, a large part by the prearticular, and a small part by the
coronoid. The bone is divided on its lower aspect by a deep groove,
as in the Pelycosaurs.

The surangular resembles that bone in AnomodonU and Thero-
cephalians.

The prearticular is a long slender splint bone. Posteriorly it
clasps the articular, and in front lies mainly on the angular, and
also articulates with the dentary, the coronoid, and the splenial.
Anteriorly it reaches to the molar region.

The articular is a short strong bone. It is supported by the
angular, the surangular, and the prearticular.

VERTEBRAE.

The proatlas is a well-developed paired element. Each half is
irregularly rhomboidal. It articulates with the atlas.

The atlas is formed as in Pelycosaurs, Dinocephalians, and
Anomodonts a pair of upper arches and a lower centrum which
forms the odontoid process of the axis. Each half of the arch has
a small zygapophysis for articulation with the axis and a transverse
process which supports the atlantal rib.

The body of the axis is 38 mm. long and 30 mm. broad. The
width across the posterior zygapophyses is 38 mm. The neural
spine is elongated anteriorly, diminishing in height in the same
direction. Its greatest length is 56 mm. and its greatest height
40 mm.

There are four other cervical vertebrae preserved, similar to each
other and characterised by a short, rounded body, a long transverse
process which forms the articular surface for a single-headed rib,
and a low dorsal spine. The third of these vertebrae, which is
typical, has the following measurements : Length of body, 29 mm. ;
greatest width of body, 37 mm. ; total height, 62 mm. ; height of
spine, 11 mm. ; width across transverse processes, 85 mm. ; width
across postzygapophyses, 29 mm.

SHOULDER GIRDLE.

Both right and left sides of the shoulder girdle are present, the
only parts missing being the right clavicle, part of the left coracoid,
and the distal end of the interclavicle. The sutures between the
coracoids and precoracoids are almost obliterated, but there seems
no doubt that the usual three elements are present, and that the

On a New Species of Scyrnnognathus (S. tigriceps). 31

girdle resembles the Therocephalian type as seen in Ictidosuclius
primaevus.

The scapula has a bi^oad lower end, and forms the upper half of
the glenoid cavity, which half is almost a plane surface bounded
by rather more than a semicircle. In its length it is curved back-
wards and inwards. The precoracoidal foramen is seen passing
down from the inner side of the scapula, where it forms a
moderately deep groove, and appears on the outside just below
the suture between the scapula and precoracoid. The right pre-
coracoid was found broken in such a way as to show the position
and direction of the precoracoidal canal. The scapula possesses
no acromion process, and there is no trace of a cleithrum. As a
loosely articulated cleithrum exists in the allied Scylacops capensis
it is more probable that it has been lost than that none existed.

The coracoid forms the lesser and lower half of the glenoid cavity,
the glenoid surface being concave, looking upwards and outwards,
and being separated from the outer side of the bone by a pro-
minent ridge. Posteriorly, the subglenoid portion is prolonged
into a process which curves slightly upwards and whose thickness
rapidly diminishes. On the left coracoid this process has been
compressed upwards and backwards.

The precoracoid is a thin plate-like bone which is anchylosed with
both the other elements. It is slightly smaller than the coracoid.
The suture between the two ventral elements appears to be a
straight one, and the precoracoid appears to form no part of the
glenoid cavity.

The chief measurements are as follows :

Greatest length of scapula 130 mm.

Width of base of scapula 80 ,,

Width of top of scapula 49

Greatest width of glenoid 43 ,,

Greatest height of glenoid 48 ,,

Length of precoracoid coracoidal suture 70 , ,

Length of coracoidal process behind glenoid 84 , ,

Distance between anterior and posterior edges of precoracoid 64 ,,

The clavicle is a slightly curved bone, flattened from above at its
upper end, oval in the middle, and expanded into a wide, flat plate
at its articulation with the interclavicle, being thickest there an-
teriorly, and thinning out to about 1 mm. behind. The two clavicles
do not meet. The total length of the left clavicle is 200 mm., and
the greatest width at the proximal end 45 mm.

32 Annals of the South African Museum.

The interclavicle is an elongated, truncated-triangular-shaped
bone about 220 mm. long and 80 mm. broad at its posterior end,
thin, slightly convex on its upper surface, and provided with a pro-
nounced ridge on its under surface. This ridge dies away distally
and proximally from an apex in the centre. From the ridge the
ventral surface slopes to left and right, and over these sloping sur-
faces the clavicles seem to have been able to move with a certain
amount of freedom.

HUMERUS.

Both humeri are complete and in a beautiful state of preservation.
The shaft is not greatly twisted, the planes of the two ends of the
bone lying at an angle of about 30 with each other ; but the lower
portion of the proximal angle is further bent until its plane makes
an angle of about 70 with that of the distal end.

The greatest length of the bone is 246 mm. ; the proximal end is
125 mm. wide; the distal end 113 mm. wide; while the narrowest
part of the shaft measures 31 mm. in diameter.

The proximal end is thickened at the articular surface, while the
lower twisted portion is comparatively thin. Anteriorly there is
a well-marked ridge running outwards from the lower end of the
articular surface ; this ridge can be traced along the shaft, and
becomes very prominent on the distal portion of the bone. The
articular surface of the proximal end is elongate-oval in shape,
widest at the middle, measuring 83 mm. in length and 38 mm.
at its greatest breadth. From a median longitudinal ridge it slopes
away on either side in a slightly convex curve. The delto-pectoral
ridge is not very pronounced, and there is a shallow concavity
above it.

At the distal end the inner side of the bone is pierced by
the entepicondylar foramen, which passes from behind down-
wards and forwards. A well-defined groove running outwards
from the upper end of the foramen gives rise to two prominent
bounding ridges to the bone in this region, the bridge being very
pronounced. The foramen, which is oval in shape and has a longer
diameter of 21 mm., is situated 76 mm. from the distal end.
Another groove runs from it along the under surface, between
the border of the bone and the inner condylar boss.

In general shape the hurnerus agrees with that figured by Owen
as " Cynodrakon major," but it differs from this chiefly in the
shorter and wider proximal portion which it possesses, in the
greater rotation of the shaft, and in the shape of the proximal
articular surface.

On a New Spades of Scymnognathus (S. tigriceps). 33

EADIUS AND ULNA.

The bones of both sides are present, those of the left being found
in sitil in connection with the humerus, and but slightly displaced
from their natural positions.

The radius is 155 mm. in length, expanded at its two ends, and
19 mm. in diameter at its narrowest. Proxirnally, the articular
surface is elongated slightly in the direction of the ulna, and is
slightly concave. It is 50 mm. long and 40 mm. broad. From the
proximal end a ridge runs down the ulnar side of the bone ; on the
opposite side the shaft is flattened ; so that in section the shaft is
markedly triangular at its middle point, with the two anterior
angles rounded. The distal articulation is almost circular in section,
measuring 40 mm. in diameter, and is convex.

Only the proximal ends of both ulnae are preserved. The articular
surface measures 44 mm. in length and about 37 in breadth. There
is no well-developed olecranon process for wrapping round the
humerus. Distally, the bone becomes thinner, and 80 mm. from
the proximal end has an average thickness of only 11 mm.

MANUS.

The manus of the' left side is almost complete, and the bones are
preserved in position and in connection with the radius and ulna.
We are thus for the first time in a position to give a complete
account of the Gorgonopsian carpus, metacarpus, and phalanges.

The most striking feature of the carpus is its resemblance to that
of the Pelycosaurs and its less striking resemblance to that of either
the Anomodonts or the Dromasaurians. This manus also sets at
rest all doubt as to the Therocephalian digital formula. Formerly it
was believed to be 2, 3, 3, 3, 3, but two years ago it was shown by
one of us that the formula almost certainly was 2, 3, 4, 5, 3, there
being two very short epiphysis-like phalanges in the 4th digit and
one short phalanx in the 3rd digit. We now see that this is correct,
and that the formula is as in the Pelycosaurs 2, 3, 4, 5, 3, and not
as in the Dromasaurians, Anomodonts, and Cynodonts 2, 3, 3, 3, 3.

The radiale articulates apparently with the whole of the distal end
of the radius. Though relatively short it is wide and deep. Distally
it articulates with both the centralia, and when the manus is flexed
also with the 1st carpal.

The ulnare is comparatively flat and much longer than narrow.
Proximately it articulates with the ulna and the intermedium, and
possibly also with a small pisiform. Distally it articulates with the

3

34 Annals of the South African Museum.

fused 4th and 5th carpalia, and its inner side articulates with the
outer centrale.

The intermedium is imperfectly preserved, but was pretty
certainly small. As preserved it lies between the radius and
ulna.

The two centralia are of about equal size. The outer is irregu-
larly cubical, and lies between the radiale and ulnare. The inner
lies between the radiale, the outer centrale, and the 1st and 2nd
distal carpalia.

There are apparently four distal carpals, but there is good reason
for believing that what is apparently the 4th is really the 4th and
5th anchylosed.

The 1st distal carpal is a comparatively flat bone somewhat
broader than long, and it gives articulation to the first metacarpal.
At its outer and posterior corner there lies a small distinct bone
which is unfortunately imperfectly preserved. This may be
regarded as a radial sesanioid or prepollex.

The 2nd distal carpal is a small cubical bone, much thicker than
the 1st, and it gives articulation to the 2nd metacarpal.

The 3rd distal carpal is when viewed from above also small, but it
is very thick. It supports the 3rd metacarpal.

The combined 4th and 5th distal carpalia is a large bone nearly
twice as wide as long. It is not so thick as either the 2nd or 3rd.
What makes it pretty certain we are dealing with two bones and not
one are the following characters : A ridge divides the bone into an
outer and inner half, and the bony fibres of the inner half at least
can be seen radiating out from the centre of this inner half, and
further the two halves have each a distinct group of nutrient
foramina. The combined bone supports the 4th and 5th meta-
carpals.

The 1st metacarpal is very imperfectly preserved. It evidently
was broad and flat.

The 2nd metacarpal is a short bone only a little longer than
broad. The distal end seems to indicate that the 1st phalanx could
be considerably overextended.

The 3rd metacarpal is nearly twice as long as broad, and the 4th
metacarpal is nearly three times as long as wide.

The 5th metacarpal is comparatively flat and broad. It is less
than twice as long as broad.

The phalanges of the 4 outer toes are almost perfectly preserved.
In the 2nd digit the 1st phalanx is comparatively short only a little
longer than broad, and it is rather flat. The second phalanx is also

On a New Species of Scymnognathus (S. tigriceps). 35

short and broad. The ungual phalanx is long and powerful, but not
greatly curved. It manifestly had a powerful claw.

The 3rd digit has the 1st phalanx short and powerful, almost as
broad as long. The 2nd phalanx is very short, its length being only
about i of its width. Though so very short it manifestly has
hinged freely on both the 1st phalanx and the 3rd. The 3rd
phalanx is a strong short bone about as long as broad. The ungual
phalanx is like that of the 2nd toe, long and powerful, and only
slightly curved.

The 4th digit has the 1st phalanx short and powerful, very little
longer than broad. Following it are two short plate-like phalanges
agreeing closely with the short phalanx in the 3rd toe. Each is
only about ^ as long as broad, and each seems to have very free
movement. The 4th phalanx is much like the penultimate
phalanges of the other toes. The ungual phalanx is also similar
to that of the 2nd and 3rd toes.

The 5th digit has only three phalanges the 1st and 3rd like the
proximal and distal phalanges of the other toes, and the middle
phalanx like the penultimate phalanges of the others.

It will thus be seen that though the digital formula is 2, 3, 4, 5, 3,
the 3rd and 4th toes have been shortened by the 2nd phalanx in the
3rd toe, and the 2nd and 3rd phalanges in the 4th toe becoming
very much reduced in size.

Considering the width of the foot the toes are short. The extent
of movement of the bones is not quite clear. Probably the animal
walked with the foot mostly on the ground after the manner of the
bear. Considerable movement seems to have been possible between
the carpals and metacarpals, and more between the metacarpals
and phalanges. Considerable flexion of the claws on the palm was
possible, but the whole hand can only be regarded as a very imper-
fect instrument of prehension.

As it is pretty certain that both the Anornodonts and the Cyno-
donts are descended from a Therocephalian ancestor, it would be
very interesting to find out exactly how the 2, 3, 3, 3, 3 digital formula
was derived from the 2, 3, 4, 5, 3 formula of the Therocephalian. We
see clearly one stage the great reduction in length of the 2nd
phalanx of the 3rd toe and of the 2nd and 3rd phalanges of the 4th
toe, but at present we have no evidence as to whether these reduced
phalanges became further steadily reduced or suddenly disappeared.
The latter seems the more probable alternative.

(36)

7. On Two New S-pecies of Dicynodon. By E. BBOOM, M.D., D.Sc.,

and S. H. HAUGHTON, B.A.

DICYNODON TESTUDIEOSTRIS, sp. nov.

THE type of this new species (No. 2354 S.A.M.) was collected at
Dunedin, Beaufort West, just south of the Homestead, and is prob-
ably from the Cistecephalus zone. It is a small skull of a female,
measuring in greatest length 85 mm., or, allowing for slight crushing,
probably originally 90 mm. The greatest breadth is 56 mm. From
the snout to. the front of the orbit is only about 18 mm. when the
skull is viewed directly from above. The orbit is about 20 mm. in
length, and from the front of the postorbital arch to the back of the
squamosal is about 47 mm.

Two specimens were discovered near the same spot, which agree
closely. Both are tuskless, and would formerly have been placed in
the genus Oudenodon, but we now know that this is the female of
Dicynodon. In a number of respects the present species differs
remarkably from the Dicynodons of a lower horizon, and it will
probably be necessary to break up the genus later on. At present,
however, we do not know enough safely to do so, and for a time we
may conveniently place all in the old genus Dicynodon, even though
the number of species is becoming large.

The beak is very short, and the caniniforni process lies under the
middle of the orbit. The arrangement of bones round the nostril is
remarkable. The premaxilla is, as always in Dicynodon, unpaired.
It is flattened in front and rounded at the sides. It passes up
between the nostrils and between the nasals. The maxilla, though
not large, forms practically the whole of the cheek. It almost
reaches to the orbit, hiding the jugal and lachrymal except just at
the orbital margin. It forms the anterior margin of the nostril and
has a fairly long articulation with the nasal.

There is no evidence of a septomaxillary, at least on the outer side
of the skull. Whether there may be one hidden underneath it is,
impossible to say without damaging the skulls. We incline, how-
ever, to think that the septomaxillary is absent. Formerly it was

On Tivo New Species of Dicynodon. 37

believed to be absent in Dicynodon. Then it was clearly seen in one
or two specimens. Probably we shall find that in some species it is
present and in others absent.

The nasal is fairly large, and has a large articulation with the
frontal, the maxillary, and the premaxillary.

The lachrymal and prefrontal are much reduced. Probably the
lachrymal is considerably hidden by the nasal and maxilla, but the
prefrontal can only be regarded as quite rudimentary. In many
species of Dicynodon it is a large bone, and the great reduction here
seems to suggest an affinity with Cistecephalus, where it is also
rudimentary.

The frontal is large and forms most of the upper margin of the
orbit. The postfrontal is present, but very small though fairly long.

The postorbital is very large, and besides forming the whole of the
outer part of the postorbital arch it forms almost the whole of the
upper margin of the temporal fossa. The pair of postorbitals form a
large part of the upper surface of the skull, meeting each other
behind the pineal foramen and covering most of the parietals.

The preparietal is fairly large and almost entirely in front of the
pineal foramen, only a minute portion of the bone touching the front
of the foramen.

The parietals are well seen on each side of the foramen and in
front of it, but are mainly hidden behind by the postorbital.

The squamosal is of usual type and presents no specially note-
worthy features. The quadrate- jugal is very large, and apparently
not anchylosed to the quadrate, as is usually the case.

The palate has not been fully displayed in either of the specimens,
but so far as can be seen is of the normal Dicynodon type.

DICYNODON ALTICEPS, sp. uov.

The skull on which the following description is based was collected
about one mile west of Oudeberg, in the division of Graaff Beinet, on
the main road cutting between Murraysburg and Graaff Beinet.
Although crushed and slightly distorted, and imperfect in the
zygomatic arches and the tip of the snout, the skull is in a good
state of preservation.

In general appearance the skull resembles Dicynodon leoniceps,
but besides being much smaller, it differs from Owen's species in a
number of important particulars sufficient to warrant us in the
erection of a new species.

The snout is inclined at an angle of about 50 to the plane of top
of skull.

38 Annals of the South African Museum,

The suture between the maxillaries and the premaxillary is inde-
terminable. The maxillae are fairly large bones, forming most of
the snout behind the nostrils, which are roughly circular and about
24 mm. in diameter. Posteriorly the maxillae have suborbital
portions which articulate with the jugal and meet the anterior
process of the squamosal. The caniniform processes are found just
behind the plane of the back of the nostrils, and the tusks, which are
incomplete, curve downwards, forwards, and very slightly inwards
the general direction, however, being slightly more forwards than in
D. teoniceps.

The nasals form the border of the upper posterior quadrant of the
nostrils, and, passing backwards, are apparently separated from one
another, save near the frontals, by the internasal process of the
premaxilla.

The sep torn axillary cannot be discerned.

The lachrymal forms a portion of the face about 35 mm. long and
12 mm. broad, and near its lower end forms a small boss on the
border of the orbit. Anteriorly, the lachrymal advances almost to
the nostril.

The prefrontal is larger than the lachrymal and forms a prominent
supraorbital boss, which is the anterior part of the supraorbital ridge.
The frontal forms about 25 mm. of the upper border of the orbit,
and apparently passes backward in a slender prolongation to behind
the plane of the pineal foramen. The postfrontals are not well-
defined, but they appear to be fairly large bones.

The postorbital forms almost the whole of the outer part of the
postorbital arch, besides extending over the length of the inner wall
of the upper margin of the temporal fossa. Nowhere, however, do
the pair of bones meet.

The pineal foramen is fairly large, situated a short distance behind
the front of the temporal fossa, and is wholly enclosed by the pre-
parietal. Immediately behind the foramen is a small, elongate boss.
The preparietal is about 30 mm. long, and extends as much behind
as before the pineal foramen.

The parietals extend forward to a point just anterior to the back
of the pineal foramen. The parietal region consists of a fairly deep
groove bounded by the outer margins of the parietals and the post-
orbitals. The total length of the parietal is 37 rum.

The squamosal is a large bone, not wholly perfect, in which the
curve between the zygornatic and downward portions is more regular
and not so markedly angular as in many other species. The relations
of the squamosal with the occipitals is not displayed. Quite a large

On Two New Species of Dicynodon. 39

proportion of the upper and inner portion is covered by the post-
orbital. The zygomatic portion passes forward to a point below the
orbit, where it comes to an end between the maxilla and the jugal.
The downward portion is attached at its lower end to the quadrate,
and also supports the large quadrato- jugal plate.

The palate, although not fully developed, appears to be of the
normal type.

The following are some of the chief measurements :

Greatest length 242 mm.

Greatest breadth probably 170 ,,

Snout to front of orbit 67 ,,

Length of orbit 48 ,,

Minimum width between orbits 42 ,,

Minimum width of temporal roof 18 , ,

Front of postorbital arch to back of squamosal... 127 ,,

Maximum height of skull 115 ,,

Maximum width of lateral plate of squamosal

probably 35 ,,

The chief features, therefore, lie in the height of the skull, in the
rather large size of the orbit, the extended postorbital, and the large
ratio (7 : 3) between the frontal and temporal widths.

(40)

8. On a Skull of Tapinocephalus atherstoni, Owen. By S. H.

HAUGHTON, B.A.

EECENTLY the South African Museum was presented with a skull,
without the lower jaw, by Mr. P. Le Eoux, of Uitkyk, in the Gouph
district of Beaufort West, by whom other Dinocephalian bones have
been donated in the past. This skull almost certainly belongs to
Tapinocephalus atherstoni, and is the only skull we yet possess.

FIG. 4. TapinocephaliLS atherstoni, Owen.
Top view of skull, x &

Tapinocephalus atherstoni was described by Owen in 1876 from
an imperfect snout sent by Dr. W. G. Atherstone from Jan Willem's
Fontein, Prince Albert District. A large number of other bones were
sent by Atherstone at the same time, but these bones have apparently
got mixed, and some of them were figured by Owen as bones of
Pareiasaurus. From the Brit. Mus. Catalogue there would appear
to be nearly a whole skeleton of Tapinocephalus all bones sent by
Atherstone in 1872. Some of the bones are said to come from
Gats-Plaatz, others from Vers Fontein, but as there is admittedly
some doubt about some of the localities, and as no bone occurs in

On a Skull of Tapinocephalus atherstoni, Owen.

41

duplicate, one suspects that all the bones are from one locality and
of one individual. In 1909 Broom described a Dinocephalian skull
in the possession of the British Museum under this same specific
name. Dr. Broom now informs me, however, that it is almost
certain that this latter skull, howbeit a Dinocephalian, is not
Tapinocephalus atherstoni, although closely allied to that species.
The greatest length of the skull under description is 530 mm.,
while the distance from the occipital condyle to the front of the
snout is 435 mm. The maximum width of the skull is equal to its
length, while its height in the temporal region is 285 mm. It con-
sists of a large, high, broad, rounded temporal and posterior portion
and a flat, broad snout. The frontals rise abruptly from the nasals
almost at right angles.

FIG. 5. Tdpinocephalns atherstoni, Owen.
Side view of skull, x

Owing to the weathered condition of the surface of the skull, and
the hardness of the matrix, the sutures between the bones are not
easy of determination. Fortunately, however, on arrival at the
Museum the skull was found to be broken in several pieces, a fact
which has materially assisted in the working of the skull. The
palate is very well displayed.

The premaxilla was comparatively small, and bore teeth, as did
the maxilla. Most of the teeth are lost, and the number is indeter-
minable. Just behind the junction between the premaxilla and
maxilla of the left side there is a portion of a large tooth, and,
internal to it, is a small secondary replacing tooth, circular in section,
5 mm. in diameter.

The maxilla is large, moderately flat, and carried teeth. Its

42 Annals of the South African Museum.

posterior limit cannot be seen. The septomaxillary is well displayed.
It lies along the floor of the nostril and ascends to form a small part
of the face. As pointed out by Broom, this is the bone which was
described by Owen as the palatine.

The orbit is large and round, and looks outwards, upwards, and
slightly forwards. The supraorbital border overhangs it to a certain
extent, and behind, the postorbital arch is very strong. The limits
of the lachrymal and prefrontal are not well-defined, but both seem
to be present, and the lachrymal the larger of the two. The jugal,
which forms the lower border of the orbit, is a flat bone, relatively
small, which passes backwards to meet the squamosal.

It is impossible to define the quadrato-jugal.

The temporal fossa is small, and is directed outwards and back-
wards. Its anterior, and almost the whole of its superior, border
appear to be formed by the postorbital, while the squamosal bounds
the inferior and posterior sides. The squamosal is of the normal
Dinocephalian type, meeting the parietal, the quadrate, and the jugal
by means of three processes.

The frontals and parietals are very much thickened, and the
whole of the top of the skull is very broad. Just behind the plane
of the back of the orbit the temporal region is slightly hollowed, with
a slightly raised boss in the centre of the hollow. This is probably
the pineal region ; but the pineal foramen is apparently almost, if
not entirely, covered with bone.

The occiput is fairly flat and large and slopes backwards and
upwards. The condyle is rounded, 55 mm. wide, and on its upper
side has a deep groove for the medulla. This groove is about 25 mm.
broad and half as deep. In the middle line below the medullary
groove is a pit 25 mm. broad. There is a large exoccipital which
touches the squamosal and quadrate. The stapes is elongated
posteriorly in the direction of the quadrate.

The structure of the palate is essentially that given by Broom
in describing the British Museum skull. There is no trace of a
transpalatine.

In addition to those given above, the following are some of the
chief measurements :

Distance from occipital condyle to front of snout 435 mm.

Length of snout 220 ,,

Distance from quadrate to front of snout 290 ,,

Breadth of palate at posterior end of prevomers 280 ,,

Length of prevomers 145 ,,

(43)

9. On a New Species of Propappus. By S. H. HAUGHTON, B.A.

THE pelvis which forms the type of this new species was
discovered by me at the drift across the Zak River on the main
Beaufort road, on the farm Dunedin, in the division of Beaufort
West. In conjunction with it were found a number of vertebrae and
ribs, together with some dermal ossicles.

A large portion of the right innominate is preserved, including the
whole of the ilium, the acetabulum, and the anterior portions of
the ischium and pubis. The ilium differs considerably in shape from
that of any other South African Pareiasaurian. The crest is short,
with its anterior part curved outwards in a quarter- circle, in such a
manner that there is a marked concavity at the anterior end of
the ilium. The straight part of the crest measures 120 mm. in
length, while the whole length is about 170 mm. The greatest
height of the ilium is 115 mm., while the distance from the top
of the acetabulum to the most anterior part of the pelvis is 185 mm.
The neck of the ilium is greatly constricted, measuring only 43 mm.
in height and 23 mm. in thickness at its narrowest part.

The acetabulum is distinctly oval in shape, the longer axis lying
along the suture between the ilium and the posterior bones, shallow
except for the development of the upper iliac and upper ischial
border. Its height is 90 mm., and its greatest width 57 mm. More
than half the acetabulum is apparently formed by the ilium, about
one-sixth by the pubis, and the remainder by the ischium ; but the
sutures between the bones can be traced only with difficulty and in
part. The acetabulum looks downwards and outwards.

The ischium displays a shape very different from that of any
other Pareiasaurian. In Pareiasuchus there is a well-marked
tuberosity at the posterior corner, and there is a smaller one in
Pareiasurus serridens ; but in this specimen the tuberosity is a
large, thickened process, the anterior border of which is at right
angles to the upper border of the ischium. The shape and size
of this can best be understood from reference to the figure. The

44 Annals of the South African Museum.

lower half of the posterior border of the ischiurn. is missing, but
below the protuberance the border thins rapidly.

The pubic foramen occupies relatively the usual position. The
thickening for the pre-pubic cartilage, so prominent in other
Pareiasaurians, is not present here. Downwards from the aceta-
bulum the pubis is bent slightly backwards, but its thickness
reaches only 10 mm.

The sacral ribs, apparently four in number, fitted into a very
shallow groove on the inside of the ilium. The first, which was
much expanded distally, seems to have given support by curling
round the lower border of the ilium. The second and third ribs
were fairly stout.

The shape of the ilium and of the acetabulum, the large ischial
protuberance, the absence of the thickened pubic brim, and the
diminutive size are all new features displayed by this pelvis, and
serve to distinguish it from any hitherto described Pareiasaurian.

The vertebral remains consist of a series of six dorsal vertebrae,
and a number of connected caudal vertebrae.

The dorsal vertebrae are possibly the 12th to 17th inclusive, and
are characterised mainly by the shortness of the neural spines,
and the general size, which is less than that of any known
Pareiasaurian vertebrae, with the exception of those of Anthodon.
Those preserved differ but little from one another. The centra are
as long as they are broad, but constricted considerably in the middle,
so that they have an elongated appearance. On the lower side both
ends of the body are truncated obliquely, so that, although no inter-
centra are preserved, they were probably well developed. The
following measurements of what is possibly the 13th may be taken
as typical of these dorsal vertebrae : Total height of vertebra,
107 mm. ; width across transverse processes, 120 mm. ; length of
centrum, 41 mm. ; height of spine above junction of postzyga-
pophyses, 18 mm. The transverse process is continued downwards
and forwards to the front of the body, and gives rise to a surface,
52 mm. long, for the articulation of a single-headed rib. From
the transverse process the anterior zygapophysis is formed, lying
at about the level of the top of the body.

There is a continuous series of eleven caudal vertebrae,
comprising probably the 5th to the 15th inclusive. The bodies
of the first three of the series differ from those of the praesacral
vertebrae in that they are not medially constricted. The total
height of the 2nd of the series is 56 mm. ; the body is 24 mm. long ;
the width between the points of the postzygapophyses is about

On a New Species of Propappus. 45

40 mm. ; and the spine is 15 mm. high. The vertebrae diminish in
size backwards, the height of the penultimate being 40 mm., and the
maximum width across the transverse processes 29 mm. The spines
are short, and directed backwards. Chevrons begin after the 3rd of
the series and are continued to the end.

The dermal ossicles are comparatively large, and were very
plentifully scattered over the back and ribs of the animal. The
largest of them reached a diameter of 55 mm., which is a greater
size than is seen in the larger animal Propappus omocratus. They
have the typical shape and sculpture of the normal Propappus
ossicles.

Dr. Broom has kindly examined the specimen for me, and
confirms my opinion that it must be regarded as a new species.
For it 1 propose the name Propappus parvus, sp. nov.

(46)

EXPLANATION OF PLATES.

PLATE I.
FIG.

1. Caruichthys ornatus, Broom, x J.

2. Atherstonia caintcrossi, Broom, x .

PLATE II.

1. Palaeoniscus capensis, Broom, x J.

2. ElonicJithys u-haitsi, Broom, x J.

PLATE III.

1. Pareiasuchus periii/iucyi, Broom and Haughton. Posterior view of the left

shoulder blade, x 3.

2. Pareiasuchus p6ringueyi. Outer view of left shoulder blade, x J.

3. ,, ,, Anterior view of right humerus. x ^.

4. ,, ,, Posterior view of right humerus. x .

5. ,, ,, Anterior view of right radius, x .

6. ,, ,, Anterior view of proximal half of right ulna, x .

PLATE IV.

1. Pareiasnchu* pi'ru>u/u<yi, Broom and Haughton. Posterior view of left

femur, x .

2. Pareiasuchus p6ringueyi. Anterior view of left femur, x .

3. ,, ,, Outer view of left side of pelvis with sacral rib. x

4. ,, ,, Lower view of left side of pelvis with sacral rib. x .

5. Propappus parvus,lELa,ugliton. Outer view of right side of pelvis, x J.

PLATE V.
Parc-lanHchus pi'rinyueyi, Broom and Haughton. Mounted skeleton, x y 1 ^.

PLATE VI.

1. Scymnognathus tigriceps, Broom and Haughton. Side view of skull, x J.

2. ,, ,, Side view of left side of shoulder girdle, left clavicle,

and interclavicle. X i nearly.

3. ,, ,, Eight side of shoulder girdle, x J nearly.

4. ,, ,, Anterior view of left fore-limb, x J nearly.

5. Scylacops capensi>i, Broom. Top view of skull, x ^.

PLATE VII.

1. Dicynodon alticeps, Broom and Haughton. Side view of skull, x J nearly.

2. ,, ,, Top view of skull, x .

3. Dicynodon testiidirostris, Broom and Haughton. Top view of skull, x |.

4. ,, ,, Side view of skull, x |.

Annals S. Afr. Mus. Vol. XII.

Plate I.

CARUICHTHYS ORNATUS, BKOOM.
ATIIKHXTONIA CAIRNCROSSI, BROOM.

mals S. Afr. Mus. Vol. XII.

Plate II.

\

PALJUONISCUS CAPEXSIS, BROOM.
ELONICHTHYS WHAITSI, BKOOM.

Annals S. Afr. Mus. Vol. XII.

Plate III.

PAREIASUCHUS PERINGUEYI, BKOOM & HAUGHTON.

Annals S. Afr. Mus. Vol. XII.

Plate IV.

>;

PAREIASUCHUS PERINGUEYI, BROOM & HAUGHTON.
PROPAPPUS PARVUS, HAUGHTON.

Annals S. Afr. Vol XII.

Plate V.

o

H

B
O
p

K

o
o

K]

i

nnals S. Afr. Mus. Vol. XII

Plate VI.

SCYMNOGNATHUS TIGRICEPS, BROOM & HAUGHTON.
SCYLACOPS CAPENXIti, BBOOM.

Annals S. Afr. Mus. Vol. XII.

Plate VII.

DICYNODON ALTICEPS, BROOM & HAUGHTON.
DICYNODON TESTUDIROSTR1S, BKOOM & HAUGHTON.

(47)

10. Investigations in South African Fossil Reptiles and Amphibia
(Parts 1-4). By S. H. HAUGHTON, B.A., F.G.S., Assistant
Director.

1. On a New Species of Trematosaurus (T. sobeiji).
(Plates VIII., IX.)

THE remarkably fine skull upon which the following observations
are based was recently presented to the South African Museum
by Messrs. Sobey Bros, through the medium of the Queenstown
Municipal Authorities. It was found in the building-stone quarries
belonging to these gentlemen, and when received at the Museum
the top had been wholly freed from matrix. The palate was em-
bedded in a slab of hard, homogeneous calcified sandstone, and the
occipital condyle had been broken off, and has not been recovered.
I have cleaned one side of the palate to show the sutures between
the bones. In doing so the skull fractured across in three places,
and thereby enabled some knowledge of the relations of the median
bones to be obtained. The skull obviously belongs to a hitherto
undescribed species, and I have pleasure in naming it after the
donors.

The skull is large, triangular in outline, the snout gradually taper-
ing. The nostrils are oval in shape, approximate, some distance
behind point of snout. Orbits are small, placed midway between
front and back of skull. The snout is rugose. The sculpturing
of the cranial bones is of the usual Labyrinthodont type, pitted at
the centre with radiating grooves at the periphery. The supra-
squamosal, postfrontal, and postorbital are relatively less grooved and
more pitted than the other bones.

The chief measurements of the skull are as follows :

Greatest length along border 517mm.

Greatest breadth 305 ,,

Tip of snout to occipital plate 411 ,,

Snout to plane of front of nostril 65 ,,

4

48 Annals of the South African Museum.

Length of nostril 27 mm.

Internasal breadth 31

Preorbital length 217 ,,

Length of orbit 24-5 , ,

Interorbital width 90 ,,

Length of frontal 160 ,,

Length of nasal 149 ,,

Length of parietal 114 ,,

The premaxillae are divided from one another by a median suture,
and posteriorly they are separated for half their length by the nasals.
They form most of the snout, and the whole of the anterior borders
of the nostrils. Midway between the front of the nasals and the
snout the premaxillary suture is pierced by a small fenestra, which
apparently contains no bone corresponding to the internasal described
by Watson in Micropholis stowi. A fracture obliquely across the
nostrils shows the premaxilla lying outside the septomaxilla and
below and within the maxilla, articulating with the latter by a
complex suture.

The maxilla extends for a length of 308 mm. from the premaxilla
back to the quadrato-jugal. It is a narrow bone, tapering out
posteriorly from a maximum height of 27 mm. just behind the
septomaxilla.

The septomaxilla appears on the surface of the face for a distance
of 11 mm. behind the nostril, and forms almost the whole of the
posterior border of the nostril, the nasal just entering into the border
on the superior edge, The septomaxilla meets the maxilla and the
nasal, but not the lachrymal. Its cheek portion has a triangular
form. It forms the floor of the nostril, and extends downwards
inside the maxilla and premaxilla.

The nasal is a large bone articulating with the premaxilla, septo-
maxilla, maxilla, lachrymal, prefrontal, frontal, and interfrontal.
Anteriorly and between the nostrils it is comparatively narrow ;
in the middle it widens out to form the whole of the upper surface,
while posteriorly it tapers, being separated from its neighbour by the
interfrontal and frontals.

The interfrontal is a median rhombic bone, 50 mm. long and
9 mm. broad, articulating for half its length with the nasals and
for half with the frontals. Broom has recently described this bone
in Eryops, and it is known to occur in a number of small Stego-
cephalians. Unfortunately the type of Trematosaurus kannemeyeri
does not show this region. If an interfrontal did occur in that

South African Fossil Reptiles and Amphibia. 49

species, however, it must have been considerably further in advance
of the orbits than in the species under consideration.

The lachrymal is a long bone forming the inferior anterior quad-
rant of the orbital border, and passing forward to within 33 mm.
of the nostril. Anteriorly it lies between the nasal and the maxilla,
and posteriorly meets the jugal below the orbit. Superiorly it has a
long articulation with the prefrontal.

The prefrontal is a broader and stouter bone forming the superior
anterior quadrant of the orbital border. It meets the lachrymal,
nasal, frontal, and postfrontal.

The frontal is a comparatively narrow, elongate bone, 160 mm. in
its greatest length, completely shut out from the orbit by the
junction of the prefrontal and postfrontal. Together the f rentals
form one-half of the interorbital width. Posteriorly they meet the
parietal, being separated from each other for the last 35 mm. by
that bone.

The postfrontal is larger than the prefrontal. It forms at the
most but 12 mm. of the orbital border. It articulates with the
prefrontal, frontal, parietal, suprasquamosal, and poster bital.

The postorbital is considerably larger than the postfrontal, forming
15 mm. of the orbital border, and extending back for 130 mm., being
40 mm. broad in its widest part. It articulates with the jugal for
most of its length, with the squamosal, suprasquamosal, and post-
frontal. It differs from the corresponding bone in T. kanncmcyeri
in that there is no constriction behind the orbit. Indeed, from the
orbital border the bone rapidly widens.

The jugal is a large bone, forming part of the posterior border of
the orbit and extending from the lachrymal backwards for a length
of about 200 mm. It has a short vertical articulation with the
lachrymal, a long junction with the maxilla, and posteriorly lies
between the quadrato-jugal and the squamosal. On its upper
border it meets the postfrontal.

The parietal is smaller than the frontal, and lies in a marked
depression of the cranial surface. Its greatest length is 115 mm.,
and the circular pineal foramen lies between the bones 82 mm.
behind the anterior point of the elements.

The suprasquamosal is a large bone 100 mm. long and 55 mm.
broad, practically rectangular in shape. It articulates with the
parietal, postfrontal, postorbital, squamosal, tabulare, and post-
parietal.

The postparietal is placed behind the parietal, forming part of the
upper cranial surface and being bent at right angles to form the.

50

Annals of the South African Museum.

central half of the upper part of the occipital plate. Its lower
occipital border articulates apparently wholly with the exoccipital.

The tabulare forms the outer upper angle of the occiput. It
appears on the upper cranial surface, articulating with the squamosal,
suprasquamosal, and postparietal. It forms most of the upper
border of the lateral occipital foramen, and part of the outer and
lower border ; but owing to fracturing and splintering of the bones
its articulation with the paroccipital is difficult to ascertain exactly.

The squamosal is larger than the suprasquamosal. Above it articu-
lates with the postorbital, suprasquamosal, and tabulare, and below
with the jugal and quadrato-jugal.

The quadrato-jugal is a large bone forming the posterior angle of
the skull. Anteriorly it passes forward to the maxilla, articulating
with the jugal and squamosal. It is bent to form part of the back

FIG. 6. Trematosaurus sobeyi, Haughton. Diagram of section of skull just
anterior to the orbits, x .

surface of the angle, and overlaps the quadrate, which is also over-
lapped on its inner border by the squamosal and pterygoid.

Both the premaxilla and maxilla bear teeth of more or less uniform
size. Near the front of the snout the premaxilla carries two slightly
larger teeth, of which the first is 15 mm. long and recurved. There
are 10 other smaller teeth on the premaxilla. The maxillary teeth
are about 50 in number, borne in a row along the whole length of
the bone, the final teeth being slightly the smallest. The average
length of the larger teeth is about 10 mm., and they are arranged
fairly evenly along the bone, 10 teeth below the orbit occupying
.a space of 26 rnm.

The palatal view of the skull shows some interesting features.
The premaxilla extends back for a distance of about 50 mm. from
the front of the palate, and articulates behind with the prevomer.

The prevomer is a large bone, forming the inner and anterior walls
of the posterior nares, and most of the front of the palate. It articu-

South African Fossil Reptiles and Amphibia. 51

lates with the premaxilla in front, and with the palatine and para-
sphenoid behind. Arising from a depression in front of the posterior
nares is a large tusk, with a basal diameter of 20 mm. and a height
of about 33 mm. the largest tooth in the skull. Lying along the
wall of the posterior opening is a series of 8 smaller teeth, elliptical
in section, with the long axis of the ellipse lying at right angles to
the mid-plane of the palate. These teeth are separated from each
other by elliptical pits, now filled with calcite crystals. There is no
evidence of minute prevomerine teeth.

The palatine forms the hinder border of the posterior nares, and
articulates with the prevomer, the maxilla, and the transpalatine. It
carries three large tusks in an antero-posterior line, of which the
first is the largest.

The transpalatine lies along the inside of the maxilla, articulating
with the pterygoid behind and the palatine in front. It carries a
row of teeth of varying size.

The pterygoid is a large bone, diverging in three directions from
a central plate at the back. Interiorly it is loosely articulated with
the flat portion of the median bone which is presumably the para-
sphenoid ; posteriorly it passes upwards and backwards partially
to wrap round the quadrate, and meets the squamosal ; anteriorly
and externally it meets the transpalatine in a long articulation.
Behind this last articulation it is covered with a number of minute
pointed teeth almost to the plane of the back of the large vacuity.
It also has a short articulation externally with the maxilla.

The parasphenoid has the form of a large median thin plate at the
back of the palate, which passes forward to the prevomers. These
pass backwards on both sides of the lower part of the bone for some
distance as thin bones with long articulating inner surfaces. A
section across the skull between the orbits and the nostrils shows
the parasphenoid lying above the prevomers and sending down a
small process between them. A section across the parasphenoid at
the front of the parietals shows a vertically elongated oval with two
wing-like projections superiorly not passing to the bones of the
cranium. Between these projections and the mass of the bone I
can detect no sutures. Posteriorly the parasphenoid plate sends up
a short process to meet the downward projection of the exoccipital.

The occiput is broken away in part, and neither the basisphenoid
nor basioccipital can be detected. The exoccipital seems to have
played the most prominent part in the formation of the occipital
condyle.

(52)

2. On a Neiv Dinocephalian from the Gouph.
(Plate X.)

The specimen on which the following description of a new genus
and species is based is No. 2678 in the South African Museum collec-
tion, and was collected by the Rev. J. H. Whaits near Vivier Siding,
30 miles south-west of Beaufort West, from beds in the Pareiasaurus
zone. It consists of the almost perfect skull which has undergone
considerable weathering. The general shape and features of the
skull differ sufficiently from the other Dinocephalian genera to
warrant its inclusion in a new genus and species, and for it I
propose the name Struthiocephalus whaitsi, g. et sp. nov.

The chief generic characters are as follows : Skull large ; snout
relatively long and slender ; frontal and temporal regions not so much
elevated above snout as in Tapinocephalus ; eyes look forward and
outward ; heavy overhanging supraorbital crests ; temporal fossae
larger than orbits, elliptical in shape with shorter axis parallel to
axis of skull ; teeth weak, undifferentiated, and few in number.

The following table gives some of the chief measurements :

Greatest length 580 mm.

Greatest breadth 355 , ,

Greatest height 230 ,,

Length of snout to back of internal nares 200 ,,

Breadth of snout at back of internal nares 230 ,,

Length of orbit 95 ,,

Height of orbit 60 ,,

Length of temporal fossa 80 ,,

Height of temporal fossa 148 ,,

Interorbital width at front of orbit 180 ,,

Interorbital width at back of orbit 295 ,,

Intertemporal width 130 ,,

Basal length 515 ,,

The snout is slightly distorted from its true position, the angle of
distortion being about 15 to the right. The premaxilla bears three

South African Fossil Reptiles and Amphibia. 53

teeth, which are small and of the usual type. On the maxilla two
teeth can be discerned.

The nostrils are large, and placed well forward. The nasals are
long bones, and along their junction below the well-marked frontal
boss is a fairly deep median groove 56 mm. long.

It is difficult to be absolutely certain of the sutures of the top of
the skull on account of the peculiar weathering of the surface, but
their probable positions can be assigned.

The prefrontal and lachrymal are both large bones, coming well
forward from the orbital border on to the cheek. The prefrontal
forms most of the anterior border of the orbit.

The frontals form the upper border of the orbit, giving rise to the
very pronounced supraorbital crest. Near their anterior margin,
and just above the groove in the nasals, is a large median boss, the
presence or absence of which may probably be taken as a specific
character in this genus. The postfrontals, if present, are indis-
tinguishable.

The pineal foramen is large and situated just behind the
postorbital bar. It is entirely surrounded by the parietals.

The eye is large, looking forwards and outwards. The temporal
fossa is larger than the eye, higher than long, and looks outwards,
upwards, and slightly backwards. It is almost wholly bounded by
the postorbital and the squamosal, only a portion of its superior
border being formed by the parietal.

The postorbital bar is very massive, being strongest at the junction
with the jugal. The squamosal has a broad descending portion
which passes forward, articulating with the jugal and pushing the
quadrate forward to the level of the middle of the orbit.

The quadrate has a large articular surface for the lower jaw.
Posteriorly it forms a flat, plate-like bone which articulates with
the squamosal and the paroccipital. The quadrato-jugal is well
denned, but small. Its upper border is clasped by the jugal, its
lower border rests on the squamosal, while internally it articulates
with the quadrate. Apparently it forms no part of the articular
surface for the lower jaw.

The interparietal is 70 mm. high, and narrows rapidly below. It
articulates laterally with the tabulare, and inferiorly with the bone
which may be the supraoccipital, although on account of the
imperfection of the specimen no suture can be seen between that
bone and the basioccipital. The element in the lateral part of the
occiput, which was mentioned in the description of Scymnognathus
tigriceps as occurring in Dinocephalians, is here well seen, and

54 Annals of the South African Museum.

is the tabulare. It is a large membrane bone lying behind the
squamosal almost for the whole length of the posterior border of the
temporal fossa, being thickest at its lowest end. It forms the larger
part of the occipital plate lying above the condyle.

The squamosal forms but little of the occipital plate, the par-
occipital extending almost to the border. In other Dinocephalians
the squamosal enters largely into the composition of the occipital
aspect.

The condyle is large, single and rounded.

The palatal view is similar to that of Tapinocephalus. The basi-
occipital is a large bone. The pterygoids are separated by a long
median groove. The stapes articulates with the basioccipital and
the pterygoid.

(55)

3. On Two New Tlierocephalians from the Gouph.

TROCHOSAURUS INTERMEDIUS, g. et sp. nov.

This new genus and species is founded upon a weathered and
crushed skull and lower jaw found on the farm Abraham's Kraal,
in the Prince Albert Division, in beds belonging to the Pareiasaurus
zone. The parietal region is incomplete and the occipital plate is
missing, while the zygomatic arches are very weathered. Fracturing
of the skull has displayed the structure of the palate.

The right premaxilla bears five incisors, of which only the fifth
retains the crown. They are subequal in size, and together occupy
a space of 48 mm. at the margin of the bone. The 5th, which is
slightly smaller than the others, has a diameter of 6'5 mm. at the
gums and is 15 mm. long, curving slightly backwards. The incisors
were simple, pointed teeth, apparently without serrations or longi-
tudinal grooves. There is a diastema of 7 mm. between the fifth
incisor and the first canine. There are two canines, both of which
were large and functional, the second being slightly larger than the
first. In section the teeth were apparently oval. A section across
the snout shows a smaller replacing tooth lying internal to the first
canine and parallel to it. Close behind the second canine came the
small molars, of which there do not seem to have been more than
four.

The nostrils are large and are placed almost terminally. Almost
the whole of the lower border is formed by the large septomaxillary,
which sends off a turbinal process partially dividing the nares into
two portions. Between the septomaxilla and maxilla is the usual
outer foramen.

The nasal is fairly broad and comparatively short.

The interorbital width is almost twice that of the intertemporal
region. The f rentals are large, forming 9 mm. of the upper border
of the orbit.

The limits of the prefrontal and lachrymal are not wholly
determinable, but the prefrontal was certainly large.

There is a distinct though small postfrontal, which forms no part

56 Annals of the South African Museum.

of the border of the temporal opening. The pineal foramen is large,
18 mm. behind the postorbital bar, and entirely surrounded by the
parietals.

The postorbital is rather weak, and forms the whole of the
anterior and more than half the upper border of the temporal fossa.

The palate is typically Therocephalian in structure. The pre-
vomers are narrow bones passing some distance backward behind
the internal nares, and forming the inner border of those openings.
Their sutures with the pterygoid are indeterminable.

The palatine is a plate-like bone forming a small part of the
posterior border of the internal nares, and having a long
articulation with the maxilla.

The pterygoid is large. The transpalatine is large, and articulates
with the pterygoid, maxilla, and palatine. Between it and the
pterygoid there is a very well-defined foramen.

The symphysis of the lower jaw is weak. The dentary is a strong
bone, forming about two-thirds of the jaw. It possesses a pro-
nounced mentuni. The splenial extends back as far as the plane
of the front of the orbit, lying along the inside of the dentary. It
is a thin bone. The coronoid is small. The angular lies posterior to
the dentary, wedged in between that bone and the splenial. Between
it and the dentary is a small foramen. The surangular is large, but
is more than half-hidden from view by the angular.

The chief measurements of the skull are :

Greatest length 230 rnm.

Distance from snout to front of orbit 115 ,,

Diameter of orbit 42 ,,

Interorbital width 46 ,,

Intertemporal width 28 ,,

In the possession of two large canines and five incisors this form
resembles Lycosuchus and Trochosuchus. It lies nearest to the
former genus, being distinguished from it by the shallow incisor
region of the upper jaw, by the much deeper dentary characters
in which it agrees with Trochosuchus and by the greater width
of the intertemporal region. From Trochosuchus it is distinguished
by the fact that the two canines are approximately equal in size,
whereas in the older genus the first canine is of the size of the
incisors, and the second canine is considerably larger. This form,
therefore, seems to stand in an intermediate position between
Lycosuchus and Trochosuchus.

The type is in the South African Museum (Cat. No. 2756).

Soiith African Fossil Reptiles and Amphibia. 57

TlTANOSUCHUS DUBIUS, Sp. nOV.

The portion of mandible on which this species is founded was
obtained from the farm Abraham's Kraal in the Prince Albert
Division. It is the right ramus, showing four incisors, one canine,
and eleven small molars. It differs from T. cloetei in having a
more massive symphysis, in being even squarer in the front of the
jaw, in the much larger canine and smaller molars differences which
appear to warrant the erection of a new species.

A table showing the teeth measurements in the three known
species of Titanosuchus will emphasize their differences :

T.ferox. T. cloetei. T. dubius.

First incisor Imperfect 21 x 14 Imperfect

Interval between i.l and i.2 10 10 9

Second incisor 21 x 15 18 x 13'5 19 x 14'5

Interval between z.2 and i.3 10 11 4

Third incisor 20 x 12 20 x 14 20 x 15

Interval between i.3 and z.4 18 8 6

Fourth incisor 23x17 18x14 16-5x13

Interval between iA and c ? 9 5

Canine 50x35 30x26 47x20

Four molars occupy 40 30

Eleven molars occupy 107

[All these measurements are in millimetres.]

It will be seen that the incisors are set much closer together in
this new form, although on the whole their size does not differ
appreciably from those of T. cloetei. The canine is narrower and
longer. In fact, whereas in T, cloetei it approximates in section to a
circle, in the new species it is almost oblong, with one side more than
twice the length of the other.

The molars are small and circular in section. A line drawn along
the inner side of the molars just touches the canine, and is also
tangential to the inner surface of the 4th incisor.

If a line be drawn across the front of the jaw at right angles to
the symphysis, the back of the canine lies 76 mm. from it, and the
back of the 4th molar 111 mm. ; whereas in T. cloetei the figures for
the same measurements are 67 mm. and 116 mm. respectively.

(58)

4. On Some Neiu Anomodonts.
(Plate XI.)

DlCYNODON MUSTONIS, Sp. n.

This species is founded on an almost perfect small skull and lower
jaw collected by me in 1913 at Dunedin, Beaufort West. The only
parts lacking are the posterior extension of the right squamosal, the
right postorbital arch, and the left articular end of the lower jaw.
Most of the matrix has been cleared away, and the sutures between
the bones are for the most part beautifully displayed.

In general shape the specimen approximates most nearly to
Diictodon Kolbci, although it is but half the size ; but it is generi-
cally distinct in that the small preparietal does not entirely surround
the pineal foramen, but forms only its anterior border. The anterior
half is roughly of the shape of an equilateral triangle, the posterior
half almost square. The skull is flat and of graceful proportions.
The antorbital portion is short, the orbits large and directed mostly
upwards, lying wholly in the anterior half of the skull.

The premaxillary is large, the internasal process passing back
almost to the frontals. The maxilla is comparatively small, although
the caniniform process descends considerably below the level of the
beak and zygomatic arch. The nostril is large, and is overhung by a
prominent rugose nasal boss. There appears to be a small septo-
maxillary at the back of the nostril, which does not, however, form
any part of the cheek.

The anterior frontal region is flat, but posteriorly the region
between the postorbital arches and extending between the post-
orbitals is markedly concave. The frontals are long, passing back
almost to the pineal foramen, each separating the two anterior
processes of the parietal. There is a marked supraorbital ridge
which passes forward into the prefrontal, the ridge becoming less
pronounced towards the lachrymal.

The postfrontals are large, elongate, triangular bones extending
back between the parietals and postorbitals to the level of the pineal
foramen. The postorbital arch is comparatively slender, and in its

South African Fossil Reptiles and Amphibia. 59

outer half the postorbital is supported by the jugal, which also
extends along two-thirds of the inner side of the zygoma.

The pineal foramen is situated some distance behind the post-
orbital arch. It is 6 mm. long, and its anterior border is formed by
the narrow, elongate preparietal. The parietal sends a process
forward between the frontal and preparietal to the level of the post-
orbital arch, and extends back for about half the length of the
temporal fossa. The postorbital forms almost the whole of the
upper border of the fossa.

The squamosal is large, and has a large boss on its expanded
portion. It extends back for some considerable distance behind the
occipital plate, which slopes well forward in its upper half.

The interparietal comes well on the top of the skull and is greatly
developed laterally. The quadrato-jugal is large, and the quadrate
well displayed.

The lower jaw has a very deep mentum, the lowest point coming
a little behind the plane of the caniniform process, i.e. just below the
front of the orbit.

The following are the chief measurements of the skull :

Greatest length 150 mm.

Maximum width 112 ,,

Maximum width of parietal region 24 , ,

Maximum width of frontal region 19 ,,

Maximum width of nasal overhang 29 ,,

Minimum width across pterygoids 15 , ,

Length of pineal foramen 6 , ,

Width of pineal foramen 3

Distance of pineal foramen behind postorbital

bar 10 ,,

Basal length 126 ,,

Width between inside of caniniform processes... 24 ,,

Maximum depth of lower jaw 33 ,,

I have much pleasure in naming this species after Mr. J. A. Musto,
the former owner of Dunedin, from which place so many interesting
and varied types have been obtained.

Type. Female skull. (S.A.M. Cat. No. 2674.)

DlCYNODON BREVICEPS, Sp. n.

This new species is founded upon a somewhat imperfect and
weathered skull and lower jaw (S.A.M. Cat. No. 2366) from the

60 Annals of the South African Museum.

farm Voetpad in the Division of Murray sburg, Cape. The chief
features lie in the shortness of the skull compared with the width,
the great height, the size and shape of the orbit and of the temporal
fossa. Although all the features are not visible, enough is seen to
show that the skull differs considerably from any hitherto-described
species.

The greatest length is 170 mm., while the maximum width across
the squamosals was probably 180 mm. The front of the snout is
weathered away, but was nearly vertical and very high, the nasal
region being very convex and the nostrils almost at the front of the
skull. The preorbital portion is short.

The premaxilla is small. The nasals are narrowed anteriorly the
nostrils being close together. There is a well-marked septo-maxillary
forming the posterior wall of the nostril and a small portion of the
cheek-surface.

The eyes are large, rhomboidal in shape, looking mainly outwards.
The frontal region widens considerably posteriorly, the minimum
width 25 mm. being well towards the front of the orbit. The
frontal region is flat and there is a slight supraorbital ridge.

The postorbital portion is not complete. The bar was apparently
comparatively slender, while the postorbital extended back to form
the whole of the upper border and part of the posterior border of
the temporal fossa. The parietal region has a maximum width of
32 mm., and is very short.

There is a long and slender preparietal, supported along the pos-
terior part by the parietals, and in its anterior half by the frontals.
It extends only so far back as just to touch the anterior border of the
pineal foramen, which lies 10 mrn. behind the postorbital bar, and is
11 mm. long and 6 mm. wide.

The postfrontal is well seen on the right side. It is well
developed, very narrow in its posterior half and articulates with
the frontal, parietal, and postorbital.

The squamosal has a very long downward process supporting a
large, plate-like quadrato-jugal. Anteriorly it is overlapped by the
jugal, which is a very stout bone and extends half-way along the
lower border of the temporal opening.

The palate is short and wide, the width between the bases of the
caniniform processes being about 40 mm. There is no trace of

tusks.

The lower jaw is displaced from its true position. Its total length
is probably 110 mm., and the depth at the symphysis 40 mm. The
front of the jaw is bluntly rounded and strong, and there is no

South African Fossil Reptiles and Amphibia. 61

upwardly projecting beak and no pronounced mentuna descending
below the level of tbe bottom of the jaw.

LYSTROSAURUS OVICEPS, sp. n.

There is in the collection of the South African Museum the skull
and lower jaw (No. 641) of a small specimen of Lystrosaurus which
can be correlated with no hitherto-described species, and which is
therefore described here as a new species. It is said to have come
from the " Tarka Eiver, Cradock District," and is in a fine-grained
grey sandstone.

The greatest length of the skull when viewed from above is
163 mm. The greatest breadth is 129 mm. The minimum
interorbital width is 46 mm. and the minimum intertemporal
width 25 mm. The orbit is elliptical with the long axis parallel
to the nasal plane ; its length is 36 mm., its height 33 mm.

The snout is slightly convex with a median ridge, which is very
prominent in the middle of the premaxilla, but dies away at the
extremities of that bone. The premaxilla passes up as a narrow
prolongation between the nasals. Its greatest length is 76 mm., and
its width at the mouth measured round the bone is 56 mm.

The maxilla is a rectangular bone, with a suborbital prolongation
supporting the jugal. The tusks are circular in section, and directed
slightly inwards. Their diameter at the base is 9 mm. ; length
probably 25 mm. The distance between the inner sides of their
bases is 46 mm.

The septomaxillary forms a distinct part of the cheek, besides
being the posterior wall of the nostril.

The fronto-parietal plane makes an angle of about 120 with the
general plane of the snout, but the change of slope is not very
abrupt owing to the convexity of the snout. The transverse ridge
across the nasals is not nearly so pronounced as in the other species.
The prefrontal forms the superior anterior quadrant of the orbital
border, but the supraorbital boss is very slight.

The frontal forms the remainder of the upper border of the orbit,
and at the frontal junction is a slight median ridge. In each bone is
a slight but well-marked central boss from which lines of sculpture
radiate. These bosses are further from the median ridge than the
corresponding protuberances in L. declivis.

The preparietal is large, circular, excavated, concave, and has a
maximum length of 19 mm. It forms the border of the anterior
third of the pineal foramen, which is of the keyhole shape common

62 Annals of the South African Museum.

to species of Lystrosaurus and 8 rnm. in length. The boundaries of
the postfrontal are not seen, but it must have been a small bone.

The parietal region is wide and flat. The parietals are small, the
postorbitals forming the whole of the upper borders of the temporal
fossae. These are transversely oblong in shape, 39 mm. broad and
25 mm. long.

The squamosal has a pronounced ear-shaped expanded portion at
the outer border of the temporal opening, the projection being more
prominent than in L. latirostris. The zygomatic portion of the
squamosal is thin and flat. There is a large quadrato-jugal which
apparently forms part of the articular surface for the lower jaw.

The development of the posterior part of the palatal surface shows
one or two interesting features. The pteiygoid is of the usual
Lystrosaurus type, having a long articulation w r ith the basisphenoid,
and an outward and backward process which extends to the stapes.
The basisphenoid is bent at right angles to the plane of the palate,
and is pierced by the two carotid foramina.

Hitherto in Lystrosaurus the whole of the lateral bone extending
from the condyle to the squamosal has been called exoccipital ; but
this specimen shows that the so-called " exoccipital " is in reality
made up of two bones, the true exoccipital and the paroccipital
(opisthotic). The exoccipital is a small bone closely united to the
paroccipital, forming part of the condyle and of the wall of the
foramen magnum. The paroccipital is a large bone stretching from
the squamosal to the small exoccipital and having an inner down-
ward projection articulating with the basioccipital. It is pierced
near the exoccipital suture by the opening for the IXth-XIIth
nerves.

The depth of the lower jaw at the symphysis is 34 mm. The
lowest point of the mentum is directly below the point of the tusk.

This new species comes nearest to Lystrosaurus latirostris, from
which it differs, however, in the convexity of the snout, in the angle
between the fronto-parietal and premaxillary planes, in the ratio
between the interorbital and interternporal widths, in the shape of
the orbit, and in other minor features. When viewed posteriorly
the skull is seen to differ from both L. declwis and L. latirostris.
The parietal region is not grooved so deeply as in the type species,
and the zygomatic portion of the squamosal is much wider in
L. latirostris.

(63)

EXPLANATION OF PLATES.

PLATE VIII.

1. TREMATOSAURUS SOBEYI, sp. n.

PI6.

1. Top view of skull, x ^ nearly.

2. Diagram showing arrangement of bones of top of skull, x J nearly.

PLATE IX.

2. TREMATOSAURUS SOBEYI, sp. n.

1. Palatal view of skull. c J nearly.

2. Diagram showing arrangement of bones and teeth of palate. c J nearly.

PLATE X.

STRUTHIOCEPHALUS WHAITSI, g. et sp. n.

1. Side view of skull. : |.

2. Palatal view of skull. X {

3. Top view of skull. X 5.

PLATE XI.

1. Top view of skull of Dicynodon mmtonis, sp. n. x J.

2. Diagram showing arrangement of bones round pineal foramen in Dicynodon

breviceps, sp. n. x 1.

3. Top view of skull of Lystrosaurus oviceps, sp. n. x T 7 ^.

4. Side view of skull of Lystrosaurus oviceps, sp. n. x .

EXPLANATION OF LETTEEING.

P., Frontal; I.F., Interfrontal ; J., Jugal ; L., Lachrymal; MX., Maxilla; N.,
Nasal; P., Parietal ; Pal., Palatine ; Pa.S., Parasphenoid ; P. for., Pineal foramen ;
Pmx., Premaxilla; Po.F., Postfrontal; Po.O., Postorbital ; Po.P., Postparietal ;
Pp., Preparietal ; Pr.F., Prefrontal ; Pt., Pterygoid ; P.Vo. , Prevomer ; Q.,
Quadrate; Q.J., Quadratojugal ; Smx., Septomaxilla ; Sq., Squamosal ; S.Sq.,
Suprasquamosal ; Tab., Tabulare ; T.P., Transpalatine.

nn

. S. Afr. Mus. Vol. XII.

Plate VIII.

\

TREMATOSAURUS SOBEYI, HAUGHTON. x NEARLY.

Ill

. S. Afr. Mus. Vol. XII

Plate IX.

TREMATOSAURUS SOBEYI, HAUGHTON. x 1 NEARLY.

!. S. Afr. Mus. Vol. XII.

Plate X.

STRUTHIOCEPHALUS WHAITSI, HAUGHTON. i-

Ann. S. Afr. Mus. Vol. XII.

Plate. XI.

1. DICYNODON MUSTONIS, HAUGHTON.

2. DIGYNODON BREVICEPS, HAUGHTON. 1.
3,4. LYSTROSAUBl'S OVICEPS, HAUGHTON.

(65)

11. Investigations in South African Fossil Reptilia and Amphibia
(Parts 5-9). By S. H. HAUGHTON, B.A., F.G.S., Assistant
Director.

5. On the Genus Khinesuchus, Broom, with Notes on the described

Species.

THE genus Hhinesuchus was first described by Broom in the Annals
of the South African Museum (vol. iv., p. 376, 1908) from some frag-
mentary remains of the skull of a presumably temnospondylous
Labyrinthodont found near Prince Albert, Cape Colony, to which
the specific name R. whaitsi was given. The genus was said then
to include also the species named by Lydekker Eryops africanus and
the European species called Macromerion Giimbcli by von Ammon.
At first the genus was thought to have no affinities with Eryops, but
later (1910, Bull. Amer. Mus. Nat. Hist., xxviii.) Broom modified
this view, concluding that the two genera were apparently closely
allied. In 1911 van Hoepen (Ann. Transvaal Museum, iii., 2)
described part of a skull of a large Labyrinthodont from Senekal,
O.F.S., as Myriodon senekalensis, while in 1912 Broom described
part of another skull from the same place as Rhinesuchus major.
Recently, thanks to the courtesy of the authorities of the Bloem-
fontein Museum, I was enabled to examine the beautiful Labyrintho-
dont skeleton which they obtained from Senekal. In addition to
that specimen I have also examined a skull from Beaufort West,
a crushed skull from the Nieuweveld (both collected by the Rev. J. H.
Whaits), and a portion of lower jaw from Senekal. These three
latter specimens are in the collection of the South African Museum.

The original description of the genus Ehinesuchus as given by
Broom is as follows :

" Skull probably like that of Capitosaiirus in general shape ;
maxillaries and dentaries each bearing a row of uniform teeth ;
inside of maxillary teeth are large numbers of very small teeth,
probably borne by palatines, and covering much of the pterygoids
and continued across the back part of the parasphenoid : similar
small teeth are found inside of the teeth borne by the dentary ; they

6

66 Annals of tlie South African Museum.

are probably on the splenial bone ; the median anterior part of the
parasphenoid is narrow."

The following is a translation (the original paper is in Dutch) of
van Hoepen's description of the genus Myriodon :

" Skull triangular, rounded in front, almost as long as broad.
Maxilla and dentary bear each a row of large, conical, pointed,
pleurodont, flattened teeth, becoming smaller posteriorly, probably
labyrinthodont in section. Within the row of maxillary teeth,
probably on the palatine, is a row of large teeth, similar to the
maxillary and dentary teeth. On the remaining bones of the palate
are large numbers of small teeth, of which the biggest is far smaller
than the large teeth already mentioned. Similar small teeth are
found on a ridge which runs along the row of teeth on the dentary."
Van Hoepen further says that his genus differs from Rliinesuclius
in that it has but one row of teeth on the palatine, and lacks the
small teeth on this bone.

The fragment of skull upon which Broom based his description of
the palatine bone is in a crushed condition, and it is probable that
these supposed palatine teeth are on the prevomer. In the complete
skull of R. whaitsi which I describe in this paper there is a row of
medium-sized teeth lying on the inner border of the internal nares
which are almost certainly carried by the prevomer. I am unable
to find in the type fragments any justification for the statement,
" Inside of maxillary teeth are large numbers of very small teeth,
probably borne by palatines." The only fragments which show small
teeth are portions of the lower jaw, which bear the usual coronoidal
teeth lying internal to the larger teeth of the dentary.

For these reasons the two genera must be grouped together under
the one generic name Rhinesuckus, Broom for which the following
short diagnosis may stand :

" Medium-sized to large ternnospondylous Labyrinthodonts.
Skull triangular, rounded in front, median length slightly greater
than breadth. Eyes wholly in posterior half of skull. Otic notch
present. Bones of skull-roof complete. Maxillary and dentary
each carrying row of uniform teeth, slightly decreasing in size
posteriorly. Prevomer carrying one or more large tusks, a few
medium-sized teeth, and covered together with the major parts
of the parasphenoid and the pterygoid with minute denticles.
Palatine with row of teeth similar to those on maxilla. Small
transpalatiue with teeth present. Coronoid carries number of
denticles on upper surface."

Besides having the eyes larger and set more nearly together, and

Investigations in South African Fossil Rcptilia and Amphibia. G7

in being more pointed, the skull of Ehinesuchus differs from that of
the American genus Eryops in lacking the interfrontal bone. I have
searched carefully for evidence of this bone in all the species, but
can find none. Moreover, although the lower jaws I have examined
are not in the best condition for showing sutures, I am not able to
find any tripartite division in the coronoid bone.

EHINESUCHUS WHAITSI, Broom.

(Plate XII., figs. 3, 4. Text-fig. 7.)

1908. Broom, Ann. S. African Mus., iv., 8, p. 373; pi. xlvi., fig. 3.

An almost complete skull and lower jaw (S.A.M. Cat. No. 3009)
collected by the Eev. J. H. "Whaits at Beaufort West (Endothiodon
zone of Beaufort Beds) enables me to give more details concerning
this species.

In its general outline the skull has a shape like that of Eryops
megaccphalus and Rhinesuchus semkalensis. The upper surface of
the skull and the outer surface of the lower jaw are covered with
a pitted sculpturing. In the snout and mid-regions of the skull the
pits are roughly circular; in the jugal, quadratojugal, and squamosal
regions the pits are more elongated. The orbits are wholly in the
posterior half of the skull and are comparatively more nearly set
together than in Eryops. The articular region extends well behind
the occiput. The following are some of the principal measurements
of the skull :

Maximum length 312 mm.

Length on median line 265 ,,

Maximum width 258 ,,

Interorbital width 40 ,,

Internasal width 41 ,,

Length from back of nostril to front of orbit 125 ,,

Length from plane of snout to front of orbit 158 ,,

Length of orbit 33 ,,

Width of orbit 34

Comparison with the type specimen of R. whaitsi shows that the
skull is flattened, but the width and general size of the parasphenoid
and the pterygoids, as far as they can be compared, leave no doubt that
this skull belongs to the species already described. Unfortunately
the sutures on the top of the skull are not visible, but the structure
of the palate can be made out fairly well.

The parasphenoid has a flat posterior portion similar to that

68 Annals of the SoutJi African Museum.

figured by Broom in his description of this species, and passes
forward between the large vacuities to meet the prevomer. As in
the type the anterior part of the plate bears a number of tiny
denticles. Posteriorly the bone covers a large part of the basi-
occipital. The lateral articulations with the pterygoids are of the
type seen in Eryops.

The pterygoid forms most of the posterior and outer walls of the
large vacuity. Behind the parasphenoid the pterygoid passes back-
wards and downwards, articulating with the squamosal and overlying
the inner half of the quadrate. Laterally it passes forward outside
the large vacuity to meet the prevomer, articulating on its outer
border with the palatine. From the posterior corner of this bone,
just behind the level of the parasphenoid, the surface is covered with
small denticles save for a narrow band on the inner side along the
wall of the vacuity. This non-dentigerous band is about 50 mm.
long, and at its widest at the posterior end has a breadth of
13 mm. In the type specimen there is also an outer non den-
tigerous band. As the bone narrows anteriorly the denticles tend
to become arranged in well-defined rows.

The palatine lies between the maxilla on the outside and the
pterygoid and prevomer internally. It bears one row of large
teeth, of which the first two are larger and more tusk-like and
the remainder small and flattened, so that the breadth is greater
than the length. There is a slight decrease in size of the teeth
posteriorly. The few that are preserved entire show that the teeth
were simple, pointed, and conical, with very slight longitudinal
grooves. Just behind the internal nares, internal to the row already
described, and possibly on the palatine, is a secondary row of three
or four similar teeth.

I am unable to say definitely that a transpalatine is present ; but
behind and outside the palatine is a small mass of bone slightly
below the level of the palatine bearing three or four medium- sized
teeth. This may be the transpalatine.

The prevomer is large. The pair of bones lie between the posterior
nares forming most of the front of the palate. On the outside of the
large vacuity they pass back to meet the pterygoids, and between
the openings they send back a long process to meet the parasphenoid.
In front of the posterior nares are two large tusks. Between the
hinder pair of these is a transverse row of smaller teeth, each pre-
vomer bearing three. Between this row and the posterior vacuities
is a very well-defined dentigerous area coered with denticles, which
area passes backwards almost, if not quite, to the pterygoids.

Investigations in South African Fossil Kept ilia and Amphibia.. 69

The premaxilla carries teeth along its border, as does the maxilla.

The quadrate is a triangular bone lying between the pterygoid and
the quadrato-jugal and squamosal. It appears mostly on the back
of the skull, the apex of the bone lying 32 mm. above and some
distance internal to the articular surface. This latter is weakly
concave, the concavity being directed backwards and outwards
from the anterior inner angle of the bone.

Passing outwards and backwards from the posterior lateral curved
border of the parasphenoid is a small, thin, apparently plate-like
bone. This may be the stapes.

FIG. 7.

Sections across left ramus of lower jaw of RhinesucJius
a. 85 mm., behind point of jaw.
fi. 185 mm., behind point of jaw.
y. 220 mm., behind point of jaw.

D, dentary; C, coronoid ; Sp, splenial ; A, angular; P. Art, Prearticular ;
S.A., surangular.

The whole of the left side of the lower jaw is preserved, together
with the articular region of the right. The length of the jaw is
about 322 mm.

The dentary carries a row of labyrinthodont teeth. It has a
narrow posterior elongation extending towards the articular surface
for some distance behind the last tooth.

Lying along the inside of the dentary and extending almost from
the front of the jaw to the anterior border of the supra-Meckeliaii
fossa is the " coronoid," which throughout nearly the whole of
its length bears a large number of small teeth. In the jaws of
Trimerorachis and Eryops, as described by Broom and by Williston,
this "coronoid" consists of three elements the coronoid proper,

70 Annals of the South African Museum.

the intercoronoid, and the precoronoid. Unfortunately the bones
of the lower jaw in this specimen are somewhat crushed ; but,
except for a possible line of suture between a precoronoidal portion
and the remainder of the bone, I am unable to distinguish the three
elements each from the other. Moreover, the whole series of coro-
noidal teeth is evenly distributed ; no portion of the upper surface
of the bone bears an augmented share, and none is without teeth,
whereas in the figure given by Williston of the jaw of Trimerorachis
alleni the " coronoidal " teeth are grouped in three well-defined areas,
the edges of the upper surface of each of the three elements being
non-dentigerous.

The lower jaw is of much more slender build than that of Eryops
or of Trimerorachis, and the supra-Meckelian fossa considerably
smaller in proportion. Viewed from within, the posterior flattened
expansion of the prearticular completely hides the fossa and sur-
angular from view.

A fragment of the right jaw shows what is apparently a distinct
suture between the splenial and the postsplenial, the latter of
which is the preangular of Broom. This suture occurs below the
front margin of the small anterior fossa.

EHINESUCHUS SENEKALENSIS (v. Hoepen).
(Plate XII., figs. 1, 2.)

1911. Myriodon senekalensis, van Hoepen. Ann. Transvaal Mus.
iii., 2, p. 103 ; pis. 1, 2.

1912. llliinesuchus major, Broom. Trans. Geol. Soc. S. Africa, xiv.,
p. 79 ; pi. xiii., 1-2.

The types of van Hoepen and Broom were obtained at Senekal.
From the same place came the almost perfect specimen now in the
Bloemfontein Museum, which I was able to examine by permission
of the authorities of that Museum, and of which a cast was made for
the South African Museum. In addition there is in the South
African Museum a portion of the left mandible and the left maxilla
of the same species, also from Senekal, presented by the late H.
Kynaston, Esq.

The following description of the species is based mainly on the
Bloemfontein specimen. This shows the whole of the upper side of
the animal crushed flat on a slab of sandstone, together with some
of the ventral armour. The front legs lie bent backwards along the
sides of the body, while the hind legs are spread out at right angles
to the trunk. Unfortunately the sutures of the skull can scarcely

Investigations in South African Fossil Eeptilia and Amphibia. 71

be made out among the sculpturing of the surface, and in this
connection I have made use of the facts given by Dr. Broom. The
specimen described by him seems to be a smaller representative
of the species than the Bloemfontein one ; but from the general
similarity and from the fact that all the known examples come from
the same locality, I think there can be no doubt as to their specific
identity. Compainson of the photograph of the Bloemfontein
specimen with the restoration of the skull given by Broom shows
that in the latter the snout is somewhat too long, the nostrils being
nearer the front of the skull.

As preserved, the animal measures nearly 7 feet in length. There
are only five caudal vertebrae showing, so that the length may have
been nearly 8 feet in life, supposing that the tail was short as
in Eryops,

The skull is moderately long with a rounded snout and practically
straight sides, flat, and broadest at the back. It has the pitting
characteristic of this genus and Eryops, the pits being roughly
circular in the middle of the bones and lengthening out towards
the edges. The otic notch is not large. The nasal-frontal region
is furnished with a median groove, and on either side is a bow-shaped
channel running from the plane of the back of the nostrils to that of
the front of the orbits, concave towards the median line.

The greatest length of the skull is 580 mm., the greatest breadth
444 mm. The nostrils lie 33 mm. from the front of the snout, and
the orbits 308 mm. There is a small pineal foramen 407 mm.
behind the snout. The internasal width is 53 mm., the interorbital
width 57 mm. (Broom's specimen has an interorbital width of
50 mm.). From the orbit to the nostril the distance is 235 mm.
(Broom's specimen gives 203 mm.).

No sutures can be seen in the preorbital region save parts of the
boundary of the prefrontal. This is a very large bone. Broom
says : " The frontal bones are long and narrow, and separated from
the orbits by the forward extension of the postfrontals, agreeing
in this with Archegosaurus. The prefrontal is very large, stretching
more than half-way to the nostril. The nasal bone is also very
large. The lachrymal is narrow and placed w r ell forward. The
jugal is large and forms about a quarter of the side of the head. It
only forms a small part of the orbital margin. Behind it is the large
prosquamosal, or, as it is more often called, the supratemporal. The
postorbital comes well down behind the orbit."

In the Bloemfontein specimen it is possible to delimit most of
the posterior cranial bones, partly from the sutures and partly from

72 Annals of the South African Museum.

the sculpturing. The quadrato-jugal forms the outer posterior angle
of the skull, articulating with the squamosal, jugal, and probably
with the maxilla. The squamosal is a fairly large pentagonal bone
forming the outer boundary of the auditory notch, having a well-
defined junction with the suprasquamosal, and probably meeting the
postorbital in front. The suprasquamosal forms a small part of
the boundary of the auditory notch, the inner wall of which is formed
by the tabulare. The pineal foramen lies wholly in the parietals,
which are rectangular bones.

The postorbital apparently extends further out from the orbit
than in Eryops. The postfrontal is small, and the frontal is
completely excluded from the orbital border. The jugal forms
about 30 mm. of the outer border of the orbit, and is the largest
bone on the surface of the skull. The postparietal has an almost
square cranial portion, and is larger than the corresponding bone
in Eryops. The superficial cranial bones completely hide the
exoccipital condyles in a view from above.

Nothing can be seen of the palate. Several teeth are seen along
the edge of the maxilla, each about 15 mm. long. They are simple,
pointed, unserrated.

Vertebral column. There are 29 vertebrae between the occiput
and the sacrum. Most of the dorsal spines are missing, and the
ventral side of the column has not been exposed. The left-hand
sides of most of the intercentra are displayed, and all the neural
arches. As in Eryops, the neural arch of the first vertebra consists
of two halves, the anterior portions of which articulate with the
exoccipital condyles, and the upper portions of which pass backwards
along the sides of the dorsal spine of the axis. Each half of the
atlantal spine has an articular width of 32 mm. and is 54 mm. long.

From the axis to the sacrum the neural arches are approximately
similar. There are well-marked transverse processes for the articu-
lation of the ribs. From the 7th vertebra to the sacrum the width
across the transverse processes gradually decreases. The following
gives the measurements of two of the best-preserved vertebrae :

13th vertebra. 22nd vertebra.

Length of arch 45mm. 52 mm.

Length of dorsal spine 36 ,, 27 ,,

Height of spine above transverse process 31 ,, 29 ,,

\Vidthbetweenextremitiesofprocesses... 88 ,, 61 ,,

All the vertebrae carry ribs. The ribs from the 2nd to the 8th
have their distal thirds strongly expanded with a well-defined

Investigation* in South African Fossil Reptilia and Amphibia. 73

superior process from the expanded portion. The ventral portion
of the expanded end is bent inwards and lies below the next
succeeding rib.

Shoulder girdle. Only the upper end of the scapula on each side
is seen. The bone is a thin flat blade with an antero-posterior
length at its extremity of 75 mm., and with an average thickness of
not more than 8 mm. The upper border is approximately straight
and posteriorly it is produced, so that the posterior angle is acute
while the anterior angle is obtuse.

The cleithrum differs in shape from that of Eryops. It has a
length of at least 210 mm., and is closely applied to the anterior
border of the scapular blade and to the outside of the anterior end
of the upper border. For at least half of its length it is overlain by
the clavicle. It is a strong bone, spatulate at its distal end when
viewed from above, and not possessing the fan-shaped expansion
seen in Eryops. It extends back beyond the scapula, not in contact
with most of the upper border of that bone, and it almost covers the
scapula when viewed from above.

The upper part of the clavicle lies in front of and upon the
cleithrum. It is expanded and somewhat spoon-shaped when seen
from above, with an inner pointed distal angle. Its maximum width
is 46 mm. Proximally it is curved inwards to meet its neighbour
beneath the interclavicle, which is not seen ; the proximal portion
of the bone is more rounded in cross-section than the distal
expanded portion.

Fore-limb. Of the humeri only the distal ends have been dis-
played, and they have been mutilated to a certain extent, so that it
is not possible to give any details of their form.

The radius and ulna of both fore-limbs are lying in position. The
radius is a single shaft with slightly expanded ends, 80 mm. long,
with a minimum width of shaft of 16 mm. The proximal end has a
width of 27 mm., and is hollowed for articulation with the humerus.
Tbe distal end has a width of 2-i mm., and has two articulating
faces. The inner one is comparatively short, while the ulnar surface
is much longer and nearly flat.

The ulna has a form similar to that figured by Case for
Eryops. The olecranon process is very blunt. The radial border
of the bone is concave, the outer border straight from the head
to the beginning of the distal expansion. The length of the
ulna is 82 mm. The outer angle of the distal end is bluntly
rounded for articulation with the pisiforme, while the main surface
is truncate.

74 Annals of the South African Museum.

The carpus of Eryops has been discussed by Cope, Emery, Broom,
and Case. In the specimen under description only two elements of
the carpus remain the "carpus" seen on the right limb being
merely lumps of matrix that have been painted black and photo-
graphed before their true nature was noticed. There were un-
doubtedly four elements in the proximal row. The element
remaining, the intermedium, articulated with the lower part of
the ulnar surface of the radius. It is a lozenge-shaped bone
20 mm. by 12 mm. in extent. Internal to it was the radiale.
Articulating with the intermedium, radius, and ulna was the
ulnare to follow Broom's interpretation of the Eryops carpus ; and
articulating with the external surface of the ulna was the pisiforme,
probably small. The other bone of the carpus remaining is imper-
fect, and is probably the 2nd carpale.

The metacarpals are more slender than those of Eryops. Only
four metacarpals and four digits are preserved on each side,
and the phalangeal formula as preserved is 2, 2, 3, 3 (?) or 4.
The following table gives the chief measurements :

Length Proximal width
in mm. in mm.

1st metacarpal 23 15

1st phalanx 15 14

Claw 11 11

2nd metacarpal 29 14

1st phalanx 21 15

Claw 12 11

3rd metacarpal 28 17

1st phalanx 19 14

2nd phalanx 17 10

Claw ? ?

4th metacarpal 23 15

1st phalanx 16 12

2nd phalanx 12 9

Pelvis. The pelvis is remarkable for the shape of the ilium. In
the possession of a backwardly directed superior process and the
absence of any superior anterior portion, the bone is more strongly
reminiscent of such reptiles as Varanosaurus and Opliiacodon than of
Ei-ijops or the Cotylosaurs. The posterior process of the ilium slopes
backwards and somewhat upwards, and has a rounded superior
margin. The anterior margin is slightly convex, and there is no

Investigations in South African Fossil Beptilia and Amphibia. 75

preacetabular process. The inner border is concave behind the
attachment of the sacral rib.

The pubes are large and plate-like, broadly expanded in front
with a convex anterior margin.

Hind limb. The hind limb of each side is preserved almost
completely, the only missing bones being one or two from the
tarsus.

The femur is approximately straight, with expanded extremities.
The total length is 170 mm. The proximal end is swollen pos-
teriorly, and apparently somewhat flattened anteriorly. The maxi-
mum width of the proximal end is 62 mm., and of the distal end
55 mm. The distal end has two prominent flattened articular sur-
faces. The shaft is moderately robust, with a minimum antero-
posterior diameter of 25 mm.

The tibia and fibula are bones with widely expanded extremities,
each bone being about half the length of the femur. The distal end
of each bone has apparently two faces, the tibia articulating with the
tibiale and intermedium, and the fibula with the intermedium and
fibulare.

Most of the structure of the tarsus and pes is satisfactorily shown,
although two of the tarsalia are missing. The intermedium is
applied closely to the inner condyle of the fibula, lying between that
bone and the tibia. It is irregularly rhomboidal in shape on the
dorsal surface ; the length of the fibular side is 25 mm., its width
is 25 mm., and its height 28 mm. The tibiale and fibulare are
both lozenge-shaped bones, each articulating with the intermedium.
The tibiale is the larger bone, having a width of 36 mm. and
a height of 19 mm., while the fibulare is 24 mm. wide and 19 mm.
high.

Of the tarsalia but two are preserved the first and what is
probably the third. The first tarsale supported exclusively the first
metatarsal. It is rectangular in shape, 16 mm. wide and 12 mm.
high. The third lies in such a position that it probably assisted in
the support of both the third and fourth metatarsals, a view which is
borne out by its ovoid shape. It is smaller than the first tarsale,
with a width of 15 mm. and a height of 10 mm. Lying between the
tibiale and the first tarsale is a small rectangular bone measuring
12 mm. by 6 mm. This is the centrale.

The foot is short and heavy, larger than that of the fore-limb. All
the metatarsals are somewhat expanded at the extremities, the first
and last being shorter than the others. The digital formula is
2, 2, 3, 4, 3. The ungual phalanges are short and very bluntly

76 Annals of the South African Museum.

pointed, as in the fore-foot. The following are the chief measure-
ments of the bones of the digits in millimetres :-

Length Proximal width
in mm. in mm.

1st metatarsal 24-5 18

1st phalanx 15

Claw approx. 10

2nd metatarsal 36

1st phalanx 26 19-5

Claw approx. 11

3rd metatarsal

1st phalanx 24-5 20

2nd phalanx 17

Claw 11 n

4th metatarsal 37

1st phalanx 24

2nd phalanx 15 15

3rd phalanx 12 10

Claw missing

5th metatarsal 23 16

1st phalanx 21 15

2nd phalanx 16 12

Claw 11

Armour. The whole of the under side of the body was covered
with an armour consisting of elongated, overlapping scales arranged
in diagonal rows. The length of each scale in the mid-ventral
region is about four times the width. Each scale, viewed from
above, has a median groove, with one or two interrupted and nar-
rower lateral grooves. Broom, in his description of a specimen
from Senekal, says : " The armour consists of elongated, imbricated
scales. One set of scales, probably from near the middle line,
are much flattened and almost identical in appearance from those
of the middle region of the abdomen of Eryops. The other series
are probably from the more lateral region, and are narrower and
less flattened."

RHINESUCHUS AFIUCAXUS (Lydekker).
1890. En/ops africanus, Lydekker. Quart. Journ. Geol. Soc., xlvi.,

p. 291; pi. xii., tig. 2.

This species was described by Lydekker from a somewhat incom-
plete mandibular ramus obtained from some unknown locality in
the Karroo, and was placed in the genus Eliincsuclius by Broom.

Investigations in South African Fossil Kept ilia and Amphibia. 77

The species is imperfectly known. A crushed and incomplete skull
jind lower jaw (S.A.M. Cat. No. 3010) collected by the Eev. J. H.
Whaits at Dunedin, Nieuweveld, Beaufort West (Cistecephalus zone),
probably belongs to the species. It shows a type somewhat smaller
than R. seneJcalensis, with a basal length of about 400 mm. The
lower jaw is more robust than that of B. whaitsi, and there is no
subdivision of the coronoid bone. More material is necessary before
the details of this species can be satisfactorily determined.

(78)

6. On a Ncic Type of Dinocephalian (Moschosaurus longiceps).

(Text-figs. 8, 9.)

The skull on which this new genus is founded was collected in
1914 on the farm La-de-da in the Division of Beaufort West from
beds belonging to probably the upper part of the Parciasaurns zone.
With it were preserved half a dozen vertebrae, almost without
doubt anterior dorsals.

The skull is long, low, and narrow, and although there is a slight
elevation in the parietal region, there is none of the tremendous
thickening of the bones which is so prominent a feature of the larger
Dinocephalians such as Tapinoceplialus and Strnthioccplialus. The
nostrils are rather far back. The eyes are wholly in the posterior
half of the skull and are larger than the temporal openings. The
quadrate is carried forward to the plane of the middle of the orbit.
The lower jaw is massive.

The front of the nostril is 60 mm. behind the tip of the snout.
The nostrils are large, longitudinally oval, and the internasal width
is 20 mm.

The premaxilla has a suture with its fellow, and together the bones
form the rather pointed extremity of the snout. Superiorly they
separate the nostrils, sending back a narrow process to separate the
anterior portions of the nasals. The suture with the maxilla is
doubtful, but each premaxilla bore three or four large teeth. Each
tooth is carried in a separate socket. The roots are long, and the
crowns are differentiated into a long anterior cusp and a smaller
posterior one. The antero-posterior width of the crown is at least
li times the width of the root, the posterior cusp of the crown being
a sort of process some distance above the level of the anterior cusp.
Similar teeth are known from other and larger Dinocephalians. In
Struthiocephalus the outside of the crown is convex and the anterior
cusp is bluntly pointed, while the interior surface is concave below the
posterior cusp which is convex on both sides. In some of the teeth
in that genus there are one or two longitudinal grooves on the
inside of the crown.

Investigations in South African Fossil Reptilia and Amphibia. 79

The maxilla is a large bone forming the lower border of the nostril
and apparently passing back to form part of the sub-orbital arcade.
It carried probably 6 or 7 teeth, of which only one or two are
preserved. The posterior teeth are smaller than the anterior. The
first may have functioned as a canine, being apparently bluntly
pointed, but it was certainly comparatively much smaller than the
canine of Titanosuchus.

The facial part of the septomaxilla is small, forming part of the
posterior wall of the nostril and separating the maxilla and nasal
for a short distance.

The nasals form a well-marked upstanding ridge between the
nostrils and orbits along the centre of the skull, a region well
marked off from the cheeks, which slope at an angle of about 50
from the jaws and are concave directly in front of the orbits. This
nasal ridge is wide in its posterior part, suddenly narrows, and then
has a uniform width of about 30 mm. throughout the anterior
two-thirds of its length.

The orbit lies wholly within the posterior half of the skull, is
longer than high, and shows very little supraorbital thickening.
The prefrontal forms the anterior upper quadrant of the orbital
margin, but its front margin is not well delineated.

The lachrymal is smaller than the prefrontal and does not reach
the septomaxillary.

The interorbital area has a width of nearly 80 mm. It is flat,
formed mostly by the frontals, which form a small part of the
superior borders of the orbits.

The postorbital bar is comparatively .weak, but the postorbitals
pass back to form a large part of the upper border of the temporal
fossa. The parietals are not distinguishable, but they must have
been small.

The pineal foramen is sub-circular in shape and very large, having
a longer diameter of 26 '5 mm. The parietal region surrounding it
was raised slightly above the level of the frontals. The foramen is
placed very far back, in the plane of the back of the temporal
fossa.

The squamosal, quadrato-jugal, and jugal occupy apparently the
same relations as in Moschops and Delphinognathus, but there is no
foramen between the jugal and quadrato-jugal such as occurs in
Delphinognathus. The quadrate is scarcely seen from the side of the
skull. It is pushed forward by the large squamosal to the level of
the middle of the orbit. From behind it is roughly boot- shaped
with the "heel" internal and the "toe" pointing outwards and

80

Annals of the South African Museum.

slightly forwards. The articular of the lower jaw is closely applied
to it.

The occipital plate is weathered and broken. There is a large
foramen magnum placed high up. The squamosal only forms the
edge of the plate and the outer boundary of the lateral post-temporal
fossa. Most of the side of the occipital plate seems to be formed by
the tabulare. Below the basioccipital condyle there is a vertical
mass of bone as in the other Dinocephalians.

The lower jaw is deep and displayed only in outer view. Its total
length is about 215 mm. The dentary covers almost the whole of
the anterior half of the jaw, the lower border being formed by the

FIG. 8. Anterior view of vertebra of Hoschosaurus longiceps. x

splenial ; and the dentary extends backwards along the upper border
of the jaw for about four-fifths of its length. This posterior process
is much more slender than that seen in Dimetrodon. The dentary
carries 5 or 6 teeth, forwardly directed and presumably diminishing
in size posteriorly.

The splenial forms a small portion of the symphysis and is only
visible along the lower border of the anterior half of the jaw.

The back of the angular is missing ; but the notch in its lower
border must have been a small one. The bone overlaps the articular
which, from below, is seen to be swollen vertically and to pass
forwards and upwards.

Six consecutive vertebrae are preserved, without doubt anterior
dorsals. The centra are short with circular biconcave ends. The
lower border is concave with a slight median ridge. The sides are

Investigations in South African Fossil Bcptilia and Amphibia. 81

more concave than the ventral surface. The transverse processes
are large, pointing slightly upwards, thicker distally than proximally.
The neural spines are high and flattened laterally.

The following table gives the measurements of three of the
vertebrae, the 1st, 3rd, and 5th of the series :

1st Vertebra. 3rd Vertebra. 5th Vertebra.

Length of centrum 30 mm.

Height of centrum 28 ,,

Total height 88 ,,

Width across transverse processes 70 ,,

Height of spine 35 ,,

Length of spine 17 ,,

33 mm.

32 mm

31

31

112

117

68

9

52

58 ,"

23

22

No intercentra are preserved.

This type, although undoubtedly a Dinocephalian, presents in its
external shape so great a contrast to the other members of the
group that I propose to erect for it a new genus, and to name it
Moschosaurus longiccps, g. et sp. nov.

Type. Skull and lower jaw. (S.A. Mus. Cat. No. 3015.)

Locality. La-de-da, Beaufort West, Cape Colony.

Horizon. Lower Beaufort Beds. (Upper part of Pareiasaurus
zone.)

TV

FIG. 9. Skull of Jloschosaurus longiccps. x

(82)

7. On Some New Gorgonopsians .

1. GALESUCHUS GRACILIS, g. et sp. nov.
(Text-figs. 10, 11.)

This new genus and species is founded on a specimen collected at
Abraham's Kraal, Prince Albert Division, that is, from the lower
half of the Pareiasaurus zone, where it occurred in conjunction with
Trochosaurus intermedius and bones of Pareiasaurus and of large
Dinocephalians. It is a weathered skull and lower jaw, lacking the
anterior part in front of the canine.

The skull is small. As preserved its greatest length is 120 mm.,
while the maximum width across the squamosals was probably
75 mm. The orbits look outwards, and the antorbital portion
seems not to have been quite so elongated as in the Gorgonopsians
from the Endotliiodon and Cisteceplialus zones. The antero-posterior
diameter of the orbit is 32 mm., and the interorbital width 30 mm.,
being equal to the interternporal. The temporal fossa looks upwards
and outwards. The occiput is very sloping, inclined at an angle of
not much more than 30 to the top of the skull.

The canine is fairly large and vertical. The molars are small,
four in number, simple, unserrated, and slightly recurved. The
maxilla is deep, with a posterior slender process passing below
the jugal to the plane of the middle of the orbit.

Both the prefrontal and lachrymal are large, while the jugal forms
part of the anterior, and the whole of the inferior, borders of the
orbit. The prefrontal forms the whole of the anterior superior
quadrant of the border.

The frontal is large, forming 5 mm. only of the orbital border.
It passes backwards, supporting the median preparietal, and
articulating also with the parietal and postfrontal.

The postfrontal is large, and forms a large part of the upper
border of the orbit, having the same general shape as in Scylacops
capcnsis.

The preparietal is 15 mm. long, and lies 4 mm. in front of the
pineal foramen, which is 5 mm. long and oval in shape.

Investigations in South African Fossil Eeptilia and Amphibia. 83

The parietals are large, and wholly enclose the pineal foramen.
Owing to weathering of the bone surface their relations with the
squamosals cannot be determined.

The interparietal forms the upper half of the occipital plate and
articulates below with the supraoccipital.

The foramen magnum is large, oval in shape. Its lateral borders
are formed by the small exoccipitals. The basioccipital condyle

FIG. 10. Galesuclnis gracilix.
Sketch of top of skull showing arrangement of bones. -,.

is rounded. The basisphenoidal tubera are far back and considerably
below the level of the condyle, so that the foramen magnum appears
to be high up on the back of the skull. The paroccipital process
is deep and strong, and the post-temporal fossa is as high as the top
of the foramen magnum, looking wholly backwards. The foramen
jugulare is not seen, but it must have been high up as in Arctops.
The condition of the matrix renders it almost impossible to develop

84 Annals of the South African Museum.

the palate, but fracturing of the specimen has shown one or two of
its features. The descending processes of the pterygoids are large.
As in Scylacops capensis and other Gorgonopsians the pterygoid
is closely united with the transpalatine, there being no evidence
of any foramen between them. The ascending pterygoidal plates
pass forward above the palatine. Superiorly the fused pterygoids
thin out into a plate slightly displaced from its true position, which
does not appear to meet the ethmoid, although the two bones may
have met further back. The ethmoid is a median element lying
vertically beneath the frontal suture, and separated from the upper
cranial wall.

The palatine carries a few simple pointed teeth.

Only the posterior half of the vomerine bone is seen. It descends
below the level of the front of the palatine, which ascends anteriorly
towards the top of the skull as a thin plate. Between this plate and the
vomerine bone is a distinct foramen. This bone is apparently single.

The lower jaw is incomplete, but agrees in external characters
with that of Scymnocjnatlms. The dentary is very large, and its
posterior process rather more powerful than in the other Gor-
gonopsians. The angular is large.

Ti/pe. Incomplete weathered skull. (S.A.M. Cat. No. 2754.)

Locality. Abraham's Kraal, Prince Albert Division.

Horizon. Beaufort Beds. (Pareiasaurus zone.)

2. GORGONOGNATHUS LONGIFKONS, g. et Sp. 11OV.

(Plate XIII., figs. 1, 3.)

The skull about to be described is No. 2671 of the South African
Museum collection, and was collected at Dunedin, Beaufort West.
from a bed of sandstone 1^ miles east of the homestead. The top of
the snout and back of the temporal region are denuded of bone by
weathering, and the whole skull is somewhat distorted. The general
features of the skull are those of Gorgonops torvus, which occurs,
however, at a lower horizon : but the head is not quite so much
depressed, and is much larger than Owen's type.

The maximum length of the skull was between 340 mm. and
350 mm. From the snout to the front of the orbit is 200 mm.
The basal length from the snout to the back of the occiput is
305 mm. The interorbital width is 77 mm., while the intertemporal
width was probably about 85 mm.

The premaxilla is divided by a median suture. The septomaxilla.

Investigations in South African Fossil Rcptilia and Amphibia. 85

and maxilla have the normal Gorgonopsian relationships. The
septomaxilla is large, and has the inwardly directed turbinal process
dividing the nares into upper and lower passages. It forms an
appreciable portion of the cheek behind the nostril, and has the
usual outer foramen between it and the maxilla. The nostrils are
nearly terminal.

The maxilla is large, forming nearly three-quarters of the cheek,
and having a sub-orbital portion supporting the jugal.

The dental formula is i5 cl ?u4. Most of the teeth are missing,
only the fourth incisor on either side being present, but the sockets
are plainly visible. The incisors are large, nearly equal in size to
one another, and closely set together, the five teeth occupying a
space of 54 mm. at the edge of the bone. At that level the
4th incisor has an antero-posterior diameter of 11 mm The roots
of the teeth are long and simple. Each tooth is implanted in a
separate socket. Behind the last incisor is a diastema of 14 mm.,
and then comes the socket of a massive canine 28 mm. in diameter.
Directly behind the canine is a series of four molars all lost
gradually decreasing in size, and occupying together a space of
33 mm.

The nasals are very large, forming the top of the snout, and
having posterior projections which separate in part the f rentals from
the prefrontals.

The prefrontal is large, forming the upper anterior quadrant of
the orbital border. The frontal is largely shut out from the orbital
border, but it forms 13 mm. of it between the prefrontal and post-
frontal. It passes forward, separating the posterior parts of the
nasals.

The orbit is small, looking forwards and outwards and possibly
slightly upwards, and lies wholly in the posterior half of the skull.
The postfrontal is large, forming nearly one-fourth of the orbital
margin, and articulating apparently with the parietal to behind the
pineal foramen.

The temporal fossa is slightly larger than the orbit. The inter-
temporal region is wide, and the pineal foramen is placed back
almost as far as the occipital crest. The preparietal is small, and
wholly in advance of the pineal foramen.

The squamosal forms the lower and most of the posterior borders
of the temporal fossa and the outer sides of the occipital plate. The
lower part of the bone and the quadrate region are missing.

The occipital plate is broader than high, and vertical. The
interparietal is slightly broader than deep, and forms a large part

86 Annals of the South African Museum.

of the plate. It has a prominent median ridge in its lower half,
which barely passes over to the bone below.

The most recent accounts of the basicranial region of the
Gorgonopsia have been given by Watson, who describes skulls of
Arctops willistoni and Scymnognathus whaitsi, discussing their
relations with Dimctrodon on the one hand and Diademodon on the
other. In Gorgonognathus the basioccipital condyle is rounded and
swollen below, and hollowed above for the reception of the foramen
magnum. The paroccipital process is short and powerful, forming
the lower border of the post-temporal fossa. It is considerably
in advance of the tabulare, and shallow when viewed from be-
hind, so that the post-temporal fossa looks almost wholly down-
wards. The exoccipitals are like those figured by Watson, and form
the lateral borders of the foramen magnum. The foramen jugulare
is at the bottom of the skull in front of the exoccipital, bounded by
that bone, the paroccipital, and the basioccipital, and looks wholly
downwards.

The whole occiput is concave and the squamosals fairly wide.
The interparietal is narrow and the tabulare large as in Scymnog-
nathus. The interparietal carries a well-marked median ridge,
which dies out below on the supraoccipital. The tabulare forms the
outer upper border of the post-tempoi'al fossa, and covers part of
the back of the inner ramus of the squamosal.

Thus this form, occurring as it does in the Cisteceplialus zone,
seems to occupy rather an anomalous position. In the width of the
parietal region, the shortness of the temporal fossae, the shape
of the basioccipital condyle, and the stoutness of the paroccipital
it agrees with Arctops. But in the reduction in height of everything
below the foramen magnum, the downward aspect of the post-
temporal fossae, the position of the foramen jugulare, and the size
of the squamosal, interparietal and tabulare it agrees with Scymnog-
nathus, indicating possibly an advance on that form.

The palatal view of the premaxilla shows a somewhat pitted
flat plate anteriorly, which forms part of the inner border of the
large internal narial openings, and articulates behind with the
" vomer."

This vomer is comparatively broad anteriorly and narrows behind.
It has ventrally a median keel and two lateral keels, with slight
intermediate grooves. Although the median suture separating the
two premaxillae can be well seen, there is no suture running down
the middle of this bone, and it is undoubtedly a single bone. For
some time Dr. Broom has considered that this median bone is a

Investigations in South African Fossil Ecptilia and Amphibia. 87

pair of fused prevomers, and Watson has recently mentioned that in
the type skull of Arctops the bone seems to be actually double when
seen in section on the anterior end of the specimen. As a contribu-
tion to our knowledge on this subject I have examined skulls of
three Gorgonopsians from different zones, viz. Galesiiclius gracilis
from the Parciasaurus zone, a snout of Goryonops sp. (S.A.M. Cat.
No. 3038) from the Endotliiodon zone of Beaufort West, and
Scymnognatlius scrratidens from the Cistecephalus zone of Dunedin,
Beaufort West. All these skulls show a cross- section through the
vomer. The bone is fairly similar in all three cases with regard to
the ventral surface, but the dorsal surface shows some variation,
best seen and understood by reference to the figures.

1. Cross-section through vomer of Galcsuchus gracilis through canine.

2. Cross-section through vomer of Goryonop* sp. immediately anterior to canine.

3. Cross-section through vomer of Scymnognathus serratidcns through canine.

All the figures x 3.

There is seen on the dorsal surface a progressive reduction of the
median ridge, and a tendency for the two grooves to become one.
None of the specimens show any signs of a median suture.

The front of the palate is considerably vaulted, the plane of the
premaxilla and "vomer" being above that of the palatine and
maxilla, whose inner portions rise vertically at right angles to the
outer portions.

The palatines approximate to each other to form the beginning of
a secondary palate considerably below the primary. The palatine is
a large bone passing forward almost to the back of the canine. On
its posterior and inner portion it carries a few strong, well-developed
teeth.

The pterygoid has a very powerful descending process, deepest at
the side of the mandible, and rising to the level of the palatine where
it meets its neighbour. The maximum depth of this process is
60 mm. and the distance between the two extremities 110 mm. In

88 Annals of tJie Soutli African Museum.

front the pterygoid meets the transpalatine and palatine. Behind
the descending processes is a long narrow median bar. The anterior
portion of this is formed by the pterygoids in contact, but for the
greater part the pterygoids seem to be separated by a thin median
forward process of the basisphenoid. This process may be the
parasphenoid, but no suture can be traced between it and the main
body of the bone. Superiorly the pterygoid forms a thin bony plate
which is not anchylosed to its neighbour, but in places is completely
separated from it by matrix.

At the back of the basisphenoid are two tubera which are strong
and but slightly in advance of the basioccipital condyle.

Type. Crushed skull in sandstone. (S.A.M. Cat. No. 2671.)

Locality. H miles E. of Homestead, Dunedin, Nieuweveld,
Beaufort West.

Horizon. Beaufort Beds. (Cistecephalus zone.)

3. SCYMNOGNATHUS SEREATIDENS, Sp. 11OV.

(Plate XIII., figs. 2, 4. Text-fig. 11.)

The type of this new species is the anterior two-thirds of a skull
and lower jaw found 3 miles WSW. of the homestead on the farm
Dunedin, Beaufort West, in a calcareous nodule. It is smaller than
either S. whaitsi or S. tigriceps, and slightly smaller than S. angusti-
ceps, although agreeing in general shape with the two former. It
differs from S. tiijriccps, S. angusticeps, and S. minor in having all
its teeth serrated behind in their lower halves. The incisors and
canines of S. tigriceps are worn down to a certain extent, and it is
just possible that the lower parts of these teeth might have been
serrated ; but the molars of the larger form certainly show no trace
of serration.

The premaxilla, maxilla, and septomaxilla are as in S. ticjriceps,
the septomaxilla having an anteriorly directed turbinal which divides
the nares almost completely into upper and lower portions.

The dental formula for the upper jaw is i5 cl w4. The first two
incisors are smaller than the others ; the largest are the third and
fourth. The five incisors occupy a space of 38 mm. They are
serrated on the posterior border. On the 3rd, which is the most
perfectly preserved, the serrations begin 7 mm. below the edge
of the premaxilla and continue almost, if not quite, to the point of
the tooth. On the same tooth in a distance of 14 mm. there are
36 serrations. The maximum width of the 3rd incisor is 7 mm., and

Investigations in South African Fossil Reptilia and Amphibia. 89

its length below the edge of the bone 25 mm. The following gives
the length of the incisors: /I, 19 mm. ; ?2, 21 mm. ; /3 25 mm. ;
4, 24 mm. ; i5, 20 mm. In section the teeth are oval. There is a
diastema of 21 mm. between the last incisor and the canine. The
canine is very long, reaching almost to the bottom of the mentum,
55 mm. below the edge of the maxilla, pointed and backwardly
curved. It is serrated behind in its lower half. Behind is a
diastema of 15 mm., and then come four molars occupying a space
of 24 mm. These are simple, pointed teeth, serrated posteriorly in
their lower halves. The first molar has 14 serrations in a distance
of 4 mm., the whole length of the crown being 10 mm. All the
teeth are implanted in very deep, distinct sockets.

The prefrontal forms the upper and anterior quadrant of the
orbital border. The frontal is 85 mm. long, and just forms part of
the orbital border, being almost shut out from it by the prefrontal
and postf rental. The relation of the frontal to the orbital margin in
this genus is a variable one. In S. angusticeps the frontal forms
a considerable part of the margin ; in this species it is almost
excluded from it ; while in S. tigriceps and S. icJiaitsi it is very
doubtful whether it plays any part in the border at all. In
Galcsuchus and Gorgonognatlius the frontal forms but a small part
of the border, while in Scylacops it is completely excluded by the
junction of prefrontal and postfrontal.

The median preparietal is bounded by the frontals and parietals,
with which it articulates by interdigitating sutures. It is oval
in shape, 30 mm. long and 17 mm. broad, and is situated at the
level of the postorbital bar. The pineal foramen is large and lies
5 mm. behind the preparietal.

The hinder part of the cheek below and in front of the orbit is
very much hollowed out. The slope from the front of the orbit
is very abrupt, while that from the maxilla is much more gentle.
This concavity is much more pronounced than in S. tirjnccps.

The nasals are long, but not convex, sloping upwards from the
face to form a prominent sharp median ridge which extends back as
far as the frontals.

The lower jaw has the mentum relatively less deep than that
of S. tigriceps, and a backward slope so that the point of the jaw
comes directly below the canine.

The chief features of the species are its comparatively small size,
the possession of serrated teeth, of a ridge on the top of the snout,
and the pronounced backward slope of the front of the lower
jaw.

90 Annals of the South African Museum.

The following are the chief measurements of the type :

Tip of snout to front of orbit 132mm.

Tip of snout to back of orbit 177 ,,

Interorbital width 66 , ,

Intertemporal width 64 , ,

Depth snout to mentum 130 , ,

Type. Incomplete skull. (S.A.M. Cat. No. 2672.)
Locality. Dunedin, Nieuweveld, Beaufort West.
Horizon. Beaufort Beds. (Cistcccphalus zone.)

(91)

8. On a Skull of the Genus Kannemeyeria.

(Text-figs. 12-14.)

The genus Kannemeyeria was founded by Seeley in 1908 (Eep.
Brit. Ass., 1908, p. 713) on a skull of a large Anornodont found
by Dr. Kannemeyer near Burghersdorp, which is according to
Watson (Ann. Mag. Nat. Hist., 1912, x., p. 575) an imperfect skull
of Dicynodon simocephalus, Weithofer. Weithofer described his
species in 1888 (Ann. K.K. Natur. Hof Mus., Wien, Bd. iii.) and
figured the type, an imperfect skull. Broom (Bull. Amer. Mus.,
1913) refers to this species as Kannemeyeria simocephalus.

The genus is characterized by having a broad frontal region, a
narrow and high parietal crest, a parietal region inclined at an
oblique angle to the frontal plane, the pineal foramen at the front of
the parietal crest situated in a well-marked depression, an absence
of the postfrontal and probably of the preparietal bones.

The hitherto described species are two :

Kannemeyeria simocephalus (Weithofer), (loc. cit.],

Kannemeyeria latifrons (Broom),

and to these I now add a third, founded upon a well-preserved and
complete skull together with most of the lower jaw and the first
20 vertebrae found by Mr. E. W. Pocock at Winnaarsbaaken,
Burghersdorp, in 1914 (S.A.M. Cat. No. 3017), to which I propose
to give the name Kannemeyeria erithrea, sp. nov.

Skull. The general shape of the skull can best be understood
from the figures. The orbits look almost entirely outwards. The
temporal fossae are large. The nostrils are near the front of the
skull. The snout is rugose as are the supraorbital borders and
the nasal overhang and is provided with a pronounced median
ridge which extends back to the plane of the front of the orbit.
From the posterior point to the plane of the back of the nostril this
ridge has a median groove, so that it has the appearance of being
double.

The nostril is large, reniform in shape, and near the front of the
snout. It has a large, overhanging bony roof. The anterior border

92

Annals of tlie Soutli African Museum.

is formed by the premaxilla, which has a median septum separating
the two nostrils. From above only the front part of the nostril can
be seen. The posterior half is overhung by the nasal, which is
nearly twice as wide as the premaxilla. The premaxilla has a long
posterior process separating the two nasals, which, although large

FIG 12. Top view of skull of Kannemeyeria eritlmn. < i nearly.

bones, only meet one another for a distance of about 40 mm.
posteriorly.

The floor of the nostril is formed by the maxilla. This carries
a large tusk, which is directed strongly forwards. Posteriorly the
maxilla sends back a long process outside and below the jugal
reaching to the postorbital bar. The bones in the nostril are slightly
crushed, and I am unable to distinguish a separate septomaxilla.
If present, it is small and certainly does not occur on the face.

s in South African Fossil Beptilia and AntpJtibia. 93

The limits of the lachrymal and prefrontal are not well displayed
in either of the two skulls we possess ; the lachrymal is certainly
small, but the prefrontal forms a large part of the inner orbital wall.
The lachrymal foramen lies wholly within the orbit.

The frontal is broad and flat, forming the posterior half of the
supraorbital border. It has a short process passing back along
the side of the parietal, between that bone and the postorbital.
There is no postfrontal.

The postorbital at its lower end rests on the jugal ; it forms the
whole of the postorbital bar and meets the parietal, forming part
of the inner border of the temporal fossa.

The pineal foramen is placed in a distinct depression. I am
inclined to believe that it is wholly surrounded by the parietals,
the preparietal being absent.

FIG. 13. Side view of skull of KanneiHeycriu i-i-itlircit. x i nearly.

The parietal crest is high and narrow. Posteriorly the parietals
are separated by a deep groove which has a shallow prolongation on
to the upper half of the occipital plate.

The occipital plate is peculiar. The lower two-thirds is vertical ;
but the interparietal and the upper halves of the squamosal parts
are bent at an angle of between 110 and 120, so that the plate
is completely hidden from view both from above and from the sides.
The bones of the occiput occupy the same relative positions as in
other members of the Anomodontia. The foramen magnum is large.
The exoccipital has, on the inner border midway between the
condyle and the supraoccipital, a prominent protuberance, and
external to that another smaller rugose knob ; the posterior corner
of the paroccipital is prolonged to form a bluntly pointed process.
The lateral occipital foramen is large, lying mostly in the exoccipital,

94 Annals of the SoutJi African Museum.

the supraoccipital forming only a minute portion of its upper border,
and the squamosal its outer border only. The condyle is tripartite.

The squamosal is a very large bone. The skull is peculiar in that
the width between the outer edges of the descending processes of the
squamosal is its greatest width.

Lying in the hollowed portion of the exoccipital just outside the
condyle is a small rounded foramen for the exit of the 9th, 10th, llth,
and 12th nerves.

No sutures are distinguishable between the basisphenoid, exocci-
pital, and quadrate, and no separate quadrato-jugai can be seen.

The basioccipital forms the lower part of the tripartite condyle,
narrows somewhat in front and then widens, forming on either side
a large process with an outer concave articular surface, each pierced
by the fenestra ovalis. The stapes is absent. The basisphenoid lies
in front of the basioccipital and forms part of the anterior surface of
the basioccipital process. The centre of the bone is hollowed out,
and on the inner sides of the ridges bounding this hollow are the
earotid foramina. Laterally and anteriorly the basisphenoid articu-
lates with the pterygoid.

The pterygoid is a long narrow bone passing from the maxilla to
the quadrate. It lies on the outer side of the palatine. For a short
distance in front of the basisphenoid the two pterygoids unite in the
median line. In front of this median pterygoidal plate, between
it and the forked posterior end of the vomer is a large pterygoidal
foramen. There is the usual posterior lateral process passing to the
front of the quadrate. The sphenoidal portion of the pterygoid is
very small.

The lower part of the columella cranii is seen, closely affixed
to the outer side of the pterygoid and basisphenoid.

The posterior nares are large. The outer border is formed almost
wholly by the palatine, which passes along the inside of the pterygoid
and meets the "vomer" posteriorly. Anteriorly the palatine lies
above the pterygoid and has a rugose surface which meets the
prernaxilla. The "vomer" is a thin median bone forming the
median septum between the posterior nares. Posteriorly it forks,
and curves on each side outwards to meet the palatine, passing
behind that bone and forming most of the hinder wall of the
posterior nares. According to Broom, this bone in the Anomodonts
is formed of fused prevomers and is not homologous with the
mammalian vomer.

In front of this median bone there is a pronounced median ridge
on the premaxilla which extends forward half the length of the bone.

Investigations in South African Fossil Reptilia and Amphibia. 95

At the front of the palate this ridge is replaced by a groove, on either
side of which is a less well-marked ridge.

The following are the chief measurements of the skull :

Greatest length 450 mm.

Greatest breadth 330 ,,

Minimum width of nasal overhang 65 ,,

Maximum width of nasal overhang 119 ,,

Interorbital width 141 ,,

Intertemporal width 15 ,,

Snout to front of orbit 167 ,,

Length of orbit 82 ,,

Basal length 332 ,,

Minimum width across pterygoids 64 ,,

Lower jaw. The most recent descriptions of the Anomodont
lower jaw have been given by Watson and van Hoepen. The
former gives a general account, basing his description of the bones
of the back of the jaw on a " fragmentary jaw of Kanncmeyeria
collected by the author at Winnaarsbaaken, Burghersdorp District "
from which farm this type also comes. In general the jaw agrees
with that figured by Watson except that, owing to the difficulty
of development, it is impossible to be sure of the presence of the
coronoid. Van Hoepen states expressly that a coronoid does not
exist either in Lystrosanrus or in Dicynodon ; but he seems certainly
to have misunderstood the position of the coronoid as defined by
Watson.

The front of the jaw projects slightly upwards, forming a beak
which fits into the concave upper jaw. The front of the mandible
has a median ridge running from the point to the mentum bounded
by two lateral grooves, one on either side, on the outer side of each
of which is a less well-defined ridge.

The dentary forms almost the whole of the front of the outer
surface. Its edentulous border is provided with a longitudinal
groove which narrows and deepens posteriorly. The posterior part
of the bone is produced into two processes, the upper of w r hich
seems to lie outside the surangular and angular. Between it and
the lower process is a deep notch. The lower process lies outside
the angular.

In the side view the splenial is only just seen. It forms part
of the median symphysis and passes inside the angular, whose thin
anterior end lies between the dentary and the splenial.

96

Annals of the South African Museum.

On the right-hand side of the jaw lying in the groove in the
dentary is a small piece of a bone which may be the displaced
coronoid.

The suture between the surangular and articular is not seen ; but
those between the angular and surangular and angular and pre-
articular are well displayed. The surangular forms most of the
upper border of the fossa, and lies within the dentary and angular.
Its upper edge forms a thickened ridge. The inner flange of the
articular is very pronounced.

FIG. 14. Sketch of odontoid and axis of Kannemeyeria erithrea. x

. Associated with the skull and lower jaw were a
number of vertebrae, which seem to form a continuous series of
20 vertebrae from the odontoid backwards.

The odontoid is the only portion of the atlas preserved. It is
of the usual trefoil shape with a height of 65 mm. and a maximum
width of 62 mm. Its anterior face is strongly convex. In the
centre of the front face is a small aperture which coincided with
the small pit at the end of the occipital condyle. The upper surface
is strongly concave for the floor of the neural canal. There is no
intercentrum between the odontoid and the axis. The atlantal arch
is not preserved. It rested, apparently, almost wholly on the
odontoid, and articulated behind with the axial prezygapophyses
and in front, probably, with the processes of the exoccipitals seen
on either side the foramen magnum.

investigations in South African Fossil Beptilia and Amphibia. 97

The axis and odontoid seem to be fused together ; possibly the
fusion may not be a true anchylosis, but due to the pressure of one
against the other.

The axis has an elongated neural spine, compressed in the middle
and swollen at either end, with a convex upper edge. The transverse
processes are fairly short, and a short distance behind the front of
the centrum and below the median line is a vertically elongated
facet for the articulation of the capitulum of the double-headed
axial rib.

The 3rd to 7th vertebrae probably all caudals are cemented
together by matrix. The centra gradually enlarge, the flattened
neural spines rapidly increase in size, as do the transverse processes,
the facets for the articulation of the tubercula of the ribs becoming
markedly oblong in shape. The central facets remain on the anterior
edges of the centra.

The remaining vertebrae were mostly isolated in the matrix, but
they seem to form a series. The neural spines from the 9th bend
backwards and gradually decrease in size. The neurocentral suture
becomes well marked, and the transverse and central facets are
connected by an oblique groove.

The following are the chief measurements :

Vertebrae No.

6

39
62

9

39 39 36 35 35
60 59 61 50 55
195 189 170 132 143

Axis 3 4

Length of centrum 42 37 38

Width of centrum 56 55 51

Total height 128 142 161

Height of spine above transverse

process 66 71 80 ?

Width across transverse process 90 97 100 100

Width across prezygapophyses... 55 37 42 38

Width across postzygapophyses 32 35 30 48

[All the measurements are in mm.]

Type. Skull and lower jaw. (S.A. Mus. Cat. No. 3017.)
Locality. Winnaarsbaaken, Albert, Cape Colony.
Horizon. Burghersdorp Beds. (Cynognathus zone.)

8 13? 17?

109

101

78

53

58

96

97

98

83

75

43

41

42

32

39

38

35

27

32

32

8

98)

9. .1 New Thccodont from the Storinbcnj Beds.

SPHENOSUCHUS ACUTUS, g. et sp. nov.

(Text-figs. 15-17.)

This exceedingly interesting new form is described from a
specimen collected by Dr. A. L. du Toit from the Bed Beds of
Paballon, Mount Fletcher. It consists of an incomplete and some-
what crushed skull, with the cervical vertebrae attached, the two

FIG. 15. Side view of skull of Sphenosuclms acutut. x .
(Slightly restored.)

scapulae, two clavicles, an interclavicle, two humeri, and some other
fragmentary bones all in one slab of rock ; and a complete tibia
and the distal third of a fibula which in all probability belong to the
same animal.

The skull is somewhat crushed but nearly whole, and shows all
the external details. It is larger than that of Euparlceria, and
is comparatively more pointed, longer, and narrower. The orbits
are rounded, and wholly in the posterior half of the skull. The
supratemporal fossa is elongate, oval in shape, and larger than that
of Euparkeria or Ornitliosuclnis. The shape of the infrateniporal
fossa is characteristic in that its anterior border, formed by the

Investigations in South African Fossil L'rptilni cnnl A /i//i'li/i>ia. 99

jugal and postorbital, passes upwards and forwards instead of
upwards and backwards as in EiipnrL-t>r/a and Ornithosuchus ; so
that the superior length of the opening is about equal to the inferior
length.

The snout is characterized by the fact that the premaxilla does
not form an anterior border to the nostrils, these being quite

FIG. 16. Top view of skull of S2>}ienosuchus acittus. < i.
(Slightly restored.)

terminal. Further, there is no trace of a median suture dividing
them. The roof of the snout is formed by the paired nasals, which
are broken posteriorly. The extreme tip of the left nasal is missing,
but the bones were obviously pointed in front. The whole of the
posterior and lower borders of the nostril are formed by the pre-
maxilla, which sends back a process separating the forward portion
of the maxilla from the nasal. The nostril seems to have had an
upper prolongation between the nasal and premaxilla. The pre-
maxilla carries apparently three or four simple pointed teeth. In

100 Annals of the South African Museum.

Ornithosuchus and Erpetosuclms the pretnaxilla is entire, and forms
an anterior border to the external nares, and Broom has considered
that the bone in Euparkeria was similar. Of the fact that in this
type the nostril was terminal and open I think there can be no
doubt. Dr. du Toit, who partially developed the front of the snout
before the fossil came into my hands, informs me that no bone was
broken away save a tiny fragment from the outer edge of the left
nasal ; and in the specimen as it is now displayed there is no sign of
any anterior vertical portion of the premaxilla.

There is no septomaxillary present on the face.

The antorbital vacuity is large, and is sunken in the face, having
borders which make an oblique angle with the sides of the face.
The whole of the lower and anterior borders is formed by the
maxilla, which extends back only so far as the front of the orbit-
not nearly so far as in Euparkeria. The maxilla carried about
12 teeth, of which 8 are preserved on the right side. Unfortunately,
not one possesses the crown ; but a small tooth in the lower jaw
shows serrations on the anterior border similar to those of the
carnivorous Dinosaurs. The teeth are flattened laterally, and vary
considerably in size. The first maxillary tooth has an antero-
posterior diameter of just over 1 mm. ; the probable 6th, which
is the largest, has a diameter at the gum of 7 '5 mm. The teeth do
not increase or decrease regularly in size from front to back of
the jaw, but are variable.

The surface of the maxilla is plentifully supplied with grooves and
small foramina for blood-vessels.

The nasal is an extremely long bone forming the upper surface of
the skull from the tip of the snout nearly to the plane of the front
of the orbit. It forms none of the posterior border of the nostril.
The greatest width across the pair of bones is 20 mm. at the back
while the length is about 88 mm.

The lachrymal forms the whole of the upper border and most
of the posterior border of the antorbital vacuity, besides forming
the larger part of the anterior orbital border.

The prefrontal is a small bone lying between the frontal, nasal,
and lachrymal. Below it has a lobe-like extension articulating with
the lachrymal, so that it forms about 18 mm. of the orbital border ;
but its width throughout most of its length is only about 6 mm.

I can see no evidence of a postfroutal. Even if one be present,
the frontal is still peculiar in that it passes back to form part of the
anterior border of the upper temporal fossa, separating the post-
orbital from the parietal. The interorbital region has a median

Investigations in South, African Fossil Eeptilia awl Ampliibia. 101

elevation, broadened at the level of the postorbital bars, and
narrowing posteriorly until it forms the median parietal crest. On
each side there is a slight supraorbital crest ; and between this and
the median ridge is a well-defined channel. The frontal forms half
of the supraorbital border. Its greatest length is in the middle
line, the sutures with the pai'ietals passing well forward from a
point one-third along the parietal crest nearly to the anterior
extremity of the supratemporal fossa.

The postorbital bar differs from that in Euparkeria and the allied
Thecodonts in that its upper end is in advance of the lower. The
descending portion of the postorbital is thus inclined backwards
instead of forwards, lying in front of the ascending process of the
jugal. The postorbital forms most of the outer border of the upper
temporal fossa, and a small portion of the upper border of the lower
opening, anteriorly articulating with the frontal, and posteriorly
overlying a portion of the squamosal. Nowhere does it meet
the parietal.

The jugal is a tripartite bone. Its anterior process forms the
inferior border of the orbit, and articulates with the lachrymal
and maxilla. It does not pass up as far in front of the orbit as
in Euparkeria. The ascending process lies behind and superiorly
interior to the postorbital and is inclined slightly forwaixls. The
posterior process forms most of the zygomatic arch, lying outside
the quadrato-jugal.

From the fact that the postorbital bar slopes in almost a pai'allel
direction to the quadrate, the lower temporal opening has a
rhomboidal shape, being bounded by the postorbital, jugal,
quadrato-jugal, quadrate, and squamosal. It is slightly bigger
than the upper opening. In this it agrees with Euparkeria,
Ornithosuchus, and the Theropodous Dinosaurs, and differs from
Erpetosuchus. In the relation between the sizes of the temporal
openings, and in the shape of the lower opening, this form is
sharply marked off from its nearest allies. Moreover, in both
Euparkeria and Omithosuchus the quadrato-jugal is a fair-sized bone
whose articulation with the jugal passes downwards and forwards,
and which passes up in front of the quadrate to meet the squamosal.
In this form, however, the quadrato-jugal is a comparatively small
flat bone which lies in the lower posterior corner of the fossa, and
whose articulation with the jugal passes downwards and backwards.
It lies under the jugal and overlaps part of the quadrate. There is
no foramen between it and the quadrate.

The quadrate is a long, strongly developed, fixed bone with a

102 Annals of the South African Museum.

somewhat expanded lower end. Its upper end is fixed between the
squamosal and the opisthotic. The external surface shows a
well-marked longitudinal depression at the lower end of the upper
half. The front edge of the bone is thin, the posterior border well
rounded.

The squamosal is a strong bone, articulating with the postorbital,
quadrate, parietal, opisthotic, paroccipital, and exoccipital. It forms
the outer posterior corner of the skull, i.e. half the outer and posterior
borders of the supraternporal fossa. It passes over on to the
occipital plate, and takes part in the border of the lateral occipital
foramen. The articulation between the squamosal and the opis-
thotic is pierced by a large oval foramen.

The parietal has a strong median crest which divides posteriorly
and forms there the upper border of the occipital plate. The bone
passes over this lateral crest and forms part of the occipital plate,
articulating with the supraoccipital and squamosal, and taking part
in the border of the lateral foramen. Anteriorly the bone articulates
with the frontal, laterally with the squamosal, and below with the
opisthotic and alisphenoid. As in Ornithosnclius there is no inter-
parietal.

The occipital plate is fairly small, and consists of the supra-, ex-,
and basi-occipitals, parietal, and squamosals. The supraoccipital
forms the upper median portion of the plate and enters into the
upper edge of the foramen magnum. The paroccipital forms no
part of the border of the foramen magnum. Its outer corner is bent
almost horizontally and rests on the posterior prolongation of the
squamosal, with which bone it has a long curved articulation. The
basioccipital is small, but it seems to form most of the incomplete
condyle. The opening for the 12th nerve is in the exoccipital about
2 mm. from the lateral border of the foramen magnum.

The palate is incomplete. The anterior prolongations of the
pterygoids are long, and there is a well-defined median groove.
The internal nares are not seen ; but they must have been con-
siderably in advance of the postpalatal vacuities. In both these
features this form agrees with the features displayed by Erpetosuchus
and Proterosiichus two forms which von Huene groups together as
the Pwtcrosuchia, and differs from Ornithosuchus and the Phytosaurs.
The palatine has a long articulation with the pterygoid and forms
the inner and anterior borders of the post-palatal vacuity, while its
outer border, and probably the posterior, is formed by the trans-
palatine. The transpalatine has a suture with the jugal on its
outer side,

Investigations in Sutttli African Fossil Heptilia ami Amphibia. 103

The lower jaw is incomplete. About 100 mm. of the right ramus
is anchylosed to the anterior 40 min. of the left ramus, while a small
portion of the back of the jaw is attached to the left quadrate. I am
thus unfortunately unable to say whether or not a fossa was present.

FIG. 17. Sphenosuehus acutus.

A. Outer view of right clavicle.

B. Inner view of interclavicle.

C. Inner view of left humerus.

D. Outer view of right humerus.

All figures x .

Remnants of 12 teeth are preserved on the right dentary, and of
5 or 6 on the left. Of these latter the 4th is small and nearly
complete. It shows that the teeth were flattened, pointed, and
simple, provided with serrations on the anterior border. The

104 Annals of the South African Museum.

dentary forms the whole of the anterior half of the outer surface
and thickens in front to meet its neighbour over the whole of the
syrnphysis. The splenial forms a large part of the inner surface
of the anterior half of the jaw, but takes no part in the
symphysis. It has a straight articulation with the dentary along
the lower border of the jaw. The fractured end shows a small
portion of the angular lying within the splenial and dentary. There
is a postarticular process passing behind the extremity of the
quadrate.

The anterior caudal vertebrae have been displaced and the centra
are missing. The elements of the atlas cannot be distinguished,
although a small curved bone lying on the right exoccipital is
probably a part of the pro-atlas. The dorsal spine of the axis
is present. It is 30 mm. long, higher in front than behind, and
overlaps the 3rd cervical. This latter shows well-developed, strong
anterior zygapophyses, shorter postzygapophyses, a straight flattened
dorsal spine, and a well-marked neural canal. The dorsal spines
of the 4th and 5th cervicals are also present. They are like that of
the third cervical, slightly expanded at the crest, with a shallow
groove running down the posterior border. The anterior ribs are
double-headed.

Both scapulae, both clavicles, and an interclavicle are present, as
\vell as both humeri.

The scapula is 81 mm. long and expanded both at its proximal
and distal ends. The width of the distal end is 43 mm., of the
proximal end 40 mm., while the narrowest part of the shaft which
occurs just above the proximal expansion measures but 15 mm.
in width. There is no acromion process.

The clavicles are comparatively large, thin bones, whose shape
can best be understood from the figure. They have a length of
77 mm., and a mean width of about 12 mm.

The interclavicle is a small narrow elongate bone with a very
prominent median ridge running throughout its whole length on
the inner side.

The humerus is 113 mm. long. The proximal end is broad
and flattened. The delto-pectoral crest is about 9 mm. long.
In the inner view the proximal expansion is seen to be hol-
lowed out between the delto-pectoral crest and a well-marked
ridge which runs down on the inner side from the proximal condyle,
which is slightly swollen and rounded. The shaft is slightly curved,
and oval in cross-section ; at the narrowest part the greatest
diameter of this oval has a length of 10 mm. The distal end

I it rt'xt /Cations of SontJi African Fossil Ec^tUni and Aiiijtliihia. 105

is but slightly expanded and shows two distinct rounded condyies,
of which the inner is much the larger.

On the slab of stone which contained the hurneri there is a
slender bone consisting of a straight shaft with slightly expanded
ends, the whole being 64 mm. long. The middle of the shaft
has a greatest diameter of 6 mm. This is probably the radius.
There are also three incomplete metacarpals in juxtaposition with
one another, and a portion of what is probably the ulna.

There is also preserved a tibia and the distal third of a fibula from
the same limb. The proximal end of the tibia is expanded and has
a pronounced swollen condyle on the inner side. The distal end is
slightly expanded, while the shaft is long and subcircular in section.
The distal end is 15 mm. wide, the shaft 9 mm., and the proximal
end 29 mm.

Typo. Skull and part of lower jaw with bones of the shoulder
girdle and fore-limb. (S.A. Mus. Cat. No. 3014.)

Locality. Paballon, Mount Fletcher, Cape Colony.

Horizon. Eed Beds. (Stormberg series.)

9

( 106 )

EXPLANATION OF PLATES.

PLATE XII.

FIG.

1. Top view of skeleton of Rhinesuchus senekalensis. x

'2. Top view of skull of Kliinesiichux senekalensis. x }.

3. Top view of skull of Rhinesuchus ichaitxi. < -,%.

4. Palatal view of skull of Rhinesuchus whaitsi. < ^.

PLATE XIII.

1. Top view of skull of Gorgonognathus loiujifrons. x f.

2. Top view of skull of Scyinnoi/nathus scrratidcns. x % nearly.

3. Side view of skull of Gorgonognathus longifrons. x f.

4. Side view of skull of Scymnognathus scrratidcns. x J nearly.

g. Afr. Mus. Vol. XII.

Plate XII

1, 2. HIIIXESUCHUS SENEKALENSIS (V. HOEPEN).
3, 4. RHINESUCHUX WHAITSI, BROOM.

Ann. S- Afr. Mus. Vol. XII.

Plate XIII.

1, 3. GORGONOGNATHUS LONGIFRONS, HAUOHTON.
2,4. SCYMNOGNATHUS KERRATIDENS, HAUGHTOX.

(107)

12. Foraminifera and Ostracoda from the Upper Cr<'fi-t'<>ns of Need's
Camp, Buffalo River, Cape Province. By FREDERICK CHAPMA.N,
A.L.S., F.R.M.S., etc.

With two Plates, XIV, XV, and Text-figure.

SOME little time since, I was favoured by Dr. L. Peringuey, Director
of the South African Museum, with samples of fossiliferous rock from
both the Upper and Lower Quarries at Need's Camp, Buffalo River,
with the request that I should examine them for microzoa.

Although the material did not promise to yield a large number of
the smaller organisms other than the polyzoa, after a lengthy search
a small but interesting series was obtained, the results of an examina-
tion of which are now given.

LIMESTONE FROM THE UPPER QUARRY.

General Characters. This rock is a fairly compact polyzoal lime-
stone with a crystalline matrix (see Text-figure 18). The polyzoa
forming the rock constitute about 50 per cent, of the whole, and
there are a few foraminiferal tests present. These can only be
examined in thin sections of the rock, owing to the compact structure
of the limestone, which prevents the extraction of the small shells by
fracture or pulverisation.

The following Foraminifera were met with in thin slices of this
rock :

MILIOLINA cf. CIRCULARIS, Bornemami sp.

Sections cut in various directions point to a comparison with this
species. It is a shallow water form, and has an e.^ensive geological
range.

MILIOLINA cf. FERUSSACII, d'Orb. sp.

An oblique section of what appears to be the test of this or a related
species occurs in the limestone. The chambers are evidently arranged
on the quinqueloculme plan. Foraminifera of this type are found in
all deposits ranging from the Lias to Recent.

10

108

Annals of ike South African Museum.

TEXTULARIA rf. GRAMEN, d'Orbigny.

The test is thin, arenaceous, and the septa alternate and strongly
arched.. The aboral end shows no tendency towards the spiral arrange-
ment" 1 as in Spiropleda, so that the above reference to d'Orbigny 's
textularian species seems most applicable. A widely distributed form.

Cf. ANOMALINA sp.

A thin-walled shell in median section of a form common in shallow
Avater sands of Cretaceous and Tertiary times (see Text-figure 18).

:-J

1'. C. 1'ltulo.

FIG. 18. Thin slice of limestone from the Upper Quarry, Need's Camp,
Buffalo River, showing fine-grained calcitic matrix with Polyzoa and
Foraminifera (c/. Anomalina sp.). x 28 diameters.

PULVINULINA sp. (of the P. eleijans group).

Vertical and tangential sections. The test shows redundant shell-
growth, as in some Cretaceous forms.

THE MICROZOA FEOM THE LIMESTONE OF THE
NEED'S CAMP LOWER QUARRY.

( T V/"'/W C In traders of the Rock. The rock from this locality is a
white, friable limestone, almost chalky in parts. Amongst the finer
s of the rubble occasional specimens of foraminifera may be

Foraminifera and Ostracoda. 109

detected. Their tests are, in many cases, badly corroded, but a
sufficient number of specimens could be secured to obtain the dia-
gnostic characters for the determination of the species.

FORAMINIFERA.

FAMILY LITUOLIDAE.

GENUS HAPLOPHRAGMIUM, Reuss.

HAPLOPHRAGMIUM NEOCOMIANUM, Chapman.
(Plate XIV, fig. 1.)

Ha/plophragmium neocomianum, Chapman, 1894, Quart. Jom'ii. Geol.

Soc., vol. 1, p. 695, pi. xxxiv, figs. 2a, b.
Idem, 1904, Annals S. African Mus., vol. iv, pt. v, p. 223 ; pi. xxix,

fig. 1.

One well-defined specimen occurs in the limestone from Need's
Camp. It has been previously recorded from the Rhaetic, Aptian, and
Cretaceous beds of Europe, and more recently from the Cretaceous of
Pondoland.

FAMILY TEXTULARIIDAE.

GENUS SPIROPLECTA, Ehrenberg.

SPIROPLECTA ANCEPS, Reuss sp.
(Plate XIV, figs. 3, 4.)

Texhdaria anceps, Reuss, 1845, Verstein. d. bohm. Kreideform., vol. i,

p. 39, pi. viii, fig. 79 ; pi. xiii, fig. 2.
Idem, 1860, Sitz. d. k. Akacl. Wiss. Wien, vol. xl, p. 234, pi. xiii, figs.

2a, &.
Spiroplecta anceps, Rss. sp., Chapman, 1892, Journ. R. Micr. Soc.,

p. 751, pi. xi, fig. 6.

This species is perhaps the most abundant foramiuifer in the Need's
Camp limestone. It is subject to great variation. Some of the short
and wide forms resemble 8. gramen, d'Orb. sp., but for their more
numerous septation. In the majority of cases the spiroplectine com-
mencement is almost concealed, as in fig. 3 ; whilst in others it is
partially unrolled and conspicuous, as in fig. 4.

10

110 Annals of the South African Museum.

8. anceps is a well-known Cretaceous species, occurring in the Chalk
of England, Westphalia, and Bohemia, and in the Gault of Folkestone.

SPIROPLECTA ANCEPS, Eeuss sp., var. INFRACTA, var. nov.
(Plate XIV, fig. 5.)

Description. Test formed in two stages, the first with an incon-
spicuous spiroplectine commencement and a short textularian series, as
in S. anceps, from which proceeds an irregularly septate and coarsely
formed test with rougly dentate margins.

Total length, T175 mm. ; length of primary test, O351 mm. ; width
of test at oral extremity, O675 mm.

SPIROPLECTA DEFLATA, sp. nov.
(Plate XIV, fig. 2.)

Description. The test of this species is rather irregularly con-
structed of arenaceous particles, but the general plan of structure can
be made out, especially when moistened ; it then shows the textularian
series with chambers much deflated and with a rude spiral series at
the aboral end of the test.

Two specimens were found, of nearly equal size. The figured
specimen has a length of - 919 mm.

This species bears the same relationship to 8. praelonga, Eeuss sp.*
that Gaudryina dispansa, Chapman f does to G. pupoides, d'Orb.

FAMILY NODOSARIIDAE.

GENUS NODOSAEIA, Lamarck.

NODOSARIA ZIPPEI, EeuSS.

(Plate XIV, figs. 6, 7.)

Nodosaria zippei, Eeuss, 1845, Verstein. bohrn. Kreideform., pt. i,
p. 25, pi. viii, figs. 1-3.

N. zippei, Ess., Chapman, 1904, Annals S. African Mas., vol. iv,
pt. v, p. 226, pi. xxix, fig. 6. (For further references see that article.)

The present examples are fragmentary but show sufficient characters
to justify their reference to the above species. The costation is rather
close and more numerous than is generally the case with the European

* Textularia praelonga, Eeuss. Verstein. bohm. Kreideform., vol. i, 1845,
p. 39, pi. xii, fig. 14.

t Journ. Eoy. Micr. Soc., 1892, p. 753, pi. xi, figs. 10a, b.

Foraminifera and Ostracoda. Ill

Chalk specimens ; although Reuss figures one example more comparable
with the present in this respect.

N. zip-pel is recorded from the chalk of Bohemia, Westphalia, the
Upper Bavarian Alps, Maastricht, and the Isle of Riigeu ; also from the
G-ault (Albian) of France and England, and in the Cambridge Green-
sand (Albian in part).

NODOSARIA SULCATA,

(Plate XIV, fig. 8.)

Nodosaria sulcata, Nilsson, 1825 (1826), K. Vet. Ak. Handl., p. 341.
Idem, 1827, Petrif. Suecana, p. 8, pi. ix, figs, la, A, B (error for 19).
N. sulcata Nilsson, Hisiuger, 1837, Lethaea Svecica, p. 33, pi. x, figs.

4a, I.

Eeuss, 1845, Verstein. bohni. Kreideform., pt. i, p. 26, pi. xiii, fig. 17,
Idem, 1855, Zeitschr. deutsch. geol. Gesellsch., vol. vii, p. 269, pi. viii.

fig. 146.
Sherborn and Chapman, 1889, Journ. Roy. Micr. Soc., p. 486, pi. xi,

fig. 24.
Egger, 1899, Abhandl. k. bayer. Akad. Wiss., Cl. ii, vol. xxi, Abth. i,

p. 75, pi. viii, fig. 24.

This is a form resembling the more extensively ranging Nodosaria
obliqua,Ij. sp., but with the striate-costate ornament disposed vertically
instead of obliquely. It is almost essentially an Upper Chalk species,
but has also been found in the Lower Tertiary (London Clay).

The Cretaceous localities for this form are the Chalk of the Paris
Basin, Lemberg, Riigeu, the Upper Bavarian Alps, and Bohemia.

GENUS CRISTELLARIA, Lamarck.

CRISTELLARIA PARALLELA, Reuss.
(Plate XIV, fig. 9.)

Cristellaria parallela, Reuss, 1862 (1863), Sitzungsb. d. k. Akad. Wiss.

Wien, vol. xlvi, p. 67, pi. vii, figs. 1, 2a, b.

Bertheliu, 1880, Mum. Soc. Geol. France, ser. 3, vol. i, Mem. v, p. 56.
Chapman, 1894, Quart. Journ. Geol. Soc., vol. 1, p. 712.
Idem, 1894, Journ. Roy. Micr. Soc., p. 647, pi. ix, figs. 5a, b.

This elongate and parallel- sided variation of the C. crepidula type is
well known from Cretaceous strata. It has occurred in the Lower
Greensand (Aptiau) of Surrey, also in the Gault (Albian) of
Folkestone, France and Germany.

112 Annals of the South African Museum,

CRTSTELLARIA INTERMEDIA, Eeuss.
(Plate XV, fig. 10.)

Cristellaria intermedia, Eeuss, 1845, Verstein. bohm. Kreideform.,pt. i,
p. 33, pi. xiii, figs. 57, 58 ; pt, ii, p. 108, pi. xxiv, figs. 50, 51.

Altli, 1850, Haidmger's Naturw., Abhandl. iii (ii), p. 267, pi. xiii,
fig. 23.

An. interesting species of the vaginuline or compressed forms of
Cristellaria, but having acute, or even keeled, margins. This is
typically a Cretaceous form, found in the Lemberg and Bohemian
Chalk ; variations of this species have been recorded by Eeuss * from
the Greensand (Cenomaniau) of New Jersey, and by Deecke t from
the Oxfordian of Moutbeliard, France. In the former the test is
inflated and the segments higher as in Eeuss' C. planiuscula, whilst
the latter has the series of chambers more closely inrolled.

CRISTELLARIA SECANS, Eeuss.

(Plate XV, figs, llrt, b.)

Cristellaria secans, Eeuss, 1860, Sitzungsb. d. k. Ak. Wiss. Wien,
vol. xl, p. 214, pi. ix, figs. 7a, b.

Cristellariae of the above type, with prominent and parallel sutural
ribs, are fairly common in the washings of the limestone from the
Lower Quarry, but in many cases so corroded that they can only be
generally identified as probably belonging to this species. The figured
specimen is a good example of Eeuss' C. secans, which that author
found in the Gault Clay of the Ehine. A variety of this species,
angulosa, has also occurred in the Folkestone Gault.

FAMILY ROTALIIDAE.

GENUS DISCOEBINA, Parker and Jones.

DISCORBINA PILEOLUS, d'Orbigny sp.
(Plate XV, figs. 13, 6.)

Valvulina pileolus, d'Orbigny, 1839, Foram. Anie'r. Mt'rid., p. 47, pi. i,

figs. 15-17.
Discorbina pileolus, d'Orb. sp., Parker and Jones, 1865, Phil. Trans.,

* Sitzungsb. d. k. Ak. Wiss. Wien, vol. xliv, 1861 (1862), p. 336, pi. viii,
figs. 2 a, b.

t Mem. Soe. d'Eumlation de Montbeliard, vol. xvi, 1886, p. 30, pi. ii, figs. 19,
19a.

Foraminifera and Ostracoda. 113

vol. civ, p. 385 ; Brady, 1884, Rep. Chall., vol. ix, p. 649, pi.
Ixxxix, figs. 2-4 ; Chapman, 1894, Quart. Journ. G-eol. Soc., vol. 1,
p. 719 ; Idem, 1896, Journ. Roy. Micr. Soc., p. 591, pi. xiii, figs.
14a, b.

The figured specimen is practically identical with] those of the
Cretaceous and Neocomian in England. It is also found in Tertiary
strata, and persists to the present day.

GENUS TRUNCATULINA, cVOrbigny.

TRUNCATULINA SCHLOENBACHI, Reuss sp.
(Plate XV, figs. 12a, 6.)

Rosalina schloenbachi, Reuss, 1862, Sitzuugsb. d. k. Ak. Wiss. Wien,

vol. xlvi, Abth. i, p. 87, pi. xi, figs. 5a-c.
Discorbina schloenbachi, Reuss sp., Egger, 1899, Abhandl. k. bayer.

Akad. Wiss., Cl. ii, vol. xxi, Abth. i, p. 164, pi. xviii, figs. 19-21.
Chapman, 1904, Annals S. Afr. Mus., vol. iv, pt. v, p. 229, pi. xxix,

figs. 16, 16a.

In some respects, as in the embracing character of the last whorl
of chambers on the inferior face, this species resembles a discorbine
form, but as already pointed out (see this publication, 1904, p. 230),
it appears naturally to belong to the genus Truncatidina on account
of its finely perforated or smooth test. It is a typical Cretaceous
form.

TRUNCATULINA UNGERIANA, d'Orbigny sp.
(Plate XV, figs. I6a, b.)

Rotalina iingcriana, d'Orbigny, 1846, Forani. Foss. Vienne, p. 15 1 /,

pi. viii, figs. 16-18.
Truncatulina ungeriana, d'Orb. sp., Brady, 1884, Rep. Chall. vol. ix,

p. 664, pi. xciv, figs. 9a-c.
Egger, 1899, Abhandl. k. bayer. Ak. Wiss., Cl. ii, vol. xxi, Abth.

p. 150, pi. xix, figs. 4-6.
Chapman, 1912, Mem. Nat. Mus. Melbourne, No. 4, p. 43, pi. vi,

figs. 2a-c.

Amongst the many modifications of this species, which ranges from
the Lower Cretaceous to recent deposits, the present examples agree
most nearly with the figured Chalk specimens in having a thicker test
and more umbonate facial aspect.

114 Annals of the South African Museum.

GENUS ANOMALINA, Parker and Jones.

ANOMALINA AMMONOIDES, Eeuss sp.
(Plate XV, fig. 14.)

Rosalina ammonoides, Reuss, 1845, Verstein. bohrn. Kreidef., pt. i
p. 36, pi. xiii, fig. 66 ; pi. viii, fig. 53.

Anomalina ammonoides, Ess. sp., Periier, 1897, Foratninifery Vrstev
Belohorskych (Palaeontograpliica Boliemiae, No. iv), p. 72.

Chapman, 1898, Journ. Roy. Micr. Soc., p. 4, pi. i, figs. 5a-c. (For
extended synonymy see last quoted paper.)

This species has been recorded from beds as old as the Neocomian.
It is one of the commonest of Cretaceous rotalines, and is found in
dredgings at the present day. The specimen before us is a perfect
and well-developed shell.

GENUS PULVINULINA, Parker and Jones.

PULVINULINA KARSTENI, ReUSS Sp.

(Plate XV, figs. 15a-c.)

Eotalia Jcarsteni, Reuss, 1855, Zeitschr. d. deutsch. geol. Gesellsch.,

vol. vii, p. 273, pi. ix, fig. 6.
Pulvinulina Jcarsteni, Rss. sp., Chapman, 1892, Quart. Journ. Geol.

Soc., vol. xlviii, p. 517.
Egger, 1899, Abhandl. d. k. Akad. bayer. Wiss., Cl. ii, vol. xxi, Abth. i,

p. 161, pi. xx, figs. 32-34.

A very common species in the present collection from the Lower
Quarry. It is characteristically an Upper Chalk form, but is also
known from the Lower Cretaceous and a few Tertiary deposits. In
recent seas a more biconvex variety is known from widely separated
areas, both in the northern and southern hemispheres. The speci-
mens now dealt with are most comparable with those figured by Reuss
from the Upper Cretaceous of Europe.

OSTRACODA.

Family BAIKDIIDAE.

GENUS BAIRDIA, McCoy.
BAIRDIA SUBDELTOIDEA, Minister sp.

Cythere subdeltoidea, Munster, 1830, Jahrb. f iir Min. etc., p. 64, No. 13 ;
1835, p. 446.

Foraminifera and Ostracoda. 115

Cytherina suldeltoidea, Miinst. sp., Ronier, 1838, Jalirb. fur Min. etc.,

p. 517, pi. vi, fig. 16.
Reuss, 1845, Verstein. bohm. Kreideform., pt. i, p. 36, pi. v, fig. 38 ;

pt. ii, p. 104.
Bnirdia subdeltoidea, Miinst. sp., Rupert Jones, 1889 (1890), Mon.

Cret. Entom., Suppl. (Pal. Soc.), p. 5, pi. ii, figs. 31-34.

Examples of the above species in the Need's Camp washings exactly
compare with the types figured from the Cretaceous of England and
"North Germany. One of the species shows a deep sinus on the ventral
side of the valves similar to that figured by Rupert Jones (loc. cit.
1890) on pi. ii, fig. 34.

B. subdeltoidea is also met with in Tertiary strata in Europe, and
a near relative is the living B. foveolata, G. S. Brady,* from Australia,
the West Indies, Crete, Serpho, Hongkong Harbour, Admiralty
Islands, etc.

BAIRDIA SUBDELTOIDEA., Minister sp. var. AEQTJALIS, var. nov.
(Plate XV, figs. 17 a, &.)

Description. This variety is distinguished from the type form in its
more ovate shape, as seen from the side of the valve ; the anterior part
being less broadly rounded and the posterior extremity less prolonged.
It thus comes nearer to B. amygdaloides, G. S. Brady.f a Miocene to
Recent species.

BAIRDIA AFRICANA, sp. nov.
(Plate XV, figs. 19a-c.)

Description. Carapace ovoid or pear-shaped, moderately tumid.
Valves seen from the side highest above the middle, anteriorly trun-
cately rounded towards the dorsal margin ; posteriorly tapering and
sub-acuminate ; ventral border widely curved and with a steep face.
Edge view of carapace subovate and with a steep face, thickest about
the middle. End view subcordate. Surface smooth or faintly pitted.

Dimensions. Length, '702 mm. ; height, '439 mm. ; thickness of
carapace, '358 mm.

The nearest form to the above is the before-mentioned B. amyg-
daloides, G. S. Brady, but which differs from B. africana in having a
more salient dorsal border.

* B. foveolata, G. S. Brady, Les Fonds de la Mer, vol. i, 1867, p. 56, pi. vii,
figs. 4-6. Id., Kep. Chall. Zool. vol. i, pt. iii, Ostracoda, 1880, p. 55, pi. viii,
figs. 8a-f, and figs. 2a-f.

t Hep. Ghall. torn, cit., p. 54, pi. ix, figs. oa-/ ; pi. x, figs. 2a-c.

116 Annals of the South African Museum.

Fam. CYTHEBIDAE.

GENUS CYTHEKE, Miiller.

CY.THEKE POSTCULTBATA, sp. nov.

(Plate XV, figs. 18, 6.)

Description. Valve subrhomboidal, rounded in front, tapering
behind to the acuminate extremity; dorsal edge straight, ventral
obliquely truncated. Surface higher at the post-ventral region, and
sloping away to the front and dorsal border. Near the ventral edge
in the posterior region is a keel-shaped prominence curving inwards
and merging into the surface about the middle of the ventral edge of
the valve. Surface relieved with a few irregularly disposed pittings.

Dimensions. Length, '527 mm.; height, '3 mm.; depth of valve

08 mm.

Relationships. There appear to be no closely related forms, either
fossil or recent, with the exception of Dr. G. S. Brady's " Challenger"
species, Cythere cytheropteroides* a form dredged at 150 fathoms from
the Cape of Good Hope. The present species, however, is more pyri-
form in outline.

SUMMARY OF EESULTS.

Upper Quarry.

The meagre series of Foraniinifera obtained from this locality does
not afford any data which can be used to decide the age of the deposit.
The genera found, viz. MiUolina, Textularia, Anomalina, and Pulvinu-
lina, are all widely distributed at the present day, and the species,
where they could be determined, denote moderately shallow water con-
ditions. Further than this, they are components of similar faunas as
far back as the Cretaceous and even earlier Mesozoic strata. In thin
sections the particular types of polyzoa appear to indicate a relationship
to the like fauna of the Lower Quarry.

Loiver Quarry.

The following list of species of Foramiuifera and Ostracoda throws
a decided light on the age of this bed.
Foraniinifera :

Haplophragmiiim neocomianum, Chapm.
Spiropleda anceps, Eeuss sp.

* Eep. ChalL, vol. i, Zool., pt. iii, Ostracoda, 1880, p. 78, pi. xv, figs. 5a-d.

Foraminifera, and Ostracoda. 117

Spiroplecta anceps, var. infracta, nov.

,, deftata, sp. uov.

Nodosaria zippei, Reuss.

,, sulcata, Nilsson.
Gristellaria parallela, Reuss.
,, intermedia,, Reuss.
,, secans, Reuss.
Discorbina pileolus, d'Orb. sp.
Tnmcatulina scJiloenbachi, Reuss sp.

,, ungeriana, d'Orb. sp.

Anomalina ammonoides, Reuss sp.
Pulvimdina karsteni, Reuss sp.
Ostracoda :

Bairdia subdeltuidea, Miinster sp.

,, var. aequalis, nov.

,, africana, sp. nov.
Gythere postcidtrata, sp. uov.

Amougst the Foraminifera of generally Cretaceous aspect may be
noted Haplopliragmium neocomianum and tfj>!rr>i>Iecta anceps. The
species which lend to the deposit an Upper Cretaceous appearance
.are Nodosaria zippei (a quite restricted form), N. snlcata (almost in-
variably Upper Cretaceous), Cristellaria /ii/nil/^ln, (_'. intermedia, and
C. secans. This latter species is in exactly the condition of growth and
size as found in the typical Chalk faunas of Europe, and is one of the
commonest species in the limestone from Need's Camp Lower Quarry.
All the rotalines, whilst having an extensive geological range, from the
Cretaceous to the present time, are typical of the Upper Cretaceous
also, and their present occurrence as to size and development favours
I he idea of their Cretaceous age.

Of the Ostracoda only one species, Bairdia subdeltoidea, is available
for comparison, as the remainder are new forms. The examples
referred to the above-named specific form (a Cretaceous and early
Tertiary species) are those of typical Cretaceous valves.

(118)

EXPLANATION OF PLATES.

(All figures magnified 36 diameters.)

PLATE XIV.

FIG. 1. Haplophragmium neocomianum, Chapman.
2. Spiroplecta deflata, sp. nov.
., 3. anceps, Eeuss sp.

4. An example with a well- developed aboral

extremity.

5. var. infracta, var. nov.

,, 6. NoJosaria zippei, Eeuss.

.. ' >? >

., 8. sulcata, Nilsson.

,, 9. Cristellaria parallela, Eeuss.

PLATE XV.

FIG. 10. Cristellaria intermedia, Eeuss.

11. secans, Eeuss : a, lateral aspect ; I, oral aspect.

12. Truncatulina schloenbachi, Eeuss sp. : a, superior aspect ; b, inferior

aspect.

13. Discorlina pileolus, d'Orb. sp.: o, superior aspect ; I, inferior aspect.

14. Anomalina ammonoides, Eeuss sp.

15. Pulvinulina karsteni, Eeuss sp. : a, superior aspect ; I, inferior aspect :

c, peripheral aspect.

16. Truncatulina imgeriana, d'Orb. sp. : a, superior aspect; b, inferior

aspect.

17. Bairdia subdeltoidea, Miinster sp., var. aeqtialis, var. nov. : a, left valve,

lateral aspect ; b, edge view, ventral aspect.

18. Cythere postcultrata, sp. nov. .- a, carapace from right side ; b, edge

19. Bairdia africana, sp. nov.: a, carapace from right side; b, edge view,

ventral aspect ; c, end view of carapace.

Ann. S. Afr. Mus. Vol. XII.

Plate XIV.

>

1

I

r 9 *

FOBAMINIFERA. NEED'S CAMP, BUFFALO E.

F.C. <lrl.

Adlard <t West Neu'tnun.

Aim. S. Afr. Mus. Vol. XII.

Plat.- XV.

16a

18,

15 a

\ 5 c

186

11 n

13

19 a

t
f

13 b

12,

19c

,

FOKAMINIFERA : FlGS. 10-16. OsTBACODA : FlGS. 17-19. NEED'S CAMP, BUFFALO K.

F.C. del.

,/ \\'fxt Xt'ii-niaii.

(119)

13. Some New Species of Anomodontia (Reptilia). By R. BROOM,
D.Sc., C.M.Z.S., and S. H. HAUGHTON, B A., F.G.S., Assistant
Director.

(With 6 Text-figures.)
GENTJS DICYNODON, Owen.

DlCYNODON COESTORPHINEI, Sp. nov.

A small skull and lower jaw collected by the Eev. J. H. Whaits at
Graaff Eeiuet (S.A. Mus. Cat., No. 3337) seems to belong to an
uudescribed species, although in general appearance it partakes some-
what of the nature of Dicynodon lutriceps.

The most noteworthy characters are the shortness of the beak, the
shape of the postfrontal, and the position of the pineal foramen. The
skull is depressed. The intertemporal bar is wider than the inter-
orbital region. The orbit is fairly large, looking more upwards than
outwards. The postfrontal extends outwards along the postorbital
bar somewhat in the manner seen in Eocyclops longus. The preparietal
is long and forms the anterior border of the pineal foramen, which is
very far back half way along the fairly long parietal bar. The
portions of the parietals behind the foramen are thus short and broad,
overlapped for more than half their width by the postorbitals and
truncated posteriorly by the interparietal. The iuterparietal in its
upper portion has a strong median ridge. The occipital condyle is of
the tripartite type.

The specimen is tuskless.
The chief measurements are :

Greatest length . . .160 mm.

Greatest width ... . ab. 150 ,,

Basal length . . . . . . . 143 ,,

luterorbital width . . . . . 25 ,,

Intertemporal width . 30 ,,

Snout to front of orbit . 40 ,,

Length of preparietal . . . 21 ,,

11

120

Annals of the South African Museum.

From Dicynodon lutriceps this type seems to differ in having the
intertemporal width greater than the interorbital, in not having the
parietals so fully covered by the postorbitals, in the position of the pineal
foramen, and in the size of the preparietal.

The form is also strongly reminiscent of D. mustoi ; but the latter
is somewhat more slenderly built, the pineal foramen is further
forward, and the postfrontal is a much larger bone.

The chief resemblance, however, is to Owen's Oudenodon laini, as
far as can be judged from the figure of the latter. The two agree in
general shape, in the relation between the interorbital and inter-
temporal width, and in the position of the pineal foramen. The species

Po.F.

FIG. IP. Dicynodon corstorpMnei, Ur. and Htn. Type, No. 3337. Tem-
poral and frontal regions. x 7.

Oudenodon laini cannot, however, stand. In the first place, we know
that Oudenodon is but the female of Dicynodon ; Owen had previously
described another specimen as the type of Dicynodon baini ; and,
lastly, Dicynodon baini is but a synonym for D. tigriceps.

Type. Skull without tusks, and lower jaw (S.A. Mus. Cat., No.
3337).

Locality. Heuning Nest Krantz, Graaff Reinet, C.P.

Horizon. Lower Beaufort Beds, Endothiodon zone (?).

DICYNODON CAVIFRONS, sp. nov.

This new species is founded on a skull from Fraserburg, C.P.,
collected by the late T. Bain, Esq.
The chief measurements are :

Some Neiv Species of Anomodontia

121

Greatest length (oblique)
Greatest width
Interorbital width .
Intertemporal width
Width across nasals
Width between nostrils .
Width between canines .
Basal lenu'th .

215 mm.

:ib. 200

31

. 37

42

ab. 25
30

. 172

The orbits are triangular, their superior borders considerably shorter
than the others, and lie entirely in the anterior half of the skull.
The frontal is considerably hollowed out, and is narrower -than the
flattened parietal region. The snout is short and weak. The tusks

P'iu. 20. Dicynodon cavifrons, Br. and Htn.

rejjion. x -J.

Type, No. 747. Pre parietal

project downwards and are midway below the nostrils and orbits.
The postorbitals are large, and almost meet in the middle line above
the parietals. The postfroutal is an elongate triangular bone. The
squamosal extends far back behind the occipital plate.

Type. Skull lacking outer arches (S.A. Mus. Cat., No. 747).

Locality. Fraserburg, C.P.

Horizon. Lower Beaufort Beds, probably Cistecephalus zone.

DICYNODON BOGERSI, sp. nov.

Some years ago Dr. Rogers collected in the Thee Kloof, Nieuweveld,
C.P., an almost complete skull and lower jaw which was regarded as a
male specimen of D. Icolbei. Recent examination has, however, led us
to consider that it may well be taken as the type of a new species.
The skull is in a good state of preservation, and shows most of the
sutures of the top of the skull.

122

Annals of the South African Museum.

The snout is slightly longer, and the nasal bosses are not so well
developed as in D. kolbei. The orbit is both relatively and absolutely
shorter, although of similar shape. The parietal bar also differs The
ridges, instead of approximating most closely to each other in the
posterior half of the bar as in D. kolbei, are closest at the front of the

FIG. 21. Dicynodon rogersi, Br. and Htn. Type, No. 2356.

X 4.

bar and diverge gradually posteriorly. The postfrontal does not
narrow so rapidly as in D. kolbei and consequently covers a greater
area. The preparietal has its two sides parallel and not convergent
posteriorly, and forms a large portion of the anterior half of the
parietal foramen. The temporal fossa is long and regularly oblong in
shape. The preparietal region is hollowed out.

A rare character displayed by the type is the feeble and anteriorly
directed tusks. They arise directly below the nostrils.

Some New Species of Anomodontia (Reptilia).

123

The chief measurements of the skull are
Greatest length (oblique)
Greatest width
Interf rental width
Intel-temporal width
Basal length .
Width across nasals
Width between nostrils .

ab. 290 mm.

228

50

45

ab. 230

48
28

FIG. 22. Preparietal region of same.

Type. Skull (S. Af. Mus. Cat., No. 2356).

Locality. Thee Kloof, Nieuweveld, C.P.

Horizon. Lower Beaufort Beds, bottom of Cistecephalus zone.

DlCYNODON PTGMAEUS, sp. 11OV.

This is one of the smallest known species of Dicynodon. As two or
three specimens were obtained from the same locality of about similar
size, it seems probable that the specimen represents a small species
rather than a young individual.

The species is characterised by the relatively great width of the
anterior ends of the parietals, which causes the preparietal region to
be nearly twice as wide as the frontal.

The frontals are large, and extend well back by the sides of the
preparietal. The postfrontals are very slender. The postorbitals are
large, the posterior portions being broad and rather flat. The
squauiosals are relatively less developed than in most species.

124

Annals of the South African Museum.

The following are the principal measurements :

Greatest length 66 mm.

Greatest width ab. 50 ,,
Interorbital width . 1 1 ,,

Intel-temporal width (minimum) . . . 15 ,,

PIG. 23. Dicynodon pygmaeus, Br. and Htn. Typo, No. 2664. Natural size.

Type. Female skull (S. Af. Mus. Cat., No. 2664).

Locality. Duneclin, Beaufort West, C.P.

Horizon. Lower Beaufort Beds, Cistecephalus zone.

GENUS EMYDOPS, Broom.

EMTDOPS PLATYCEPS, sp. nov.

This little skull, while representing a new species, is not sufficiently
well preserved to enable us to say with perfect certainty that it belongs
to the genus Emyclops. It agrees, however, sufficiently closely with
the known species of Emydops to admit of its being placed here at
least provisionally.

There is a slender tusk which is directed downwards and forwards,
and at least one slender molar tooth. The parietal region is broad and
transversely concave. The relations of all the bones in the preparietal
region are as in previously known species of Emydops, but the pro-
portions differ considerably, as will be seen from the figure given.

Some New Species of Anomodontia (Reptilia).

125

The following are the principal measurements :

Greatest length ... 53 mm.

Greatest width . . . . 37 ,,

Interorbital width ... . 10

Intel-temporal width . . 15 ,,

Basal length . . . . . . 48 ,,

Minimum width across pterygoids 6 ,,

Width across palate between tusks . . 12 - 5 mm.

FIG. 2-l.Etnydopsi>latij<-L'}>.<!, Hr. and Htn. Type, No. 2667. Natural size.

Type. Skull (S. Af. Mm. Cat , No. 2667).

Locality. Dunedin, Beaufort West, C.P.

Horizon. Lower Beaufort Beds, Cistecephalus zone.

cavifrons (Dicynodon)
corstorphinei (Dicynodon.) .
DICYNODON .
EMYDOPS .

INDEX.

PAGE

120
119
119

124

platyceps (Emydops) .
pygmaeus (Dicynodon)
rogersi (Dicynodon) .

PAGE

. 124

. 123

121

(127)

14. Investigations in South African Fossil Reptiles and Amphibia
(Part 10). By S. H. HAUOHTON, B.A., F.G.S., Assistant
Director.

10. Descriptive Catalogue of the Anomodontia, u-if/i especial reference
to the examples in the South African Museum (Part 1).

(With 3 Plates and 20 Text-figures.)

THIS first part of the Catalogue of the Anomodontia contains forms
assigned to the genus Dicynodon and to other genera without molar
teeth allied to it, but omits Lystrosaurus. In view of van Hoepen's
recent additions to the named species of this latter genus, a
revision of the genus is necessary. The South African Museum collec-
tion contains a large number of very finely preserved Lystrosaiiru*
skulls ; but until further and somewhat fuller descriptions with
figures are given of van Hoepen's forms from Harrismith it is deemed
better to allow the revision of the genus to stand over.

The genus Dicynodon is an exceedingly troublesome one to work,
on account of the large variety of forms which have been assigned
to it, forms the types of which are now deposited in collections in
widely separated parts of the world, which types are occasionally
rather fragmentary and not particularly well defined. The genus,
in its broad sense, ranges from the bottom to the top of the
Beaufort Beds ; and although certain of the described forms have
been placed in other genera such as Eocyclops, Diictodon, Kanne-
meijeria, and the like, the relations between all the types are very
close.

Of recent years the collection in the South African Museum has
been enriched by a number of very good Auomodont skulls, mainly
through the efforts of the Kev. J. H. Whaits and of the Museum
collectors ; and these enable us to obtain a partial survey of the sub-
order, as the collection contains skulls from each zone of the Beaufort
Beds. Some of them can be correlated with some degree of certainty
with already-described forms. Others have characters distinguishing
them from these forms and, at the risk of adding to the nomenclature

128 Annals of the South African Mitseum.

and of increasing the size of the genus, these have been given specific
rank.

The question of the value of certain features as species indices is
one that is not of immediate moment. The root-idea underlying
descriptive work of this sort is to give an account of the variation
among the Auomodont reptiles so that we may obtain, if possible,
some idea of the lines upon which variation has taken place. For
this purpose it is better to take notice of small variations and to give
each different form a special name than to "lump" possibly different
animals together under the one name. It is probable that future work
may pi-ove many of our supposedly different forms to be the same ;
then these new names will have to disappear. But the special features
which each of them implies will remain, and the name will have served
its purpose.

Moreover, another purpose is served by this division of species. A
genus so prolific in individuals as was Dicynodon and so abundant in
varieties forms a good genus for the purpose of zoning ; and it may be
that the few zones now known among the Beaufort Beds may be
increased and more accurately denned by the use of the species of
Dicijiwdon. Even now, only one or two " species " seem to pass from
any one zone into the next ; and it may be found that each form is
confined to a comparatively small thickness of strata.

In the following pages reference is made mainly to such forms as
are represented in the collection of the South African Museum, but to
make the catalogue somewhat more complete the literature dealing with
other described forms is given, together with a brief description of the
tvpe, culled entirely from the original papers. In the absence of
facilities for seeing and studying these types mostly now in the
British Museum or in the American Museum of Natural History -
or of specimens which can be assigned to them, no conclusions have
been drawn as to their affinities with the forms which are more fully
discussed.

GENUS DICYNODON, Owen.

DICYNODON JOUBEBTI, Broom.
1905. Broom. Kec. Albany Mus., i, p. 331.

This is apparently a well-marked species, while all the specimens in
the S.A.M. collection range in size between fairly narrow limits. The
skull is small, with fairly large orbits and comparatively broad inter-
orbital and interorbital and interfrontal regions which are roughly of

South African Fossil Reptiles and Amphibia.

129

equal width. In the region of the postorbital bar the jugal is at least
as deep as the radius of the orbit. There is a large preparietal form-
ing the anterior border of the pineal foramen, but there does not
appear to be a postf rental in any of the specimens in which the sutures
are displayed. If present, it must have been small.

Thei'e is no evidence of a septomaxilla, which, if present, does not
appear on the face. The lachrymal is small.

FIG. 25. Dicynodonjoiiberti, Br. Skull No. 577. Preparietal region.

Natural size.

The chief measurements of a typical skull are :

Greatest length . . 108 mm.

Greatest width . . 76 ,,

Interf rental width . , 20 ,,

Intel-temporal width . 21 ,,

Basal length . 89 ,,

Width of palate between canines . 30 ,,

The basioccipital region of one specimen from the type locality
(S.A.M., Cat. No. 577) has been partially displayed. The tripartite
condyle is seen in section to be made up of the basioccipital and the

130

Annals of the South African Museum.

two exoccipitals. The foramen jugulare is bounded on the inside by
the exoccipital and looks as much downwards as backwai'ds. The
basioccipital is a shallow bone and the tubera lie very little below the
level of the bottom of the condyle. Between them the basioccipital
has a broad and shallow groove.

Type. Skull and lower jaw. (S.A. Mus. Cat. No. 695.)

Locality. 1 ' The Gouph."

Horizon. Lower Beaufort Beds. (Tapinoceplialus zone.)

DlCYNODON MEGALORHINUS (BrOOni).

1904. Broom, Oudenodon megalorhimis. Rec. Albany Mus., i, p. 180.

The type skull is of interest as being the first Dicyuodont in which
a postfrontal was clearly recognised.

FIG. 26. Dicynodon megalorhinus (Br.). Type No. 640. Natural size.

The bones of the upper part of the skull are clearly shown, and a
figure is given here for the first time. Noticeable features are the
long anterior extension of the parietals and the long, narrow, post-
f rentals.

Of the occiput and palate but little is seen. It is interesting to
note, however, that the feuestra ovalis faces almost completely
laterally and but very little downwards ; it is rather small. The
sutui'e between the basispheuoid and basioccipital is clear. The basi-
sphenoid, although passing back to form the anterior portion of the
tuber, takes no part whatever in the border of the fenestra ovalis.

South African Fossil Reptiles and Amphibia. 131

The lower border of the fenestra together with part, at least, of its
anterior border is formed by the basioccipital. The posterior border
is formed by the paroccipital, which also bounds the foramen jugu-
lare. This foramen looks almost directly backwards and is placed at
the level of the condyle. The carotid foramina lie together in the
pit of the basisphenoid just behind the pterygoids.

T#pe. Incomplete skull. (S.A. Mus. Cat, No. 640 )

Locality. Prince Albert Road Station, C.P.

Horizon. Lower Beaufort Beds. (Tapinocephalus zone.)

DICYNODON CORSTOEPHINEI, Br. & Htn.
1917. Broom & Haughton. Ann. S.A. Mus., xii, 5, p. 119, fig. 19.

The occipital condyle in this species is of the tripartite type. The
foramen for the exit of the IXth-XIIth nerves looks directly back-
wards and is on a level with the uotochordal pit in the condyle. The
post-temporal vacuity lies on the occipital plate just above the level
of the top of the condyle.

The basisphenoid tubera lie above the level of the quadrate, so that
the feuestra ovalis looks more downwards than outwards. The tubera
are not greatly below the level of the condyle and the groove between
them is as high as broad.

Type. Incomplete skull and lower jaw. (S.A. Mus. Cat, No.
3337.)

Locality. Heuning Nest Krantz, G-raaff Eeinet, C.P.

Horizon. Lower Beaufort Beds. (Top (?) of Endothiodon zone.)

DICYNODON CYCLOPS, sp. nov.

The specimen to be described, although somewhat reminiscent of
Dicynodon platyeeps and D. lutriceps, seems to differ sufficiently to be
classed as a different form. It consists of a skull showing all the
sutures of the top, having the following measurements :

Greatest length . . '225 mm.

Greatest breadth . . . 145 ,,

Length from beak to front of orbit 52 ,,

Antero-posterior diam. of orbit . 44 ,,

Interorbital width 33 ,,

Intertemporal width 34 ,,

Basal length . . . . . . 167 ,,

In side view the skull agrees with that of D. pardiceps in the
concavity at the preparietal region and the convexity of the parietal

132

Annals f tie South African Museum.

bar. The skull is considerably longer than broad. The orbits are
wholly in the anterior half of the skull, are moderately large and

FIG. 27. Dicynodon cyclops, Htn. Type. Xo. 3-447. * ?.

circular, and look more upwards than outwards. The snout is
moderately long, and the nostrils are midway between the beak and

FIG. 28. Dicynodon cyclops, Htn. Typo. Preparietal region. Natural

size.

the orbit, looking whelly laterally. The supranasal bosses are feeble,
as are the caniuiform processes. Between the nasal boss and the
supraorbital border is a depression. The supraorbital borders are
elevated so that the frontal region is concave. The postorbital bar is

South African Fossil Reptiles and Ampliih'ni. 133

slender. The parietal bar is just broader than the frontal region,
and convex longitudinally. The temporal openings are large.

The premaxilla passes on to the top of the snout, separating the
nasals except for a length of 15 mm., but forming no part of the upper
border of the nostril.

The septomaxilla is well seen on the face. It separates the nasal
from the maxilla and articulates posteriorly with both the prefrontal
and the lachrymal. The lachrymal is small. The prefrontal is large
and forms a large portion of the orbital border.

The arrangement of the bones around the pineal foramen is shown
in the figure. The pi'eparietal is small and forms half the border
of the pineal foramen. The postfrontal is short, but fairly broad
anteriorly.

Fio. 29. Dicynodon cy clops, Htn. Typo. Side view of snout.

From Dicynodon platyceps the species differs in having the inter-
temporal width equal to that of the frontal region, in the small
preparietal and in the detailed arrangement of the postfrontals,
froutals, and parietals. From Dicynodon lutriceps it differs in the
relation of the width of the frontal and parietal bars and in the fact
that the preparietal in the earlier type is apparently wholly in front
of the pineal foramen and enclosed by the frontals. No adequate
figure of the arrangement of the bones in D. lutriceps has ever been
given and I have not seen the type ; so it is impossible to say here
whether the two species are actually distinct.

Type. Skull. (S.A. Mus. Cat, No. 3447.)

Locality. Dalham, Graaff Keinet, C.P.

Horizon. Lower Beaufort Beds. (Endothiodon zone.)

DICYNODON FELICEPS, Owen.

1876. Owen. Cat, Foss. Kept, S.A., p. 45 ; pi. xliii.
1889. Seeley. Phil. Trans, clxxx B ; pi. x, fig. 3.

1:34

Annals of the South African Museum.

1890. Lydekker. Cat. Foss. Rept. & Ampliib. iv, p. 20.

1913. Broom. Bull. Amer. Mus. Nat. Hist., xxxii, p. 445.

1914. Broom. Phil. Trails. B206 ; pi. v, figs. 50, 51.

1915. Broom. Bull. Amer. Mus. Nat. Hist., xxv, 2, p. 129.

Founded on a skull with lower jaw. Smaller than D. lacerticeps.
Antorbital portion relatively shorter ; canine curves more downwards.
In older skulls orbits have distinctly triangular contour. Post-
orbitals meet above the parietals. Pineal foramen on a level with the
middle of the temporal fossa. Ramus of mandible deeper than in
D. lacerticeps.

Type in British Museum.

Locality. Fort Beaufort, C.P.

Horizon. Lower Beaufort Beds. (Endothlodon zone.)

DlCYNODON GEACILIS (BrOOlll).

1901. Broom, Oudenodon gracilis. Proc. Zool. Soc., p. 162.

The skull which forms the type of this species came from Pearston,
C.P. It is rather distorted by flattening. It is a small species

FIG. 30. Dicynodon gracilis (Br.). Type. No. 590. Preparietal region.

Natural size.

whose skull is relatively long and narrow. The orbits look outwards
and are overhung posteriorly by a nasal boss. The cheek is fairly
long. The eyes are small and are directed more outwards than
upwards. The postorbital bar is halfway along the skull. The inter-
orbital width is 25 mm., the Intel-temporal 18 mm. The frontal

South African Fossil Reptiles and Amphibia. 135

region is flat with a well-marked median ridge. The pineal foramen
is placed halfway along the parietal bar. In front of it is a narrow
preparietal reaching to the level of the front of the temporal vacuity.
There is no evidence of a postfrontal. The sutm-es are clearly marked
and the postfrontal must be absent, the postorbital passing forward
along the orbital border to meet the frontal.

The palate is long and narrow. The basicranial region is not well
displayed ; but the tubera did not extend far below the level of the
occipital condyle.

The maxilla forms nearly the whole of the long and somewhat
shallow cheek. The lachrymal is small as far as can be seen ; the
prefrontal is larger, and appears mostly on the top of the skull.

Type. Skull (S.A. Mus. Cat. No. 590.)

Locality. Pearston, C.P.

Horizon. Lower Beaufort Beds. (Probably top of Endothiodon
zone.)

DICYNODON ICTIDOPS, Broom.
1913. Broom. Bull. Amer. Mus. Nat. Hist., xxxii, p. 446 ; figs. 5, 6,

A small species. Skull narrow with regularly convex upper surface.
Orbits look more outwards than upwards. The nostrils are large and
rounded. The tusk is small and directed downwards. The preparietal
is large, postfrontal small. Frontals pass back to plane of front of
pineal foramen.

A skull and lower jaw (S.A. Mus. Cat. No. 1078) were obtained at
Klipfonteiu, Fraserburg, C.P. The sutures of the top of the skull
are indeterminable, but from its general appearance the skull belongs
to this species. It has been developed to show the basicranial region.
The occipital coudyle is large and of the tripartite type. The foramen
jugulare lies at the level of the bottom of the condyle and looks
downwards and backwards. The basisphenoidal tubera are large and
descend considerably below the level of the coudyle. The foramen
ovalis looks outwards and downwards, and most of its lower border
is, apparently, formed by the basioccipital. Between the two tubera
there is a very deep narrow depression.

Type in American Museum of Natural History.

Locality. Beaufort West Commonage, C.P.

Horizon. Lower Beaufort Beds. (Endothiodon zone.)

12

136 Annals of the South African Museum.

DlCYNODON I/0TKICEPS, Br.

1912. Broom. Proc. Zool. Soc., p. 870 ; pi. xcii, figs. 14-16.

Founded on an imperfect skull. Short beak ; broad concave frontal
region ; broad flattened intertemporal region, formed almost wholly by
the postorbitals. Interorbital width : intertemporal width : : 5:4.
Supranasal ridge ; small preparietal, lying entirely in front of the
pineal foramen (?).

Type in the American Museum of Natural History.

Locality. Kuil's Poort, Beaufort West, C.P.

Horizon. Lower Beaufort Beds, 300 ft. above the level of Beaufort
West,

DlCYNODON PABDICEPS, Owen.

1876. Owen. Cat. Foss. Kept. S. Afr., p. 42 ; pis. xxxviii, xxxix.

1889. Seeley. Phil. Trans., clxxx B, p. 244 ; fig. 2.

1890. Lydekker. Cat. Foss. Kept. & Amphib., iv, p. 21.

Founded on a nearly entire skull ; large, short, abruptly deflected
muzzle ; orbits triangular ; frontals wide ; parietal bar narrow ;
temporal fossa moderately wide ; preparietal apparently fairly small.

Type in British Museum.

Locality. Fort Beaufort, C.P.

Horizon. Lower Beaufort Beds. (Endothiodon zone (?).)

DlCYNODON PSITTACOPS, Br.

1912. Broom. Proc. Zool. Soc., p. 869 ; pi. xcii, fig. 17.

1915. Broom. Bull. Amer. Mus. Nat. Hist., xxv, 2, p. 139, fig. 28.

The type skull has the following characters : " Nostrils look almost
directly upwards. Upper part of nasals thickened, and prefrontal
region of orbital margin also elevated. Frontal region broad and flat.
Parietal foramen situated in an elevated preparietal. Postorbitals
appi'oach each other behind the foramen and nearly touch, forming a
parietal ridge. The tusk is small and directed forwards and down-
wards."

A specimen from Beaufort West (Endothiodon zone) in the South
African Museum (Cat. No. 2660) almost undoubtedly belongs to this
species. It consists of a skull and lower jaw in a hard nodule. The
top of the skull has been cleared. The interorbital width is slightly
greater than the intertemporal. The postorbitals almost meet at
the back of the parietal bar. The pineal foramen is 10 mm. behind

South African Fossil Reptiles and Amphibia.

137

the level of the postorbital bar. The preparietal is large, with an
irregular anterior border. The postfroutals are apparently small.
The chief measurements are :

Length of skull (oblique) . . . ' . 100 mm.

Greatest width 64

Snout to plane of orbit in median line . . 44 ,,
Basal length . . . . . . . 83

This specimen is closely allied in general form to D. jouberti. It
differs in the possession of a postfrontal which is apparently absent in
D. jo^tbert^ and in the elevated pineal foramen. It is probable, how-
ever, that the two species are closely allied.

FIG. 31. Dicynodon psittacops, Br. Skull No. 2660. Preparietal region.

Natural size.

Another specimen (Cat. No. 3034) from the type locality is about
four-fifths the size of the foregoing, and agrees with it in general
proportions. The eyes are moderately small and rounded. The pre-
parietal is of slightly different shape, being more regularly oval,
although still considerably longer than broad. Its relations with the
frontals, the parietals, and the pineal foramen are as in the larger
specimen.

Type. Skull and skeleton in the American Museum.

Locality. Beaufort West, C.P.

Horizon. Lower Beaufort Beds. (Endothiodon zone.)

DICYNODON RECURVIDENS, Owen.
1876. Owen. Cat. Foss. Kept. S. Afr., p. 46 ; pi. Ixix, figs. 3 and 4.

Founded on a crushed skull and lower jaw. Small. Orbit wholly in
anterior half of skull ; canine projects below middle of orbit, and is

138

Annals of the South African Museum.

recurved ; parietal bar narrow, postorbitals almost meeting one
another.

Type in the British Museum.

Locality. Fort Beaufort, C.P.

Horizon. Lower Beaufort Beds. (Endothiodon zone (?) ).

DlCYNODON TRIGONICEPS (Br.).

1901. Broom, Ondenodon gracilis (pars). Proc. Zool. Soc., p. 162.
1904. Broom, Oudenodon trigoniceps. Eec. Albany Mus., i, p. 73;
pi. iv, fig. 2.

Founded on a well-preserved skull. Small. Parietal region as
broad as frontal, and comparatively flat. Occiput slopes forward.
Orbits look upwards and slightly outwards. Slight supraorbital
ridge, and less distinct median ridge between the two froutals.
Narrow, fairly long preparietal.

Locality. Pearston, C.P.

Horizon. Lower Beaufort Beds. (Probably top of Endothiodon
zone.)

DlCYNODON ALTICEPS, Br. & Htll.

1913. Broom & Haughton. Ann. S.A. Mus., xii, 1, p. 37; pi. vii r
figs. 1, 2.

FIG. 32. Dicynodon alticeps, Br. and Htn. Type. No. 2347. Parietal

and frontal regions, x A.

In the original description of this species a mistake was made with
regard to the limits of the preparietal bone, which led Dr. Broom to-

South African Fossil Reptiles and Amphibia. 139

include the form in his genus Diictodon, characterised by the enclosure
of the pineal foramen by the preparietal. It is clear, however, that the
preparietal forms only a portion of the border of the foramen, the
larger part of the opening being enclosed by the parietals, which are
fused together in their anterior half.

The figure shows the preparietal to be remarkably short and broad,
of irregular shape and with convoluted borders. Moreover, there does
not appear to be a distinct postf rental. A possible frontal-postfrontal
suture on the right-hand side of the skull is not paralleled on the other
side, and it is thus probably a crack.

The parietal bar has a deep median groove behind the pineal
foramen, flanked by the crests formed partly of parietal and partly of
postorbital.

Type. Skull. (S.A. Mus. Cat. No. 2347.)

Locality. One mile E. of Oudeberg, Graaff Reinet, C.P.

Horizon. Lower Beaufort Beds. (Bottom of Cistecephalns zone.)

DlCYNODON BOLORHINUS (Br.).

1911. Broom, Oudenodon bolorhinus. Proc. Zool. Soc., p. 1076; pi.
Ixiii, fig. 10.

Founded on a somewhat crushed and weathered preorbital portion
of a skull. Snout extremely short, bringing the front of the palate
almost under the orbit. Nasals thickened to form a rounded boss
overhanging the nostril. Orbit large. Frontals short and com-
paratively narrow.

The form, according to Dr. Broom, is comparable with D. strigiceps ;
but the latter has a much larger nostril and has the caniniform process
much farther back.

Type in the American Museum of Natural History.

Locality. Kuil's Poort, Beaufort West, C.P.

Horizon. Lower Beaufort Beds. (Top of Endothiodon zone or
bottom of Cistecephalns zone.)

DlCYNODON BREVICEPS, HtU.

1915. Haughton. Ann. S.A. Mus., xii, 2, p. 59 ; pi. xi, fig. 2.

The type and only known specimen is in an incomplete and crushed
condition, but its features are sufficiently well displayed to differentiate
it from other species. The skull is remarkable in being as broad as it

140 Annals of the South African Museum.

is long. The parietal bar is short, and the postorbitals almost meet
behind the pineal foramen. The postf rental forms a long part of the
orbital border, stretching outwards along the postorbital bar ; but it
narrows rapidly posteriorly and does not reach the level of the pineal
foramen.

The snout is short, and the caniniform process is well forward below
the nostril. The nostrils are close together and look forwards and
outwards.

Type. Skull and lower jaw. (S.A. Mus. Cat, No. 2366.)

Locality. Voetpad, Murraysburg, C.P.

Horizon. Lower Beaufort Beds. (Cistecephalus zone.)

DlCYNODON CAVIPRONS, Bl'. & Htll.

1917. Broom & Haughton. Ann. S.A. Mus., xii, 5, p. 120 ; fig. 20.

This species has a remarkably short and narrow palate compared
with the width between the jugal arches. The tusks pass downwards
below the front of the orbit ; but seen from in front the edges of the
caniniform pi'ocesses curve inwards, so that the width between the
tusks is small.

The occipital condyle is tripartite and short, The foramen jugulare
lies at the level of the top of the condyle and looks backwards. The
paroccipital process is very stout, especially laterally, and its lower
border is strongly concave. The basisphenoidal tubera are not espe-
cially prominent, but lie well below the condyle. The quadrates
set close together.

Type. Skull. (S.A. Mus. Cat. No. 747.)

Locality. Fraserburg, C.P.

Horizon. Lower Beaufort Beds. (Probably Cistecephalus zone.)

DlCYNODON GRANDIS, Sp. UOV.

A complete skull and lower jaw obtained on the farm Dunedin,
Beaufort West, C.P. (S.A. Mus. Cat. No. 2679), seems to belong to a
new species. It is somewhat crushed laterally, and the sutures do not
show very cleaiiy the specimen being presumably an old individual.
The chief measurements are :

Greatest length . .... 553 mm.

Greatest breadth ...... 453 ,,

Interorbital width ...... 123 ,,

Intertemporal width . . . . 64 ,,

Basal length . . .430

Width between caninifonn. processes . . 101 ,,

South African Fossil Reptiles and Amphibia. 141

The orbit is wholly in the anterior half of the skull. The nostrils
are large ; within the nostril can be seen a large septomaxilla which
does not appear on the face.

The postorbitals almost meet one another on the parietal bar,,
forming a marked parietal crest. In front of the crest is a small oval
pineal foramen, lying somewhat in a hollow, and not raised on a boss
of bone as in Eocyclops lonijus. In front of the pineal foramen, and
forming its anterior border, is a small preparietal, lozenge-shaped.
The iuterfrontal suture is seen along a median ridge, which terminates
somewhat in front of the pineal foramen. The postfroutal was a
narrow, inconspicuous bone lying for a short distance between the
postorbitals and the frontals.

The occipital plate is very broad, and the condyle is comparatively
small. As far as can be seen, the structure is typically Dicynodont.
The foramen magnum is very small compared with the size of the
plate.

The palate is long and narrow, the premaxilla furnished with two-
ridges, separated by a median hollow. The distance between the
caninifonn processes, which are placed below the front of the orbit, is
101 mm. The pterygoids are strong, the minimum width across them
being 43 mm.

The lower jaw is complete, and in good condition. The length from
the beak to the back of the articular is 455 mm., while the greatest
depth of the dentary is 110 mm. The front of the jaw is narrow, and
has an upwardly directed beak which fits into the front of the palate.
At a distance of 110 mm. behind the beak the rami begin to separate
more rapidly, and the distance between the outer sides of the articulars
is 280 mm. The front of the jaw is prow-shaped, inclined at an angle
of about 45 to the plane of the upper edges of the dentary.

The dentary is the largest bone und has a deep symphysis. The
spleuial is large, and forms more than half of the inner side of the
front half of the jaw and enters largely into the symphysis. Its upper
process lies on the prearticular, while the lower process is wedged
between the prearticular and the angular. The angular is not very
large. An anterior process passes between the dentary and splenial
almost to the symphysis. On the lower border of the jaw the plate-
like angular forms an overhanging, projecting flange. The pre-
articular, surangular, and articular are typically Anomodont.

Type. Skull and lower jaw. (S.A. Mus. Cat. No. 2679.)

Locality. Dunedin, Beaufort West, C.P.

Horizon. Lower Beaufort Beds. (Cistecephalus zone.)

142 Annals of the South African Museum.

DlCYNODON HALLI, Watson.

1914. Watson. Ann. Mag. Nat. Hist., ser. 8, xiv, p. 95 ; fig. 1.

This species is closely allied to D. planus, and, according to Dr.
Broom, the two names may be synonymous. It is probable that
Watson was mistaken in showing the preparietal as extending along
both sides of the pineal foramen. The line shown in his figure I take
to be a part of the V-shaped suture between the frontal and the
parietal. It would seem better to keep the two species distinct on
account of the difference in the ratios between the frontal and inter-
temporal widths. In D. planus the iuterorbital width is equal to, or
slightly greater than, the inter/temporal width and the parietal bar is
of constant width throughout. In D. halli the relative widths are
reversed, and the parietal bar gradually broadens posteriorly.

A specimen in the Museum collection agrees closely with the figure
given by Watson. It is a slightly distorted skull and lower jaw (S.A.
Mus. Cat. No. 3414) collected froni the Cistecephalus zone at Dunedin,
Beaufort West, C.P., by the Rev. J. II. Whaits. It shows the same
small face and general proportions as the type, and has apparently the
same small occipital coudyle without tripartite division. A curious
feature is the height of the basicrauium above the level of the
quadrates, which bones are also close together, lying directly below
the post-temporal vacuities. In consequence the stapes must have
been steeply inclined and the fenestra ovalis looks almost wholly
downwards. The basisphenoid tubera were not very prominent.

The bones of the top of the skull are well seen. The preparietal
forms only the anterior border of the pineal foramen and is broadest
near the front, being somewhat truncate anteriorly. The other bones
are as shown by Watson.

Type. Skull and skeleton in the British Museum.

Locality. Kuil's Poort, Beaufort West, C.P.

Horizon. Lower Beaufort Beds. (Cistecephalus zone.)

DlCYNODON INGENS, Br.

1907. Broom. Ann. Natal G-ovt. Mus., i, p. 168 ; pi. xxviii, figs. 1-4.

Founded on the well-preserved palatal portion of a skull. Large ;
palate longer than broad ; prominent median ridge on palate ; front of
snout unusually straight, with low median ridge and slight lateral one.
Nostril large.

South African Fossil Reptiles and Amphibia.

143

Type in the Natal Museum, Pietermaritzburg.

Locality. Enuersdale, Natal.

Horizon. Lower Beaufort Beds. (Probably top of Cistecephalus

20116.)

DlCYNODON KOLBBI (BrOOlll).

1911. Jaekel, Udenodon sp. Die Wirbeltiere, p. 192 ; figs. 210-212.

1912. Broom, Oudenodon kolbei. Aim. S.A. Mus., vii, 5, p. 337 ;
figs. 1-5.

1913. Broom, Diictodon Jcolbei. Bull. Amer. Mus. Nat. Hist., xxxii.
p. 454.

FIG. 33. Dicynodon kolbei (Broom). Type. No. 1886. Parietal and

frontal regions. * A.

Re-examination of the type of this form shows that the structure of
the preparietal region has hitherto been misinterpreted. A figure is
here given of this region. It will be seen that, instead of being a
long bone entirely enclosing the pineal foramen, the preparietal is
comparatively small and forms the anterior border only of the
foramen.

In other points there seems nothing to add to the description given
by Dr. Broom.
' Type. Skull. (S.A. Mus. Cat. No. 1886.)

Locality. Rhenosterfontein, Beaufort West, C.P.

Horizon. Lower Beaufort Beds. (Cistecephalus zone.)

144 Annals of the South African Museum.

DlCYNODON LACEKTICEPS, Owen.

1876. Owen. Cat. Foss. Kept. S. Afr., p. 30; pi. xxii.
1890. Lydekker. Cat. Fuss. Kept. & Amphib., iv, p. 18.

Founded on a skull with lower jaw. Comparatively small. Greatest
length of type skull about 158 nun. Orbits directed forwards and
laterally, sub-circular, almost as large as the temporal openings. Inter-
orbital width 25 mm. Temporal fossae directed upwards. Post-
orbitals almost meet on the parietal bar ; parietals narrow. Root of
tusk inclined forwards and downwards. Parietal foramen probably
on a level with the front of the temporal fossae.

Type in the British Museum.

Locality. " Tarka prolongation of the Winterberg," C.P.

Horizon. Lower Beaufort Beds. (Probably Cistecephalus zone.)

DlCYNODON LATICEPS, Broom.

1912. Broom. Proc. Zool. Soc., p. 868 ; pi. xcii, figs. 12, 13.

1915. Broom. Bull. Amer. Mus. Nat, Hist,, xxv, 2, p. 131 ; fig. 17.

Two skulls collected by the Rev. J. H. Whaits from the Graaff
Reiuet District are interesting as showing the changes which
apparently take place with the advance from youth to maturity. One
skull (S.A. Mus. 'Cat. No. 3423) is a fully grown male agreeing closely
with the type. The other (S.A. Mus. Cat. No. 3328) is a young male,
and, although it displays main features in common with the other,
shows some points of difference in detail. The latter skull is
iucompleie, but the sutures of the top of the skull are beautifully
displayed.

The two skulls are of the broad-headed variety, with a broad frontal
region and a broad parietal region narrowest in front. The orbits
are large and triangular and the snout short, The nostril is fairly
large. In the old skull the nostril is overhung by a large projecting
nasal boss ; but in the smaller skull this boss is very rudimentary.
The internasal width is large.

The postorbitals are long and slender, and, although they form the
upper borders of the temporal fossae, in neither skull do they appear
on the top of the parietal bar behind the pineal foramen. The whole
of the broad expanse of the posterior part of the bar is formed of the
parietals, the postorbitals facing entirely outwards. The preparietal
is large; but, while in the younger skull it is raised on a bony
eminence above the level of the top of the skull, in the older specimen

South African Fossil Reptiles and Amphibia.

145

the foramen is sunken but is surrounded by a slightly raised rim of
bone.

In each skull the pieparietal stands well above the level of the
frontals on either side of it. The smaller skull has a very large,
lozenge-shaped preparietal whose anterior end is considerably in
advance of the postorbital bar. The preparietal of the other is shorter.
The bone almost completely surrounds the pineal foramen.

The frontals meet the nasals in a straight suture at the level of the
front of the orbit. Posteriorly they flank the preparietal for a long
distance, and each is hollowed out at the side of that bone. The post-

r.P.

P.O.

FIG. 34. Dicynodon laticeps, Br. Skull, No. 3328 (Young). Preparietal

frontal is a long, narrow bone, which iu each specimen occupies a
somewhat sunken area between the frontal and postorbital.

The chief changes which seem to take place with increasing age,
therefore, are the development of the nasal and prefrontal bosses, the
sinkiug-in of the pineal foramen, and the decrease in size of the pre-
parietal bone. Since the size and shape of this bone in the larger skull
seems to agree exactly with that in the type specimen we can scarcely
look upon the difference in size as an individual feature, but must
reckon it as the result of a change due to age.

Other interesting features of the larger (and more complete) skull
are the short, broad palate, the large tusks, the very strong keel formed
by the pterygoids behind the posterior narial fossa, the narrowness
between the quadrates, the large tabulare seen on the back of the

146 Annals of the South African Museum.

skull, and the long upward extension of the jugal behind the post-
orbital.

Type in American Museum of Natural History.

Locality. Grootvlei (part of Paardekraal ) , Beaufort West, C.P.

Horizon. Lower Beaufort Beds. (Cistecephalus zone.)

DlCYNODON LEONICEPS, Owen.

1876. Owen. Cat. Foss. Eept. S. Afr.,p. 32 ; pis. xxiv-xxvi.
1890. Lydekker. Cat, Foss. Kept. & Amphib., iv, p. 19.

Founded on an imperfect skull ; large, long, and narrow. Orbits
directed mainly laterally ; interorbital bar very wide, parietal bar long
and narrow, postorbitals almost above parietals ; temporal fossa long
and comparatively narrow ; skull high.

A skull which I was permitted to examine through the kindness of
the Director of the Port Elizabeth Museum (No. 68, P.E. Mus.)
seems to be referable to this species. It lias the following approxi-
mate measurements :

Greatest length . . . ." . . 300mm.
Length from snout to front of orbit . 60 ,,

Interorbital width . , , 65 ,,

Intertemporal width . . 33 ,,

Width across nasal overhang 63 ,,

Orbital diameter ... 60 ,,

Basal length . . . 250

Width between base of tusks . . 32 ,,

The condition of the skull is such that sutures are difficult to deter-
mine. The bone is soft and is caked with a hard, crystalline matrix.
There appears to have been a large postfrontal ; but the preparietal, if
present, could have been but small. The pineal foramen is almost
covered by the postorbitals. The chief feature of the skull is its great
height, i. e. the length of the downward portion of the squamosal.
Type in the British Museum.

Locality. " Gats River, Sueeuwberg, Graaff Reinet," C.P.
Horizon. Lower Beaufort Beds. (Cistecephalus zone (?).)

DlCYNODON LEONTOPS, Br.

1913. Broom. Bull. Anier. Mus. Nat. Hist., xxxii, p. 451 ; fig. 12.
1915. Broom. Bull. Amer. Mus. Nat. Hist,, xxv, 2, p. 132; fig. 19.

Large. Comparable with D. leoniceps and D. pardiceps ; frontals
relatively narrower than in these types, passing back in almost parallel

South African Fossil Reptiles and Amphibia. 147

processes to plane of back of pineal foramen ; preparietal small and
narrow. Interorbital width three times the intertemporal width.

Type in the American Museum of Natural History.

Locality. Bethulie, O.F.S.

Horizon. Lower Beaufort Beds. (Top of Cistecephalug zone.)

DlCYNODON LISSOPS, Bl\

1913. Broom. Bull. Amer. Mus. Nat. Hist., xxxiii, p. 450 ; fig. 11.

Skull of medium size. Comparable with D. lacerticeps. Eye wholly
in anterior half of skull. Nostril fairly large, well forward. Large
septomaxillary which just appears on face. Snout rounded and
smooth. Frontals pass well forward. Interorbital width : intertem-
poral width : : 5 : 3. Prefrontal large. Preparietal large. Postorbitals
nearly meet on parietal bar. Postfroutal very narrow^

Type in the American Museum of Natural History.

Locality. Wilgebosch, New Bethesda, Graaff Eeinet, C.P.

Horizon. Lower Beaufort Beds. (Cistecephalus zone.)

DlCYNODON MOSCHOPS, Br.

1913. Broom. Bull. Amer. Mus. Nat. Hist., xxxii, p. 447 ; figs. 7-8.
1915. Broom. Bull. Amer. Mus. Nat. Hist,, xxv, 2 ; fig. 23.

Skull of medium size, almost as wide as long. Snout broad,
markedly bent near front of frontal. Nostrils small, roofed by marked
projection of nasal. Septomaxilla appears on face, joining with
lachrymal and separating nasal from maxilla. Frontals broad. Post-
frontals almost entirely hidden. Preparietal small. Pineal foramen
large, broader than long. luterparietal large, forming considerable
part of upper surface of skull.

Type in American Museum of Natural History.

Locality. Oudeberg, Graaff Keinet, C.P.

Horizon. Lower Beaufort Beds. (Gistecephalus zone.)

DlCYNODON MTJSTOI, Htn.

1915. Haughtou. Ann. S A. Mus., xii, 2, p. 58 ; pi. xi, fig. 1.

This form, although at first sight somewhat similar to D. kolbei,
differs sufficiently to retain separate specific rank. In the original
description given, it was considered that the preparietals of the two
forms were very distinct ; but although the revelation of the true
structure of the extent of the bone in D. Icolbei removes this point of

148

Annals of the South African Museum.

difference, the two species can still be separated by the difference in
shape of the anterior part of the postf rental, the greater relative
width between the postorbitals in D. mustoi, and the fact that in D.
mustoi the interorbital width is less than the intertemporal, whereas
in D. Jcolbei the reverse is the case.

-J

FIG. 35. Dicynudon mustoi, Htn. Type. No. 2674. x '-.

Type. Skull. (S.A. Mus. Cat, No. 2674.)

Locality. Dunedin, Beaufort West, C.P.

Horizon. Lower Beaufort Beds. (Cistecephalus zone.)

DlCYNODON PLANUS, Bl'OOm.

1913. Broom. Bull. Amer. Mus. Nat. Hist., xxxii. p. 452 ; figs. 13, 14.

A medium-sized Dicynodon with comparatively short face, and
breadth about five-sixths of the length. The interorbital width
equal to, or slightly greater than, intertemporal width. Orbits
wholly in the anterior half of the skull, looking mainly upwards.
Postfrontal large, postorbitals each forming one-third of the parietal
bar.

South African Fossil Reptiles and Amphibia. 149

A skull (S.A. Mus. Cat. No. 2364) from Dunedin, Beaufort West
(Cistecephalus zone) agrees with the type except in being somewhat
smaller. The bones of the top of the skull are beautifully displayed.
Here, as in D. halli, the quadrates are near together. The occipital
coiidyle is small, but is feebly tripartite in character. The occipital
plate is vertical. The right stapes is showing, resting medially in the
foramen ovalis, which looks outwards and downwards.

Type. Skull in the American Museum of Natural History.

Locality. Kuil's Poort, Beaufort West, C.P.

Horizon. Lower Beaufort Beds. (Cistecephalws zone.)

DlCYNODON PLATYCEPS, Br.

1913. Broom. Bull. Amer. Mus. Nat. Hist., xxxii, p. 444; fig. 4.

1914. Broom. Phil. Trans. B 206, p. 44 ; pi. v, figs. 46, 47, 48.

1915. Broom. Bull. Amer. Mus. Nat. Hist., xxv, 2, p. 138; fig. 27.

Skull and skeleton of medium size. Skull flattened, considerably
longer than broad. Orbits look upwards and outwards. Tusks
feeble. Frontal region hollowed, supraorbital margins elevated. Large
preparietal forms anterior border of large pineal foramen. Postfrontal
long and narrow.

Type in American Museum of Natural History.

Locality. New Bethesda, Graaff Reinet, C.P.

Horizon. Lower Beaufort Beds. (Cistecephalus zone.)

DlCYNODON PYGMAEUS, Br. & Htu.

1917. Broom & Haughtou. Ann. S.A. Mus., xii, 5, p. 123 ; fig. 23.

Three skulls in the South African Museum collection can be defi-
nitely assigned to this species. They are :

Cat. No. 2664. Type from Dunedin, Beaufort West, C.P. ;

Cat. No. 2668 from the same locality ; and

Cat. No. 3352 from Highlands, top of Nieuweveld, Beaufort West,

CP.
All are from the Cislecephalus zone.

The three skulls are almost of the same size ; but it is probable
that the type has been flattened by vertical crushing, so that the
width given in the oi'iginal description is greater than the skull
actually had in life. The maximum width was probably between
40 and 45 mm., and the orbit looked at least as much laterally as
upwards. The orbit is round, and the postorbital arch rather weak.
The skull from Highlands is tusked ; the others are females. The

150 Annals of the South African Museum.

tusk passes mainly downwards and its base lies just in front of the
orbit.

In the male skull the pineal foramen is slightly farther back along
the pai'ietal bar than in the type ; but the relative shapes and sizes of
the bones around the foramen remain the same.

The foramen magnum is large and the post-temporal fossa is seen
to be well up on the occipital plate (No. 3352), so that the paroccipital
processes are deep and fairly shoi't.

The only other small species known from the Cistecephalus zone is
D. testudirostris. The two species are readily distinguishable at a
glance by the natui'e of the beak. D. testudirostris is especially
characterised by the strength of the beak and the vertical inclination
of the front of the snout. The suborbital bar has its lower border
turned abruptly downwards at the level of the middle of the orbit, so
that there is a deep portion of the maxilla underlying the front half
of the eye. The two species agree in the narrowness of the frontal
region compared with the breadth of the parietal bar, in the narrowing
of the parietal bar posteriorly and in the possession of an insignificant
postfrontal. In D. testudirostris, however, the postorbitals meet above
the parietals at the back of the parietal bar, while in D. pygmaeus they
don ot.

D. pygmaeus has a fairly close ally apparently in D. ictidops from
the zone below, from which form it might possibly have been derived
by the reduction of the postfrontal and slight broadening of the
parietal crest.

Type.Sku\\. (S.A. Mus. Cat. No. 2664.)

Locality. Duuediu, Beaufort West, C.P.

Horizon. Lower Beaufort Beds. (Cistecephalus zone.)

DlCYNODON ROGERSI, Br. <t Htu.

1917. Broom & Haughton. Ann. S.A. Mus., xii, 5, p. 121 ; figs. 21, 22.

This form is closely allied to D. kolbei, and for some years the type
specimen was considered to be an example of that species. The
differences between the two forms are, however, worthy of recognition,
and have been given in the paper cited above.

One half only of the occipital plate is preserved, and the condyle is
lacking. The lateral vacuity is at the level of the middle of the rather
small foramen magnum. The tabulare is not large. The stapes is
short and its outer end is considerably lower than its inner end.

The lower jaw is very long and comparatively shallow^

South African Fossil Reptiles and Amphibia. 151

Type Skull and lower jaw. (S.A. Mus. Cat. No. 2356.)
Locality. Thee Kloof, Fraserburg, C.P.

Horizon. Lower Beaufort Beds. (Probably bottom of Cistecephalus
zone.)

DICYNODON STRIGICEPS, Owen.

1855. Owen. Trans. Geol. Soc., 2nd ser., vol. vii ; pi. vi, figs. 2, 3.
1876. Owen, Oudenodon (?) strigiceps. Cat. Foss. Eept. S.A., p. 61 ;

pi. xliv, fig. 4.
1890. Lydekker, Udenodon strigiceps. Cat. Foss. Eept. and Axnphib.,

iv, p. 30.

Founded on the anterior part of a skull. Frontal region broad ;
caniniform process descends vertically below orbit ; rostral region
descends in convex semicircular contour to beak ; orbits in fore part of
head, directly above the nares.

Type in the British Museum.

Locality. "Tarka prolongation of the Winterberg," C.P.

Horizon. Lower Beaufort Beds. (Probably top of Cistecephalus
zone.)

DICYNODON TESTUDICEPS, Owen.

1855. Owen. Trans. Geol. Soc., 2nd ser., vol. vii, p. 71.
1876. Owen. Cat. Foss. Eept, S.A., p. 45 ; pi. xliv, figs. 1-3.
1890. Lydekker. Cat. Foss. Eept. and Amphib., iv, p. 24.

Founded on the anterior two-thirds of a skull. Front of skull
short and broad ; muzzle deflected abruptly downwards. Interorbital
width 35 mm.

Type in the British Museum.

Locality. " Tarka prolongation of the Winterberg," C.P.

Horizon. Lower Beaufort Beds. (Probably Cistecephalus zone.)

Lydekker gives the locality as " from the Stormberg Beds of the
Karroo system on the Modder tributary of the Orange Eiver."

DICYNODON TESTUDIBOSTRIS, Br. & Htn.

1913. Broom & Haughton. Ann. S.A. Mus., xii, 1, p. 36; pi. vii,
figs. 3, 4.

In the photographs given with the original description of this type
the sutures were not adequately displayed, and a drawing is therefore
given of the arrangement of the bones in the preparietal region. The

13

152

Annals of the Sovith African Museum.

preparietal is seen to be a fairly large bone which touches only a
minute fraction of the pineal foramen. The postfrontal is long and
narrow, and the parietal has but one anterior process passing between
the frontal and preparietal.

Since the original paper the occiput and basicranium of the type
skull have been cleared of matrix. The occipital plate is nearly
vertical. The foramen magnum is large, its upper margin being more
than halfway up the skull. Consequently both the interparietal and
supraoccipital are somewhat shallow bones. Sutures are not well
shown on the plate, so that the exact limits of each bone cannot be
determined. The lateral vacuity lies at the level of the middle of the

FIG. 36. Dicynodon testudirostris, Br. & Htn. Type. No. 2354. Pre-
parietal region. Natural size.

foramen magnum. The occipital condyle is tripartite with a large
central pit. The foramen for the 9th-12th nerves looks almost
entirely backwards and outwards ; it is large, and most of its border
is formed by the paroccipital.

The basioccipital tubera are prominent, and wholly below the level
of the coudvle. The fenestra ovalis is almost as low as the quadrate.
Between the tubera the basioccipital is deeply grooved. The basi-
spheuoid is short. Anterior to it the pterygoids form a broad, short
plate 11 mm. wide. Just behind the iuterpterygoid vacuity is a
median knob.

The quadrato-jugal is peculiar in that it is apparently not auchy-
losed to the quadrate. On the left side it is seen in position, lying on
the downward process of the squarnosal and widely separated at the
articular end from the quadrate. On the other side it is detached from
the skull.

South African Fossil Reptiles and Amphibia. 153

The chief measurements of the type are :

Greatest length ..... ab. 86 mm.

Greatest width . 56 ,,

Snout to front of orbit . . . 18 ,,

Length of orbit . . 20 ,,

Interorbital width . . . 16 ,,

Intel-temporal width (min.) . . 12 ,,

Basal length . . 72 ,,

Width between caniniforin processes . 26 ,,

Type. Skull. (S.A. Mus. Cat. No. 2354.)

Locality. Dunedin, Beaufort West, C.P.

Horizon. Lower Beaufort Beds. (Cistecephaliis zone.)

DICYNODON TIGRICEPS, Owen.

1855. Owen. Trans. Geol. Soc., vii, 2nd ser., p. 233.
1876. Owen. Cat, Foss. Eept. S.A., p. 38.

1876. Owen, Dicynodon baini. Cat. Foss. Eept. S.A., p. 36 ; pis. xxx-
xxxv (non xxxvi, xxxvii).

1889. Seeley. Phil. Trans., 1889 B, p. 236, pi. xiii.

1890. Lydekker. Cat. Foss. Eept. and Amphib., iv, p. 24.

This species is doubtfully represented in the South African Museum
collection by an incomplete skull (No. 749) from Pearstou, C.P. The
front of the skull is very weathered and somewhat distorted, but the
occipital plate is in a good state of preservation. The basisphenoid
tubera are seen to project very considerably below the level of the
condyle, and between them the basioccipital is deeply hollowed out.
The paroccipital process is very massive and its lower border strongly
concave. The foramen jugulare looks entirely backwards, and the
post-temporal vacuity lies at the level of the middle of the foramen

niaguum.

Type. Skull in the British Museum.

Locality. Gouzia Eiver, Kaffraria. ("_D. baini " is from Fort Beau-
fort, C.P.)

Horizon. Lower Beaufort Beds.

DICYNODON TRUNCATUS (Br.).

1899. Broom, Oudenodon triincatus. Ann. S.A. Mus., i, p. 455 ;
pi. x, fig. 4.

Founded on the crushed and weathered anterior portion of a skull.
Beak almost square, with median and two lateral ridges. Premaxilla

154 Annals of the South African Museum.

forms nearly the whole of the front of the roof of the mouth. No
septomaxilla showing on face. Nostrils well advanced, large. Cauini-
forin process just behind plane of back of nostril.

Through the courtesy of the Director of the Port Elizabeth Museum
I have been enabled to see the type. It is an unsatisfactory specimen,
and its chief interest lies in the fact that it was probably obtained
from a locality from which very few Dicynodont or other reptilian
remains are known.

Type in the Port Elizabeth Museum.

Locality. Probably Hanover, C.P.

Horizon. Lower Beaufort Beds. (Zone of Endothiodon or of Ciste-
/i*. )

DlCYNODON TYLORHINUS, Bl\

1913. Broom. Bull. Ainer. Mus. Nat. Hist., xxxii, p. 448 ; figs. 9, 10.

Skull of medium size, as broad as long. Snout very broad, pre-
orbital region very short. Beak short, nostrils small. Above nostrils,
nasals form two prominent knobs which pass forward well in front of
premaxilla. Frontal region flat and broad. Interorbital width : inter-
temporal width : : 9 : 4. Preparietal large, postf rout als entirely hidden.
Postorbitals meet behind the pineal foramen.

Type in American Museum of Natural History.

Locality. Wilgebosch, New Bethesda, G-raaff Eeinet, C.P.

Horizon. Lower Beaufort Beds. (Cistecephaltts zone.)

DlCYNODON WHAITSI, Br.

1913. Broom. Bull. Amer. Mus. Nat. Hist., xxxii, p. 443 ; fig. 3.
1915. Broom. Bull. Amer. Mus. Nat. Hist., xxv, 2, p. 142 ; fig. 33.

This species is one of the largest of the Dicynodous. The original
description is as follows : " The snout is narrow and deep and the
nostrils large. The orbits are placed near the middle of the head.
The postorbital arch is powerful. The parietal region is broad, and
the posterior ranius of the postorbitals unusually well developed. The
pineal foramen is situated well behind the postorbital arch. Behind
and on about two-fifths of each side it is bordered bv the parietals.

tt

The rest of the foramen is bordered by the large preparietal. The
frontals extend back on each side of the preparietal to nearly the
plane of the back of the foramen. The postfrontals are moderately
large."

The Rev. J. H. Whaits has recently collected for the South African

South African Fossil Reptiles and Amphibia. 155

Museum from Acleiidorp Commonage, Graaff Reinet, C.P., a skull,
lower jaw and partial skeleton of an example of this species. The
skull is somewhat larger than the type. It is flattened laterally, and
the relation of the bones of the top of the skull cannot be accurately
determined. But there can be no doubt as to the specific identity of
the skull with the type figured by Broom. The species is a readily
identifiable one in general appearance. In addition to this specimen
we have another skull from Dunedin, Beaufort West even larger
than the other and similarly flattened which shows exactly the same
features. The orbit is circular, looking almost entirely outwards, and
is in the front half of the skull. In the Adeudorp specimen (S.A.
Mus. Cat. No. 3714), however, the pineal foramen seems to be nearer
the postorbital arch than in the type. Otherwise tlin general pro-
portions of the top of the skulls are the same. The postorbital arch is
exactly midway along the skull in this specimen, and slightly in
advance of the middle line in the Dnnedin skull (S.A. Mus. Cat.
No. 2681).

The importance of the Adendorp specimen lies in the fact that with
the skull and lower jaw are a number of associated limb-bones,
vertebrae, and portions of the girdles. The postcranial skeleton of the
Aiiomodonts is well known in its general features ; but hitherto there
has been wanting detailed knowledge concerning the features of the
various species, so that it is impossible almost to correlate the many
skeletal portions of Dicynodouts found with the known skull types.
For example, the bones named by Owen as Platypodosaurus robustus
have been variously conjectured to be bones of Eocyclops tnagnus, E.
lonyus, and Dicynodon gril!x, but not one of the three types is known
from more than the skull, so that the point at issue cannot yet be
settled. Hence any light shed upon the subject by the discovery of
skeletal remains in connection with skulls is welcome.

In this specimen the bones are preserved in too hard a matrix to
permit of their being very satisfactorily cleaned, and many of them are
incomplete, having apparently suffered a certain amount of fracture
and rubbing before their final entombment. They agree with the
normal type of Dicynodont limbs, and thus their measurements alone

are given.

Scapula. The upper border of the left scapula is missing. The
bone as preserved has a length of 325 mm. The basal breadth is
125 mm. and the minimum breadth of the bone is 80 mm. Superiorly
the blade widens. The anterior point of the acromioii process is
100 mm. above the base of the bone. At the glenoid cavity the bone
is strongly thickened to a thickness of 80 mm.

156 Annals of the South African Museum.

Hwnerus. The right humerus is nearly complete, lacking the
proximal end, which is present in a more fragmentary left humerus.
The bone is larger than that of Platypodosaurus, but has the same
characters except that it does not seem to have the " hook " at the
distal end of the deltoid crest. The length of the bone is 295 mm. ;
the breadth of the proximal end is 190 mm., and of the distal end
175 mm. The minimum breadth across the shaft is 75 mm., and the
distance of the top of the ^picondylar foramen from the distal border
of the bone is 95 mm.

Radius. The greatest length of the radius is 215 mm., the breadth
of the proximal surface 90 mm., and of the distal 83 mm. The
proximal articular surface is an elongate oval, its inner half concave.
The distal end of the bone is relatively much stouter than in D.
trigoniceps, the articular surface having a thickness of 62 mm. The
middle of the shaft is thin.

Ulna. The ulna is longer than the radius, having a total length of
237 mm. The olecranon process is not at all strongly developed,
therein showing a difference from that of D. trigoniceps. The greatest
breadth proximally is 112 mm. At its narrowest the shaft is 43 mm.
broad, and distally the bone is slightly expanded to a breadth of
67 mm. The distal articular surface is about, 35 mm. thick. The
whole bone is somewhat thinned by flattening.

Type. Larger part of skull in the American Museum of Natural
Historv.

/

Locality. Uitspanfontein, Beaufort West, C.P.
Horizon. Lower Beaufort Beds. (Cist ecephal its zone.)

GENUS CHELYRHYNCHUS, gen. nov.

CHELYRHYNCHUS LACHRYMALIS, gen. et sp. nov.

The specimen under consideration consists of an almost perfect

uncrushed skull collected by the Eev. J. H. Whaits on the Graaff

Reinet Commonage (S.A. Mus. Coll. 3334).

It is a medium-sized skull having a maximum length of 200 mm.,

and a greatest breadth across the squamosals of 180 mm. The other

chief measurements are :

Interorbital width 30 mm.

Intel-temporal width . 37

Basal length . . 179 ,.

Distance from snout to front of orbit . 50

Distance between nares and orbit . . 17,,

Width between caniuiforui processes . . 60 ,,

South African Fossil Reptiles and Amphibia. 157

The snout is narrow and the face short.

The maxilla passes back to below the middle of the orbit and forms
most of the cheek. It has a small palatal portion upon which there is
no trace of any molar teeth.

The septoniaxilla lies wholly within the nostril, forming the
posterior lower border, and articulates with the nasal, lachrymal, and
maxilla.

The lachrymal extends from the orbit to the nostril as in Tropido-
stoma. It forms a large portion of the anterior border of the orbit,
but is much shallower on the cheek. At the nostril it abruptly
narrows, and it has a short articulation with the septoniaxilla.

The nostril is overhung by a small nasal boss.

The frontal reaches forward to the plane of the back of the nostril,
separating the nasals from each other posteriorly.

The chief features of the skull-top are the slightly thickened supra-
orbital border, the width of the intertemporal region, the absence of a
preparietal, and the appearance of the interparietal on the top of the
skull.

The postorbital bar is long, but comparatively slender.

The occipital plate is broad, flat, and vertical. The occipital cou-
dyle is small and not tripartite, similar to that of Tropidostoma.

The foramen jugulare lies on the back of the skull and looks wholly
backwards. The paroccipital process is very massive in its outer
portion and its outer border inclines strongly medially below. The
basisphenoid forms nearly half of the tubera on the inner surface ; but
it takes 110 part in the border of the fenestra ovalis, which is bounded
posteriorly, medially, and anteriorly by the basioccipital. The fenestra
ovalis looks downwards and outwards. The palatal portion of the
basisphenoid is short.

In size and shape the skull agrees most closely with Dicynodon
planus, Tropidostoma microtrema, and the form described by Owen as
" Oudenodon bainii." From each it is distinguished by one or more
characters.

From Tropidostoma microtrema, with which this form agrees in the
primitive state of the lachrymal, the skull differs in the absence of
molar teeth, in the fact that the postorbitals and parietals are not
developed into lateral crests with a groove between them, and in the
absence of a preparietal.

The skull has many affinities with the type of D. planus, but there
is no preparietal. One of the most striking features of the skull is the
extension of the lachrymal to the narial border, a feature displayed by
Pareiasaurus and other skulls of a primitive type. This condition is

158

Annals of the South African Museum.

closely paralleled in a skull in the S.A. Mus. collection (No. 2364),
which has hitherto been considered as a specimen of Dicynodon planus,
somewhat smaller than the type. In this skull (2364) the lachrymal
has a wide orbital end, and narrows rapidly anteriorly, passing forward
just to touch the posterior border of the nares, and articulating with a

FIG. 37. Chelyrhynchus lachrymalis, Htn. Type. No. 3334. x '8.

bone within the nostril, presumably the septomaxilla. Certainly the
nasal is separated completely from the maxilla. Moreover, skull 2364
agrees fairly closely in shape with that under discussion (3334) ; the
palate is rather longer and narrower, and the occipital condyle shows
somewhat, but very little, more of the tripartite division. But, on
the other hand, it has a preparietal closely comparable with that of
the type of D. planus and the whole features of the top of the skull

South African Fossil Reptiles and Amphibia.

159

are those of D. planus. (Unfortunately, I am not aware of the exact
relations of the lachrymal and the state of the occipital conclyle in the
type of D. planus.}

The question then arises Can the relations of the bones in the pre-
parieta region be taken as factors of generic, or even of specific,
impoi'tance among the many forms of Dicynodon and its allies ?
There can be little doubt that the skull (S.A. Mus. 2364) referred to
D. planus and that under discussion closely resemble one another in
all points save in these relations. Nevertheless, following Dr. Broom's
classification, they would be placed in different genera according to

SM-

FIG. 38. Chelyrhynchus laclirymalis, Htn. Type.

of skull. x - 8.

No. 3334. Side view

the presence or absence of the preparietal bone. To settle this
question, it is necessary to show that the preparietal is constant in
any one species, /. e. that forms which agree in all other characters
agree also in the possession, shape, .and position of the preparietal.

In describing the following Auomodont species, Dicynodon platyceps,
Dicynodon ictidops, Emydops longiceps, and EmydorhyncJnis palustris,
Dr. Broom was able to base his descriptions upon a number of skulls
in each case, and in each instance he remarks upon the close agree-
ment of the individuals within any one species. Moreover, a long
series of skulls in the S.A. Mus. collection referred to Dicynodon
jouberti have the preparietal very constant. On the other hand, the
two closely-allied skulls 2364 and 3334 differ in that the former has a

i/

long narrow preparietal while the latter is without.

This skull can be taken as the type of a new genus and may be

160 Annals of the South African Museum.

called Chelyrhynchus lacliry mails, gen. et sp. nov., characterised by the
following main features :

Skull almost as broad as long. Snout rather short. Orbits large.
Intertemporal width greater than interorbital width.

No preparietal, lachrymal extending from orbit to nares, completely
separating nasal from maxilla. Septomaxilla within nostril, articu-
lating with lachrymal. Condyle small, and not tripartite. Palate short
and broad.

Type. Skull. (S.A. Mus. Cat. No. 3334.)

Locality. Commonage, Graaff Iteinet, C.P.

Horizon. Lower Beaufort Beds. (Endotldodon zone.)

GENUS DIICTODON, Br.
1913. Broom. Bull. Amer. Mus. Nat. Hist., xxxii, p. 453.

Differs from Dicynodon in that the pineal foramen is entirely
surrounded by the large preparietal.

DIICTODON GALEOPS, Br.

1913. Broom. Loc. cit. fig. 15.

1915. Broom. Bull. Amer. Mus. Nat. Hist,, xxv, 2, p. 145 ; fig. 33.

Founded on a small skull. Orbits large, beak and nostril like those
of Dicynodon ictidops. No septomaxilla on face. Tusk very slender,
directed mainly downwards. Prefrontal small. Preparietal large,
with broad anterior third and narrower posterior two-thirds. Pineal
foramen small. Postf rental absent.

Type in American Museum of Natural History.

Locality. Slachter's Nek, C.P.

Horizon. Lower Beaufort Beds. (Probably top of Endothiodon
zone.)

GENUS EOCYCLOPS, Broom.

EOCYCLOPS LONGUS, Broom.
1913. Broom. Bull. Amer. Mus. Nat. Hist., xxxii, p. 441 ; figs. 1, 2.

Broom described in 1913 a skull from the Nieuweveld which he
called Eocyclops lone/us, and which he said was closely allied with the
type of Owen's form Oudenodon IIHHJHH*. Both agree in the structure

South African Fossil Reptiles and Amphibia.

161

of the top of the skull, possessing no preparietal and having the
pineal foramen surrounded by the parietals.

Recently the Museum became possessed of a beautiful skull col-
lected at Graaff Reinet bv the Rev. J. H. Whaits, which agrees fairly

FIG. 39.Eoryclops lonr/us, Br. Skiill No. 3425. c i nearly.

closely with Eocyclops, but which has a distinct preparietal forming
most, if not all, of the boss around the pineal foramen. Of this fact
I think there can be no doubt. The sutures between the bones are
plainly shown. The in terf rental suture is seen plainly on top of the
skull ; at its posterior end it divides with two widely diverging arms,

162

Annals of the South African Museum.

which siirround a single bone undivided by any median suture.
Sutures are not traceable in the parietal boss ; but a short distance
behind it a median suture between the two parietals is visible. The
evidence thus points strongly to the conclusion that there is a single
median bone between the frontals and the parietals, forming at least
part of the boss around the pineal foramen. In the type of Eocyclops
longus there is no doubt that a median suture occurs just in front of
the pineal foramen ; and the question again arises therefore How
much stress can be laid upon the presence or absence of the pre-
pai'ietal ? It is just possible, of course, that in the South African
Museum specimen the suture in front of the foramen between two

FIG. 40. Eocyclops longus, Br. Skull. No. 3425. x $ nearly.

parietals may have been obliterated ; but the other sutures on the
skull stand out so clearly, and that between the parietals in their
posterior part is so obvious, that this does not seem a feasible solution
of the matter. It would seem much better to suppose that, although
in the smaller Dicynodonts the preparietal is a constant within any
one species and is constantly present or absent in the genus, in the
case of these very large and presumably somewhat aberrant forms
the bone is inconstant in character and is not of importance sys-
tematically. Certainly the three skulls S.A. Mus. 3425, type of
" Oudenodon magnus " and type of Eocyclops lonyus agree so closely
in other regards, as far as can be seen from the somewhat incomplete
descriptions, that this one difference can scarcely be held to be
sufficient to separate them generically.

The skull from Graaff Reinet is in very fine condition and is worthy

South African Fossil Reptiles and Amphibia. 163

of description. In general outline it resembles closely Eocyclops
lonyus and Eocyclops magnus, as also iu the very large size of the
temporal openings. From the former it differs in having a broader
intertemporal region, due to the fact that the postorbitals are not so
close together, a large portion of the parietals showing between them.
From the latter form it differs in having a relatively shorter autorbital
portion, and in the fact that the parietal bar is much wider and is not
provided with a crest.

The chief measurements are as follows :

Greatest length . ... 502 mm.

Greatest breadth . . . 406 ,,

Interorbital widlh .... 105 ,,

Intel-temporal width . 69 ,,

Snout to front of orbit . . . . 150 ,,

Basal length . . . 406

The nostril is very large, the interuasal width rather small. The
snout is fairly long and narrow, and is abruptly truncated anteriorly.
Above the nostril is a small nasal boss.

There is no septomaxilla seen on the surface. As usual the maxilla
forms most of the cheek, and, as in E. longus, passes back to below
the back of the orbit, being separated for some distance from the
jugal by an anterior prolongation of the squamosal.

The lachrymal is fairly small. The lachrymal foramen is well seen
within the orbit.

The prefrontal is large, almost reaching the back of the nostril and
passing well on to the top of the skull. It is strongly thickened in the-
supraorbital region.

The nasals are not separated by the frontals, the anterior boundary
of the latter running straight across the skull at the plane of the front
of the orbit. The frontal forms the whole of the upper border of the
orbit and has a backwardly directed process by the side of the pineal
boss. In front of that boss each frontal is somewhat excavated. The
whole interorbital region is slightly concave.

The orbits are large and distinctly triangular, looking mainly
forwards and outwards.

The postfrontal is not well defined, but it was doubtless as in E.
longus. The postorbital is large, forming the whole of the postorbital
bar, articulating at its outer end wholly with the jugal.

The relations of the preparietal and parietals have already been
discussed. Behind the parietals is the interparietal, whose anterior
end is split up by an intrusion of the parietals. The bone appears
partly on top of the skull and partly on the occipital plate, the

164 Annals of the South African Museum.

transition from one to the other taking place gradually by a regular
bending and not abruptly. There is no transverse ridge separating
the occiput from the parietal bar.

The occiput is not well displayed and the sutures between the
bones are difficult of determination. There is, however, a large
tabulare lying outside the interparietal and articulating with the
squamosal and the supraoccipital. The occipital conclyle is large and
of the tripartite type.

The articular surface for the lower jaw is not very much in advance
of the occipital condyle. It is very large, and although no suture can
be seen running across there is certainly a large quadrato-jugal
present. Between it and the quadrate on the anterior face of the
descending plate is a large oval foramen.

The palate is long and comparatively narrow. There is no trans-
palatine. The jugal passes back within the maxilla to meet the
palatine, which articulates anteriorly with the maxilla.

Type in American Museum of Natural History.

Locality. Grootvlei (part of Paarde kraal), Beaufort West, C.P.

Horizon. Lower Beaufort Beds. (Cistecejjhalus zone.)

GENUS MYOSAURUS, gen. nov.

MYOSAURUS GBACILIS, gen. et sp. nov.

Two beautiful skulls form the co-types of this new genus. They
are of the same size and were obtained from Harrismith, O.F.S., in
the same calcareous nodule of Middle Beaufort age, in beds associated
with numerous Lystrosaurus skulls. The form is allied to Emydops,
but differs in being absolutely free of molar or canine teeth.
The following are the chief measurements :

Greatest length (oblique) . 48 mm.

Greatest width . . . 36 ,,

Interorbital width . . 11 ,,

Intel-temporal width . 17 ,,

Length of orbit . 13 ,,

Width of orbit . 12

Snout to front of orbit (oblique) . . 12 ,,

Width of palate . . 10

Basal length . . . . 40 ,,

The snout is feeble and narrow, and the nostrils are almost ter-
minal. They are close together and look practically wholly forwards.
At the front of the orbit the skull broadens rapidly. The orbit is

South African Fossil Reptiles and Amphibia.

165

large and circular, and looks forwards, upwards and outwards. The
postorbital bar is feeble. The temporal fossae are fairly long, and, on
; account of the breadth of the parietal region, look mainly outwards.

The premaxilla is very feeble, and apparently provided with a short
internasal process.

The nasal forms most of the upper and half of the posterior borders
of the nostril. It passes back along the top of the snout to meet the
frontal in a long suture, and its lower edge articulates with the pre-
frontal, lachrymal, and maxilla. There is no septomaxilla visible.

The maxilla forms most of the small cheek. It passes back under
the jugal not quite to the postorbital bar. The lower edge of the bone

FIG. 41. Myosaitrus gracilis, Htn. Type. No. 3526. Natural size.

is very thin in the anterior half and the cauiniform process extremely
feeble almost non-existent.

The lachrymal and prefrontal are both fairly small triangular bones
whose bases form portions of the antorbital boi'der.

The frontal is fairly broad and extends back to the level of the
parietal foramen, i. e. half way along the parietal bar, forming there a
wedge dividing the anterior part of the parietal into two portions.
The orbital borders are not raised, and there are no supraorbital'
bosses.

I can find no evidence of a postfrontal. The frontal overlaps the
postorbital from the orbital border back to the parietal.

The postorbital forms the whole of the upper and anterior borders
of the temporal fossa. Its downward pi'olongation is weak, but it
expands slightly at its articulation with the jugal.

The preparietal is a narrow rhomboidal bone extending from the
parietal foramen to just in front of the plane of the postorbital arch.

166 Annals of the South African Museum.

The piiieal foramen is far back on the broad parietal bar, its posterior
end being only 2 mm. in advance of the occipital plate. From either
side of the foramen a groove passes forward to the orbital border, the
preparietal region and the lateral portions of the parietal bar standing
out as prominences.

The parietal is a short, broad bone, an outer narrow prolongation
passing between the postorbital, interparietal and tabulare. It forms
no part of the border of the fossa. At the back of the parietal bar
the bone is bent abruptly and lies vertically in front of the inter-
parietal.

The occipital plate is broad and shallow and almost vertical. Its
upper portion is formed by the broad interparietal, which has a

' EO "Par.Occ.
B.o.

FIG. 42. Myosaunts <jracilis, Htn. Sketch of occipital plate. Natural size.

vertical median keel. Below the interparietal lies the supraoccipital,
whose upper border is semicircular, and which forms the border of the
upper half of the foramen magnum. Below it meets the exoccipital
and the paroccipital, but takes no part in the border of the post-
temporal fossa.

Lateral to the interparietal and the supraoccipital lies the tabulare,
a Ion"-, narrow bone which extends down almost to the lateral vacuity,
but which takes no part in forming the border of the vacuity.

The condyle is tripartite. The exoccipitals form the lateral portions,
while the basioccipital forms the lower median third. This latter lies
in advance of the exoccipital portions, so that the plane of the back of
the condvle is inclined and not vertical. The exoccipital forms the
half of the lower border of foramen magnum. It articulates laterally
with the paroccipital, the two being separated by a well-defined

suture.

The paroccipital is short and stout. It forms more than half the
border of the lateral vacuity and also encloses most of the foramen
jugulare. Below this foramen it articulates with the basioccipital.

South African Fossil Reptiles mid Amphibia. 167

The foramen jugulare lies below the level of the foramen magnuni and
looks downwards and backwards.

The palatal area is long and narrow, the basisphenoid region short
and broad. The minimum width across the pterygoids is under 4 mm.
There is a small but distinct transpalatine.

Types. Two skulls in nodule. (S.A. Mus. Cat. Nos. 3526, 3526a.)

Locality. Harrismith, O.F.S.

Horizon. Middle Beaufort Beds. (Lystrosaurus zone.)

GENUS PKOLYSTROSAURTJS, gen. nov.

PROLYSTROSAURUS NATALENSIS, gen. et sp. nov.

A skull from Loskop, Natal, is made the type of this new form.
It was found in beds of undoubted Middle Beaufort age associated
with typical skulls of Lystrosaurus. Details of the skeleton are obtain-
able from an almost complete specimen discovered by Dr. du Toit also
at Loskop, the skull of which is slightly smaller and slightly different
in shape from the type. Nevertheless, it is advisable to put the two
specimens in the same species.

The skull is intermediate in form between the normal Dicynodon
and the normal Lystrosaurus. The snout is bent down as in the latter
genus but is not elongated to anything like the same extent. The
interorbital region is broad, the intertemporal narrower. The nostril
is large and the distance between it and the orbit short. The temporal
openings are comparatively hrger than in Lijstrosaurus. The pineal
foramen is oval, as in Dicynodon.

The following table gives the chief measurements of the skulls of
the two specimens. It is probable that the type has been slightly
flattened laterally.

Type. Co-type.

Greatest length (front of snout to squamosal) . 110 mm. 118 mm.
Greatest width . . 85 ,, 105 ,,

Interorbital width 35 ,. 43 ,,

Intertemporal width . 22 ,, 23 ,,

Length of orbit . . . . 36 ., 38 ,,

Height of orbit 21 ,. 26

Height of suborbital border above edge of

maxilla ... 41 ,, 45

Width between tusks 30 33

Length from snout to front of pineal foramen . 57 ., 57 ,,

The chief features of the skull can be seen from the figures given.

14

168

Annals of the Soutli African Museum.

The premaxilla forms the whole of the lower and inner borders of the
nostrils and covers the front of the snout, passing up as a wedge-
shaped bone to lie in front of the nasals, appearing thus to separate
them from each other for part of their length.

The maxilla is a square bone supporting the jugal below the orbit
and carrying the tusk which passes downwards and appears below the
front part of the orbit. No definite septomaxilla can be seen on the
face, although it is probable that one is present.

'--PO.F

FIG. 43. Prolystrosaurus natalensis, Htn. Type. No. 3715. x f .

The nasals form a straight suture behind with the frontals.
Anteriorly they lie behind the premaxilla. They also form the insig-
nificant nasal bosses. Their width is greatest in front, as posteriorly
thev are narrowed bv the encroachment on the top of the snout of the

V

prefrontal.

The prefrontal is large, the lachrymal smaller. The frontals are
broad and do not pass behind the postorbital bar. The relations of
the bones round the pineal foramen is shown in the figure.

The outer bar of the temporal vacuity passes backwards and inwards
so that the greatest width across the skull is measured at the post-

South African Fossil Reptiles and Amphibia.

169

orbital bar and not as in Dicynodon. The downward processes of the
squamosals are also considerably expanded laterally.

The occipital plate slopes backwards, so that the basal length of
the larger skull is 100 mm. The foramen magnum is fairly small,
the condyle tripartite and large. The lateral vacuities lie far apart
and at the level of the bottom of the foramen magnum.

The lower jaw is short and the mentum is far back, the beak sloping
forwards gradually and fitting within the upper jaw.

Associated with the larger skull is most of the postcranial
skeleton.

FIG. 44. Prolystrosanrus natalensts, Htn. Type. No. 3715. x

Of the vertebral column there are 34 vertebrae preserved 25 cervical
and dorsal, 6 sacral, and 3 caudal, The number of presacral and
sacral vertebrae is similar to that given by Watson for Lystrosaurus.
The total length of the column from the occipital condyle to the back
of the posterior sacral is 410 mm.

The shoulder-girdle is almost entire. The sternum and interclavicle
cannot wholly be seen, the clavicles are slightly displaced, and the right
scapula has been crushed flat on to the ribs. The scapula is 85 mm.
long, 45 mm. broad at the upper end, 25 mm. broad at the acromion
process, and 35 mm. at the proximal end. The articular surface for
the precoracoid is 21 mm. long. Anteriorly it narrows to a point, but
posteriorly it is 17 mm. broad. The precoracoid is a rounded bone
25 mm. long and 27 mm. broad. It has a large notch on its upper
border. Posteriorly it forms apparently a small portion of the glenoid

170 Annals of the South African Museum.

cavity. The coracoid is of the normal Auoinodout type. The clavicles
are slightly curved and are 77 nun. long.

The humerus is 85 mm. long, has a breadth at the head of 47 mm.,
a width at the distal end of 49 mm., while the shaft has a minimum
width of 10 mm.

The radius is 54 mm. long, 18 mm. broad proximally, and 26 mm.
broad distallv. The bone is twisted so that the longer diameter of

o

the distal end is not continuous with that of the distal end of the ulna,
but is beut at an angle to it passing inwards and backwards.

The ulna is 57 mm. long, i. e. slightly longer in proportion to the
humerus than in Lystrosaurus. As in that genus there is no bony
olecranon process ; but on ihe uluar side of the facet for the humerus
the top of the bone is broad and flattened, possibly for the support of
cartilage acting as a cap. The maximum width at the proximal end
is 19 mm., at the distal end 17 mm., and the minimum width across
the shaft is 9 mm . The lower arm is carried at right angles to the
upper ann.

Part of the right fore foot is preserved. The carpals are not
ossified, but metacarpals I-III are there and some of the phalanges of
the digits, which are twisted inwards. The preserved portion agrees
closely with the hand of Lystrosaurus described by Watson (Geol.
Mag., 1913, p. 256). The third metacarpal is larger than the second.
The terminal phalanx of the thumb is a very broad, thin bone. The
following are the measurements of the bones seen on the dorsal surface,

in millimetres :

Proximal
breadth.

Metacarpal I . . 5 - 5 . 1O5

Phalanx I 911

II . . 12 12

Metacarpal II . . 11 '5 . 14

Phalanx I 7 12

II . 9 11

Metacarpal III . 14'5 . 14'5

Phalanx II . 6 11

II . 7-5 . 11

The ilium has a large preacetabular portion as in Lystrosaurus, but
is somewhat higher than in that genus. Its upper border carries two
notches, corresponding to the posterior two of the three figured by
Watson. The total length of the bone is 88 mm., the greatest height
63 mm., and the width at the acetabular end 30 mm. Compared with
Lystrosaurus the upper border of the bone ascends more rapidly from
the postacetabular process to the highest point.

South African Fossil Reptiles find Amphibia. 171

Neither ischium is complete, but the bone is greatly produced back-
wards. The pubis is not seen.

The femur is 95 mm. loug, has a narrow proximal width of 40 mm.,
and a distal width of 31 mm. The shaft has a minimum width of
16 mm. The median coudyle is somewhat smaller than the lateral,
and the medial surface of the bone is produced lower than the lateral.
The groove between the condyles on the posterior face is a little deeper
than that on the anterior.

Of the lower part of the limb only the proximal half is preserved.
The tibia has a strongly expanded end. The fibula is a slender bone
expanded proximally.

This form may be considered as a link between Dicynodon and
Lystrosaurus, approaching the latter rather closely. It has close
affinities with the form from Harrismith described by Broom as
Dicynodon strigops, but is at least specifically distinct in the fact that
the plane of the snout is at right angles to the froiito-parietal plane
and not inclined at 45. Moreover, the anterior border of the orbit is
rounded and not angular. These differences I am inclined to consider
specific and thus place Bi'oom's species in this new genus under the
name Prolystrosaurus strigops (Broom).

It is possible that the bending down of the snout and lengthening
of the premaxilla and maxilla which must have, taken place in the
evolution of Lystrosaurus from the earlier Auomodonts is foreshadowed
in such a form as Dicynodon testudirostris from the Cisteceplialus zone.
At the same time as the lengthening of the prenasal portion of the
snout took place there was a broadening of the frontal region and a
shortening of the temporal openings. In these features Prolystro-
saurus agrees with Lystrosaurus ; and the discovery of further forms
from the upper part of the Cistecephalus zone may enable us to trace
out gradual changes between D. testudirostris or some similar form and
Lystrosaurus through Prolystrosaurus.

Type. Skull and'lower jaw. (S.A. Mus. Cat. No. 3715.)

Go-type. Skull, lower jaw, and nearly complete skeleton (331</).

Locality. Loskop, Natal.

Horizon. Middle Beaufort Beds. (Lystrosaurus zone.)

PROLYSTEOSAURUS STRIGOPS (Br.),

1913. Broom, Dicynodon strigops. Eec. Albany Mus., ii, 5, p. 400.
1915. Broom, Dicynodon strigops. Bull. Amer. Mus. Nat. Hist., xxv,
2, p. 140 ; fig. 29.

Skull rather small. Skull unusually flat, orbits directed more

172 Aimed* of the South African Museum.

upwards than outwards ; tusks passing as much forwards as down-
wards. Beak elongated, upper surface at, 45 with fronto parietal
plane. Anterior side of orbit comes to an angle of 70-80. Parietal
region 23 mm. broad, frontal 33 mm. Large preparietal. Pineal
foramen bounded by parietals on the side and by preparietal in front.
Parietals unusually large. No opisthotic.

Type in the American Museum of Natural History.

Locality. (?) Harrismith, O.F.S.

Horizon. Middle Beaufort Beds. (Lystrosaurus zone.)

GENUS KANNEMEYERIA, Seeley.

KANNEMEYEEIA ERITHREA, Htu.
1915. Haughton. Ann. S.A. Mus., xii, 3, p. 91 ; figs. 12-14.

Kannemeyeria is the large Dicynodont which occurs in the Upper
Beaufort Beds. The skull is characterised by the high, narrow, convex
parietal crest which rises well above the interorbital plane, the broad
frontal region, the strongly ridged and rugose snout, and the large
nostrils. In these features it resembles to some extent Dicynodon
ijrandis, Eocyclops, and similar large forms from the Cistecephalus
zone, and also some of the species of Endothiodon. From the large
Lower Beaufort Dicynodons it differs in the size of the temporal
openings and in one or two other details. The parietal crest is more
pi-onouucedly convex. From Endothiodon it differs, of course, in the
lack of niolar teeth.

The beautiful skull which forms the type of this species has been
described fairly fully as far as the external features go. It does not
show clearly whether a preparietal is present, but a second fragmen-
tary skull from the type locality seems to indicate that a small pre-
parietal formed the anterior border of the pineal foramen. The
foramen lies in a deep excavation as in Dicynodon </r<ni<li* and on one
side of the wall of this hollow two sutures can be seen joining in front
of the level of the pineal foramen, forming presumably the borders of
an anterior wedge of the parietal lying on one side of the median pre-
parietal.

This same fragmentary skull has been sectioned longitudinally and
the brain-case between the foramen magnum and the opening for the
5th nerve cleaned. The cranial cavity is higher than wide. The basi-
occipital and basispheuoid are thick bones even in the middle line ;
and the basioccipital tubera lie considerably below the level of the

South African Fossil Reptiles and Amphibia. 173

midline of the basicranium. The floor of the brain-case is horizontal
in front of the foramen magnum with a median swelling flanked by a
slight groove on each side. Just in advance of the inner opening for
the 12th nerve the floor dips suddenly, and then rises as suddenly to
the opening for the 7th nerve. The sides of the brain-case are
vertical.

All the foramina for the nerve-exits are low down in the cavity.
They agree closely in position with those figured by Broom for Dicy-
nodon (Proc. Zool. Soc., 1912, PI. LVI). Between the small opening
for the 12th nerve and the larger one for the 9th-l 1th nerves there
is a broad transverse ridge of bone which crosses the floor of the

o

cranial cavity. The opening for the 9th-llth nerves is connected
with the upper end of the vestibule by a narrow groove. The upper
end of the vestibule is in ;i pit in the floor and lies considerably lower
than the other openings. The opening for the 7th nerve is small.
The vestibule for the ear was very long, but nothing can be seen of the
osseous canals.

Type. Skull and lower jaw. (S.A. Mus. Cat. No. 3017.)

Locality. Winnaarsbaaken, Albert, C.P.

Horizon. Upper Beaufort Beds. (Cynognathus zone.)

KANNEMEYEUIA LATIFRONS (Br.).

1899. Broom, Dicynodon latifrons. Ann. S.A. Mus., i, p. 452 ; pi. x,
tigs. 1-3.

Founded on an imperfect and crushed skull. Maxillary breadth
just larger than frontal breadth. Orbits directed laterally. Frontal
region broad and flat. Nasals greatly developed. Parietal crest
narrow. Parietal region inclined obtusely to the frontal. Pineal
foramen placed just behind postorbital bar. No postf rental. Ap-
parently no preparietal. Frontal width four times as great as the
intertemporal. Postorbitals do not touch one another. Skull deeply
hollowed in front of the pineal foramen.

Type in the Port Elizabeth Museum

Locality. Burghersdorp, C.P.

Horizon. Upper Beaufort Beds.

174

Annals of the South African Museum.

INDEX.

alticeps (Dicynodon) .

bolorhimis (Dicynodon)
breviceps (Dicynodon)

cavifrons (Dicynodon)
CHELYRHYNCHUS .

corstorphinei (Dicynodon) .
cyclops (Dicynodon) .

DICYNODON
DIICTODON

EOCYCLOPS .
erithrea (Kannemeyeria)

feliceps (Dicynodon) .

galeops (Diictodon)

gracilis (Dicynodon) .

gracilis (Myosaurus) .

grandis (Dicynodon) .

halli (Dicynodon)

ictidops (Dicynodon) .
ingens (Dicynodon)

jouberti (Dic}*nodon) .

KANNEMEYERIA .

kolbei (Dicynodon)

lacerticeps (Dicynodon)
lachrymalis (Chelyrhynchus)

laticeps (Dicynodon) .

PAGE
138

139

139

140
156
131
131

128
160

160
L72

133

160
134
164
140

142

135
142

128

172
143

144
]56
144

PAGE

latifrons (Kannemeyeria) . .173

leoniceps (Dicynodon) . . 146

leontops (Dicynodon) . .146

lissops (Dicynodon) . . 147

longus (Eocyclops) . . . 160

lutriceps (Dicynodon) . 136

megalorhinus (Dicynodon) . . 130

moschops (Dicynodon) . 147

mustoi (Dicynodon) . . 147

MYOSAURUS . . 164

natalensis (Prolystrosaurus) . 167

pardiceps (Dicynodon) 136

plazius (Dicynodon) . . 1-48

platyceps (Dicynodon) . 149

PROLYSTROSAURUS . 167

psittacops (Dicynodon) . 136

pygmaeus (Dicynodon) . 149

recurvidens (Dicynodon) . . 137

rogersi (Dicynodon) . .150

strigiceps (Dicynodon) . 151

strigops (Prolystrosaurus) . . 171

testudiceps (Dicynodon) . . 151

testudirostris (Dieynodon) . .151

tigriceps (Dicynodon) . . 153

trigoniceps (Dicynodon) . 138

truncatus (Dicynodon) . ].",>:;

tylorhinus (Dicynodon) . 154

whaitsi (Dicynodon) . .154

EXPLANATION OF PLATES XVI XVIII.

PLATE XVI.

Dicynodon grandis, Htn.

Fig. 1. Upper view of type skull. c j- nearly.
Fig. 2. Side view of type skull, x i.

PLATE XVII.
Eocyclops longus, Br.

Fig. 1 Upper view of skull. S.A. Mus. No. 3425. x % nearly.
Fig. 2. Side view of skull. S.A. Mus. No. 3425. x J- nearly.

PLATE XVI II.
Upper view of skeleton of Prolystrosaurus natalensis, Htn. $.

Ann. S.A. Mus., Vol. XII.

Plate XVI.

jf

DICYNODON <1RANDI8, HAUGHTON.

Aitlnrd <( Son 4- Went Newman, T.td:

Ann. S.A. Mus., Vol. XII.

Plate XVII.

EOCYCLOP8 LONGUS, BROOM.

Adlard $ Son <f" W ?st Neirmati, Lfil.

Ann. S.A. Mus., Vol. XII.

Plate XVIII.

PROLYSTROSAURUS NATALEN8IS, HAUGHTON.

Adlai-d J- Sun J- Weft XPIFIMHH, Ltd.

(175)

15.-- Investigations in South African Fossil Reptiles and Amphibia
(Part 11). By S. H. HAUGHTON, B.A., F.G.S., Assistant
Director.

11. Some New Carnivorous Therapsida, with Notes upon
the Brain-Case in Certain Species.

THERAPSIDA.

MACROSCELESAURUS JANSENI, n.g. et n.sp.

During the year 1917 the Museum became possessed, through
the generosity of Mr. F. J. Jansen, formerly Resident Magistrate
of Victoria West, of a slab of sandstone showing the impression
of the almost complete skeleton of a small reptile. The slab was
found built into the wall of a kraal at Gemsbokfontein, 12 miles
from the town of Victoria West, and there is no reason to suppose
that it had been brought from any place other than the nearest
sandstone ridge on the farm Gemsbokfontein. The known fossils
from the Victoria West District are few in number, and seem to
consist in the main of types not hitherto known from other parts
of the Karroo Beds, and this fossil therefore is of some interest.

Unfortunately, all the bone substance is weathered away, and
the mould in the sandstone which remains is not very clear-cut ;
and, in addition, the majority of the skull is altogether missing.
What is seen is a mould of the ventral side of the skeleton. The
backbone is curved ; both fore-limbs are thrown to the right
hand side and are bent ; and the hind limbs are flexed at the
knee, the femora lying thrown forward.

Of the skull but little remains. The head was obviously tri-

j

angular and sharply pointed in front. The teeth were small and
differentiated. There are remains of a canine and 8 molars in
the maxilla, the canine being preceded by small incisors, of which
the last, at least, remains. The canine is a larger tooth than
the others, and is set outside the line of the molars, so that the
skull must have had a considerable swelling at the level of the

15

176 Annals of the South African Museum.

canine or, in other words, the snout was constricted behind
the canine. Each tooth is elliptical in section, the somewhat
longer axis lying in the line of the jaw. The longer axis of the
cross-section of the canine is 3.5 mm. long, while the first 7 molars
occupy a length of 11 mm.

In the right ramus of the lower jaw 8 molars are to be seen,
each placed directly beside the corresponding tooth of the upper
jaw, the teeth of the maxilla biting outside those of the dentary.
The point of the tooth is bluntly rounded, and not sharply pointed.
It is probable that there were 3 or 4 more molars present in the
complete set. The posterior molars are slightly smaller than
the anterior ones in both upper and lower jaws.

There is a faint impression of one of the bones of the palate on
the right side having a curved outer border ; the bone is probably
thr palatine, forming the inner border of an oval suborbitul vacuity.
As preserved the skull is 56 mm. long ; but it was probably some-
what longer when complete. The front of the lower jaw was
narrow, fairly high, and sloped backwards.

The neck has almost disappeared ; only traces of the cervical
vertebrae can be seen, with the exception of the last. The neck
was fairly long and contained apparently 7 vertebrae. The whole
skeleton lies in position, and there seems to have been no dis-
placement of the neck. If this supposition of 7 cervicals be correct
then there are in all 27 presacral vertebrae. The centra of the
cervicals are longer than those of the dorsals. Every centrum
is constricted in the middle and expanded at the articular ends.
In the last cervical the centrum has a longitudinal ventral median
keel ; in the dorsals the ventral surface is rounded and very
slightly flattened. All the dorsals are very similar in size. None
of the vertebrae show the transverse processes or the zygapophyses.
Judging from the position of the ribs, the transverse processes
were fairly short. There are no intercentra present, nor is it
likely that any occurred in life ; the articular surfaces of the centra
show no bevelling on the ventral side.

There are three coalesced sacral vertebrae, of almost the same
size as the presacrals. The centra are rounded on their under
surface. The last cervical is 7 mm. long. The 3rd dorsal centrum
is 6.5 mm. long and has a maximum width of 6 mm. The three
sacrals are 6 mm., 6.5 mm., and 6.5 mm. in length respectively ;
the anterior end of the first is 7.5 mm. wide.

South African Fossil Reptiles a?id Amphifitu.

Ill

All the presacral vertebrae carry ribs. The presacral ribs are
possibly double-headed ; the articular end is long and broadened.
The ribs are long, slender, and curved ; their ventral surface is
longitudinally grooved. The anterior sacral rib is strong and
fairly long. It arises from the anterior portion of the centrum
and is directed strongly backwards. It has a swollen end ioi
articulation with the ilium. Only a very small portion of the

SCAR

FIG. 45. Impression of type. Macroscelesaurus janseni. x 0.48.

2nd sacral is preserved ; it was very much more slender than the
first, but articulated with the ilium. The 3rd sacral rib is a fairly
slender rib arising on the anterior half of the centrum and directed
outwards and slightly forwards to articulate with the ilium. Its
distal end is in contact with that of the 2nd sacral rib. There
is no evidence of abdominal ribs.

The pectoral girdle is not well preserved and the details of the
structure are difficult to determine. The scapula is only seen
in its lower .half. It has a fairly narrow shaft and a broad proximal
end, but apparently had no distinct acromion process. The
precoracoid is partially seen. It is a medium-sized bone with a

178 Annals of the South African Museum.

rounded anterior border ; there is in all probability a small
rounded foramen lying near the. suture with the coracoid. The
bone takes no part in the formation of the glenoid cavity, which
is bounded wholly by the scapula and the coracoid. The anterior
half of the coracoid only is seen. The border is notched just below
the glenoid cavity. A piece of rod-like bone lying near the coracoid
is probably a portion of the clavicle.

The humerus has a length of 60 mm. -The delto-pectoral crest
is not strong, but it extends nearly half-way down the bone. The
proximal half is somewhat broadened and is 15 mm. wide at its
broadest. The upper border of the articular end is bowed, and
there is a slight swelling at the head. The shaft of the bone is
slender, and the distal end slightly expanded.

The radius and ulna are both long, slender bones, very slightly
expanded at the ends. The ulna is 57 mm. long, and the radius
possibly 1 mm. shorter. The former has no olecranon process.
Both bones are nearly straight. The proximal surface of the
radius was slightly cupped. Distally the radius is somewhat
more expanded than the ulna. Nothing can be seen of the carpus
or manus.

The pelvis was broad. The pubis and ischium, although present,
have not been developed ; they were probably of the plate-like
type. The ilium had a pronounced posterior process which ex-
tended some way behind the articulation with the 3rd sacral rib.
The upper border of the ilium was probably about 20 mm. long.
The distance between the inner surfaces of the ilia is 27 mm.

The femur is long and slightly curved, with a maximum length
<>f 71 mm. The ends are not much swollen. The tibia and fibula
are long, slender bones, almost straight, with slightly expanded
ends. The tibia is stouter than the fibula. Each is as long as,
or slightly longer than, the femur.

Portion of the left tarsus is preserved. The fibulare is a larger
bone than the tibiale. Its proximal articular surface is strongly
concave, the outer edge being higher than the inner. The tibiale
articulates closely with the fibulare on the outer surface except
that between them in the distal half is a circular foramen such
as is seen in Broomia. The other elements are difficult of de-
termination. In the tarsus there seems to be a distal row of
tarsalia, and one centrale, lying distal to the tibiale. The re-
mainder of the foot is lying below the impression of the fore-limb
and so cannot be developed.

South African Fossil Reptiles and Amphibia. 179

From its general features this form seems to have affinities
with the Dromasauria, but it certainly differs from any of the
described forms. The most noticeable feature is the length of
the limbs, of which "the hinder are considerably longer than the
front legs. In Galechirus scholtzi the femur is only slightly longer
than the humerus ; in Galepns no measurements of the femur
have been given ; and in Galeops it is unknown. In all these
forms, moreover, the humerus is much longer than the radius
and ulna, while in the new form the upper arm is roughly the
same length as the lower arm.

The presence of a distinct enlarged canine brings the form
nearer to the Therocephalia than to the Dromasauria. The
absence of intercentra in the dorsal region is a feature shared
with the Dromasauria and other Therapsida. The pectoral girdle,
as far as can be seen, shows no difference from those of the
Dromasauria. There is the same absence of a distinct acromion,
the same two large coracoidal elements. The humerus is of some-
what different form. The delto-pectoral crest extends further
down the shaft than in Galc^ns or Galeops. The absence of an
olecranon process to the ulna is paralleled in Galechirus. The
sacrum is not known in any of the hitherto-described Dromasauria.
The posterior process to the ilium is seen in Galepus. The structure
of the tarsus is also typically Dromasaurian as far as can be seen.
It must be noted, however, that some of the features such as
the presence of two coracoidal elements and the possible tarsal
structure in which this form parallels the known Dromasauria
are primitive features common to most of the South African
Therapsida. Indeed, the differences between the postcranial
skeleton of this form and that of the Therocephalia such as Icti-
dosuchus as far as the latter group is known are not very striking ;
and the nature of the dentition allies the form to the Therocephalia.

The other long-limbed South African Permian reptiles are
Heleosaurus, Hcleophilns, Brooinia, and Mesosaurus. With none
of these is the form so closely allied as it is with the Dromasauria
and the Therocephalia, and it may be concluded that it is a Droma-
saurian specialised in the development of the limbs for swift
movement over the ground and akin to the Therocephalia in the
specialisation of the teeth. In the absence of a well-preserved
skull, however, it is impossible to dogmatise upon the systematic
position of the form.

180 Annah of the South African Museum.

It would seem advisable, therefore, in the absence of definite
correlation with any hitherto-described forms, to make the form
the type of a new genus which can be called Macroscelesaurus,
with the full name of Macroscelesaurus janseni in honour of the '
gentleman who transferred the specimen from a sheep-kraal wall
to the Museum collection.

Type. Sandstone slab with almost complete skeleton (in mould)
S.A.M. Cat, No. 4004.

Locality. Gemsbokfontein, Victoria West, C.P.

Horizon. Lower Beaufort Beds (Tapinocephalus zone ?).

ALOPECOGNATHUS MINOR, n.sp.

The following description is based upon the almost perfect
skull of a medium-sized Therocephalian found by the Rev. J. H.
Whaits at Klipbank Siding, south-west of Beaufort West, in beds
which belong to the upper part of the Tapinocephalus zone. The
matrix is extremely hard, but it is possible to clear the bone, and
I have been able to discern most of the features of the occiput
and basicranium as well as of the upper part of the skull.

The skull is long and narrow, the orbit lies in the middle of
the skull, the parietal crest is narrow and fairly high, and the
dental formula of the upper jaw is i6clm4. The skull, in its
general appearance and in the possession of 6 incisors, displays
affinities with Alopecognathus an(/usticeps. There are four molars
in the skull under description instead of six, the eyes are placed
further forward, and there are differences in the relative positions
of the teeth. Of the basicranial region of Alopecognathus angus-
ticcps no description has been given. In spite of the differences
mentioned I deem it advisable to keep the two forms in the same
genus, and I propose for the new form the name Alopecognathus
minor, sp. n.

The premaxilla is small, and carries 6 incisors which occupy a
space of 31 mm. The 5th and 6th incisors are slightly smaller
than the others.

The single large canine lies 10.5 mm. behind i6, and has a diameter
of 12 mm. Then follows a diastema of 8 mm., behind which are
4 small molars occupying 26 mm. The molars are short, but
robust, and flattened. All the teeth were probably finely serrated
along their posterior borders, the incisors and molars certainly so.
The snout is bluntly rounded. The nostrils are almost terminal,
and look nearly entirely upwards. They are longer than wide,

South African Fossil Reptiles and Amphibia.

181

and the anterior ends approximate to one another The whole
of the lower border is formed by the septo maxillary which also
foims the upper border of the small external foramen, and passes
back some distance behind the nostril.

The maxilla is long and narrow. The nasal is long and com-
paratively narrow, broader at its two extremities than in the
middle. Posteriorly the two bones are separated for a short
distance by the frontals.

FIG. 46. L'fpsr view of type skull of Aloffcogmit-lrus tinnm . x J.

FIG. 47. Ventral view of type skull of Alopecognathus minor as
developed, x ^.

The prefrontal is fairly large ; the lachrymal smaller. There is
a pronounced antorbital depression.

The orbit is longer than wide, almost wholly in the posterior
half of the skull. The postorbital arch is comparatively weak.

There is a pronounced triangular-shaped postfrontal. The
postorbital does not pass far along the upper border of the
temporal fossa, most of the border being formed by the parietal.

There is a large pineal foramen 10 ram. behind the postorbital
arch, wholly surrounded by the parietals There is no preparietal

182 Annals of the South African Museum.

Unfortunately the sutures of the occiput are not clearly shown,
but some points in the structure can be discerned. The basi-
occipital condyle is large and rounded, and projects considerably
behind the occipital plate. The lateral margin of the foramen
magnum is formed by the exoccipital, which also forms the upper
margin of the foramen jugulare. The exoccipital is a larger bone
than that described by Watson in the Gorgonopsians Arctops and
Scymnognath'us. The two exoccipitals do not meet, but the upper
border of the foramen magnum is formed by the supraoccipital.
The post-temporal fossa is of moderate size, on a level with the
foramen magnum. The bone which forms almost the whole of
its upper border is, I think, the supraoccipital, which is very wide
and comparatively shallow. Superiorly there is a pronounced
horizontal backward extension of the squamosal, so that the
tabulare is hidden when the skull is viewed from above. This
bone, the limits of which I cannot determine, lies laterally to the
supraoccipital and forms part, at least, of the outer border of
the post-temporal fossa. In the Therapsida as far as it has hitherto
Ix-cn described it almost invariably extends down outside this
f<>ssa to meet the paroccipital process ; but the lower limit in
this form is indeh'nable. It is very doubtful, however, whether
the tabulare and the paroccipital process meet. The paroccipital
process is a stout, high bone forming the whole of the lower border
of the post-temporal opening. Externally it articulates for most
of its height with the squamosal. Internally it meets the ex-
occipital and the basioccipital. The extreme inner and front
corner of the bone forms part of the border of the fenestra ovalis.
The paroccipital process has a considerable forward extension
laterally, and the outer part of its under surface has a groove
ending in what is apparently a foramen between this bone and
the squamosal.

The fenestra ovalis is a medium-sized hole, apparently not
bounded by bone on its outer side. Its outer posterior border is
formed by the paroccipital process, the anterior half of the border
is formed by the basisphenoid tuber and the prootic, and the
remainder by the basioccipital. It lies well in front of, and a
little below, the basioccipital process.

Seen from below, the anterior part of the basioccipital between
the fenestrae ovales is strongly concave, and the median depression
is continued forward to the basisphenoid. Anteriorly it ends
abruptly and gives place to a deep and narrow median basisphenoid

South African Fossil Reptiles and Amphibia. 183

keel formed by the junction of the ridges from the tubera, and
having its lower border considerably below the level of the basi-
occipital condyle. This condition is also seen in the primitive
Gorgonopsian Arctops and in the Therocephalian Scymnosaurus
watsoni. The basisphenoid keel passes forward almost to the
lateral process of the pterygoid, lessening in height anteriorly and
being clasped anteriorly and superiorly by the vertical pterygoid
plates.

On the median line these pterygoid plates pass back almost to
the level of the back of the basisphenoid keel. They are at first
vertical, then turn outwards and finally downwards, so that in
section they have the appearance of a curved arch. The outer
ramus passes back presumably to the quadrate.

In front of the basisphenoid keel there is a narrow, somewhat
elongate pterygoid vacuity, at the sides of the back of which the
powerful pterygoid flanges pass out to meet the transpalatine.
The transpalatine is comparatively small.

There is a fair-sized suborbital vacuity. The inner posterior
border is formed by the pterygoid, which passes forward with the
same relation to the palatine as in Scymnosaurus watsoni. I am
not able to distinguish a median vomer. The pterygoid has a small
ridge bearing a few teeth.

The front of the palate is not displayed, but it is probably
typically Therocephalian in structure. Its chief feature is its great
length compared with the breadth ; in fact, the whole palatal
aspect is long and narrow.

The lower jaw is practically complete. The symphysis is loose
and low, the front of the jaw rapidly retreating. There is a large
and wide coronoid process to the dentary. The angular is large
and passes well forward to below the middle of the orbit. The
splenial plays no part in the symphysis.

The following are some of the chief measurements of the type
specimen :

Greatest length . . .... 240 mm.

Greatest width .. (about) 110 ,,

Snout to front of orbit .. .. .. ..Ill ,,

Interorbital width . . . . . . . . . . 30.5 ,,

Intertemporal width .. .. .. .. 11.5

Length of temporal opening . . . . 58

Type. Skull and lower jaw (S.A.M. Cat. No. 3415).

Locality. Klipbank, Beaufort West Division, Cape Province.

Horizon.- Lower Beaufort Beds (Tapinocephahis zone.)

184 Annals of thr. So-itli African

WHAITSIA PLATYCEPS, gen. et sp. nov.

The extremely interesting type described here is founded on
two skulls collected by the Rev. J. H. Whaits at Zuurpoort in
the District of Graaff-Reinet, C.P., at a height of 4,000 feet above
sea-level, and is therefore probably from the middle or upper
part of the Cistecephdlus zone of the Lower Beaufort Beds. One
skull has suffered lateral pressure ; the other a more perfect
one has been slightly flattened from above. The latter No.
4006, S.A. Mus. Catalogue may be taken as the type ; the other
-No. 4330 as the co-type.

In external appearance the skull might be mistaken for that
of a fairly broad-snouted Cynodont. The nostrils are small,
almost terminal, and look mainly forwards. The snout is rounded
at the end, broad, flat, and of a medium length. The orbits are
small and widely separated. The temporal openings are very
large and the parietal region is narrow, crested as in the Cynodonts
and Therocephalia and not broadened as in the Bauridae and
Gorgonopsia.

The maximum length of the type skull was about 280 mm.,
the maximum width about 220 mm. The interorbital width is
66 mm.

The internasal septum is formed in the lower half by the pre-
maxilla and in its upper half by the nasals. The premaxilla
forms the lower border of the nostrils. In the type the premaxilla
carries no teeth, with the possible exception of one fragment on
the right-hand side ; but in the co-type there are 4 incisors on
each side of which the first three are equal in size, the 4th smaller.
The roots are long : the crowns were apparently long and sharply
pointed, but not serrated.

The top of the snout is formed almost entirely by the nasals.
These are broad anteriorly, contract in the middle where the
maxilla encroaches on the top of the skull, broaden again, and
then narrow to their junction with the frontals. There is a small
boss above the outer side of the nostril which may be formed
by a septomaxilla as in the Gorgonopsia, but the limits of the
bone, if present, are not seen. External to the nostril are two
foramina corresponding in position to the foramen found in the
Therocephalia and Gorgonopsia.

The maxilla forms the side of the snout, extends below the
jugal to the level of the postorbital bar, and in its posterior half

South African Fossil Reptiles and Amphibia. 185

has a gradually widening broad palatal portion. Anteriorly it
carries a large canine tooth which does not fill the alveolus ; but
behind the canine I can see no trace of any molar teeth nor of
alveoli for such teeth in either skull.

The prefrontal is large and forms a large portion of the upper
border of the orbit. The frontal is fairly short and seems to form
portion of the supra-orbital border and meet the postorbital

FIG. 48. Upper view of type skull of Whaitsia pliii/ceps. x f.
FIG. 49. Ventral surface of type skull of Whaitsia plati/ceps. x f-.

posteriorly. It may be, however, that part of the bone is in
reality postfrontal. There are several irregular sinuses and pittings
in the bone, but no definite suture is visible. If this be so, then
the condition of the frontal would be similar to that in the Gorgon-
opsia.

There is no preparietal.

The postorbital bar is long and the postorbital extends some
distance along the parietal crest, being interlocked with the parietal.
This crest is narrow as in the Therocephalia and. the Cynodontia.
There is an elongate pineaf foramen 25 mm. behind the post-
orbital bar.

186 Annals of the South African Mn^tuin.

The parietal forms the main portion of the parietal crest and
is almost entirely a vertical bone. Lying behind it on the occipital
plate is a large interparietal which articulates below with the
supra-occipital and laterally with the tabulare.

The jugal forms the lower border of the orbit and extends back
to meet the squamosal.

The occipital condyle is large and moderately thick and does
not extend very far up the sides of the foramen magnum. It
is shallower than that of Arctops and thicker than that of Scym-
noc/xatkus figured by Watson. It is not certain to what extent
the exoccipitals participate in the formation of the condyle, but
certainly it is for the most part formed of the basioccipital.

The exoccipital differs from that of Arctops it is a small tri-
angular bone forming the upper border of the foramen jugulare
and lying behind the paroccipital process. It plays no part in
the boundary of the post-temporal fossa, and is well marked off
from the paroccipital process.

The suture between the basioccipital and basisphenoid can be
seen a limit 31 mm. in advance of the back of the condyle. The
mass of the basisphenoidal tubera, which are not extremely large
although quite prominent, is formed bv the basioccipital. the
basisphenoid lying below as in the Dicynodonts. Jjetween the
tubera the basioccipital is flat. Laterally the bone presumably
articulates with the opisthotic, but the suture is not visible.

The foramen jugulare lies at a level just above the bottom of
the condyle and faces entirely backwards. Its upper border is
formed by the exoccipital, but in the main it lies in the paroccipital.
whose posterior face is hollowed out horizontally on the outer
side of the foramen.

The post-temporal fossa lies at the level of the middle of the
foramen magnum ; consequently the paroccipital, which is long,
is more massive than in Scymnognathus tchaitsi and less so than
in Arctops.

The fenestra ovalis is an irregular-shaped opening, unbounded
by bone on the outer side and lying some distance out from the
basisphenoidal tubera, below the level of the occipital condyle.

Just in front of its suture with the basioccipital the basisphenoid
forms a strong median keel which is continued forwards between
the pterygoid vacuities by the pterygoids. The exact junction
between the basisphenoid and the* pterygoids is difficult of de-
termination on account of fracturing in this region ; but the

South African Fossil Reptiles and Amphibia. 187

posterior part of the ridge presumably basisphenoid is pierced
on either side by a large foramen, the upper border of which seems
to be formed by the periotic. This is presumably the fossa identified
by Watson in Diademodon as the pituitary fossa. A fracture
across the basisphenoid of the co-type just in front of the median
ventral ridge shows that the dorsal surface of the bone is broadly
rounded.

The pterygoid is a large bone attached to its neighbour throughout
its length except where it forms part of the median keel, where
the two are separated by the anterior portion of the basisphenoid.
There is no interpterygoid vacuity. In front of the median keel
the bones form a broad plate which is hollowed out in the middle.
This hollowed portion occupies the position which is taken up by
the vomer in the Gorgonopsia and the Cynodontia ; but here are
no sutures to divide this portion from the rest of the pterygoid,
and the vomer, in consequence, seems to play no part in the
formation of this part of the palate. Moreover, the suture between
the two pterygoids can be traced almost throughout the length
of the plate-like portion.

Laterally the pterygoid articulates with the transpalatine. It
forms the anterior boundary of the pterygoid vacuity 'and also
thickens to form the fairly small downward process or pterygoid
flange. The lower surfac" of this flange, however, is formed
entirely by the transpalatine.

Anteriorly, the pterygoid narrows and passes forward as a narrow
bar between the palatines to meet the prevomer. The lateral
posterior ramus of the pterygoid passes outwards towards the
quadrate and lies in front of a thin inwardly-directed process of
the squamosal. A section across this ramus in the co-type shows
the ramus lying below a thin plate of bone with which it is slightly
interlocked. This thin plate is presumably part of the quadrate.
The pterygoid is separated fairly widely from the paroccipital.
Attached to the outer side of this ramus is the epipterygoid or
columella cranii. This is a broad bone lying pressed against the
pterygoid below and the parietal above. Its upper articulation
is long ; its anterior upper end lies considerably in advance of
the lower end ; and its anterior border is concave. Posteriorly
its lower half extends far back, while the upper half articulates
with the periotic. The bone occupies the same position and is
of similar shape to the pterygoidal portion of the side-wall of

188

Annals of the South African Museum.

the brain-case figured by Watson in Diademodon ; but it differs
in being obviously marked off by suture from the pterygoid.
Broom figures the same bone as the alisphenoid.

The transpalatine articulates anteriorly with the palatine and
maxilla and medially with the pterygoid. In the suture between
the pterygoid and transpalatine there is a very small pit which
may be looked upon as the relic of the suborbital vacuity of the
Therocephalia.

WHAITSIA PLATYCEPS Htn.

FIG. 50a. Transverse section just behind "transverse bar" of
type.

FIG 50b. Section across distorted snout of co-type.

FIG. 50c. Section of prevomer just in front of "transverse bar '
in type.

The palatine articulates \\ith the maxilla laterally, with the
pterygoid medially, and anteriorly it meets the prevomer. The
larger portion of the bone is vertical. Laterally it has a narrow
horizontal portion lying alongside the maxilla and approximating
to the bone on the other side to foreshadow the formation of a
secondary palate. Then it bends abruptly at right angles, especially
in its anterior half, and is again bent to form with the pterygoid
and prevomer a plate considerably higher than the level of the
edge of the maxilla.

The palatal portion of the maxilla broadens rapidly behind
the canine and passes back to form part of the anterior border
of the pterygoid vacuity. Its outer process lies under the jugal,
which bone is completely shut out from articulation with the
pterygoid by the maxilla and the transpalatine.

South African Fossil Reptiles and Amphibia. 189

The prevomer forms part of the anterior median bar and lies
between the palatine process of the premaxillae and the anterior
portions of the pterygoids. In shape it is cruciform, the transverse
arms forming with the palatines a bar separating two vacuities in
the anterior part of the palate. Posteriorly it unites with the
pterygoid. In cross-section just behind the transverse portion
the prevomer is seen to be a triangular bone with its apex above,
and undivided by any median suture. Superiorly it supports
a thin plate of bone which has been crushed in the specimen. This
thin plate passes back to meet the upper portion of the pterygoids
the exact method of articulation is not perceptible.

There are no palatal teeth, nor does the palatal surface of the
maxilla carry any molars.

The palate shows many features of interest, and is seen to differ
considerably from any described South African form. In the
first place there are the two anterior vacuities to be accounted
for. Fortunately it has been possible to develop the type specimen
In nly completely, and the main structure can be ascertained.
The median bar lies considerably above the level of the dentigerous
border of the premaxilla and the palatal portion of the maxilla ;
in fact, the front of the palate is highly vaulted. The inner border
of the anterior vacuity is formed by the premaxilla and the pre-
vomer the median bar. On the palatal surface half of this bar
is formed by the prevomer ; but on the nearly vertical side, of the
bar the premaxilla plays by far the major part. The side- wall
of the bar does not ascend to the roof of the skull, but stops abruptly
about half-way between the palate and the under side of the roof.
The front border of the cavity is formed by the premaxilla, the
outer border by the maxilla, the posterior border by the palatine
and prevomer, and it is roofed by the cranial bones of the region
of the nostril. Into this cavity the outer foramen, seen lying
external to the nostril in outer view, leads directly.

The posterior cavity is bounded by the transverse bar anteriorly,
bv the palatine laterally, and the pterygoid and prevomer medially.
A vertical section fortunately obtainable shows that the two
cavities anterior and posterior are connected with the external
nares. The "transverse bar' has its upper surface sloping
downwards and backwards, forming \vhat seems to be the floor
of the narial passage, leading one to conjecture that the posterior
vacuities are the internal nares. The anterior cavity must then
be the anterior palatine foramen (foramen incisivum). Although

190 Annals of the South African Museum.

the prevomer, the palatine, and the premaxilla each plays a small
part in the formation of its border, it is mainly surrounded by the
premaxilla. It is very large, the size being its most striking feature.
It leads directly up into the nasal cavity. A section across the
snout of the co-type shows that the nostril is divided into two
unequal parts by a nearly horizontal turbinal, probably formed
by a process of the septomaxilla. The upper portion is very
much larger than the lower, whose floor is concave with a slight
median ridge formed along the junction of the dorsal surfaces
of the two premaxillary bones. The evidence seems to connect
the lower portion with the anterior vacuity of the palate, while
the upper part the true nostril is connected by a more gradually
sloping tube with what has been identified as the internal narial
opening.

Hitherto, the anterior palatine vacuity has been observed in
very few Therapsids, and never to the extent displayed here.
Broom (1911) mentioned its occurrence in Gomphognathus, and says
' In the anterior palatine vacuity there is a pair of narrow bones
showing what I suggested a good many years ago were probably
prevomers." Here the opening seems to have been a single
median one ; its size is not mentioned, and it is not figured. The
same author (1914) has figured a pair of openings surrounded by
the premaxillae in Lycochampsa ferox. These two forms are both
( Vnognathids, and the vacuity does not seem to have been recorded
in any of the Therocephalia or Gorgonopsia.

Another interesting feature concerns itself with the vomer. In
Diademodon there is a large median vomer lying between the
pterygoids and palatines. Watson considers that in Gorgonops
there is a large mammalian vomer in the posterior part of the
palate as in Diademodon and a pair of fused prevomers anteriorly.
In the Therocephalian Scymnosaurus ivatsoni (described by Watson
as Lycosuchus sp.) there is a narrow vomer lying in the same position
as in Gorgonops, and a pair of prevomers anteriorly. In Scylaco-
saurus sdateri there is no median vomer on the palate. From
the general features of the skull under discussion one would expect
to find in it some trace of a posterior vomer on the palate. I can,
however, find no trace of sutures where such a bone is likely to
occur, and, moreover, the pterygoids are seen to be separated
from one another almost throughout their length by a median
longitudinal suture. On the other hand, the bone separating the
premaxilla from the pterygoids is a single bone, as in Dicynodon.

South African Fossil Reptiles and Amphibia. 191

Such a bone in the Gorgonopsia is said by Watson to be a fused
pair of prevomers and has been described by Broom as a true
unpaired vomer. A section across the median bar just in front
of the 'transverse bar" shows the upper surface of this bone
with a median ridge, bounded on each side by a broad shallowly
concave depression and a lateral rounded ridge. Lying to one
side of the median ridge is a narrow spike of bone which was
doubtless a vertical median septum before crushing. It may be
that in this skull we have a fused pair of prevomers and that the
thin median bone seen above it in section which passes back to
articulate with the pterygoids is the vomer. This latter may,
however, be the mesethmoid.

Then the nature of the dentition is peculiar in that no molars
are present. It might be supposed that all the teeth had been
lost during life ; but both skulls show the same features, and
there are not even remnants of the alveoli still preserved, nor of
roots when seen in section.

A portion of the lower jaw was found crushed on to the back
of the type skull. It is part of the right dentary. Near the
anterior end is a canine tooth, oval in cross-section. Just in front
of this is seen in cross-section a much smaller tooth, occupying the
position of an incisor. The outer surface of the anterior part of
the dentary is rugose. The dentary is long and slender. Behind
the canine the upper edge is regularly concave right back to the
coronoid process, so that when the jaw is placed in position in the
skull there is a wide gap between the upper and lower jaws behind
the canines. Posteriorly the upper border of the dentary narrows.
There is no distinct mentum, the dentary being thus more remi-
niscent of the Therocephalia or the Cynodontia than of the
Gorgonopsia.

The lower jaw is more fully preserved in the co-type. The
symphysis is long and loose. The front of the jaw is prow-shaped
and there is no distinct mentum. Each dentary carries one large
canine, and, fairly closely in front of it, one incisor.* The dentary
is large and long, but not very deep, and has a very large coronoid
process. The bone forms almost as large, a portion of the lower
jaw as in Cynognathus, and certainly a greater portion than in a
Gorgonopsian such as Scymnognathus. The splenial is a small

* In a specimen consisting of the front halves of both dentaries from Kruger's
Kraal, Graaff-Eeinet, 3 incisors occur in front of the canine. This specimen
probably belongs to the species under discussion.

16

192 Annals of the South African Museum.

bone playing little or no part in the symphysis. The post-dentary
part of the jaw is seen partly in external view and partly as a
cast of the inner side. The structure is somewhat difficult to
determine as a certain amount of crushing has obliterated the
sutures. There, is a long posterior process lying ventrally as in
Cynognathus presumably part of the angular, which is much
shallower anteriorly than in Scymnqgnathus. The prearticular
and surangular are seen occupying their normal positions, but
the actual articular surface is missing.

Although in the general character of its palate and occipital
plate the form approximates to the Gorgonopsia, it differs from
that group in the narrow parietal crest, the absence of the pre-
parietal, and the absence of the vomer on the palate. It has
been shown, too, that it differs also from the typical Theroce-
phalia, Cynodontia, and Bauridae ; and I have therefore placed
it in a new genus under the name WJiaitsia platyceps named in
honour of the Rev. J. H. Whaits, to whose zeal, energy, and
knowledge as a collector South African palaeontology owes such
a debt of gratitude. This new genus may be considered as the
type of a new family of Therapsida, the Whaitsidae.

Type. Skull and partial lower jaw. S.A. Mus. Cat. No. 4006.
Co-type. Skull and lower jaw. S.A. Mus. Cat. Xo. 4330.
Locality.- Zuurpoort, Graaff-Reinet District, C.P.
Horizon. Lower Beaufort Beds, Cistecephalus zone.

AKIDNOGNATHUS PARVUS n.g. et n. sp

The type skull of this species shows some interesting features.
It was collected from the Cistecephalus zone of the Lower Beaufort
Beds at Zuurpoort, Graaff-Reinet, C.P., by the Rev. J. H. Whaits.
(S.A. Mus. Cat. No. 4021.)

The skull is small, the snout broader than high. Its greatest
length is 107 mm., the greatest breadth 54 mm. The distance
from the snout to the front of the orbit is 50 mm., the anterior-
posterior diameter of the orbit 25 mm., and the interorbital width
22 mm.

The dental formula is doubtful ; it may be i5 c2 m7 or i6 cl m7.
The sixth tooth from the centre line is smaller than the other
incisors, and it is not possible to determine absolutely whether
it is carried by the premaxilla or the maxilla. In Scaloposaurus
and Ictidoynathus the number of canines is greater than one, and

South African Fossil Reptiles and Amphibia.

193

there is thus no inherent improbability in supposing that this
allied form has two canines. The 5 incisors, supposing the sixth
tooth to be a canine, occupy a space of 22 mm. The second
canine is the largest tooth, oval in cross-section, and having a
longer diameter of 5 mm. There is a diastema between the 1st

FIG. 51. Palatal view of type skull of Akidnognathus parvus.

canine and the 5th incisor of 2 mm. ; and one of equal size between
the large canine and the 1st molar. The molars are much smaller
than the incisors, and the 7 teeth occupy a space of 12 mm. The
teeth are not serrated ; the incisors are sharply pointed, back-
wardly curved, with longitudinal grooves. The molars are pointed,
with anterior and posterior borders. Where the crowns are fully
preserved, they are seen to be somewhat grooved,

194 Annah of the South African Muteum.

The nostrils are large, close together, almost terminal, and
look upwards and forwards.

The bones of the skull surface are similar to those of Scal<>/><'-
saurus. The frontal forms part of the orbital border. The parietal
crest is ridged, but the parietal region is fairly broad ; the temporal
openings are short. There is no postfrontal and no preparietal ;
but a pineal foramen is present. The postorbital arch was possibly
incomplete.

The basicranial region is essentially similar to that of Xcalopo-
saurus. The occipital condyle is single. The large jugular foramen
looks almost entirely downwards. The basisphenoidal tubera
lying in front of the fenestrae ovales are stronger than in the
specimen figured by Dr. Broom. The two carotid foramina lie
one on either side of the posterior end of a short basisphenoid
keel. Anterior to this the pterygoids are separated by a V<T\-
long interpterygoid vacuity. The suborbital vacuity is present
as in the Therocephalia and is bounded by the transpalatine,
pterygoid, and palatine.

A fracture across the interpterygoid vacuity shows a small
bone lying in the median line about 5 mm. above the level of the
basisphenoid keel. In cross-section it is seen to have a roundly-
keeled ventral surface and a shallowly concave broadened upper
surface. This bone is not seen as a separate bone in a section
taken a little further back, across the basisphenoid keel ; but the
upper surface of the basisphenoid is similarly grooved. On the
ventral surface of the latter bone there is a narrow groove on
either side of the median keel.

Anterior to the interpterygoid vacuity the palate is slightly
vaulted and rises slightly anteriorly. The structure seems to be
similar to that of Scylacosaurus except that the distance between
the posterior nares and the transverse process of the pterygoid
is much shorter. The palatine occupies its usual position, forming
part of the posterior border of the internal narial opening and of
the anterior and inner borders of the suborbital vacuity, and
articulating with the pterygoid and prevomer medially. Its
inner portion has a longitudinal canal, covered both dorsal I y
and ventrally with a very thin layer of bone. I cannot be certain
of a median suture between the pterygoids. That between the
prevomers is very definite in the posterior half of the bones, which
reach very far back, but between them and the interpterygoid
vacuity it is not definite. On the other hand, the presence of

Soittli African Fossil Reptiles and Amphibia.

a median vomer on the palate separated by sutures from the
pterygoids is equally uncertain, and the structure of that region
must remain doubtful. The bone is cracked in a rather sym-
metrical manner, which increases the difficulty of determining
sutures.

n

c.

UT Pr VAC

AKIDNOGNATHUS PARVUS n.g. et n. sp.
FIG. 52a. Section across skull just behind internal nares.
FIG. 52b. Cross-section at middle of interpterygoid vacuity.
FIG. 52c. Section across basisphenoid at front of keel.
FIG. 52d. Section across prevomers near anterior end.

As mentioned, the prevomers are separated by a definite suture
posteriorly ; but anteriorly the suture becomes obliterated, and
the bones broaden out and become thinner. A section across
the front of the bone shows them forming a single median septum,
thin below, but broadening dorsally to form a grooved upper
surface. The palatal portion of the bone is thin and concave
dorsally, as in Ictidognathus parvidens. In the upper part of the
median septum there is a small canal.

196 Annals of the South African Museum.

There is an incipient secondary palate. The palatal surface
of the maxilla has a groove terminating in a foramen which leads
to a canal passing backwards through the bone. There is also
an oval foramen in the palatine just behind its suture with the
maxilla.

The lower jaw is slender and carries 4 or 5 incisors, 1 canine,
and 6 molars. The dentary is long but not particularly powerful,
and is not furnished with a distinct mentum. A section across
the jaw at the level of the back of the orbit shows a splint of bone
lying internal to. and separated from, the upper portion of the
dentary. This splint is probably the coronoid. The hinder
portion of the jaw is very reduced in depth and length. The
angular and surangular can be seen, but their relations are not
determinable.

The form is closely allied to Scaloposaurus and Ictidognathus,
and may be safely placed in the Scaloposauridae. This family
shows some affinities with the Bauridae, agreeing in the following
characters :

1. The short temporal region.

2. The nostril directed more forward than outward.

3. The frontal forming part of the orbital margin.

4. The small squamosal.

5. The interpterygoid vacuity.

(i. The great distance between the posterior ramus of the
pterygoid and the paroccipitul process.

7. The suborbital vacuity.

8. The absence of the long narrow pterygoid-basisphenoid

median bar.

9. The presence of a quadrate ramus of the pterygoid.
10. The feeble postorbital arch.

Features 3, 5, 7, and 9 are common also in the lower Therocephalia.
It differs from the Bauridae in :

1. The nasal being slightly widened posteriorly.

2. The nature of the molar teeth.

3. The larger size of the interpterygoid vacuity, and the

consequent great reduction of the basisphenoidal keel.

In the latter feature the Scaloposauridae differ more from the
Therocephalia than do the Bauridae ; but in the nature of the
molars they approximate to the Therocephalia.

Type. Skull and lower jaw (S.A. Mus. Cat. No. 4021).

Locality. Zuurpoort, Graaff-Reinet District, C.P.

Horizon. Lower Beaufort Beds (Cistecephalns zone).

South African Fossil Reptiles and Amphibia. 197

CYNOSUCHUS WHAITSI, n.sp.

The form Cynosuchus suppostus was founded by Owen on the
preorbital portion of a skull and lower jaw " from a clavstone
nodule in a (triassic ?) formation of the Sneeuwberg mountain
range." He considered that it belonged with Cynochampsa to a
'' single-nostrilled group of Theriodontal Saurians." Both Owen's
figures and description show the dental formula of the upper jaw
in the type to be i4clm8 there being 8 molars on the right hand
side and 6 on the left.

Lydekker (1890) gave the number of upper molars as seven.
He noted that the skull has been flattened from above, giving the
muzzle an abnormal width. He also considered the absence of
the internarial septum to be due to crushing.

Broom in 1915 made the following remarks: "Cynosuchus
suppostus is known only by the imperfect type skull. It is a most
remarkable form, having cusped molars and a secondary palate like
the typical Cynodonts, but in other respects differing from all
known Cynodonts and resembling more the Gorgonopsians. The
dental formula is probably i^ciinf, and to whatever sub-order a
more complete skull may show it to belong, it must be placed in
a distinct family the Cynosuchidae."

The discovery within the last few weeks of an almost complete
skull by the Rev. J. H. Whaits enables some further observations
to be made. Although, owing to the imperfect nature of the
type, the type species is somewhat indeterminable, it seems probable
that this new skull is closely allied to it, and should be considered
as a member of the genus Cynosuchus known under the name
Cynosuchus u-haitsi. The skull was found at Weltevreden, in
the Sneeuwberg range, GraafT-Reinet, at a height of 5,000 feet
above sea-level, and is therefore from the top portion of the Ciste-
cephalus zone of the Lower Beaufort Beds.

The skull is somewhat obliquely crushed, but the larger portion
of it has been freed of matrix and the details of the structure
beautifully displayed.

The greatest length was probably 125 mm., the antorbital length
50 mm., the interorbital width 27 mm., and the basal length
120 mm.

The dental formula is i4clm8. The four incisors are closely
set together and occupy a space of 14 mm. They are simple
pointed conical teeth without serrations and curve slightly back-
wards. Each tooth has a few longitudinal grooves which do not

198

x of tin South African Museum

fix

B.Occ

AL.S.

SPH.

CYNOSUCHUS UHATTSI n. sp.
FIG. 53a. Palatal view of type skull, x 1.
FIG. 53b. Canine and molars of right side
FIG. 53c. Section across skull at back of notch in alisphenoid

South African Fossil Reptiles and Amphibia. 199

continue to the point of the crown. Behind the 4th incisor is a
diastema of 5 mm. which is followed by the large canine tooth.
This has a compressed oval cross-section with a longer antero-
posterior diameter of 8.5 mm. It, too, is unserrated. Closely
f olio wing the canine are the 8 molars (only 6 are preserved on
the left side). These occcupy a space of 25 ram. Speaking
generally, they decrease in size backwards ; but they are not of
equal age, the 2nd and 4th on the right side being probably im-
mature, as they are smaller than their neighbours on either side.
The 1st molar is to all intents and purposes a simple pointed tooth
with a convex anterior edge and a concave posterior one. On
the posterior edge there is a slight " step "' but no true cusp.
The 2nd has a slight protuberance on the posterior edge at about
midway down, so that the crown might be said to be formed of a
long pointed anterior cusp and a short blunt posterior cusp. On
the - r )th tooth the posterior cusp is much more pronounced, while
the 6th and 7th seem to possess two posterior cusps. The last
molar is a short simple tooth. A figure showing the details of the
molar dentition is given.

The nostrils are somewhat higher than wide and are separated
l)\ a thin internasal septum formed by the premaxillae and the
nasals. The floor of the nostril is formed by a small septomaxilla
which appears only to a slight extent on the face. Between the
septomaxilla and the maxilla is an external foramen, seen in many
of the carnivorous Therapsida.

The top of the skull is very similar to that of Diademodon. The
nasals are narrow anteriorly but broaden at their junction with
the lachrymals, and narrow again slightly between the prefrontals.
The prefrontal forms the anterior half of the supraorbital margin.
Posteriorly it unites with the postorbital to shut out the frontal
completely from the orbit. The frontal is a narrow bone broadest
anteriorly, and sending forward a narrow anterior lateral process
which separates for some distance the nasal from the prefrontal.

The maxilla occupies most of the cheek andis pierced by a number
of small foramina. It passes back below the jugal to below the
middle of the orbit. ^

The lachrymal forms most of the anterior border of the orbit.
It is pierced just behind its suture with the maxilla by the lachrymal
foramen which communicates by a short canal through the bone
with a foramen within the orbit. Within the orbit there is also
another smaller foramen above the one referred to.

200 Annals of the South African Muxtuui

The jugal is incomplete, but does not seem to have an inferior
process.

The parietal region is narrow and crested. The pineal foramen
is small.

The squamosal is crushed, but appears to approximate fairly
closely to that of Diademodon. It has the groove on the posterior
surface connected with the external auditory meatus, which does
not, however, pass over on to the top of the skull. There is a process
directed inwards separating the end of the paroccipital process
from that of the pterygoid ; this plate bears one deep notch below,
instead of two as in Diademodon. The quadrate is not preserved.

The basicranial axis has been cleared on its ventral surface,
and the side-walls of the brain-case are visible ; and both bear a
striking resemblance to the corresponding parts in Diademodon.
Anterior to the pterygoid flanges the palate has not been wholly
cleared, but posterior to them nearly all the details of structure
can be seen.

The median ridge of the pterygoids is shorter than in Diademodon.
Posteriorly it splits into two which form the lateral borders of the
triangular area bounded behind by the occipital surface.

The occipital condyle is imperfect ; but there is no doubt that
it was double the left half is wholly preserved and formed by
the basioccipital. At the side of it, facing laterally, is a small
foramen for the exit of the Xllth nerve. The suture between
the basioccipital and the basisphenoid is seen to pass across the
basicranium between the two fenestrae ovales. Laterally the
basioccipital articulates with the massive paroccipital process
but the suture between the two is not visible. The front face of
this process is excavated at its inner end to form the posterior
border of the fenestra ovalis. Laterally to this the front face is
pierced by the pterygo-paroccipital foramen, whose anterior border
is formed by the posterior ramus of the " pterygoid.''

Behind and medial to the fenestra ovalis is the foramen jugulare
which is a deep pit looking entirely downwards. Piercing the
anterior wall of this pit is a small foiamen, corresponding to one
of those seen in Cynognathus.

The under surface of the basisphenoid is slightly concave with
a low median ridge and two lateral ridges which die out before
reaching the fenestrae ovales. There are no distinct basisphenoidal
tubera. On either side of the median ridge just behind the suture
with the basioccipital are two small pits in the latter bone which

South African Fossil Reptiles and Amphibia. 201

may lead into foramina. Anteriorly the basisphenoicl narrows
and sends a short process forward between the pterygoids. On
each side is a short basi-pterygoid process which does not seem
to have the long slender splint lying on the inner side of the posterior
ramus of the pterygoid described by Watson in Diademodon.
The upper surface of the basisphenoid has a median groove.

The side-wall of the brain-case does not differ in structure from
that of Diademodon. The suture between the prootic and the
epipterygoid (alisphenoid of Broom in Cynognathus) runs upwards
from the middle of the foramen which gives exit to the 2nd and
3rd branches of the Vth nerve to the sinus canal. The anterior
edge of the epipterygoid the suture between it and the pterygoid
is not easily seen is notched deeply by the opening transmitting
the II, III, IV, V and VI nerves. Superiorly it articulates with
the parietal.

Fractures show cross-sections of the skull above the incisura
prooticum and just behind the orbit. In the former section are
seen two shallow plates of bone lying vertically between the
parietals and close together. In the second these two plates have
become much deeper, thinner, and somewhat further separated.
These are presumably crushed sphenoids or orbito-sphenoids.

The front part of the palate has not been wholly cleaned ; but
there was certainly a nearly complete secondary palate, even if
the maxilla did not wholly meet its neighbour in the middle Hue.
The palatine is mainly a vertical bone. The ventral edge of the
bone medial to its suture with the maxilla is rugose. The presence
of a median vomer on the palate is not certain. The pterygoid
flange is large and descends considerably below the level of the
upper jaw.

As far as can be seen this form is extremely closely allied to
Diademodon. The structure of the brain-case, the foramina for
the exit of the nerves, the double condyle, and the general " look ''
of Gynosuchus all bring it close to Diademodon, from which it is
separated stratigraphically by the whole thickness of the Middle
Beaufort Beds. The most important differences are that in
Cynosuchits

1. the foramen jugulare is more widely separated from the

fenestra ovalis ;

2. the basipterygoid process of the basisphenoid does not

send out a long splint to lie along the inner side of the
posterior ramus of the pterygoid ;

202 Annals of the South African Museum.

3. the groove for the external auditory meatus does not

continue on to the top of the skull ;

4. the squamosal has apparently but one deep groove for the

reception of the quadrate ;

5. the molars are only cusped in a simple manner ; and

6. in the anterior portion of the basisphenoid just medial to

the end of the large groove between the bone and the
posterior ramus of the pterygoid there is on each side of
the bone a small foramen which may be a Vidian foramen.
This has been noted by Watson in a single Gorgonopsian
skull, but is not present in Diademodon.

In all these features Diademodon can be considered a direct
advance upon Cynosuchus. The brain-structure of the two closely
approximates ; the nature of the dentition is not so close. The
latter is probably a secondary advance, although there is not very
much difference between the molars in Cynosuchus and in
Nythosaurus.

Type. Somewhat crushed skull (S.A. Mus. Cat. No. 4333).

Locality. Weltevreden, GraafT-Keinet, C.P.

Horizon. Lower Beaufort Beds Cistecephalus zone (near top of).

NOTES UPON THE BRAIN-CASE IN CERTAIN THERAPSIDA.

DINOCEPHALIA. -Hitherto comparatively little has been known
about the detailed structure of the brain-case of the Dinocephalia.
Watson, in his paper on the Order, has given the fullest account
of the region and, inter alia, says : <: There is a relatively small
brain-cavity. This is throughout higher than wide and has not
-been cleaned in any specimens ; fractures, however, show that it
possessed the characteristic Therapsid character of having the
large opening to the vestibule placed very low down in the skull.
The floor of the brain-cavity rises considerably towards the front."
Fortunately it has been found possible to obtain a sagittal section
of the back part of a large Tapinocephaloid skull from the Gouph
in the collection of the South African Museum, and to trace
the position of the foramina opening into the brain-cavity. It
has not been possible to work out, however, the structure of the
inner ear on account of the intractability of the matrix. The follow-
ing account contains incorporated in it the account published by
Watson.

South African Fossil Reptiles and Amphibia. 203

The most striking feature is the smallness of the brain-cavity
compared with the immense thickness of the roofing-bones'of the
skull and of those of the occipital plate. The foramen magnum
is small, oval in shape, higher than wide. In front of it the brain-
cavity is not much enlarged, sloping gently upwards as an almost

VT.

fir Poss

FIG. 54. Median section through the back part of skull of a large
Tapinocephaloid. (S. A. M. Cat. No. 750.) x 1 nearly.

uniform tunnel to the exit for the Vth nerve. This cavity is
throughout higher than wide. The posterior portion of the con-
dyle is weathered away ; but just anterior to the posterior face as
preserved there is a small foramen in the side wall of the brain.
This probably transmitted a vein. Anterior to it the floor of the
brain-case is excavated on each side. The hollow thus formed is
divided into two parts by a bony plate protruding from the lateral

204 Annals of the, South African Museum.

*

wall and lying obliquely forwards in the hole. This divides the
posterior foramen for the IX-XI nerves from the anterior opening
into the vestibule. A short distance in front of this opening is a
small oval foramen for the passage of the facial nerve. Above
and slightly in advance of the vestibule there is a deep depression
in the side-wall of the brain-case which is probably a floccular
recess. Slightly behind that is a small foramen which may be
the opening of the canal for the ductus endolymphaticus situated
as figured by Watson in Diademodon.

The pro-otic forms the side wall of the case in advance of the
vestibule. It is pierced above its anterior inferior process by
two large foramina which are widely open laterally. The two
are only separated by a thin flange of bone. The lower is for
a branch of the Vth nerve, the upper possibly venous. In de-
scribing Lamiasaurus, AVatson said : ' The prootic, of course,
contributes to the fenestra ovalis, above which it is perforated
by the aquaeductus fallopii for the Vllth nerve."

The hypophysis is very deep and short, descending far below
the level of the occipital condyle. Watson has figured the pituitary
notch in external view in Lamiasaurus showing the ind-like
epipterygoid partially covering it. The posterior wall of the
pituitary fossa has a slight median ridge with a shallow pit on
either side in the upper half. Laterally it curves round to form
the posterior edge of the lateral opening, and has an anterior
process above. The fossa, however, is not separated from the
posterior part of the brain-case by any transverse wall. Above
it the roof of the brain-case is pierced by a large hole which com-
municates with the pineal foramen by a very long, circular, slightly
curved tube.

The bones of the brain-case have been described by Watson.
In this form there is no deep vertical area below the basioccipital
condyle the structure approximates more to that of Lamiasaurus
than to that of Mormosaurus. The fenestra ovalis does not lie
very much below the level of the bottom of the condyle and sections
show the stapes to be firmly fixed in it.

The paroccipital process is seen to be composed of a fairly thin
plate of bone forming the posterior occipital face and a massive
anterior portion which is probably the fused opisthotic and pro-otic.
The suture between the basisphenoid and prootic is not visible.

The parasphenoid is a median plate of bone forming the front
wall of the pituitary fossa. From the upper part of its posterior

South African Fossil Reptiles and Amphibia. 205

border there is a process which projects into the fossa on a level
with the floor of the posterior part of the brain-case. At the
level of this process the side-face of the bone is provided with
a broad shallow horizontal channel

The epipterygoid is not visible, but Watson has described it
as a slender rod rising very high in the skull with its posterior
edge nearly in contact with the front of the prootic, arising from
a small process of the pterygoid which curves round so as to shield
the front of the fenestra ovalis.

The occipital plate is weathered but the limits of some of the
bones can be made out. There is a fairly small supra-occipital
forming the upper border of the foramen magnum. The inter-
parietal is a larger bone lying above the supra-occipital. The
upper portion of its lateral edge articulates with part of the parietal
which overlaps on to the side of the occipital plate ; the lower
portion with the tabulare. The post-temporal fossa is very small.

GORGONOPSIA.- In view of the generally accepted belief
that the Gorgonopsia are nearly related to the Cynognathidae,
and consequently close to the line of mammalian descent, it be-
comes important to elucidate as much of their detailed structure
as possible. Watson, in a study of the external aspects of the
brain-case, has traced a gradual change from Dimetrodon through
An-tops and Scymnognatkus to Diademodon, without, of course,
suggesting that these genera lie actually on the one line of descent.
It has recently been possible to study the interior of the brain-
cases in one or two Gorgonopsia, and the results obtained are
given here.

The first skull examined was a large skull the largest Gorgon-
opsian skull known to me allied to Scymnognathus tigriceps. It
was collected by the Rev. J. H. Whaits at a height of 3,500 feet
on Zuurplaats in the Graaff-Reinet district, and is possibly thus
from the bottom of the Cistecephalus zone. The interior of the
brain-case has been completely cleared.

Watson in 1914 gave a short description of the brain-cavity
of Scymnognathxs whaitsi ? accompanied by a figure of a sagittal
section. The specimen showed '' a distinct resemblance to
Diademodon owing to the comparatively thin basioccipital and
the large opening to the vestibule through the posterior end of
which the tenth nerve has its exit. The general features of the
vestibule recall Diademodon, but there is no visible trace of a
cochlea. The chief differences from the more recent animal are

206

Annah of the South African Museum.

that the whole cavity for the cerebellum is much smaller, and
that its base rises very rapidly in front, very much as it does in
Dimetrodon. The pro-otic, although it extends further forward
than in Anomodonts and Dimetrodon, has more of the greater
anterior projection which, occurring in Diademodon and also in
OrnithorhyncJtus, has a long suture with the parietal."

SCYMNOGNATHUS TIGRICEPS ?

FIG. 55a. Diagrammatised sagittal section through back part of
skull No. 4334.

FIG. 55b. Section across vestibule of same. Both figures x ^.

In the skull of Scymnognathus mentioned above (S.A. Mus. Cat.
No. 4334) the occipital condyle is crushed, so that seen from
behind it forms but a half-ring below the small foramen magnum.
It is actually a large condyle with a central notochordal pit. The
basioccipital is fairly thin. It seems to play little or no part in
the border of the foramen magnum.

The paroccipital process has a concave lower border. It is
higher in front than behind. Its upper surface forms the lower
border of the small post-temporal fossa which lies on the level
of the middle of the foramen magnum. The foramen jugulare
is bounded above by the paroccipital and below by the exoccipital
and looks almost entirely downwards. The front face of the par-
occipital process is fairly flat and forms the posterior border of
the fenestra ovalis, which lies at about the level of the notochordal

Soutli African Fossil Reptiles and Amphibia. 207

pit in the occipital condyle. The front face is lightly channelled
horizontally by two small grooves, each ending in a foramen for
a vein.

The inside of the brain-case has been wholly freed from matrix,
and shows the positions of the foramina for the exit of the cranial
nerves. In front of the foramen magnum the opening is narrow
for a short distance, being bounded laterally by the exoccipitals.
In front of these it broadens and its plane is horizontal for about
20 mm., when its base slopes abruptly upwards. The exit for
the Xllth nerve is seen to pass through the exoccipital almost
at the top of the condyle. Anterior to the exoccipitals the floor
of the brain-case falls away laterally to the foramen jugulare,
nerves IX-XI passing out through an ill-defined broad groove
in the floor, which expands laterally. Anterior to this, the hori-
zontal portion of the floor has a strongly marked rounded median
ridge which separates the two inner ear-openings from one another.
These lie fairly close together in pits excavated wholly in the
floor of the brain-case. There is no abrupt line of demarcation
between this opening and the proximal end of the course of the
IX-XI nerves ; and one may consider that there is one large
excavation shallow behind and deeper in front the former
passing to the foramen for the exit of nerves IX-XI and the anterior
portion being the opening into the vestibule.

Anterior to the vestibule the, floor of the case slopes upward
forming a plate about 40 mm. long in the middle, longer at the
sides, and thinner above than below. The front of this plate
slopes backwards and downwards until it forms the base of the
pituitary fossa, in front of which it rises again almost vertically.
In a specimen which may be an immature skull of Gorgonognathus
this plate is high and vertical behind the pituitary fossa and is
divided by an open median vertical suture.

A fracture shows the inner ear to be connected with the fenestra
ovalis by an hour-glass shaped passage about 25 mm. long. This
passes mainly outwards and slightly downwards. The constriction
in the middle is very pronounced and nearer the inner than the
outer end. The fenestra ovalis is very wide, wider than the upper
end of the vestibule. It is separated from the foramen jugulare
by the process of the basioccipital which forms its anterior border.
The fenestra ovalis is entirely surrounded by bone, thus differing
from that of Diademodon. At the upper end of the ear the course
of the semicircular canals cannot be distinguished. The foramen
for the Vllth nerve has not been seen.

17

208 Annals of the South African Museum.

The foramen for the exit of the Vlth nerve is seen considerably
in advance of the vestibule, more than half-way up the sloping
portion of the floor. It is a small foramen, but the course of the
passage of the nerve has not been determined.

The notch at the top of the sloping portion of the floor is saddle-
shaped. On either side of it the bone is prolonged forwards by
two short processes the processi anteriores inferiores of the
prootics.

The side-wall of the posterior part of the brain-case is nearly
vertical seen from within. Just above the vestibule it is somewhat
excavate possibly the depression is the fossa sub-arcuata. On
the wall are two large oval foramina separated by a thin splint
of bone. The anterior of these lies just behind the anterior wing
of the sloping floor, and with its longer axis inclined. The side-
wall of the case is here very thin, and the canal from the foramen
is short, passing outwards and downwards. This foramen is
probably the incisura prooticum for the exit of branches 2 and 3
of nerve V. Slightly posterior to and above the foramen is a
more irregularly-shaped opening, which is probably the equivalent
of the venous foramen figured by Watson in Diademodon.

The basioccipital and basisphenoid are separated by an ill-
defined suture occupying the position seen in a well-preserved
skull of Scylacops capensis. The suture runs from the middle
of the lower border of the fenestra ovalis slightly forwards to the
median line of the deeply-vaulted region between the basisphenoid
keel and the basioccipital condyle, and then back to the fenestru
ovalis of the other side

Posteriorly the under surface of the basisphenoid is deeply
vaulted ; the under surface suddenly bends vertically downwards
and becomes a median keel passing forward to meet the pterygoids,
between which the anterior portion seems to be clasped. The
lateral ventral ridges are thick, but are not swollen into tuberosities.
They form the anterior borders to the fenestrae ovales. Above
the fenestra ovalis the suture between the pro-otic and the basi-
sphenoid runs upwards and forwards to the lower posterior border
of the pituitary fossa. The lower and anterior borders of this
high, short opening are formed by the basisphenoid which continues
forward as a vertical median sphenoid clasped by the vertical
median plates of the pterygoids. Below the pituitary fossa and
posterior to the suture between the pterygoid and the basisphenoid

South African Fossil Reptiles and Amphibia.

209

is a small foramen leading into a canal which passes up through
the body of the bone into the hypophysial fossa. This is possibly
a carotid foramen.

The pro-otic forms the posterior and part of the ventral borders
of the pituitary fossa. Superiorly it is pierced by the incisura
pro-oticum and posteriorly it forms with the inner end of the
opisthotic the border of the. oval foramen lying between the post-
temporal fossa and the incisura pro-oticam.

The epipterygoid is preserved on the left side of the specimen.
Its ventral end is a broad thin plate and is supported by the posterior
horizontal ramus of the pterygoid, from which it seems to be dis-
tinctly separated. Superiorly the bone soon becomes a small rod,
oval in cross-section, and covering the incisura pro-oticum.

ft Or

GORGONOGNATHUS LONGIFRONS Htn.

FIG. 56. Back half of type skull sectioned vertically, seen from
right and showing some of the foramina, x ^ nearly.

FIG. 57. Back half of type skull sectioned vertically near median
line, seen from left, x ^ nearly.

Gorgonognathus longifrons. The type-skull of this species shows
one or two further details. The back half of the skull has been
split by a slightly oblique longitudinal vertical section through
the middle of the occipital condyle, but it has been impossible
owing to the extreme hardness of the matrix to clear the
cavities.

The posterior part of the brain-case is of the same type as in the
previous skull, but the process of the pro-otic forming its anterior
boundary is rather more vertical and somewhat longer. Posterior
to its upper end on the side-wall is seen the incisura pro-oticum
and, separated from that by a thin splint of bone, is the large
irregular foramen which has been considered venous. Superiorly
the prootic meets the descending plate of the parietal ; but just

210 Annals of the South African Museum.

,

in front of the meeting place of the two bones the parietal lies
above a plate of bone which forms the upper border of the hypo-
physial fossa and passes down some distance on its front border
to meet the basisphenoid. Anteriorly this bone which occupies
the position of the parasphenoid of the Dinocephalia, at least in
its lower half meets the pterygoids.

The vertical ascending plates of the pterygoids are separated
from one another in two places. The first is seen in cross-section
just above the palatal surface at the level of the lateral flanges.
The other separation is the passage for the olfactory portion of
the brain which passes upwards and forwards through the opening,
doubtless swelling out anteriorly to form the olfactory lobes.

Scylacops capensis. A skull of this species (Cat. No. 3444) from
Well wood, Graaff-Reinet, C.P., which was fractured longitudinally,
has the bone so brecciated that the structure of the brain-case is
indeterminable. It can be seen, however, that the posterior portion
is somewhat longer and not so high as in the larger forms and the
transverse pro-otic plate is not so vertical. The stapes is seen
lying in position between the fenestra ovalis and the quadrate.
The bone at its outer end, is not rod-like, as it frequently seems
to be when the under surface only is cleaned, but plate-like with a
slightly convex anterior face and a parallel concave posterior face.
The upper surface is expanded with a flat horizontal posterior
projecting plate.

THEROCEPHALIA. Alopecognathus minor. Although the matrix
of the type of this species which is from the upper part of the
Tapinocepkalus zone is too resistant to allow of much develop-
ment, the region of the brain-case has been broken through in
several directions so that a certain amount of its structure has
been made out.

In front of the foramen magnum the brain-case swells out laterally
and vertically. The basioccipital forms most of the condyle. On
the ventral surface of the skull it forms the anterior border of
the fenestra ovalis and articulates with the basisphenoid in a
jagged suture lying between the fenestrae ovales. In sagittal
section, however, the basisphenoid is seen as a thin plate underlying
the basioccipital which extends forwards above it to well in front
of the level of the internal auditory meatus.

South African Fossil Reptiles and Amphibia.

211

The posterior part of the busisphenoid is thin, forming a plate
lying below the tubera basisphenoidales. Anteriorly it thickens
and narrows until, at the pituitary fossa, it is a deep narrow plate
which passes forward to be clasped ventrally between the pterygoids.

At its posterior end the basisphenoid forms, with the basioccipital,
paroccipital, and prootic, a large lounded opening which is appar-
ently not wholly enclosed by bone, being open on its outer side.
The inner portion of this pit is the fenestra ovalis, which is partially
walled in on the outer side by an anterior process from the par-
occipital (opisthotic) which stretches forwards towards the suture
between the basisphenoid and the prootic.

PoOT

H-XL

ALOPECOGNATHUS MINOR Htn.
FIG. 58a. Median longitudinal section showing wall of brain-case

from within.
F IG . 58b. Horizontal section across right side of brain-case just

above the condyle.

The foramen jugulare lies on the back of the skull at the side
of the occipital condyle. Its upper border is formed by the ex-
occipital, its lower by the basioccipital and the paroccipital. The
inner opening for the IXth-XIth nerves lies low down on the
side-wall of the brain-case, and the passage between it and the
foramen jugulare passes backwards and slightly outwards and
downwards. This canal is separated from the posterior canal
of the vestibule by a forward and inwardly-directed thin wall
formed from the inner end of the paroccipital process. Behind
the inner foramen for the IX-XI nerves is a small foramen opening
into a narrow canal which pierces the basioccipital and opens
into the foramen jugulare.

212 Annals of the South African Museum.

It has not been possible to clean the inner ear, but the region
is somewhat fractured and some of the details can be seen. The
lower portion of the side-wall of the brain-case is perforated by
a large hole, partly divided by the process of the paroccipital
above-mentioned. Behind this wall of bone is the opening for
the IX-XI nerves. Anterior to it the opening leads laterally
into two elongate cavities separated from one another, as seen
in cross-section, by a swelling of bone partly composed of prootic
and partly of what seems to be an opisthotic part of the par-
occipital. Ventrally the opening passes down, doubtless, to the
fenestra ovalis.

Superior to the internal auditory meatus the wall of the brain-
case is provided with a broad rounded ridge, above which is a
fairly deep fossa subarcuata, bounded anteriorly and superiorly
by the prootic. Just anterior to the lower border of the vestibule
opening is a small canal passing downwards and slightly outwards
the aquaeductus fallopii for the Vllth nerve.

The upper part of the prootic is pierced laterally by one large
foramen and possibly by another. The larger anterior one is for
part of the Vth nerve ; the possible posterior one venous. In
front of the larger foramen the pro-otic is carried forward and
upward as a processus anterior prootici. This, on one side of
the skull, is pierced by the foramen for the Vlth nerve. Superiorly
the prootic articulates with the parietal.

The prootic forms the posterior border of the pituitary fossa ;
but it shows a striking difference from the bone in the Gorgonopsia
in that it does not meet its neighbour in the middle line to form
a high ascending transverse plate. The portion of the bone pierced
by the Vlth nerve is merely a lateral anterior process ; between
it and its neighbour the bone is deeply notched as in Diademodon.

In front of the pituitary fossa there is no vertical plate of the
basisphenoid such as is seen in the Gorgonopsia I have examined,
but the basisphenoid is continued forward as a shallow, thin,
slightly upwardly inclined splint of bone whose outer side is
broadly grooved. This median splint does not articulate with
the vertical plates of the pterygoids, which are separated from
each other to accommodate a median canal,

A medium-sized Therocephalian skull from Wilgebosch (Lammer
Kraal), Prince Albert District, C.P., shows some additional features.

South African Fossil Reptiles and Amphibia.

213

The specific identity is doubtful, as the dentition is not well dis-
played, but the form is from the middle or top of the lower half
of the Tapinocephalus zone. The pro-otic has similar features
to that of Alopecognathus minor except that the anterior processes
are more vertical and longer.

Pr

FIG. 59. Irregular longitudinal section of Therocephalian No.
4332.

A fracture has shown the posterior ramus of the pterygoid, a
thin arched plate of bone which posteriorly forms the outer border
of the fossa which opens into the brain-cavity laterally through
the pituitary fossa.

Articulating with the outer face of this plate and rising directly
upwards from it is a fairly broad thin slightly curved lamina of
bone which articulates above with the parietal. The posterior
limit of this bone, which must be the epipterygoid, is not seen ;
but the plate appears to have extended sufficiently far back to
have covered the pituitary completely from side view.

There is a fairly large elongate interpterygoid vacuity. A
median section shows a thin median vertical plate of bone lying
some distance above this which plate may be an anterior pro-
longation of the basisphenoid or a parasphenoid. Above this is
a median bone whose cross-section is an inverted V ; and in contact
with the dorsal ridged surface of this is a hollow cylinder of bone
which may be open at the top below the roofing bones of the skull

214 Annals of the /South African Museum.

This is reminiscent of the sphenethmoid surrounding the anterior
part of the brain in Pareiasaurus described by Watson. Un-
fortunately, the complete course of the bone is not seen ; the
bony walls of the cylinder are very thin. The anterior prolongation
of the basisphenoid is sheathed by the inverted V-shaped bone,
which may be formed of the ascending plates of the pterygoid.

General Considerations. Taken in conjunction with the accounts
already published of the brain-case in Dimetrodon and Diademodon
and its allies, this study shows that in the essentials of structure
there is a striking similarity throughout the whole of the carnivorous
Therapsida. All have the inner ear lying well down in the brain
in many cases the vestibule pierces the floor and not up in the
side-wall, thus showing a mammalian affinity rather than one
with modern reptiles. In this respec't, the Dinocephalia seem to
be quite as far advanced as forms like Diademodon from much
higher zones.

It can be seen that the Gorgonopsia at least as far as the
larger types are concerned do not approximate so closely to the
Cynognathidae as a study of their external characters would
indicate. In the first place the sloping, sometimes almost vertical,
wall separating the posterior part of the brain from the hypophysis
and pierced by the foramen for the Vlth nerve finds no counter-
part in Diademodon nor in the Therocephalia from the lower zones,
where one simply finds the lateral portions remaining as processi
inferiores anteriores prooticorum, separated by a deep notch
whose base does not lie above the level of the bottom of the foramen
magnum. In Dimetrodon, however, a similar transverse wall is
seen, and in the Dinocephalia it may be present in some forms.
Further, the epipterygoid in the Gorgonopsia as in the Dinocephalia
is a rod-like bone with an expanded base ; whereas in Diademodon
and in at least some of the Therocephalia it is a plate.

Thus the Gorgonopsia seem to be more closely allied to the
Pelycosauria and Dinocephalia than to any other sub-orders.
This relationship has previously been noted by Broom and others
from a study of the external characters ; and it is important to
find that evidence from the brain-case affirms their conclusions.
It is unfortunate that no investigations have yet been made upon
the Gorgonopsia such as Galesuchus gracilis from the lowest portion
of the Beaufort Beds ; but the palatal portion of a medium-sized
Scymnof/nathus ? from the Endothiodon zone of Beaufort West
accords well with the larger forms from higher up.

South African Fossil Reptiles and Amphibia. 215

The discovery of the close relationship between Diademodon
and Cynosuchus whaitsi from the Cistecephalus zone renders it
probable that the line of descent of the Cynognathidae, if it passed
through any of the Gorgonopsia at all, must have passed through
some early form ; the larger forms from the Cistecephalus and
Endothiodon zones cannot be as close to that line as has hitherto
been supposed.

The position of the Therocephalia is uncertain. The brain-
case in the Bauridae and the Scaloposauridae is not known in
detail ; but it seems probable that some Therocephalian of the
lower zones gave rise to the Scaloposauridae of the Cistecephalus
zone and these in turn to the Bauridae of the Upper Beaufort
Beds. Further, the Therocephalia studied seem to approximate
to the Cynognathidae in the absence of the transverse wall behind
the pituitary fossa and in the presence of a plate-like epipterygoid.

216

Annals of the South African Museum.

INDEX.

PAGE

AKIDNOGNATHUS . . .192
ALOPECOGNATHUS . 180,210

BAUR1D.E 196

capensis (Scylac-ops) . . 208,210
CYNOSUCHUS . . . 197,215

DICYNODON . . . .190
DIMETRODON . . . .214
DINOCEPHALIA . . .202

GALECHIRUS . . . .179

GALEPUS 179

GOMPHOGNATHUS . .190
GORGONOGNATHUS . .207
GORGONOPSIA . . .205

ICTTDOGNATHUS . . 192,195
ICTIDOSUCHUS . .179

janseni (Macroscelesaurus) . 175

PAGE

LAMTASAURUS . . 204

longifrons (Gorgonognathus) . 207

.MACROSCELESAURUS . . 175
minor (Alopecognathus) . 180,210

parvus (Akidnognathus) . .192
platyceps (Whaitsia) . . .184

SCALOPOSAURID.*] . . HMi

SCALOPOSAURUS . . 192,194

sclateri (Scylacosaurus) . .190

SCYLACOPS . 208.210

SCYLACOSAURUS . . 190,194

SCYMNOGNATHUS . . . 205

suppostus (Cynosuchus) . .197

THERAPSIDA .... 175

tigriceps (Scymnognathus) . 205

whaitsi (Cynosuchus). . 197,215
WHAITSIA . 184

(217)

16. On Cretaceous Cephalopoda from Zululand. By L. F. SPATH,

M.Se., F.G.S.

(With a Sketch Map, 4 Text-figures and Plates XIX to XXVI.)

CONTENTS. PAGE.

I. INTRODUCTION . 219

II. THE UMKWELANE HILL FAUNA . 224

DESCRIPTION OF SPECIES :
A. AMMONOIDEA :

Family DESMOCERATIDJS :
Gen. Parapuzosia, Nowak.

1. P. sp. nov. ? hid. . . 224
Gen. Parapachydiscus, Hyatt.

2. P. sp. nov. aff. colligatus, Binkhorst sp. . . 226

3. P. cf. wittekindi, Schliiter sp. . . . 229

4. P. sp. hid. . 231
Family PRIONOTROPID.E :

Gen. Mortoniceras, Meek.

5. M. ivoodsi, sp. nov. . . . 232

6. M. aff. umkivelanense, Crick ..... 234

7. M. sp. aff. soutoni, Baily sp. . . 235
Gen. Pseudoschloenbachia, nov. ... . 236

8. P. umbulazi, Baily sp. ...... 240

9. P. umbulazi (Baily) var. acuta, nov. . . 241
Gen. Diaziceras, nov. ...... . 242

10. D. tissotiaeforme, nov. ...... 245

Family PLACENTICERATIDJE .-

Gen. Placenticeras, Meek.

11. P. sublMffrarium, sp. nov. . . . 247
Family NOSTOCERATID^ :

Gen. Nostoceras, Hyatt.

12. N. ? natalense, sp. nov. .... . 248

13. N. ? subangulatum, sp. nov. . . 250
Gen. Bostnjchoceras, Hyatt.

14. B. } sp. nov. . . 252

15. B. ? sp. ind. . . . .255
Gen. Diplomoceras, Hyatt.

16. D. ? indicum, Forbes sp. . . 256

19

218 Annals of the South African Museum.

Family BACULITIDJE :

Gen. Baculites, Lamarck ....... 257

17. ft. ccipensis, Woods . ..... 257

18. B. sp. aff. capensis, Woods . . . 258

19. B. cf. aspero-anceps, Lasswitz ..... 259

20. B. cf. brevicosta, Schluter . 260
2t. B. sp. of. sulcatiis, Baily . . 260

22. B. ballyl, Woods . . 261
B. NAUT*[LOIDEA :

Gen. Eutrephoceras, Hyatt.

23. E. aff. delcayi, Morton sp. . 262

24. E. cf. sublaevigatiim (d'Orbigny) var. indica

(Spengler) . . 263
Gen. Cymatocera-s, Hyatt.

25. C. ? sp. juv. aff. ml inln ij a /((isc. Blanford sp. . . 264
OBSERVATIONS ON THE UMKWELANE HILL FAUNA . . 264

III. THE MANUAN CREEK FAUNA . . 273

DESCRIPTION OF SPECIES :
A. ALBIAN :

1. AMMONOIDEA . . 273

Family PHYLLOCERATID.S; :
Gen. Phylloceras, Sness.

1. P. velledae, Michelin sp. . . . 273
Family DESMOCERATID^; :

Gen. Puzosia, Bayle.

2. P. cf. Ihima, Stoliczka sp. . . 274
Gen. Uhligella, Jacob.

3. U. ? sp. nov. aff. stolicskai, Kossmat sp. . . 275
Family DIPOLOCERATIIXE :

Gen. Dipoloceras, Hyatt.

4. D. cristatum, Deluc sp. . . . . '211

5. D. quadrat um, sp. nov. . . . . 278

6. D. sp. nov. . . . . 280
Gen. Pseudopliacoceras, nov.

7. P. manuanense, nov. ..... 281

Gen. Sulischloenlachia, nov.

8. S. prerostrata, nov. . . 284

9. S. cf. trinodosa, Bose sp. ..... 285

10. 8. ligpinosa, nov. . . 285
Family LYELLICERATID.E :

Gen. Stoliczkaia, Neiiniayr.

11. S. sp. ind. . . . 286
Family HAMITID.S: :

Gen. Torneutoceras, Hyatt.

12. T. sp. ind. ... . 288
Family ANISOCERATIDJE :

Gen. Anisoceras, Pictet.

13. A. sp. ind. . ... 288

On Cretaceous Cephalopoda, from Zululand. 219

Family TURRILITID^E :
Gen. Turrilites, Lamarck.

14. T. cf. gresslyi, Pictet & Campiche . . .289
2. NAUTILOIDEA :

Gen. Cymatoceras, Hyatt.

15. C. manuanense, G. C. Crick sp. . . . 290
36. C. cf. manuanense, G. C. Crick sp. . . . 291

17. C. cf. albense, d'Orbigny sp 292

18. C. sp. ind. 293

19. C. sp. nov. ? ind 293

20. C. cf. kossmati, Spengler sp 294

21. C. ? cf. clementinum, d'Orbigny sp. . . . 295
B. SENONIAN:

22. Peroniceras cf. dravidicum, Kossmat sp. . . 295

23. Kossmaticeras (Madrasites) lhavani, Stoliczka

sp. . . ... 299

24. Placenticeras cf. siibkaffrarium, sp. nov. . . 300

25. Cymatoceras ? sp. cf. justum, Blanford sp. . 301
OBSERVATIONS ON THE MANUAN CREEK FAUNA .... 302

IV. AMMONOIDEA FEOM ISOLATED LOCALITIES . 307

A. NORTH-WEST SHORE OF FALSE BAY (SENONIAN).

1. Mortoniceras vanuxemi, Morton sp. ..... 308

2. Bostrychoceras ? sp. . . ..... 308

B. POWELL'S CAMP, UPPER CATEMBE (APTIAN) . . . 309

1. Aconeceras nisoides, Sarasin sp. ... . 311

2. Cheloniceras gottschei, Kilian sp. ..... 312

3. Cheloniceras (Acantliopliies ?) delagoense, Krenkel sp. . 316

I. INTRODUCTION.

THE Cephalopoda that form the subject of the present paper were
included in a collection of Cretaceous fossils, sent by the Director of
the South African Museum, Cape Town, and the writer is greatly
indebted to Mr. Henry Woods, F.R.S., for entrusting him with the
study and description of these Cephalopoda. They comprise some
eighty specimens, and their study has proved of the greatest interest.
Aptian fossils are described from a new locality in South Africa,
and, of the later forms, some are quite new to science, whereas
others have not before been recorded from this continent. It may
suffice to point out here that, e. g., a near ally of " Schloenbachia acuto-
carinata (Shumard) var. -nmltifida" Steinmauu, i. e. of a group hitherto
recorded chiefly from localities in North and South America, is re-
presented in this collection. But the main interest of the collection
lies in the demonstration of the occurrence, together, at some of the
previously known fossil localities of Zululand, of deposits of varying
dates, hitherto confused on account of similarity of matrix.

220 Annals of the South African Museum.

Some time ago, when re-examining Mr. Crick's types of Zululand
Cephalopoda, in connection with the description of an Albian fauna
from Angola,* it became clear to the writer that the collection from
the South Branch of the Manuan Creek included Albian Ammonites.
Mr. Crickf had considered that they " most probably represented a
somewhat higher horizon (possibly Seuouiau) than that indicated by
the False Bay fossils," which latter were regarded to be Cenomanian.
Since EtheridgeJ already had described a Douvilleiceras and a Lyelli-
ceras ? from the neighbouring Umsiuene Kiver, and since Mr. Crick ||
himself recorded, from the Middle Branch of the Manuan Creek, two
undoubted Gault Ammonites, namely " Hysteroceras " [Brancoceras^
sp. and " Schloenbachia" [Dipoloceras^\ sp., the presence of the Albian,
at the South Branch also, was to be expected. In fact Mr. Crick
identified four examples from this South Branch as ? Beudanticeras
beudanti, Anisoceras sp., Douvilleiceras sp., and as " Schloenbachia"
aff. delarnei, d'Orbiguy sp. ; the latter, a typical Dlpoloceras of the
cristatum- group (s.7.) was worked out of the matrix of one of the large
Cymatoceras (referred to on p. 244) by Mr. Crick. But probably, on
the one hand, Mr. Crick was doubtful about his identifications, for
he did not mention these four important specimens in his paper ; on
the other hand, he may have been reluctant to assume different
horizons for what appeared to be the fauna of one single formation.
The Albian, Cenomauiau and Senouian Ammonites may be preserved
in a very similar brownish, friable matrix, and since there were as
many Senonian as Albian forms (in addition to one Cenomanian
Acanthoceras) present in the fauna from the South Branch, Mr. Crick
described the whole as " possibly Senonian."'

In a later paper, Mr. Crickf" stated that the occurrence (in the
Mauuau Creek district) of Cretaceous beds of an age younger than
Cenomaniau was somewhat doubtful. On the other hand, Mr. E. B.
Newton, in his paper on " The Cretaceous Gastropoda and Pelecypoda
from Zululand, 1 '** discusses the evidence in favour of a Senonian

* Bead before the Eoyal Society of Edinburgh, December 6th, 1920. (See
'Nature,' vol. cvi, No. 2669, December 23, 1920, pp. 554-5.)

I " Cretaceous Fossils of Natal," pt. iii, No. 2 .- " The Cephalopoda from the
Tributaries of the Manuan Creek, Zululand," ' Third Kep. Geol. Surv. Nat. and
Zulu!./ 1907, p. 249.

I Ibid., pt. ii, " The Umsinene River Deposit," p. 87.

Gen. nov. (type, A. lyelli, Desh. in Leym. ; d'Orbigny, 'Pal. Franc. Ter.
Cret.,' pi. Ixxiv, figs. 1 and 2) dealt with in the writer's Angola paper.

|| Loc. cit., pp. 247-8.

[ " Cretaceous Kocks of Natal and Zululaud," ' Geol. Mag.,' N.S., dec. v, vol. iv
(1907), p. 347.

** ' Trans. Eoy. Soc. S. A.,' vol. i, pt. i (1909), pp. 94-5.

On Cretaceous Cephalopoda from Zululand. 221

horizon for the Manuan Creek Fauna, and though he points out
that " several of the species bear a distinctly older appearance," he
goes on to say : " Taking into consideration . . . that there is
no great difference in the lithological character of the matrix accom-
panying the various specimens from the Manuan Creek, it would seem
that we are dealing with a fauna of one geological age which may be
regarded as Emscherian or Lower Senonian, since it includes Veniella
forbesiana, a characteristic pelecypod of the Upper Trichinopoly beds
of Southern India . . ."

The Ammonites of the present collection include a number of typical
forms that confirm the presence, at some of the localities, of deposits
of various geological ages. The collection includes :

(a) A typical Albian fauna from " Manuan Creek " and the
"Middle Branch, Manuau Creek" (19 specimens).

(6) Albian and Senonian Ammouitids from the " south side of
Manuan Creek Valley," from " high ground on north side of United
Manuan Creek and Umsiueue River, almost opposite junction " ; and
from " Low Ridge about 3 m. east of foot of Lebombo Mountains,
north of M'Kusi River, due east of Ubombo " (10 specimens).

(c) Senonian examples from the " North-West Shore of False Bay "
(2 specimens).

(d) Aptiau Ammonites from Powell's Camp, Upper Catembe
(3 specimens).

(e) A Senonian fauna from Umkwelane Hill. This includes over
forty specimens (half of which number ax&Bacvlites), mainly from Um-
kwelane Hill itself ( = d on the accompanying sketch-map), but
comprising a few examples from the three neighbouring localities :
Lake Itesa (Eteza) (= a on map), the railway cutting (== 6 on map),
and the Uinfolozi Valley, east of railway (= c on map). This fauna,
first made known by Etheridge,* probably corresponds in age with
the Pondoland fauna described by Woods. t

The following note by Dr. A. L. du Toit, dated March 20th, 1920,
describes the nature of the deposits :

" At Umkwelane Hill, the Cretaceous rests directly upon Stormberg
basalts (Liassic?), the strata being hard shelly limestones and softer
buff -coloured sandstones and other softer beds, rarely exposed ;
pebbles of older formations are found occasionally in them. The first
fossiliferous limestone comes in a few feet from the base. These bands

* " Cretaceous Fossils of Natal," pt. i : " The Umkwelane Hill Deposit, Zulu-
land," ' Second Eeport Geol. Surv. Nat. and Zulul.,' 1904, pp. 69-93, pis.

t " The Cretaceous Fauna of Pondoland," ' Ann. S. Afr. Mus ,' vol. iv, pt. vu,
No. 12 (1906).

222

An naif <>f tlif South African Museum.

Q)

On Cretaceous Cephalopoda from Zululand. 223

are intensely hard and are crowded with lamellibranchs, etc., much
broken shelly matter, but the fossils are difficult to extract.

" Along the railway cutting, the section shows dark-brown to khaki-
coloured sandy stuff, without good bedding, with thin sandstone-ribs
and concretionary limestone-nodules, in which Ammonites are found.
Apparently, the same horizon is represented N. of the Umfolozi River at
point X (see plan), where Ammonites are common. . . . The whole
thickness of strata involved is probably not more than 250 feet, the dip
being S.E. at 1 or H ; and since the strata must have accumulated in
shallow water, it is unlikely that several distinct zones are represented .
all the fossils may therefore be considered to come from one horizon'
and the beds and their fauna can therefore be regarded as a whole. 1 '

It will be advisable to describe the Umkwelaue Hill fauna separately
from that of the Manuan Creek and the other localities. There is a
large collection of Ammonites from the Umzamba (" TJmtamvuna '')
Beds of Pondoland in the British Museum, approximately corresponding
in age with the Umkwelaue Hill fauna, as stated by Woods * and by
Crick 1 in his very useful general account of the " Cretaceous Rocks
of Natal and Zululand." Crick was at work describing this fauna
already before 1906 + but his MS. is still unfinished, and the writer
hopes to complete and revise it as soon as facilities for publication are
offered. This collection in the British Museum includes a number of
species not known to Baily, Griesbach, Woods and Van Hoepeu, and
not represented at Umkwelane Hill ; and reference to some of these
will be made in the specific descriptions when necessary. A new
collection of Pondolaud Ammonites, kindly sent to the writer by Mr.
Henry Woods, includes a further series of undescribed forms. In his
account of this fauna, to be published shortly in the ' Annals of the
Durban Museum/ the writer is drawing attention to the improbability
of such faunas representing only "one horizon." The great majority, if
not all, of the Pondoland and Umkwelane Hill forms are of Campaniau
and Maestrichtian age, a possible range of at least five zones. The
Pondoland strata are only twenty feet thick and of a sandy facies,
suggesting rapid deposition : but the new collection contains doubtful
or long-lived species that might even be pre-Campanian in age. The
assertion, thus, is not justified that the corresponding beds at
Umkwelaue Hill, of a much greater thickness, can be regarded as
belonging to " one horizon."

The third part of this paper will deal with the Mauuan Creek

* Loc. cit., p. 347.
t Loc. cit. (' Geol. Mag-.'), p. 313.

J Woods, loc. cit. (1906), p. 337. Kossmat, "Die Bedetitung d. Siidind.-
Kreideform.," ' Jb.K.K.Geol.K.A.,' vol.xliv (1894), Heft 3 and 4 (1895), pp. 463-4

224 Annals of the South African Museum.

fauna ; the few specimens from isolated localities, namely (c) and (d)
above, are described separately under IV.

By the kindness of Dr. A. Smith-Woodward and Dr. F. A. Bather,
of the British Museum, the writer has been able to make the fullest
possible use, for comparison, of the rich collections under their charge.
Similar facilities were accorded him by Prof. W. J. Sollas, Mr. C. J.
Bayzaud, and Mr. J. A. Douglas, at the Oxford University Museum ;
by Prof. A. M. Davies at the Imperial College of Science ; by Dr.
F. L. Kitchiu at the Museum of Practical Geology. To all these
gentlemen, and especially to Mr. Henry Woods, of the Sedgwick
Museum, Cambridge, the writer expresses his cordial thanks.

II. THE UMKWELANE HILL FAUNA.

DESCRIPTION OF SPECIES.

A. AMMONOIDEA.

FAMILY: DESMOCERATID^ :

GEN. PAEAPUZOSIA, Nowak.

1. PAEAPUZOSIA sp. nov. ? hid.

(PI. XIX, fig 2; PI. XX, figs. 1, !,.; PL XXIV, fig. 3.)

? 1906. Desmoceras (Pnzosia) yaudama (Forbes) Boule, Lemoiue &
Thcveuin, "Pal. de Madagascar," III, "Ccphal. Cret. Diego-
Suarez." Ann. de Pal., vol. i, fasc. iv, p. 20, pi. iv, fig. 5 only.

The writer has not seen the specimen upon which this description is
based ; but in addition to the measurements given below and to the
two photographs here reproduced, a plaster cast of a portion of the
inner whorls (PI. XXIV, fig. 3) was forwarded, the specimen itself being
too bulky to be sent. The measurements, according to Mr. S. H.
Haughton's note, accompanying the photographs, are as follows :

Diameter . . . 670 mm.

Height of last whorl . 325 [=48'5 per cent, of the diameter].

Thickness . 225 [== 33 ].

Umbilicus . 120 [=18 ].

The specimen was also characterised as " looking very similar to
one or other of the species of Desmoceras (Puzosia} described from
Madagascar " ; and since the inner whorls, according to the plaster
cast of the dorsal impression, show the ribbing and compressed aspect
of P. gaudama, Boule, Lemoine and Thi'venin * non Forbes, the
* Loc. cit., fig. 4 a of pi. iv, which, however, is more compressed.

On Cretaceous Cephalopoda from ZulultnnL 225

specimen is compared with the gigantic Ammonite cited above, and by
these authors doubtfully classed with Forbes' species, and with the
smaller figured example. The suture-line seems to agree very well,
judging from the photographs only ; but the bulges on the inner
portions of the lateral area, reminiscent at once of Parapuzosia lepto-
pltylla (Sharpe) and of certain Pacliydiscus and Parapachydiscus, are
not apparent in the photographs of the form here described. P.
gaudama itself (B.M. G-eol. Soc. Coll. 10487) is different, and P. corbarica?
Grossouvre,* with a thickness of only 27 per cent, of the diameter,
is too compressed. The ornament of the inner whorls of the example
here described, however, is very similar to that of this species, as it
also is to that of the more coarsely ribbed P. daubreei, Grossouvre sp.,|
though, owing to the absence of the inner half of the lateral areas, the
primary costse are only just indicated, so that comparison with this
species, the presumed genotype, is difficult. The sectional outline of
Grossouvre's species given by Nowak + is more compressed than that
of the Zululaud form. On the other hand Nowak's figure of the
suture line apparently shows good agreement, as does that of P.
leptophyUa, Sharpe sp.|| The fine example of P. daubn'ei figured by
Miiller and Wollemann^" has a larger umbilicus and very strong
primary costation.

Nowak is inclined to unite these two Santouian species, and considers
P. tannenbergica, Fritsch and Schloeubach* to be closely related,
but he also quotes, as an example of Parapuzosia, Stoliczka's A.
denisonianiis, which is pre-Senouian, like P. <///*//'///, Sharpe sp.,tt a
form much nearer the ancestral Pitzosia-type. P. stobae Nilssou,++

* "Koch. s. 1. Craie Sup.," II., Pal., " Les Amm. d. 1. Craie Sup." 'Mem.
C.-irto Geol. France,' 1893 (1894), p. 174, pi. xxvii, figs. 1 a, b.

t Hid., p. 154, pi. xxviii. (" Sonneratia," in Grossouvre.)

j " Unters. ii. d. Ceph. d. Ob. Kreide Pol.," iii, ' Bull. Ac. Sci. Cracovie
ser. B (1913), pi. xliii, fig. 32, p. 363.

Hid., pi. xliv, fig. 40.

|| ' Moll. Chalk England,' III, " Cephal." (1856), pi. xxi, fig. 2.

H" " Moll. Fauna d. Unter-Senon v. Braxinschweig," II, " Ceph.," ' Abh. Preuss.
L.A.,' N.F., Heft 47 (1906), p. 8, pi. v.

** ' Cephal. d. Bohm. Kreideform.,' Prague, 1872, pi. ix.

ft Loc. cit., II, 1854, p. 28, pi. xii, figs. 1 a, l>.

It ' Petrif. Suec. form, cret.,' p. i, London, 1827, p. 5, pi. i. Moberg, ' Ceph.
i Sverig. Kritsyst.,' II, " Artbeskrifn.," Sver. Geol. Unders., ser. C, No. 73, 1885,
p. 18, pi. ii, figs. 1-5. Nilsson's figure is somewhat diagrammatic, but Moberg's
example (la) represents a form apparently similar to the specimen here
described, if more compressed. The suture-line, however, stamps P. stobae to
be a Parapachydiscus, connected with such forms as P. colligatus by P. exilis,
Binkhorst, which Schliiter (loc. cit., p. 56) thought perhaps belonged to P. stobae.

226 Annals of the South African Museum.

which is a pachydiscoid development (Parapachydlscus) later than
P. denisoniana, and which has Pwaosm-like inner whorls, is also grouped
here by JSTowak. But the forms that continue the Pusosia-type into
the Senonian (P. ijaitdama, Forbes, P. indopacifica, Kossmat, etc.), and
once more produce transitional forms to Kossmaticeras (P. danvini,
Philippi in Steiumann), cannot be classed in the same genus with
P. (It'iiixnuinna- which developed a " Pachydiscus-st&ge " already in the
Turonian.*

The inclusion of the Zululand example in this polyphyletic genus,
then, is only provisional, and based on the assumption that
P. da^(,breei is the genotype of Parapuzosia . The ornamentation of
the inner whorls shows that it is not a Parapachydiscns and the
writer believes that it has nothing to do with the gigantic forms of
the type of P. sepjoem*aden.sis, Landois,t which Nowak,^ in the writer's
opinion wrongly, also considers possibly to belong to Parapnzosla.

Locality. Railway cutting, Umfolozi. Coll. Dr. A. L. du Toit.
S.A.M. Cat, No. 5513.

GEN. PARAPACHYDISCUS, Hyatt,

2. PARAPACHYDISCUS sp. nov. aft'. COLLIGATUS, Binkhorst sp.
(PI. XXII, figs. 1 a, 6.)

1861. Ammonites colligatus, Biukhorst. Mou. d. Gast. & Ci'ph. de la
Craie Sup. d. Limbourg, ii, p. 25, pi. viii a only.

1894. Pachydiscus colligatus, de Grossouvre. Amm. Craie Super., p. 202.

1908. Pachydiscus colligates, de Grossouvre. Descr. d. Amm. d. Crc't.
Sup. du Limbourg, etc. Mem. Mus. Roy. d'Hist. Nat. Belg.,
vol. iv (1908), p. 28, pis. iv-viii.

' f Forbes ('Trans.,' 2nd ser., vol. vii, 184(5, p. 114) stated that there was a
" fragment of a very large Ammonite, but undeterminable, among the specimens
from Trichinopoly." This example, now in the British Museum (Geol. Soc.
Coll.), probably belonged to a form of this group of " Puzosia," but consists of
one camera only (H. = 170 mm. and Th. just a little less). The extremely
complex lateral lobe and short siphonal lobe are like those in P. denisoniana,
as figured by Kossmat (loc. cit., pi. xiv, fig. (3), but there is no ornamentation
remaining- at this pachydiscoid stage.

t " Die Eiesen-Amm. v. Seppenrade," 23. Jahresb., ' Zool. Sect., Westfal.
Prov.-Ver. f. Wiss. and K.' (1895), pp. 99-108, pis. i and ii.

t Loc. cit. (1913), p. 365.

Kossmat (loc. cit., p. 166 [101]) considered the two species, colligatus and
otacodensis, to be closely allied, but the latter is quite different, judging by
Kossmat's own identifications in the British Museum collections.

On Cretaceous Cephalopoda from Zulidand. 227

1913. Pachydiscus collujatns, Nowak. Unters. Cephal. ob. Kreide
Poleu, iii, Bull. Ac. Sci. Cracovie, ser. B, p. 361, pi. xliii, fig. 30,
pi. xliv, fig. 39.

A completely septate specimen (No. 5489) of 145 ram. diameter has
an umbilicus of about 20 per cent, of the diameter and a whorl thickness
of about 60 per cent. The ribbing is obscure, the example being only
an internal cast ; but there are about 25 costae on the outer whorl,
weakened at the grooved siphonalline and with a slight forward sinus.
Some of these costae do not reach to the rounded umbilical border ;
those that do are not distinctly tuberculate, and the inner whorls, as
shown in the deep umbilicus, are almost unornamented. There are
only nine septa on the last whorl, but these attain an extreme degree
of complication, equalled, perhaps, only by such forms of Parapachy-
(//./'* as P. quiriquincue (Philippi), Steinmann sp.

The specimen shows close resemblance with the examples figured by
Biukhorst on pi. viii a, and which were selected as typical by
Grossouvre, though Pervinquiere * was of opinion that the large
form figured by Binkhorst on pi. viii, and which was renamed
P. van den broecJci, should be taken as type of Binkhorst's species, f

The present example differs from the typical P. coHigatus (pi. iv,
fig. 3, and pi. v, fig. 1, in G-rossouvre) only in having, at a radius of
75 mm., a thickness of 75 mm., not 52 mm. as given by Grossouvre:
in other words it retains the globosity of the inner whorls of
P. collujatiix to a larger diameter. The costation, possibly, also is a
little too distant in the Zululand example.

One of Schliiter's j examples of Parapachydiscu'S witteJdndi has
this more distant costation and thick whorls, but judging by a
number of Westphalian examples of this species in the British
Museum, the wide and strongly costate umbilicus separates Schliiter's
species from the form here described.

Para i>a <-li inl^r a x '? portlocki, Sharpe sp., similarly differs from the
latter in having a wider umbilicus, surrounded by tubercles. In
whorl-shape, however (thickness -.- 60 per cent, of the diameter), the

* "Et, de Pal. Tunis," I, " CY-ph. d. Ter. Sec.," 'Carte Geol. d. 1. Tunisia'
(1907), p. 175 (footnote).

t This "species" was withdrawn in 190S by A. de Grossouvre, who then
figured Binkhorst's original (in the Berlin Museum) a poorly preserved exam pie.

t ' Ceph. d. Ob. Deutsch. Kreide./ I, Palaeontogr., vol. xxi (LS72), c. g. example
2 of p. 68 (A. robustus), with thickness = 57 per cent, of the diameter, pi. xxi,
figs. 5 and (5 ; pt. ii (1870), p. 160. Boule, Lemoine and Thevenin (loc. cit., II,
1907, p. 22) wrongly consider this species of Parapachydistus to be the type of
Pachydiscus, Zittel.

"The Fossil . . . Mollusca . . . Chalk of England," II, 'Ceph.,'
Pal. Soc, Mon., 1854., p. 30, pi. xiii, figs. 2, 3.

228 Annals of the South African Museum.

two forms are closely comparable, though the adult portloclci
develops ventral tuberculation.

P. epiplectns, Redteubacher,* included by Clrossouvre and Nowak
in the synonymy of P. colligatus, differs both from the latter and from
the present example in the pronounced peripheral sinus in the costation.

P. quiriqitinae (Philippi), Steiumann sp.f resembles the Zululaud
example in suture-line, and has a similar straight principal lobe, but
is too thin and too closely costate.

" A. newberryamis," Meek sp., as figured by G-abb has a
similar suture-line, but the costatiou is finer and closer, and the
whorl-section is more compressed than in the specimen under
description. On the other hand, Whiteaves figures as A. new-
berryanus, Meek, an Ammonite which, judging by comparable
examples from Vancouver Island in the British Museum, || belongs
to a group of forms quite different from the Parapachy discus here
discussed and related to the isculensis-gToup of Nowak and to Kossmati-
ceras. Of Japanese forms, P. teshionensis, Jimbo,^ is less globose and
less involute than the African example ; it also has umbilical tubercles
and a less complex suture-line. P. naumanni, Yokoyama,** resembles
the specimen here described in globosity, but has varices with close
costation, and a comparatively large umbilicus.

* "Ceph. Fauna d. Gosau-Sch.," ' Abh. K.K.E.A.,' vol. v (1873), p. 121,
pi. xxviii, fig. l.

t " Beitr. z. Geol. and Pal. S. Amer.," Ill, " D. Alt. und d. Fauna d. Quiriquina
Sch. in Chile," N. Jb. f. Min., etc., Beil. Bd. x (1895), p. 74, pi. vi, fig. 3, Text-
fig. 5 on p. 77.

t 'Pal. of Calif./ vol. i (1864), p. 61, pi. xxvii, fig. 199, and pi. xxviii,
fig. 199 . Whiteaves (' Mesoz. Foss.,' p. 107), included this form in A.
complexus var. suciensis, Meek ; later (ibid., p. 344) renamed Pachydiscus
suciensis, Meek sp. (The reference is [wrongly] to A. brewerianus, Gabb, in
both cases.) Three examples, doubtfully referred to Pachydiscus complexus,
var. suciensis, by Kossmat, from Vancouver Island in the British Museum
Collection, are quite different from the Californian Ammonite, which also is
quite different from Meek's original A. complexus ?, var. suciensis ('U.S. Geol.
and Geogr. Surv.,' Bull. No. II, 1870, p. 369, pi. v, fig. 2).

' Mesoz. Foss.,' I, pt. ii. " Foss. Cret. Eocks Vancouver, etc.," ' Geol. Surv.
Can.,' 1879, p. 109, pi. xiv, figs. 1 and 1 a.

|| Associated with P. otacodensis, Stoliczka (Whiteaves, loc. cit., I, V, 1903, p.
340, pi. xlvi, fig. 1, and 'Trans. Eoy. Soc. Can.,' 2nd ser., vol. i, sect, iv, 1895
[1896], p. 131), and other species of Parapachydiscus labelled by Kossmat (see
' Jb. K.K. Geol. E.A.,' vol. xliv [1894], p. 472).

*|[ " Beitr. z. Kenntn. d. Fauna d. Kreidef. v. Hokkaido," ' Palseontol. Abh.
Dames & Kayser,' vol. vi, pt. iii (1894), p. 30 (176), pi. iii, fig. 1.

** " Verstein. a. d. Japan Kreide," ' Palseontograph.,' vol. xxxvi (1890), p. 187,
pi. xxii, fig. 1.

On Cretaceous Cephalopoda from Zululainl . 229

P. arrialoorensis, Stoliczka,* is very close to the Zululand example
in shape, umbilicus and number of costae but differs in the character
of the periphery, unless this difference, well seen on comparing
fig. 1 c of Grossouvre's pi. v with fig. 3 a of Stoliczka's pi. Ixiii, is due to
the presence of the shell in the Indian form, which seems doubtful
in view of its sandy matrix. However, the larger and somewhat
different example figured by Yokoyamat shows distinct and sinuous
costatiou on the ventral area both of the cast and of the shell. The
suture-line of the Indian species also is simpler than that of the example
here described.

P. colligatus has been recorded from Tullear, on the West Coast of
Madagascar, and other species of " PacJiy discus," including Jacobites
from other parts of the island. Woods stated that Pacliy discus
was absent in Poudoland ; but there are four specimens from the
Umtamvuua River in the British Museum, || including two large
examples that may belong to forms of the colli gat us -group. The
presence of " Pacliy discus " both in Pondolaud and in Zululand thus
forms a further point of resemblance with the Indo-Malgascan fauna.
. Umkwelane Hill. Coll. Dr. A. L. du Toit.

3. PARAPACHYDISCUS cf. WITTEKINDI, Schliiter, sp.
(PL XXIV, fig. 1.)

1872. Am. robustus, Schliiter, "Ceph. d. Ob. Deutsch. Kreide,' 1 Palaeon-

togr., vol. xxi, p. 67, pi. xxi, figs. 5 and 6, pi. xxii, figs. 1-3.
1876. Am. ivittekindi, Schliiter (ibid.), vol. xxiv, p. 40 (160).

* In Blant'ord and Stoliczka, " Foss. Ceph., Cret. Rocks of S. India," ' Mem.
Greol. Surv., India, Pal. Indica ' (1865), p. 126, pi. Ixiii, fig. 3 only.

t Loc. cit. (1890), i, p. 186, pi. xxi.

J See Boiile and Thevenin, ' Bull. Soc. Geol. France,' ser. iv, vol. iii, 1903,
p. 436; Boiile, Lemoiiie and Thevenin, loc. cit. (1907), pp. 23-25; Kilian and
Eeboul : " Les Ceph. Neocret. d. lies Seymour et Snow-Hill," ' Wiss. Erg.
Schwed. S. Pol. Exp. 1901-03,' III, 6 (1909), p. 25.

Loc. cit. (1906), p. 346.

|| Two of these, named P. umtafunensis by Crick (MS.), Nos. C19434-5, and
compared with P. tweenianus, Stoliczka (loc. cit., 1865, p. 107, pi. Iv only), one
of the species found in Madagascar, are evolute, like P. conduciensis, Choffat,
and with comparatively simple suture-line, thus differing from the typical
Parapacliy discus. They are similar to certain South- American " Pachy discus "
(Paulcke "Die Ceph. d. Ob. Kreide Siidpatagoniens," 'Ber. Naturf. Ges. Freiburg
i. B.,' vol. xv (1907), e. y. pi. xix [x]). The other two gigantic specimens, not
described by Crick, may belong to Parapachy discus of the colligatus-supremus
type, but one has an umbilicus of 19 per cent, and a thickness of 45 per cent, of
the diameter, the other at a whorl-height of 260 mm. a thickness of 200 mm. ;
both are more compressed than the Zululand specimens here described, and
intermediate in sectional outline between figs. 30 (P. colligatus) and 31 (P.
oldhami) in Nowak (loc. cit., 1913, pi. xliii).

230 Ann al# <>/ the South African Museum.

A gigantic example (No. 3969) resembles the form last described,,
but has a wider umbilicus. Its dimensions are :
Diameter - . 390 mm.

Height of the last whorl . . 46 per cent, of the diameter.
Thickness ,, ,, . 49

Umbilicus .... 25

The costatioii disappears on the ventral area and remains distinct on
the inner half of the side, which is just the reverse of what takes
place in P. supremus, Petho sp.,* included in the synonymy of
P. enlligaius, Binkhorst sp., by G-rossouvre and Nowak.f P.
f'rt'xrHlrHsis, Seunes,J and P. epiplectus, Redtenbacher, show a similar
change, but one of the examples of the former species, figured by
Seunes (pi. xii (iii), fig. 1), agrees with the specimen here described in
having a somewhat reniform whorl-section, with the greatest thickness
near the umbilical border. The latter character distinguishes the
Zululand example from P. teshionensis, Jimbo,|| which apparently
weakens the ornament of the periphery and retains the principal
lateral costae after the manner of P. ecjertonianus (Forbes), Stoliczka
sp.,^~ which latter species, however, is far too compressed.

P. haradai, Jimbo,** and the larger example referred to the same
species by Whiteaves,ft have more compressed whorls at a stage when
the present form is still very depressed. It is also doubtful whether
large examples of this species would develop the secondary ornamenta-
tion of the typical forms of this group. This equally applies to the
evolute P. steinmanni, Paulcke,++ that may or may not develop these
obscure bulges at a large diameter, but resembles the form here
described in the roundness of the inner whorls.

Schliiter's A. wittekindi differs from the form here described in
whorl-section, though the inner whorls of the South African form
apparently are more depressed than is its outer whorl. The West-
phalian form also appears to lose its first costatioii at an earlier stage,

* " Kreidefauna d. Peterwardeiner Geb.," 'Palaeontogr.,' vol. Iii (1906), p. 88,
pi. v, fig. 1.

t Loc. cit. (1908), p. 29 and (1913) p. 361.

I "Contrib. a 1'Et. d. Cephal. Cret, Sup. France," ' Mem. Soc. Geol. Fr.,' Pal.,.
vol. ii, fasc. iii (1891), p. 3, pi. i, fig. 1.

Loc. cit., p. 121, pi. xxviii, fig. 1.
i Loc. cit. (1894), p. 30 (176), pi. iii, fig. 1.

^ Loc. cit. (1865), p. 104, pi. liii, fig. 1.

** Loc. cit. (1894), p. 29, pi. ii, fig. 2.

ft "Vancouver Cret, Foss.," 'Trans. Roy. Soc. Can.,' 2nd ser., vol. i (1895),,
sect, iv, 1896, p. 132, pi. iii, fig. 6.

H Loc. cit. (1907), p. 230 (64), pi. xviii (ix), fig. 1.

On Cretaceous Cephalopoda from Zululand. 231

and the costae of the secondary ornament (if the large and thin
examples belong to the same species) are too pronounced on the
venter. P. seppenradensis, Landois,* which was compared with P.
wittekindi (and Pachydiscus /r/r/x/V//s/x) by Zittel, retains its costation
to a very large diameter, but apparently belongs to the true Pachydiscus
of the peramplus group, with comparatively simple suture-line.

P. lei-yi, Grossouvre sp.,t probably is a close ally of the Zululand
form, but has pronounced umbilical tubercles instead of mere swellings
of the primary ribs, as indicated in Schliiter's fig. 5 of pi. xxi.

The large forms of Pardpachydiscus from Pondoland, referred to
under P. sp. nov. cf. colligatns, are more compressed than the present
example.

The suture-line of P. witteJcindi is less complex than that of the
Zululand form, which resembles in this respect the suture-line of
P. iiniriijultiae (Philippi), Steimnann sp., especially in the straight
line formed by the main stem of the principal lobe. The distance of
this line from the parallel siphonal line is 95 mm. There is a slight
groove on the periphery, marking the position of the siphuncle, as in
P. colliyatus.

The comparison of this species with P. wittekindi cannot be taken to
indicate affinity with the European fauna, for they certainly are not
specifically identical, and the present form is attached to Schliiter's
species only because of Indo-Pacific species, that may be more nearly
related, corresponding large stages are unknown, so that comparison
is difficult.

Locality. Umfolozi River, East of Railway. Coll. Mr. Illiugworth.

4. PARAPACHYDISCUS, sp. iud.

Portions of a gigantic specimen (4985), at least half as large again
as the example last described, and therefore probably of a diameter
of about 600 mm., differ slightly from this other large specimen
(No. 3969) in the suture-line. Only the peripheral portions of two
camerae, including the greater part of the external saddles, are preserved ;
the median saddle in the ventral lobe alone is 46 mm. wide. The
details of the ventral lobe differ somewhat from those shown in the
peripheral aspect of P. colligatus, Binkhorst in Grossouvre (with
which the small example (No. 5489) shows good agreement).

* " Die Eiesen-Amrn. v. Seppenrade," 23. Jahresb., ' Westfal. Prov.-Ver. f.
Wiss. and Kunst.,' 1895, p. 104, pis. i and ii.
t Loc. cit. (1894), p. 178, pi. xxi.
I Loc. cit. (1895), text-fig. 5, p. 77.
Loc. cit. (1908), pi. vi, fig. la.

232 Annals of the South African Museum.

The shell is partly preserved and 3 mm. thick in places, but
less than 2 mm. in others. It is in two thick layers, fibrous and of
a white porcellauous aspect, like certain Inoceramus shells in the
chalk, with a thin inner and outer coating. The specimen may belong
to a fat form of ParapacJiydiscus, like the two examples previously
described, but is too incomplete for specific determination. The large
ParapacJiydiscus from Pondoland are far more compressed.

Locality. Lake Itesa (Eteza), Umfolozi. Coll. "W. J. Wybergh.

FAMILY : PRIONOTROPID.E.
GEN. MORTONICERAS, Meek.

5. MORTONICERAS WOODSI, Sp. 110V.

(PI. XXI, figs. 1 a-fZ.)

The single specimen (No. 5451) upon which this species is based
has the following dimensions :

Diameter . . . . .80 mm.

Height of the last whorl . . 45 per cent, of the diameter.

Thickness ,, ,, . . 40 ,, ,,

Umbilicus . . . . . ? 25

The important characteristics of this new species are (1) the decline
of lateral and peripheral ornament on the last whorl, which is still
septate, so that the specimen represents the inner whorls of an
Ammonite that appears to lose altogether the typical Mortoniceras
features ; (2) the projection of the tubercle at the overhanging
umbilical edge in an inward, not a lateral direction ; (3) a com-
paratively small umbilicus.

Among a large number of Morton In- rax of the type of M. soutoni
(Baily)* from the Umtamvuua River, in the British Museum, there
are some transitional forms to the present species, showing decline
of tuberculatiou on the outer whorl (at a cousidei-ably larger diameter)
and a decrease in the size of the umbilicus. On the other hand, one
of the forms figured by Stuart Wellerf as Mortoniceras delawarense,

* The example figured by Woods (loc cit., 1906, p. 337, pi. xliii, fig. 1 ) repre-
sents a more evolute shell. Baily's type in the British Museum (Geol. Soc.
Colin. No. 11365) has the decline of tuberculation more pronounced, but at a
diameter of close on half a metre is still costate. Its small umbilicus brings
it closer to the new species here described than is Woods' more evolute
example.

t "Keport on Cret. Pal. of New Jersey," vol. iv (Pal. Ser.), 'Geol. Surv. of
New Jersey,' 1907, p. 837, pi. civ, figs. 1-3 only. (See under Mortoniceras
Vanuxemi, Morton sp., p. 308.)

On Cretaceous Cephalopoda from Zululand. 233

Morton sp., is considerably closer to the present species, as is one of
the specimens figured by Whitfield,* which latter example, however, is
of a considerably larger size. These forms of the delatvarense-group,
however, neither have the overhanging umbilical edge, nor the close
costation of the present species, though they are nearer to the latter
than is any other of the very numerous species of Mortoniceras
described.

Perviuquicref doubtfully recorded M. delawarense (with varieties)
from Tunis, but his specimens are much more strongly tuberculate
than the new form here described and thus resemble M. campaniense,
Grossouvre, which species subsequently was united by its author
with 3/. delawarense.

One characteristic feature of these forms of the delawarense group
is the weakening of the wide and low keel, carried to extremes in the
fragment figured as M. delawarense by Julia A. Gardner.* It is
difficult, from an inspection of the figure, to form an opinion as to the
exact relationship of this form, for it has an almost flat, Hoplites-like
ventral area. At any rate, its convergence towards such a form as the
Upper Campanian A. marroti (Coquaud), Grossouvre, is striking.

None of the species of Mortoniceras from the European Senoniau
resemble the form here described. The suture-lines of Grossouvre's
Coniacian species are considerably simpler than are those of the Cam-
panian delawarense-gronp. It has been possible to develop the
internal portion of the suture-line of the present species, and a com-
parison with that of M. texanum (Burner) as figured by Schliiterj| is
interesting as showing not only great increase in complication, but
accommodation to a different whorl-shape. The suture-line of an

* " Gast. and Ceph. of the Earitan Clays, etc.," ' Mon. U.S. Geol. Surv.,' vol.
xviii (1892), p. 252, pi. xliii, figs. 1, 2 only.

t " Etudes de Pal. Tunis," I, ' Ceph. d. Ter. Second,' 1907, p. 243, pi. XL, figs. 21
and 22. Pervinquiere, in his note (1) on p. 244, somewhat misrepresents
Whitfield, for the remark in the latter author's description of M. vanuxemi
(p. 254) refers to the compressed specimen he figures (pi. xlii, figs. 3 and 4),
not to Morton's type. The latter is somewhat doubtful. It would be advisable
to take as type of M. delawarense, Morton sp., figs. 6 and 7 in Whitfield, and as
type of M . vanuxemi (Morton em. Whitfield), figs. 3 and 4 of the same plate (xlii)
in Whitfield. Stuart Weller (loc. cit., p. 839) unites the two species again in
the writer's opinion unjustly.

t " Up. Cret. Dep. of Maryland," ' Maryland Geol. Surv. Baltimore,' 1916,
p 391, pi. xii, fig. 7.

Hoplites vari, Schliiter var. mar rot i in Grossoxivre, loc. cit. (1894), p. 119,
pi. viii, fig. 36= Hoplitoplacenticeras, Paulcke.

| "Cephal. d. Ob. Deutsch. Kreide," ' Palaeontogr.,' vol. xxi, part 2 (1872),
p. 41, pi. xii, fig. 3.

20

234 Annals of the ftouth African Museum.

example of M. sontoni (Baily), very close bo Baily 's type, with small
umbilicus, but comparatively smooth outer whorl (B.M. No. C19441,
after G-. C. Crick), also is given for comparison (Plate XX, fig. 4),
since it differs considerably from that figured by Woods and from the
original (faulty) drawing in Baily.

Locality. Umkwelane Hill, Umfolozi. Coll. Dr. A. L. du Toit.

6. MOETONICEEAS aff. UMKWELANENSE, CT. C. Crick.

(Text-fig. D 2, p. 297.)

1907. M. nmkwelanense, G-. C. Crick, Joe. cit. (Third Eeport), p. 228,
pi. xv, figs. 9, 9 o.

Crick briefly characterised this species, which he considered to be
related to M. sovtoni and M. stangeri, as having a " subquadrate trans-
verse section, a little wider than high." The example to be described
(No. 5491) differs from the holotype (B.M. No. C18134) in several
respects, but in view of the great variability of the nearly related M.
soutoni, of which a large number is available, it is considered in-
advisable to separate the present example from the specimen figured
by Crick. The dimensions of the specimen in the present collection are :
Diameter . . . . 250 mm.

Whorl height . . 82

Whorl thickness . . . . 69

At a diameter of 160 mm. the measurements for height and thick-
ness are 60 mm. and 62 mm. respectively ; at 90 mm. diameter they
are 33 mm. and 39 mm. Whereas, thus, the inner whorls are wider
than high, and at a diameter of 160 mm. the whorl-height almost
equals the thickness, as in the holotype, on the outer whorl the
proportions are reversed and the whorl is higher than wide. It may
be added that at 250 mm. diameter the specimen is still septate.
Owing to the presence of an additional, if slight, tubercle on the lateral
area of the specimen here described, the whorl-section is more rounded
than that of the holotype, a feature still further accentuated by the
removal, away from the umbilicus and higher up on the side, of the
umbilical tubercle.

At a diameter of 90 mm. there are only fifteen coarse costae per
whorl, as in the holotype. The suture-line is of the same general type
as that of M. soutoni; the position of the umbilical tubercle, however,
in the latter, corresponds with the second lateral saddle, whereas in the
present example it coincides with the inner branch of the bifid first
lateral saddle (compare Text-fig. D 2 with fig. 4 of Plate XX).

The presence of a fifth tubercle, smaller than the others, as it also is

On Cretaceous Cephalopoda from Zululand. 235

in examples of M. texanina (Romer), makes the Ammonite here dis-
cussed somewhat of a transition between the compressed form next
described and compared with M. soutoni, and Crick's holotype. The
inner whorls, however, are different in the two developments, and there
are other distinctions, as pointed out in the description below. In M.
soutoni, as well as in M. texanum (Romer) and M. quinquenodosum
(Redtenbacher), the umbilical tubercle is near the edge, and M. cam-
paniense, de Grossouvre, :1: though the young has a similar squarish
section and low keel, differs in its dichotomous costation.

It may be added that in the holotype of M. umkwelanense the keel
has quite disappeared near the end a feature of great significance, but
not sufficiently apparent from the original figure. In the present
example the two outermost tubercles are not quite so close, the keel
between them not quite so feeble and not quite lost at the end.

Locality. Umkwelane Hill. Coll. Dr. A. L. du Toit,

7. MORTONICERAS sp. aff. souTONi, Baily sp.

A large but somewhat fragmentary example (No. 5492), of 180mm.
diameter, agrees with the evolute specimen figured by Woods,t and
has fairly smooth inner whorls, with the lateral tuberculatiou only just
indicated. The ribs also are, then, quite indistinct. The five
tubercles become as conspicuous as they ai'e in Woods' example and in
Redtenbacher's M. quinquenodosum,* only just before the beginning of
the body-chamber (at a diameter of about 100 mm.). The body-
chamber, however, develops increasingly strong tubercles, which
character separates the example here described from the typical and
more involute M. soufnn! ,- for Baily's type shows decline of tuber-
culation at a stage when the present example develops its strong
tuberculation, and in the specimen of M. soutoni mentioned under M.
woodsi (see supra, p. 234) the outer whorl becomes almost smooth.
There are twenty-seven costae, as in the specimen figured by Woods ;
near the end of the specimen the whorl-height is 64 mm. as compared
with a thickness of 55 mm. The whorl-section, thus, is considerably
thicker than that of M. texanum, Romer, which, however, it greatly
resembles in the spacing of the tuberculation.

The writer agrees with Woods || in considering M. bontanti, de

* Loc. cii. (1894), pi. xiii, figs. 1 and 3.
t Loc. cit. (1906J, p. 337, pi. xliii, fig. 1 a.

t " Cephal. Fauna d. Gosau-Sch. i. d. N.6. Alpen.," ' Abb. K.K.R.A.,' vol. v
(No. 5), 1873, p. 108, pi. xxiv, fig. 3.

' Die Kreidebild. v. Texas, etc.,' Bonn, 1852, p. 31, pi. iii, fig. 1 Z> (and 1 ?).
|| Loc. cit., p. 338.

236 Annals of the Soiiih African Museum.

Grossouvre, to be quite different from Baily's species. On the other
hand, the specimen figured as M. texanum by de Grossouvre* agrees
with the example here described, both in the character of the
ornament and in whorl-section, though the inner whorls appear to be
quite different in so far as the fragmentary condition of the present
example permits of comparison. De Grossouvre's example may be iden-
tical with Hauer's A. texanus'f = A. quinquenodosus, Redtenbacher,+ as
Lasswitz thinks, if the thinness of the latter species is due to crushing,
but the writer cannot admit the same author's identification with M.
texanum of de Grossouvre 1 s M. campaniense, a form near to M. dela-
warense, with which, indeed, de Grossouvre || subsequently united it.

M. umlcwelanense, Crick,^[ has much thicker whorls, with a square
section, but the specimen here compared with Crick's species,** owing
to the presence of a slight fifth (lateral) tubercle, is, perhaps, closer to
the example under examination than is the type. In both, however, the
umbilical tubercle is further away from the umbilical edge than it is
in M. sp. aff. soutoni, and whereas in M. umliivelariense and in the
close ally, described in this paper, the inner whorls are relatively more
coarsely ornamented than the outer, the reverse development is found
in the example here compared with M. soutoni.

It may be added that the great variability of the species here dealt
with is shown in a large series of Mortoniceras from Pondoland in the
British Museum, referred by Crick to Baily's two species and to an
" intermediate " group.

Locality. Umkwelaue Hill. Coll. Dr. A. L. du Toit.

GEN. PSEUDOSCHLOENBACHIA, nov.
Genotype : A. umbulazi, Baily, 1855, pi. xi, fig. 4.

A. umbnlazi, Baily, described as Schloenbachia by Woodsff and
Crick, ++ has been considered by Kossmat to belong to the " narrowly -

* Loc. cit. (1894), pi. xvii, figs. 1 a, b, non pi. xvi, figs. 2-4.

t "Ceph. d. Gosati-Sch." ' Beitr. z. Pal. v. Osterr.,' vol. i, 1858, pt. i, p. 10,
pi. ii, figs. 4-6.

J Loc. cit., p. 108, pi. xxiv, fig. 3.

" Kreide-Amm. v. Texas," ' Geol. und Pal. Abb. Koken,' N.F., vol. vi
(1904), p. 31.

l| ' Recherclies s. 1. Craie Super.' (1901), p. 379.

1 Loc. cit., p. 228, pi. xv, fig. 9.

** See above, p. 234.

ft Loc. cit. (1906), p. 336.

JJ Loc. cit. (1907), p. 250.

" Die Bedeut. d. Siidind. Kreideform., etc.," ' Jb. K.K.E.A.,' vol. xliv (1894),
p. 464.

On Cretaceous Cephalopoda from Zululand. 237

umbilicated Schloenbachia (Prionocyclus) forms that appear in the
Lower Senonian," aud to be closest to A. paon, Eedtenbacher, and
to A. haberfellneri, Hauer. The genus Barroisiceras, however, to
which these two species belong, is characterised by a simplifying
suture-line ; and whereas its typical branch develops a concave
periphery, another group (with acute periphery) tends towards certain
Tissotids. Solger ,* who doubtfully classed A umbulazi in M^mier^ceras,
appears to have been much nearer the mark. The young A. umbulazi
reproduces the ornament of Muniericeras lappareuti, Grossouvre.f and
has chevrons on the venter, but no keel (Fig. B2). But before this
stage, the young A. umbulazi reproduces Puzosia, with constrictions
(Fig. B 3), and the suture-line throughout shows great resemblance
to that of many Desmoceratids. There is, however, at least an
equally great similarity, shown by the apparently continuous series
Sonneratia-Cleoniceras and of Hoplites-Pleurohoplites +-8chloenbach la
with certain contemporary Desmoceratids ; also, possibly, Munieri-
ceras itself includes Prionotropid forms (Grossouvre's species, M.
yosauicum, Hauer sp., M. dresdense, Petraschek) and Desmoceratid
species, e.g. the lower Seuonian A. clypealin, Schliiter and Brauns,||
and " Desmoceras " clypealoides, Leonhard,^" apparently connected
with the Turoniau Puzosia hernensis, Schliiter sp. Thus the presence
of constrictions on the inner whorls of P. umbulazi, and a suture-line
that resembles that of certain Desmoceratids, are not definite proof
that Pseudoschloenbachia be derived from Puzosia rather than from
Prionotropidae, via Muniericeras. Constrictions may appear in
Schloenbachia as well as in Pseudoschloenbachia and Gauthiericeras and
other genera (e. g. Hystatoceras, Anahoplites) apparently quite spon-
taneously, and one example of Schloenbachia varians in the writer's
collection shows constrictions, like Gauthiericeras four nieri, Grrossouvre
sp., with deep chevrons across the venter, accompanied by decline
of ornament and loss of keel.** At any rate, if comparison is at
all permissible with the rather distant Hatiericeras, a Desmoceratid

* " Foss. Mungokreide," ' Beitr. z. Geol. v. Kamerun, II ' (1904), p. 205.

f Loc. cit. (1894), p. 158, especially pi. xxxv, fig. 3.

t Gen. nov. for studeri group, Jacob em. Spath (genotype, A. renauxiamis,
d'Orbigny, 'Pal. Frang. Ter. Crct./ pi. xxvii).

Fcmcms-group only.

|| de Grossouvre (loc. cit., p. 175) has pointed out that Brauns' form is
different from Schliiter's type.

f "Fauna d. Kreidef. Oberschles.," ' Palseontogr.,' xliv (1897), p. 57,
pi. vi, figs. 2 a, b.

** This excludes any possibility of a connection between Prohauericeras or
Pseudoschloenbachia and the true Schloenbachia.

238

Annals of the South African Museum.

genus with acute periphery, the group of " Schloenbachia" here
discussed would be post-Hauericeras, not pve-Hauericeras, speaking
phylogeuetically, and irrespective of age in other words, it would have
travelled farther away from the Desmoceratid stock than Hauericeras.
Now Nowak* proposed the new genus Prohauericeras for those
forms of Schloenbachia that showed a decided tendency in the direction
of Haut'rii-f-rax. The derivation of the latter genus from Schloenbacliia
(via Prohauericeras) can on no account be admitted. Hauericeras has
a truly Desmoceratid suture-line, as a comparison of its internal
sutures (Fig. A 7) with those of Puzosia p/fl?<?rtYa(Sowerby),Kossmat,t

TEXT-FIG. A. Hauericeras gardcni, Baily, sp. Umtamvuna Kiver, Natal.
(B.M. No. C 18528 [17 and 9] and C 18531 [8]). 1. Suture-line at
diameter == 4 mm. si = siphuncle ; u = umbilicus. 2. Ditto at 6 mm.,
without external saddle. 3. Second lateral saddle and auxiliaries at 14
mm. D. 4. Ditto at 20 mm. D. 5. Ditto at 50 mm. D. 6. Internal
(dorsal) suture-lino at diameter = 6 mm. ; AL = antisiphonal lobe ; u =
umbilicus. 7. Ditto at 50 mm. D, with second and third auxiliary saddles
of external suture-line, and umbilical saddle (u). A.L. = antisiphonal lobe.
8. Whorl-fragment at D = 90 mm., K == hollow keel, on barely carinate
venter of cast. (Photo, by Gr. C. Crick.) 9. Outline whorl- section atD =
75 mm., showing hollow keel and rounded inner whorls. All the figures
are enlarged.

will show. The inner branch of the second lateral saddle, that is so
conspicuous a feature of the development of the suture-line of
Hauericeras gardem (see Fig. A 1-5)+ is similarly developed in many
forms of Puzosia (e. g. subplanulata, Schliiter, compressa, indopacijica,

* " Unters. Ceph. Ob. Kreide Polen., Ill," ' Biill. Ac. Sci. Cracovie,' June,
1913, p. 370.

t Loc. cit. (1898), pi. xvi, fig. 4.

J The specimen (B.M. No. C18528) from which these figures were drawn,
agrees in all respects with Baily's type (B.M. Geol. Soc. Coll., No. 11370),
and the larger of the two fragmentary co-types (No. 11371) from which Baily's
figure of the suture-line was taken. Some Japanese examples of H. gardeni in
the British Museum (Geol. Soc. Coll.) also show this peculiarity of the suture-
line, figured by various authoi-s, in exactly the same manner.

On Cretaceous Cephalopoda from Zululand. 239

Kossmat, yaudaiua, Forbes, etc.)- When ornament appears in
Hauericeras (H. buszii, var. nodosa and var. costata, Wegner*), it does
so at a late stage ; and the periphery of the cast is barely sharpened
in H. yardeui at a diameter of 60 mm., which accounts for the fact that,
e. (j., Pervinquiere,t who, unlike Nowak, clearly recognised the hollow
keel, found the casts of his small examples perfectly rounded.

The assemblage mentioned by JSTowak includes Albian, Cenomaniau,
Turoniau and Seuouiau forms and is most heterogeneous. Sharpe's
A. goupilianus (nun d'Orbiguy) is only a smooth variety of Scldoen-
bachia varians, Sowerby sp. Whiteaves' newly-created ScJiJ. propinqua
has as little to do with Stoliczka's earlier ^4. propinquus, as
Anderson's Sell. ^rnpiu^ua% (Stol.), possibly a Turoniau Prionotropid,
is related to the Indian species. Such Albian forms as A. acuto-
carinatus, Shumard, belonging to the group of A. roissyanus,
d'Orbigny, $ again, arc as little related to the true Ceuomaniaii
Schloeribachia or to " Prohauericeras," as the Neocomian Oosterella
cultrataeformis, Thlig sp., is to either.

Since Prohauericeras has been proposed, it may be convenient to
retain it|| for A. youpilianus, d'Orbigny, which apparently was intended
to be the type, since Nowak mentions it first, and refers to it again in
connection with "Schloeribachia "fotirnieri, G-rossouvre, and "$.'' obesa,
Stolic/ka sp. In suture-line, as well as in other characters, this restricted
Prohauericeras differs both from the trueCeuomanian Schloeribachia and
from its own (Turonian) contemporaries among the Prionotropidae. On
the other hand, Pseudoschloeribachia is quite different again from any of
these developments. Its internal suture-line (Fig. B 5 and 6) is figured
for comparison with that of Schloenliackia varians (Fig. B 10) ; and
it will be seen that it differs very considerably, especially from that of
Hauericeras. As, however, more involute forms of Puzosia and other
Desmoceratids, including the keeled "P." stigata, Forbes sp., show, the
stretching out of the auxiliary elements may only be the result of
adaptation of a suture-line to wider sides, and the raising of the
umbilical portion is often found in younger developments. The suture-
line by itself thus is as unsatisfactory a character in many cases as, say,
the carination would be, if taken as the only basis for classification.

* "Die Granulat. Kr. .1. AY. Miiusterland.," ' Zeit. D. Cleol. Ges.,' vol. Ivii
(1905), p. 20S, pi. viii, figs. 1 a and b.

t Loc. cit. (1907), p. 165.

J Loc. cit. (1902), p. 128, pi. ii, figs. .34 -38. Anderson (p. 63) also compared
AVhiteaves' " 8." propinqun with the Utatur form.

See under Pseudophacoceras (p. 283).

|| The name is no more unsuitable than is, e. g., Freeh's Paralytoceras for a
Devonian Clymenid.

240 Annals of the South African Museum.

The hollow keel of Hauericeras, appearing comparatively late in
ontogeny, cannot be a development of the solid keel of Schloenbachia,
in which genus it appears after the costation, and, occasionally, tuber-
culation, and after the shell had become fairly involute. Similarly,
the ibex-like periphery of the young Pseiidoschloenbachia is not in favour
of a possible connection of this genus either with Schloenbachia or
with Prohauericeras. The various Natal forms indicate that this stock
had its own involute and smooth developments (P. griesbachi, Crick MS.)
and highly tuberculate and constricted forms (P. papillata, Crick MS.),
and transitions between these extremes. There are oxycone develop-
ments with simplifying suture-lines (Eulophoceras and Spheniscoceras),
and a hitherto unknown group (Diaziceras) that shows a superficial
resemblance to the South American Lenticeratids, and is interpreted
as a link connecting Pseudoschloenbachia with those genera (JEulopho-
ceras-Spheniscoceras) that carry the simplification of suture-line and
specialisation of whorl-shape to a higher degree. It is probable
that these clearly allied forms are only indirectly connected with
Desmoceratids, and that Pseudoschloenbachia via Muniericeras (and
like Gauthiericeras, to which probably 8. bertrandi, S.fournieri,Grros-
souvre,* and the forms of the Syrian Senouian have to be added) is
derived from Prionotropidae.

8 and 9. PSEUDOSCHLOENBACHIA UMBULAZI, Baily sp.
(PI. XX, figs. 2 and 3 ; Text-fig. B 2-7).

1906. Schloenbachia uinbulazi (Baily). Woods, " Cret. Fauna of
Poudolaud," Ann S. Afr. Mus., vol. iv, part vii, No. 12, p. 336.
(See there for synonymy.)

Two specimens are referred to this form, the smaller one (No. 5494)
(PI. XX, fig. 2) agreeing particularly well in costation with Baily's
type-figure. Its dimensions are :

Diameter . . .43 mm.
Height of last whorl . 51 per cent, of the diameter
Thickness .28

Umbilicus ... 13

The larger specimen (No. 5459) is less coarsely costate at the same

* A specimen of " Schloenbachia " boreaui, Grossouvre, from the Middle
Coniacian of the Charente Inferieure (B.M. No. C7387) shows that the group
of " S." fournieri is more nearly allied to Gauthiericeras, and has nothing to do
with the probably Campanian Pseudoschloenbachia.

On Cretaceous Cephalopoda from Zululand.

241

diameter, but develops the typical ornameut, with distinct umbilical
tubercles, on the outer whorl. It has the following dimensions :

Diameter . . .59 mm.

Height of last whorl . 54 per cent, of the diameter

Thickness . 27

Umbilicus 13

Both the specimens show the suture-lines well and have portions.
of the body-chambers preserved, the former rather less than the last
half of the outer whorl, the. larger specimen a little over half a whorl.

TEXT-FIG. B. 1. Pseudoschloenbachia umbulazi, Baily sp., Umtamvuna River,
Natal (B.M. No. C19427). Sectional outline of inner whorls (at diameter
= 32 mm.). 2-6. Pseudoschloenbachia umbulazi, Baily sp., Umkwelane
Hill, Zululand (Specimen No. 5494) (p. 240). 2. Ventral aspect of inner
whorls showing chevrons at D = 12 mm. 3. Lateral view of innermost
whorls at D = 5 mm. 4. Ditto, suture-line at D = 2 mm. 4A. Ditto,
suture-line at D = 4 mm.. 5. Ditto, internal suture-line at 4 mm.
6. Ditto, internal suture-line at 16 mm. 7. Pseudoschloenbachia umbulazi
(Baily) var. acuta, nov. Suture-line of type-specimen (No. 5450) from
Umkwelane Hill (p. 241). 8. Pseudoschloenbachia griesbachi (Crick MS.),
Umtamvuna River, Natal (B.M. No. C19428). Suture-line of type-specimen
at D = 60 mm. (reduced 5). The edge should be minutely frilled.
9. Aconeceras nisoides, Sarasin, sp. Suture-line of specimen No. 5119 from
Powell's Camp, Upper Catembe (p.311). 10. Schloenbachia i-arians(Sowerby),
Cenomanian, Warminster, Wilts. (Coll. L. F. Spath). Internal suture-
line at diameter = 15 mm. All the figures except 8 are enlarged.

An immature third example (No. 5450) (PI. XX, fig. 3) of 24 mm.
diameter, and wholly septate, agrees with the first specimen described
and with Baily's type-figure in the distinct costation, :;: but has a thinner
section at an equal diameter and an acutely fastigate periphery. It
may be separated as-

var. ACUTA, uov.

Mr. Woods stated that P. umbulazi appeared to be confined to
Poudoland. In addition to the present specimens from Zululand,

* There is no tubercle on the ribs at the middle of the side, as might wrongly
be inferred from the side-view given in PI. XX, fig. 3.

242 Annals of the South African Museum.

there is an immature, evolute specimen from the Umpeuyati Eiver,
Natal, recorded by Crick" ; on the other hand, five specimens from
the Umtamvuna Eiver, Natal, in the British Museum, identified by
Crick, represent more strongly ornamented varieties, with increasing
tuberculation round the umbilicus. One of these examples (No.
C19425) is slightly constricted and transitional to a highly tuber-
culate species named " Schloenbachia" papillata by Crick.

Specimens of the latter species, also of the transitional form, from
Zululand and belonging to the Albany Museum, were sent to the
writer in 1914 through the kindness of Mr. Woods, but since
Crick had been at work on his paper on this fauna for many years,
and since there were only a few specimens, they were not described.
P. p(ti>ill<tta, G. C. Crick MS. sp., is not represented in the present
collectiou,f nor are the transitional forms, referred to above.

The three specimens were collected at Umkwelane Hill, Umfolozi,
Zululand (Coll. Dr. A. L. du Toit).

GEN. DIAZICEKAS, nov.
Genotype : I), tissotiaeforme, sp. nov., p. 245, PI. XIX, figs. 1 a-lf.

This genus is created for one form in the collection that cannot be
included in any of the known genera of keeled Seucnian Ammonites.
Its suture-line stamps it as being near to the genera Eulophoceras,
Hyatt, and Splieniscoceras, Crick MS.,J and in general outline this

* Lnc. cit. (1907), p. 250.

t Two additional specimens of tliis species were included in the collection
from the Durban Museum, already referred to, and will be figured.

This genus was created for the three species S. africanum, S. minor and
5. tenue, G. C. Crick MS., which obvioxisly are the " other species of Eulopho-
ceras " referred to by Woods (loc. cit., p. 337). Crick, in his diagnosis of the new
genus, does not in any way mention the genus Eulophoceras, which, however, is
very close, as will be seen on comparing the suture-lines given in Text-fig. C J
with that of Eulophoceras natalense, Hyatt (Text-fig. C 2, after Woods, loc. cit.,
pi. xlii, fig. 8), and with Hyatt's type (Pseudoceratites, 1903, p. 86, pi. xi,
figs. 2-6). On the other hand, Crick considered the new genus to be " inter-
mediate between Placenticeras and Sphenodiscus," which statement is meaning-
less, even from a purely morphological point of view, for Spheniscoceras has a
thickened keel, after the style of that of the remarkable form figured as A.
roissyanus, d'Orbigny, varietas, by Ooster (Catalogue, etc., 1860, pi. xxvi, fig. 7),
or of certain Dipoloceras and Pseudophacoceras described in this paper (see, e. </.
PI. XXV, fig. 1 d, and PI. XXVI, fig. 56), placed on a very thin and acute whorl.
This is an exaggeration of the feature shown in Hyatt's fig. 6 a of pi. xi, but of
doubtful generic importance. Haiifriceras rembda, Forbes, shows a similar
feature occasionally. In fact the writer is not convinced that Crick was right in

On Cretaceous Cephalopoda from Zululond.

243

suture-line agrees with the South American Lenticems and Paralenti-
ceras included by Hyatt in the family Euloplwcerat'ulae* These
genera, however, include more or less smooth forms, whereas the new
genus here proposed is characterised by strong ornament. The latter
consists of very prominent umbilical tubercles, each connected by
obscure broad ribs with about five outer tubercles. This type of
ornament is found in the Senoniau in certain Pseudoschloenbachia
(P. papillata, G. C. Crick MS.), in JRnrrolslceras (desmoulinsi) and in
certain Tissotids. The first genus has a highly complex suture-line,
characterised by a very deep principal lobe, whereas the suture-line of
the new genus shows signs of simplification in the peculiar rounding

separating the forms he described from Enln^lmn-i-as. Eulophoceratidae maybe
distantly related to the contemporary Sphcnodiscidae (not the Turonian Ccnlo.
poceratidae, often confused with Sphenodiscidae), but there is no connection

TEXT-FIO. C. 1. Sijhenisrocervs, G. C. Crick (MS.), Upper Senonian,
Umtanivuna River, Natal. (After drawings, of the natural size, by the
late G. C. Crick.) 1 a. S. <//"//><( /MO/I, Crick MS. (genotype), B.M. No.
C194.21. 1 b. S. minor, Crick MS. (B.M. No. C19422). 1 c. S. tenue. Crick
MS. (B.M. No. C19423). 2. Eulophoceras natalense, Hyatt (after Woods),
Pondoland. For comparison.

whatever between Spheniscoceras or any other Eulophoceratid and the
Placenticeratidae.

* Loc. cil., 1903 (Pseudoceratites), p. 16, also wrongly including Tegoceras,
Hyatt, which is a Hystatoceratid.

244 Annals of the South African Museum.

of the terminal folioles, the short and simple lateral lobe, aud the
beginning development of adventitious lobes.

Barroisiceras includes shells (e.g. B. desmouUnsi, Grossouvre sp.)
that are near the new genus in external shape, though less so in
suture-line, aud these forms with persistent keel, at any rate, may
have to be excluded from Barroisiceras, since the typical group of this
genus develops a concave periphery.

Tissotidae have a pseudoceratitic suture-line, quite different from
that of the genus here described, though certain forms, e. g. Meta-
tissotia fourneli, Bayle sp., somewhat resemble it in external shape.
The earlier Psendotissotidae have a less simplified suture-line, but
cannot have given rise to a highly ornamented descendant in the
Senonian such as the form here described.

Derivation from the main stock of the normally-lobed Prionotro-
pidae, through Pseudoschloenbachia, is most probable, and it seems that
whereas Diaziceras is nearer to the ancestral Psendoscliloenbacliia in
ornamentation, Splieniscoceras and Eidophoceras are, perhaps, closer to
it' in suture-line. It might be advisable to retain Hyatt's family
Eulophoceratidae for these four genera, all presumably of Campanian
age, but though there is a certain family resemblance with the suture-
lines of Eulophoceras&ud Spheniscoceras (cf. Fig. C la) and less so with
Psendoscliloenbachia, the similarity of the suture-line of Diaziceras
with those of Lenticeras, Paralenticeras and other pre-Campauiau
genera makes it doubtful whether in the present state of our know-
ledge a subdivision of the Seuouian Prionotropidae can yet be
attempted. Moreover, an undescribed form, closely resembling
"Barroisiceras naberfellneri '" from Madagascar, as figured by Boule,
Lemoiue aud Tlu'veniu,* but possibly nearer to the Upper Chico
" Schloenbachia " chicoensis, Trask sp. (Audersouf) occurs in Pondoland,
associated with many Campauian species, but also with " Puzosia "
sugata, Forbes sp., which occurs in the Lower Chico Beds of California,
and in Lower and Upper Senouian beds in India and elsewhere.
The likeness of these presumed Campaniau forms with Barroisiceras
may only be accidental, and since the median row of ventral tubercles
in the new Pondoland form is very high and acute, it may well be
assumed to lead from e. g. Muniericeras to Pseiidoschloenbachia and
Diaziceras. On the other hand, it looks as though a revision of
the many forms of " Schloenbachia " in the Lower and Upper Chico
Beds were most likely to throw light on the possible connec-
tion between Diaziceras and Barroisiceras on the one hand, and

* Loc. cit. ii (1907), p. 43, pi. xi, fig. 3.
f Loc. cit. (1902), p. 116, pi. ii, fi<?s. 23-25.

Oti Cretaceous Cephalopoda from Zulu! and, 245

Pseudoschloenbachia and Ganthiericeras and Muniericeras on the
other.

The uncertainty as to the presence of pre-Campanian horizons in the
Senoniau of Pondoland and Umkwelane Hill is an additional difficulty.
Though, thus, the close resemblance of the suture-line of Diaziceras
with that of, e. g., Lenticeras andii, G-abb sp., and L. baltai, Lisson,*
might be explained by the mechanical laws that govern the formation
of the Ammonite-septum and its edge,f yet it is curious that as
Lenticeras is associated with Mortoniceras te.ranum, so Diaziceras
occurs together with the comparable M. umkwelanense, whereas the
presence of Peroniceras in Zululand, and of " Pnzosia " suyata and
the above new form in Pondolaud, suggests that the new genus may
be closer to Coniaciaii genera than is here assumed. Unfortunately
the writer has no comparable material for dissection.

10. DIAZICERAS TISSOTIAEFORME, nov.
(PI. XIX, figs. 1 a-*.)

This species is based on a completely septate specimen (No. 5478)
having the following dimensions :

Diameter . . . .80 mm.

Height of the last whorl . 50 per cent, of the diameter.

Thickness ., . 59

Umbilicus .14 ,, ,, .,

The small and deep umbilicus is surrounded by four very prominent
tubercles, increasing in size with age, and each connected by faint ribs,
with about five smaller rounded tubercles on the ventro-lateral edges.
The prorsoradiate processes of these tubercles towards the very sharp
ventral edge are very faint, so that the roof-shaped periphery is
almost smooth. Where the shell is preserved on the ventral edge,
near the end of the specimen, it follows the shape of the fastigate
periphery of the cast, but at the beginning of the last whorl, where
the peripheral character of the inner whorl is well shown, the shell
rises in a distinct keel above the less acute ventral edge of the cast.
The whorl section is polygonal, with the greatest whorl-thickness at
the umbilical tubercles, and the two ventral and the two lateral faces
concave, but the umbilical slopes convex.

* Loc. cit. (1908), pis. xiii and xiv.

t See the writer's " Notes on Ammonites," ' Geol. Mag./ 1919, January to
May numbers, and compare, e.g., the suture-lines of Pseudophacoceras (PI. XXV,
fig. 1 b and c), and Oxynoticeras (Pia, 1914, pis. viii-xi), or of Aconeceras
nisoides (Fig. B 9, p. 33) and Pseudoschloenbachia grieslachi (Fig. B 8).

246 Annals of the South African Museum.

There are twelve septa on the last whorl. .The most striking
characteristics of the suture-line are the very deep external lobe and
reduced first lateral lobe, the trifid external saddle and the peculiar
rounding' of the terminal folioles of all the saddles. These characters
are found in the suture-lines of Lenticeras audit, G-abb sp., L. baltai,
Lisson, and of Paralenticeras sieversi, Gerhard t sp.,* and of the two
close allies Eulophoceras and Splieniscoceras, which here are assumed
to represent developments of the same stock. On the other hand, the
suture line of " Barroisiceras " desmoulinsi, Grossouvre sp., as figured
by Solger,f though showing a superficial likeness, differs considerably
in its short external lobe, broad- stemmed external saddle and very
simple outlines, even of the internal portion. This species, however,
shows the greatest resemblance to the present species in its polygonal
whorl shape. There are differences in ornament, but the important
distinction is the presence of ventral tubercles in Grossouvre's form,
which, when worn, may present the appearance of an entire keel.
Solger's Cameroons specimens had an entire keel from the youngest
stage, but differ from the Zululand example in proportions, ornament,
and in having the peripheral tubercles elongated longitudinally.
One of Solger's specimens also becomes quite rounded ventrally in the
adult, so that these Couiaciau forms may belong to quite a different
stock from the presumably Campaniau Diaziceras.

PseuJofchloenbarJiia papillata, G. C. Crick (MS.) sp., is a considerably
thinner form, with eight umbilical tubercles, each corresponding to
about four outer tubercles. It has a higher keel than is shown in
the sectional views of Gauthiericeras bertrandi and of G. fonrnieri in
Grossouvre, J and has much more prominent umbilical tubercles. But
in Pseudoschloeribachia the suture-line is quite complex and charac-
terised by a large and deep first lateral lobe. A comparison of
fig. B 4 a (p. 241) with fig. 1 A- of PI. XIX will show that whereas
Pseudoschloeribachia at 4 mm. diameter is much more advanced than
Diaziceras, as regards elaboration of the septal edge, the latter genus
shows a deeper ventral lobe, with the whole suture-line rising
towards the umbilicus generally a feature of later developments.
The Muniericeras stage of Pseudoschloeribachia cannot be traced in
Diaziceras, and it has already been pointed out that there is a
possibility of the keel of these two genera being a development of the

* " Beitr. z. Kenntn. d. Kreideform. in Venezuela und Peru," in Steinmann,
" Geol. und Pal. v. S. Amer.," V, ' N. Jb. f. Min., etc.,' Beil-Bd. xi, 1897, p. 82,
text-fig. 6, and p. 79, text-fig. 5.

t Loc. cit. (1904), pp. 168 and 169, text-figs. 53 and 54.

| Loc. cit. (1894), pi. xxix, fig. (56, and pis. xxxv, figs. 1 b and 1 c.

On Cretaceous Cephalopoda from Znl/ihunl. 247

crenulated keel of certain Upper Chico forms that cannot safely be
referred to such known genera as Muniericeras or Barroisiceras, and
probably are later than either.

Locality. Umkwelaue Hill. Coll. Dr. A. L. du Toit.

FAMILY : PLACENTICERATID^.

GEN. PLACENTICEEAS, Meek.

11. PLACENTICERAS SUBKAFFRAKIUM, sp. nov.

(PI. XXI, figs. 2a-d.)

This species is based on a fairly well-preserved specimen (No. 5106)
of the following dimensions :

Diameter . . .85 mm.

Height of last whorl . 50 per cent, of the diameter

Thickness . 37 ,,

Umbilicus . . .15 ,, ,,

About one-half of the outer whorl belongs to the body-chamber,
but the mouth-border is not preserved. The suture-line given in
Fig. 2 d was taken at the beginning of the last whorl.

The species differs from P. kaffrarium, R. Etheridge fil.,* in having
a smaller umbilicus (surrounded by tubercles that are very prominent
already on the inner whorls, as shown in the umbilicus), in having a
wider ventral area, and in the great thickness, caused by the prominent
umbilical tubercles. The very conspicuous lateral ribs of P. liaffrarium,
on the other hand, are not found in the example here described, but
it should be mentioned that near the end of the specimen, where the
shell has been removed, the lateral folds are more distinct on the
internal cast than they are on the shell. The latter only shows very
indistinct ribbing between the seven umbilical tubercles and the
twenty-eight elongated outer prominences (unpaired) that border
the flat and smooth ventral area. In addition to this distant ribbing,
there are fine sigmoidal striae, both on the shell, as in P. tamulicum
(Blauford) Kossmat,f and on the cast of the body-chamber, as in
P. stantoni, var. bolli, Hyatt. J.

P. tamulicum, which, like the present species, belongs to the group
of P. syrtale, Morton, is distinguished from the form here described

* Loc. cit. (1904, Second Report), p. 89, pi. iii, fig. 16.
t Loc. cit. (1895), p. 174, pi. xxii (viii), figs. 1 a-c.

" Pseudoceratites of the Cretaceous," ' Mon. U.S. Ueol. Sui-v./ vol. xliv
(1903), pi. xli, fig. 7, p. 214.

248 Annals of the South African Museum.

by being much thinner, by having more delicate ornamentation, a
narrower siphonal area, with less prominent and closer nodes, and
slight differences in the suture-line, e. y. the first adventitious lobe is
smaller than the second one in P. subkaffrarium, whereas the reverse
proportions are noticeable in the Indian species.*

P. stantoni, var. bolli, Hyatt, f also is a close ally of the present
species, as is P. intermedium, Johnson, J the latter only distinguished
from the Zululaud form by a narrower ventral area and a wider
umbilicus.

Locality. Umkwelaue Hill, Umfolozi, Zululand. Coll. J. S. Hedges.

FAMILY: NOSTOCERATKLE.
GEN. NOSTOCERAS, Hyatt.

12. NOSTOCEKAS ? NATALENSE, sp. 11OV.

(PI. XXII, figs. 2 , 6.)

A depressed turricoue (No. 2746) with the apical portion missing,
like the somewhat similar Didijmoceras ? newtoni, Whitfield sp., cannot
definitely be referred to either Didijmoceras or Nostoceras until more
complete specimens are known. The example is distinct enough,
however, to justify a new specific designation.

The two and a quarter whorls preserved are septate throughout,
and though the suture-lines are too indistinct for delineation, they
appear to be of the same general plan as that of Nostoceras ? sul-
.angulatum, uov., with the external lobe and its small median (siphonal)
saddle between the two rows of tubercles. These are very prominent,
elongated, and continued on the under surface of the whorls into
simple and strongly forwardly inclined ribs, somewhat like those of
D. umbilicatum, Meek, but more oblique, and with a very steep
backward edge. On the upper surface of the whorls the costae
bifurcate at the tubercle, as in D. ? tricostatitm, Whitfield, || which
Hyatt^[ considered to be the possible geroutic stage of D. ? newtoni.

* P. syrtale, Morton, var. tamulicum (Blanford), Kossniat in Boule, Lemoine
and Thevenin (loc. cit., II, 1907, p. 47, pi. xii, figs. 3 and 4), is very close to the
Zululand specimen, but apparently possesses the outer tubercle of P. syrtale.

t Loc. cit., " Pseudoceratites," 1903, as above, also, e. g., pi. xliii, fig. 1.

J ' The Geol. of the Cerrillos Hills, New Mexico,' part ii, ' Palaeont.,' School
of Mines Quarterly, vol. xxiv, no. 2, 1903, p. 206, pi. viii, figs. 27 a, b.

" Invert. Cret. and Tert. Poss.," ' U.S. Geol. S. Territ.,' vol. ix (1876),
pi. xxii, fig. 5.

|| "Pal. Black Hills, Dakota/" U.S. Geol. S.' (1880), pi. xv, fig. 7.

H " Phylogeny of an Acquired Characteristic," ' Proc. Ani. Phil. Soc.,'
vol. xxxii (1894), p. 574.

On Cretaceous Cephalopoda from Ztiliilcnnl. 249

Only in the Zululaud species, there generally is only a thick posterior
and a fine anterior branch, without the intermediaries found in both
the Dakota forms. The new form of Didymoceras, figured by Schliiter
on pi. xxxv, figs. 1-4, as jlet&roceras polyplocum?, showing bifurcation
only, is, perhaps, closer to the form here described in this respect,
but the tubercles are far too fine. The whorl-section is as compressed
as that of Schliiter 1 s loosely coiled form,* and less rounded than that
of D. '? newtoni, which has a larger umbilicus. That of the present
species is as small as the umbilicus of D. pauper, Whitfield sp., or that
of D. arcliiaciannm, d'Orbiguy sp., which latter also shows a similar
forward sweep of the costatiou on the under surface.

Since the apex of the spire is not preserved, it is impossible to state
whether the early whorls were closely coiled, as seems probable from
the presence of a contact furrow on the upper surface of the highest
whorl, or whether the apex was an irregular spiral, as is the case in so
many forms of this group (Didymoceras). The coiling of the portion
that is preserved is the same as that of Tnrrlliti^ urnftis, Passy, also
recorded from Zululand by Crick. t " Turrilites" tridens, Schliitcr^
also shows similar coiling, but the drawing appears deceptive, and the
writer is inclined to consider this species to belong to Hyphantoceras,
to judge by the uuder-surface of the whorls.

If the form here described is a Nostoceras, it probably represents a
less specialised type than N. stantoni, Hyatt || ; for the single costae
with two lines of tubercles, so characteristic of Exiteloceras, are found
in the young of Nostoceras stantoni and of N. lieUcinum, Shumard sp.lj"
The latter form has a very short spire like the Zululand form, with a
deeply impressed suture, but the costatiou is quite different.

Locality. TJmfolozi Valley, East of Railway. Coll. Mr. Illing-
worth.

* This shows close resemblance to a beautiful specimen of Emperoceras
simplicostatum, Whitfield sp. (larger than the example figured by Whitfield,
' Bull. Am. Mus. Nat. Hist.,' vol. xvi, 1902, pi. xxvii), in the B.M. (No. C10808),
and though the earlier whorls apparently are quite different from those of
Nostoceras and Didymoceras, the ornament of the last volution strikingly
recalls that of D ? newtoni, Whitfield sp., and of D ? cooperi (Gabb) Whiteaves
(B.M. from Vancouver).

f Loc. cit. (1907), p. 176, pi. xi, figs. 3 and 4.

J Loc. cit. (1872), pi. xxxv, fig. 9.

The trituberculate (?) Turrilites peramplus Lasswitz (loc. cit., p. 14, pi. ii,
fig. 1) also resembles the present form somewhat in shape, but may be a true
Turrilites, though Lasswitz compares it with Schltiter's "Turrilites" tridens.

|| Loc. cit. (1894), p. 570.

If "Descr. of New Cret. Poss. from Texas," Proc. Boston Sou. Nat, Hist./
vol. viii (1861), p. 190.

21

250 Annals of tie South African Museum.

13. NOSTOCERAS ? SUBANGULATDM, Sp. UOV.

(PL XXII, figs. 3 or-c.)

A sinistrally coiled, fairly elevated but fragmentary turricone
(No. 2746A), consisting of just over two whorls of body-chamber and
of a small portion of the septate and possibly more loosely coiled
earlier whorls, is doubtfully referred to the genus Nostoceras. There
is great resemblance to the dextrally coiled Didymoceras ? stevensoni,
Whitfield sp.,* and the helicoid character of the earlier whorls, if
proved by the discovery of more perfect examples, may necessitate the
transfer of the new form to the genus Didymoceras ; on the other
hand, the beginning of the example here described already shows an
impressed zone of contact, so that the reference to Nostoceras seems
most proper. Like the species last described, the present example
with its strong, simple costation and double row of ventral tubercles
recalls the ornamentation of the genus Exiteloceras, and the fragment
of Ex. artfjulatnm figured by Meek,t shows a close resemblance to the
septate portion here described, though there is a considerable difference
in size. The openly helicoid or irregular coiling of the young of Evite-
loceras,% however, is quite distinct.

The whorl-section is almost rounded, except for the double row of
tubercles on the ventral area, slightly below the middle, and the
impressed zone on the upper surface, indicating affinity with N.
stantoni and " IV. ? " cf. stevensoni (Whitfield) in Hyatt. The costation
is very irregular ; on the small septate portion, the costae have a very
steep forward edge and a gentle slope backward, and are continuous
between the slight tubercles. On the under-side they are projected
strongly forward, towards the umbilicus, as in the basal view of
Didymoceras? stert'itx<>n!, Whitfield sp.|j On the upper surface they
describe first a backward curve, and then, in the contact furrow,

* " Note on a Very Fine Example of Helicoceras stevensoni, etc.," ' Bull. Am.
Mus. Nat. Hist.,' vol. xiv (1901), p. 219, pis. xxix and xxx.

f Loc. cit., 1S76, p. 484, pi. xxi, fig. 3 (perhaps a fragment of a Didymoceras ?).

j Hyatt (loc. cit., p. 577) calls Hamites fremonti, Marcou (holotype in B.M.,
Geol. Soc. Coll., No. 12667), probably a gerontic stage of some species of
Exiteloceras, which is doubtful, the former probably being of Albian age, and
close to "Anisoceras alternatum," Pict. & Camp, non Mantell (loc. cit., 1861,
pi.