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ANNALS

OF THE

SOUTH AFRICAN MUSEUM

VOLUME XII

ANNALS

OF 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. Jonn Xavier Merriman, P.C.

Sir Toomas Morr, Kt., M.A., F.R.S., F.R.S.E., C.M.G., LL.D.

The Hon. Jonn WiitiamM JacaGer, 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.

KepreL Harcourt BARNARD, M.A., F.L.S., F.R.S.S8.Afr., Assistant Director, in

Charge of Fish and Marine Invertebrate Collections.

Srar GARABEDIAN, B.A., Assistant in Charge of the Herbarium.

REGINALD FrepERICK LAWRENCE, B.A., Assistant in Charge of Arthropoda (Insects

and Crustacea excluded).

ALBERT JOHN Hesse, B.Sc., Ph.D., Assistant in Charge of Insects.

Artuur Lewis Hatt, M.A., Hon. Keeper of the Geological and Mineralogical

Collections. ,

Sipney Henry Haveuton, B.A., D.Se., 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. : ; ]

On a new South African Stegocephalian (Phrynosuchus whaitsi) . é 6

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

Man contemporaneous with Extinct Animals in South Africa 0 2 dle}

R. Broom and §. H. HauGuton.

On the Skeleton of a new Pareiasaurian (Pareiasuchus peringueyi g. et sp.

nov.) ; : é ; ; F ; : j 17

On a new species of Se patiopnathig (S. tigriceps) : : ; = | 26

On two new species of Dicynodon : : . : ; : . 36

Some new species of Anomodontia (Reptilia) ; : : F oS)

FF. CHAPMAN.

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

Buffalo River, Cape Province. , : 5 : ; 7 OF

S. H. Havucuron.

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

On a new species of Propappus : : . 43

Investigations in South African Fossil Reptiles id PRINTS

1. On a new species of T'rematosaurus (T. sobeyi) : 5 . 47

2. On a new Dinocephalian from the Gouph : : : . 62

3. On two new Therocephalians from the Gouph . 55

4. On some new Anomodonts . , ; 58

5. On the genus Rhinesuchus, Broom, with Notes on Ne eecnbea

Species 5 2. é : . 3 : : OO

6. On a new type of Dinpoophalian . 5 : ; 3 a Ue}

7. On some new Gorgonopsians . : : : : : eZ:

8. On a Skull of the genus Kannemeyeria . co OI

9. A new Thecodont from the Stormberg Beds (Sanenonuenis weutue

g. et sp. nov.) . : : 5 Be

10. Descriptive Catalogue of the comodontia! w ith Sec reference

to the examples in the South African Museum. Part 1. o> ALTKS)

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

case in certain species 3 175

12. On some Gorgonopsian Skulls in ihe Gollestion of the isonth

African Museum : : : : . 499

The Fauna and Stratigraphy of the Stormbare Series ; : : oS

L. I. Spatn.

On Cretaceous Cephalopoda from Zululand - ; < : 2 2G

LIST OF NEW GENERA PROPOSED IN THIS

VOLUME.

PAGE

Aelurognathus (gen, nov.) Gorgonopsia (Reptilia) Haveuton 503

Akidnognathus (gen. nov.) Therocephalia (Reptilia) 0 192

Archaegryllodes (gen. nov.) Gryllidae (Orthoptera-Insecta) 0 336

Caruichthys (gen. nov.) Platysomatidae (Pisces) Broom 4

Chelyrhynchus (gen. nov.) Anomudontia (Reptilia) Havcuton 156

Diaziceras (gen. nov.) Prionotropidae (Cephalopoda) SpatH 242

Galesuchus (gen. nov.) Gorgonopsia (Reptilia) Havucuton 82

Gorgonognathus (gen. noy.) Gorgonopsia (Reptilia) 3 84

Lycorhinus (gen. nov.) Cynodontia (Reptilia) x 343

Macroscelesaurus (gen. nov.) Therocephalia (Reptilia) 0 175

Melanorosaurus (gen. nov.) Plateosauridae (Reptilia) . 428

Moschosaurus (gen. nov.) Dinocephalia (Reptilia) 3 78

Myosaurus (gen. nov.) Anomodontia (Reptilia) 50 164

Pareiasuchus (gen. nov.) Cotylosauria (Reptilia) Broom and 17

HAUGHTON

Phrynosuchus (gen. nov.) Stegocephalia (Amphibia) Broom 6

Prolystrosaurus (gen. nov.) Anomodontia (Reptilia) Haveuton 167

Pseudophacoceras (gen. nov.) Dipoloceratidae (Cephalopoda) SpaTH 281

Pseudoschloenbachia (gen. nov.) Prionotropidae (Cephalopoda) DS 236

Scylacops (gen. nov.) Gorgonopsia (Reptilia) Broom 8

Spheniscoceras (gen. nov.) Prionotropidae (Cephalopoda) Crick MSS. 242

SPaTtH

Sphenosuchus (gen. nov.) Pseudosuchia (Reptilia) Havucuton 98

Striatotegmen (gen. nov.) Mesoblattinidae (Orthoptera- - 331

Insecta)

Struthiocephalus (gen. nov.) Dinocephalia (Reptilia) 35 52

Subschloenbachia (gen. nov.) Dipoloceratidae (Cephalopoda) Sparr 284

Sycosaurus (gen. nov.) Gorgonopsia (Reptilia) Haventon 509

Trochosaurus (gen. noy.) Therocephalia (Reptilia) 3 55

Whaitsia (gen. nov.) Therocephalia (Reptilia) A 184

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.

vii

4 9),

ERRATA.

(Part 8.)

Page 348, line 33, for “‘ from” read “ form.”

== >

» 355, ,, 6, ,, ‘ Ornitosuchus ” read “ Ornithosuchus.”

» 355, ,, 21, ,, ‘““Spenosuchus” ,, “‘Sphenosuchus.”

ODIs sos ss, GOTACOId.. s coracoid.”

» 357, ., 38, ,, ‘““Spenosuchus” ,, “‘ Sphenosuchus.”’

SD S3 esl Osment broene » - Broom.”

ty o9 BIB ay nollie Pm DUDIC:

55) (8983. 65 09s 4, ton ag ero

ee 4 OOM ee WI bb ate

,. 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.”

ee 4300) 240) ee second sO eadas lsc:

,. 444, last line, for “* place” read “ plane.”

,, 457, last word, for “ microline ”

» 464, line 5, for “it,” read “if.”

» 464, ,, 25, ,, “tick” read “ thick.”

» 468, ,, 24, ,, ‘‘ pruvial” read “ pluvial.”

» 475, ,, 13, ,, “‘transportion ”’ read “ transportation.”

ay LD, Pa Bh a Sake ea cake

read ‘“* microcline.”

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 acquairited 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. Havaeuron, 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 Museum.

(With 8 Text-figures. )

vill

Ib Jet (Qin) dedlbpsed ots.

Scymnognathus serratidens, Haughton.

Haplophragmium neocomianum, Chapman.

Spiroplecta deflata, Chapman.

PLATE

I if Caruichthys ornatus, Broom.

: Atherstonia cairncrossi, Broom.

J Palaeoniscus capensis, Broom.

ins : ror

Elonichthys whaitsi, Broom.

NE Pareiasuchus peringueyi, Broom and Haughton.

IV. Jf Pareiasuchus peringueyi, Broom and Haughton.

‘L Propappus parvus, Haughton.

V. Pareiasuchus peringueyi, Broom and Haughton.

VL { Scymnognathus tigriceps, Broom and Haughton.

Seylacops capensis, Broom.

VII Dicynodon alticeps, Broom and Haughton.

, { Dicynodon testudirostris, Broom and Haughton.

VII. Trematosaurus sobeyi, Haughton.

IX. Trematosaurus sobeyi, Haughton.

X. Struthiocephalus whaitsi, Haughton.

Dicynodon mustoi, Haughton.

XI. Dicynodon breviceps, Haughton.

Lystrosaurus oviceps, Haughton. .

= Rhinesuchus senekalensis (v. Hoepen).

XII { : ae

Rhinesuchus whaitsi, Broom.

XTIT { Gorgonognathus longifrons, Haughton.

Spiroplecta anceps (Reuss).

XIV. Spiroplecta anceps (Reuss) var. infracta, Chapman.

Nodosaria zippei, Reuss.

Nodosaria sulecata, Nilsson.

|. Cristellaria parallela, Reuss.

Cristellaria intermedia, Reuss.

Cristellaria secans, Reuss.

Truncatulina schloenbachi (Reuss).

Discorbina pileolus (d’Orbigny).

Anomalina ammonoides (Reuss).

Pulvinulina karsteni (Reuss).

Truncatulina ungeriana (d’Orbigny).

Bairdia subdeltoidea (Munster) var.

Cythere postcultrata, Chapman.

Bairdia africana, Chapman.

List of Plates.

PLATE

XVI. Dicynodon grandis, Haughton.

XVII. Eocyclops longus, Broom.

XVIII. Prolystrosaurus natalensis, Haughton.

XIX Diaziceras tissotiaeforme, Spath.

Parapuzosia sp. nov. ? ind.

xx , Pseudoschloenbachia umbulazi (Baily).

Pseudoschloenbachia umbulazi (Baily) var. acuta Spath.

lL Mortoniceras soutoni (Baily).

XXL Mortoniceras woodsi, Spath.

\. Placenticeras subkaffrarium, Spath.

Parapachydiscus sp. aff. colligatus (Binkhorst).

XXII. Nostoceras ? natalense, Spath.

Nostoceras ? subangulatum, Spath.

Peroniceras cf. ezornigi (Redtenbacher).

XXTIL Mortoniceras stangeri (Baily).

Mortoniceras vanuxemi (Morton).

Diplomoceras ? indicum (Forbes).

( Parapachydiscus cf. wittekindi (Schluter).

Bostrychoceras ? sp. ind.

Parapuzosia sp. ind.

Baculites cf. aspero-anceps, Lasswitz.

XXIV. Baculites cf. brevicosta, Schluter.

| Baculites capensis, Woods.

Kossmaticeras (Madrasites) bhavani (Stoliczka).

Subschloenbachia bispinosa, Spath.

Subschloenbachia prerostrata, Spath.

Pseudophacoceras manuanense, Spath.

XXV. { Dipoloceras cristatum (Deluc).

Dipoloceras quadratum, Spath.

Subschloenbachia cf. trinodosa (Bose).

Cheloniceras gottschei (Kilian).

Cheloniceras delagoense (Krenkel).

Cheloniceras hambrovii (Forbes).

c Aconeceras nisoides (Sarasin).

Dipoloceras sp. nov. ?

Dipoloceras cristatum (Deluc).

Anisoceras sp. ind.

XXVI

Acanthoceras

Aconeceras

Adolphia .

Aelurognathus .

Aetonyx :

Akidnognathus .

Alopecognathus.

Anisoceras

Anomalina

Archaegryllodes

Arctognathus

Aristosaurus

Atherstonia

Baculites .

Bairdia

Barroisiceras

Bostrychoceras .

Bubalus

Caruichthys

Cheloniceras

Chelyrhynchus .

Cobus

Connochaetes

Cristellaria

Cymatoceras

Cynosuchus

Cythere

Cytherina .

Cyzicus

INDEX OF

180, 210

256, 2

108,

264, 290, 301

114,

all

GENERA.

Diaziceras.

Dictyopyge

Dicynodon

Diictodon .

Diplomoceras

Dipoloceras

Discorbina

Douvilleiceras

Dromicosaurus .

Elonichthys

Emydops .

Kocyclops.

Equus :

Erythrochampsa

Estheria

Eucnemesaurus .

Eulophoceras

Kuskelesaurus

Eutrephoceras

(ep)

Galesuchus

Gauthiericeras

Geranosaurus

Gigantoscelus

Gorgonognathus

Gryponyx.

Gyposaurus

Hamites

Haplophragmium

Hauericeras

Helichthys

Heteroceras

Hippopotamus .

Holcodiscus

Hortalotarsus

Kannemeyeria .

Kossmaticeras .

Lenticeras

Lepidurus.

Lycorhinus

Lystrosaurus

Macroscelesaurus

Madrasites

Massospondylus.

Melanorosaurus .

Miliolina .

Mortoniceras

Moschosaurus

Muniericeras

Myosaurus

Nautilus

Nodosaria.

Nostoceras

Notochampsa

Oppelia

Index of Genera.

PAGE

256,

370,

91,

264, 290,

288

109

237

340)

308

299

379

299

244

328

345

dll

PAGE

Pachydiscus 226

Pachygenelus 340

Palaeoniscus i

Paralenticeras 244

Parapachydiscus 226

Parapuzosia 224

Pareiasuchus lg

Pedeticosaurus . 366

Peroniceras 295

Phacochoerus 14

Phrynosuchus 6

Phthartus. 331

Phylloceras . 273

Placenticeras 247, 300

Plateosaurus 406

Prohauericeras . 238

Prolystrosaurus. 167

Propappus 43

Pseudophacoceras 281

Pseudoschloenbachia . . 236

Pulvinulina 108, 114

Puzosia 2o1, 204.

R \

Rhinesuchus ; 65

Rosalina 113, 114

Rotalia 114

Rotalina 113

Schloenbachia 240, 281, 285, 295

Scymnognathus. 26, 88, 205, 503

Scylacops . . 8, 210, 501

Semionotus 338

Spheniscoceras . . 242

Sphenosuchus 98, 344

Spiroplecta 109

Stoliczkaia 286

Striatotegmen 331

Struthiocephalus 52

Subschloenbachia 284

Sycosaurus 509

Tapinoce phalus

Taurotragus

Textularia

Thecodontosaurus

Titanosuchus

Torneutoceras

Trematosaurus .

Tritheledon

Trochosaurus

Truncatulina

Turrilites .

Index of Genera.

PAGE

108, 109

370

288

Uhligella .

Valeulina .

Whaitsia

xill

PAGE

275

112

184

ANNALS

OF THE

SOUTH AFRICAN MUSEUM.

(Vou. XII.)

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

R. Broom, M.D., D.Se.

WirH 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.

PALAIONISCIDAE.

PALHONISCUS CAPENSIS, Sp. Noy.

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.

Annals of the South African Museum.

The dorsal fin is placed more anteriorly than in Pal@oniscus

macropomus, 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 and 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

fulera.

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, Fraserburg 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 CAIRNCROSSI, Sp. Nov.

This new species of Atherstonia is founded on the greater part of

a fish found by Mr. J. L. Cairneross at Coleskop in 1906, and

presented by him to the South African Museum. It was regarded

at the time as a small specimen of Atherstoma scutata, 8.-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 57mm. In other

words, the depth of the body in the new species is half the length

without the head and tail; in A. scwtata 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. scuwtata, and

the fulera 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.

Famity PLATYSOMATIDAK.

CARUICHTHYS ORNATUS, g. et sp. n.

The type of this species is a single specimen found at Doorn

River, 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 Hurynotus, but not so deep as in

Platysomus. 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 Hury-

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

fulera. 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 Hurynotus. 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 Stegocephalian (Phrynosuchus whaitsi).

—By R. Broom, M.D., D.Se.

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 squamosal 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 New 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.

Fic. 1.—Skull of Phrynosuchus whaitst. 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 Rev. J. H. Whaits, has

been placed in the South African Museum.

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

—By R. Broom, M.D., D.Sc.

Tue skull here described was found by Mr, 8. 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. Rogers, 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 torvus, 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 Hndothiodon zone, not more than a few hundred feet

above the Pareiasaurus zone. Another Gorgonopsid occurs in the

same zone, viz. Scynunognathus whaitst 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 6. These molars are only about 4 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

process 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 Cistecephalus, 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 South 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 1s 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 4 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 squamosals and

inferiorly 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 les 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 Neariy Perfect Skull of a New 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 Cynognathus, 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, XIth,

and XIIth 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

Scymnognathus 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 Scynunognathus. 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 Therocephahan

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-

12 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 :—

PELYCOSAURIA

Baplosaure,

Aneel

‘. Mammata =

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

sp. noy.

(13 )

4.—Man contemporaneous with Extinct Animals in South Africa.—

By RB. 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 Hquus 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 baint 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 haying 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 Hquus—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, ete. I do not think the Hippopotamus

differs from the present-day species, and some of the specimens

must have been of great size.

Fic. 2.—Connochaetes antiquus. Broom. x 4

Phacochoerus aethiopicus—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. antiquus 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

Fic. 3.—Cobus venterae. Broom. x 4.

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 24 inches in greatest width, and

about 2 inch thick at its thickest part. Im 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, ete.—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

péringueyi, g. et sp. nov.) By R. Broom, M.D., D.Sc., and

S. H. Havauton, B.A.

THE almost complete skeleton described in this paper was found

early in the year 1912 at the drift across the Zak River on the

main Beaufort road on the farm Dunedin, in the division of Beaufort

West. The skeleton was found in sitéi 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 11th 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 Parevasaurus 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

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 the 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 Parerasaurus

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 Cvstecephalus 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 Pareiasuchus, which is also

probably as high as the Cistecephalus 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 New 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 Procolophon, Dvadectes,

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 Procolophon, 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

Pareiasaurus 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 basicranium 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 Musewm.

Remains 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 11th 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, beimg 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 were certainly small.

The 8rd 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 mm. ;

3rd, 62 mm.; 4th, 63 mm.; 5th, 74 mm.; 6th, 94 mm.; 7th,

102 mm.; 8th, 120 mm. The Sth 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 Pareiasaurus 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 illum 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 lst is 110 mm. high, the 11th 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 rogersi,

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

rogerst. 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 elenoid=s1s-....0.nsea2ccesametee sn nee seeemeee S25;

Height of scapula above acromion ..............0000...«ese-sers Piatt OA

Width of scapula immediately above acromion ............ U2) 35

Thickness of scapula immediately above acromion ......... 319) 35

Width across top of scapula... .c5-ccse-seseee ee acencecs ese 124 ,,

Weng thOl aCrOmmOmpe ceases neces -mastsns tere anc ceee cet ane eee 13 45

Maximum heightiol@cromion o2..s-0 cc. ye sae eeneseaneenae O25;

Distance between anterior and posterior borders of cora-

GoldvandeprecOracold tara ea osoee ness lauren ee aereee ee OM

The clavicles and interclavicle are almost complete. As in

Pareiasaurus 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 cleithrum.

HuMERUwvs.

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

45mm. xX 40mm. 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 rogerst. 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.

RapDius 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-

pled 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 rogers: 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 rogerst. 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 rogers?.

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

péringueyr. omocratus. rogerst.

Bengt of femiur .-soscs00ces caries 320 mm. 335 mm. 290 mm.

WWatctin- of head: .cahes-secosaaseeeos oan (foe Fe CO> 35

emetn of Head: .... ss. ceaectoosase 1OGoe i Laney jee

Widthof shaft at narrowest part 70 ,, GOT: 60 ..;

Width of distal endof bone ... 152 ,, TS: 14T 5;

The chief differences, therefore, le 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.

6.—On a New Species of Seymnognathus (S._ tigriceps).—By

R. Broom, M.D., D.Se., and 8. H. Hauaeuron, B.A.

THE magnificent specimen which forms the type of this new species

was discovered about 5 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 interclavicle. 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 Cuistecephalus 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, Gorgonops 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 Seymnognathus (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 Hndothiodon zone, and with it

is a larger allied form, Scymnognathus whaitst. In the Parecvasaurus

zone a small Gorgonopsian has been found by the Rey. 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 Hndothiodon 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 Scylacops, 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 Scymnognathus whaitsi, 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 intertemporal 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 whaitse. 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 1 5, ¢ 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

4 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 South 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 péringueyt 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 postfrontal 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. Whetherit 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 asmall 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 Seymnognathus (8. 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. Hach

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 broad.

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 Anomodonts 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. Hach 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 Seymnognathus (S. tigriceps). 31

girdle resembles the Therocephalian type as seen in Ictidosuchus

primaevus.

The scapula has a broad 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 he a

straight one, and the precoracoid appears to form no part of the

glenoid cavity.

The chief measurements are as follows :—

Greatest lenebhy ol Scapula,.......sicee nice vsssereccseinesestnadearees 130 mm.

Vici wots bsisevon SCAU AL. 205.55 sn0asi.ssivone-nnat'sasnien ene neeeee SOR;

NGM OL TOprOtasCePUll di... .hiecc.aiscisisccis sec «cis are seaccamemmemener 49 ,,

Gren be st widthvor OlOMO1d © <2... ...0.sics scisicacnaede arise anion emaranmetnerer A ae

Greatest hechtivat Glenoid: i: 260% descssuatcsaeve coeeeuecocesaten Aor i,,

Length of precoracoid-—coracoidal suture .................006: Ole

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 Musewm.

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 humerus agrees with that figured by Owen

as “Oynodrakon 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 Species of Seymnognathus (8. tigriceps). 33

RapDiIus AND ULNA.

The bones of both sides are present, those of the left being found

wm sitt 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. Proximally, 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 thé 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

34 Annals of the South African Museum.

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

outer centrale.

The imtermedium 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 lst 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

dth 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 sesamoid or prepollex.

The 2nd distal carpal is a small cubical bone, much thicker than

the Ist, and it gives articulation to the 2nd metacarpal.

The 38rd 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 dth 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 Seymnognathus (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 4 of its width. Though so very short it manifestly has

hinged freely on both the Ist 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 4 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 38rd 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 Anomodonts 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 Species of Dicynodon.—By R. Broom, M.D., D.Sc.,

and S. H. Haveuron, B.A.

DiIcYNODON TESTUDIROSTRIS, Sp. Nov.

THE type of this new species (No. 2354 8.A.M.) was collected at

Dunedin, Beaufort West, just south of the Homestead, and is prob-

ably from the Crstecephalus 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 56mm. 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 caniniform 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 Two 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 Cvstecephalus, 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 quadrato-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. Novy.

The skull on which the following description is based was collected

about one mile west of Oudeberg, in the division of Graaff Reinet, on

the main road cutting between Murraysburg and Graaff Reinet.

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 Dieynodon 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. leoniceps.

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 septomaxillary 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 mm.

The squamosal is a large bone, not wholly perfect, in which the

curve between the zygomatic 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 :—

GreatestMen ety wicca, scacss- asewascsscoreiodeecssslsecss 242 mm.

Greatest breddtlivnasc sce dcont-scansseeeeencene probably 170 _,,

Snow totront: Om oOnbiimesearenetaess ictus tes ace esos Olnen

MemethcOlOnbitannaestthen errata sce stsetese estas case AS:

Minimum width between orbits .................666. 42 ,,

Minimum width of temporal roof .................. IISE ss,

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

Maxima, herein oiselomliie tcc tceseceetc uses sete a+s< lb] eee

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.

Havucuton, B.A.

Recentiy the South African Museum was presented with a skull,

without the lower jaw, by Mr. P. Le Roux, 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.

Fic. 4.—Tapinocephalus atherstoni, Owen.

Top view of skull. x 4

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 435mm. 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.

Fic. 5.—Tapinocephalus atherstoni, Owen.

Side view of skull. x 4

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

49 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.

Gengthyof gwouliys.. acceccesee we sen eeee cose mcmre tetra 220 ,,

Distance from quadrate to front of snout ......... 290 3

Breadth of palate at posterior end of prevomers 280 _,,

Length of prevomers ss c.-c:s0oasescciqeenensoeavecereccle 145 ,,

( 43 )

9.—On a New Species of Propappus.—By 8. H. Hauauron, 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 ilum. 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

Parevasurus 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 Musewn.

lower half of the posterior border of the ischium 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.

PuatE I.

Fic.

1. Caruichthys ornatus, Broom. x 4.

2. Atherstonia cairncrossi, Broom. x $.

Puate II,

1. Palaeoniscus capensis, Broom. x 3.

92 Sue Ge

bo e

Sh So

wm 09 bo

. Pareiasuchus péringueyt. Outer view of left shoulder blade.

. Elonichthys whaitsi, Broom. x $.

Puate III.

. Pareiasuchus péringueyi, Broom and Haughton. Posterior view of the left

shoulder blade. x 4.

x ¢.

Anterior view of right humerus. x ¢.

Posterior view of right humerus. x 4.

a + Anterior view of right radius. x 4.

Anterior view of proximal half of right ulna. x 4.

Puate LY.

. Pareiasuchus péringueyi, Broom and Haughton. Posterior view of left

femur. x 4.

. Pareiasuchus péringueyi. Anterior view of left femur. x 1.

” 9

Outer view of left side of pelvis with sacral rib. x 4-

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

% 9

. Propappus parvus, Haughton. Outer view of right side of pelvis. x 4.

Puate V.

Pareiasuchus péringueyi, Broom and Haughton. Mounted skeleton. x 4.

Piatt VI.

. Scymnognathus tigriceps, Broom and Haughton. Side view of skull. x 4.

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

and interclavicle. x $ nearly.

Right side of shoulder girdle. x 4 nearly.

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

2? ”

9 99

9 FS)

. Scylacops capensis, Broom. Top view of skull. x 4.

PuateE VII.

. Dicynodon alticeps, Broom and Haughton. Side view of skull. x 4% nearly.

5 Top view of skull. x 4.

: Dieunaden testudirostris, Broom and Haughton. Top view of skull. x 3.

Side view of skull. x 3.

9 9

Annals S. Afr. Mus. Vol. XII.

Plate I.

CARUICHTHYS

ATHERSTONIA

ORNATUS, Broom.

CAIRNCROSSI, Broom.

‘

hnals §. Afr. Mus. Vol. XII. Plate IT.

PALAIONISCUS CAPENSIS, Broom.

ELONICHTHYS WHAITSI, Broom.

Annals S. Afr. Mus. Vol. XII. Plate IIT.

PAREIASUCHUS PERINGUEYI, Broom & Havcuron.

Annals S. Afr. Mus. Vol. XII.

PAREIASUCHUS PERINGUEYI, Broom & Hauauron.

PROPAPPUS PARVUS, Havanton.

Plate V.

Annals S. Afr. Vol XII.

‘NOLHOOVA, VY Noouqd “TATQONIVAd

SQHOASPIAUFd

nals 8. Afr. Mus. Vol. XII. ae mem eons Plate VI.

SCYMNOGNATHUS TIGRICEPS, Broom & HauGcuton.

SCYLACOPS CAPENSIS, Broom.

Annals S. Afr. Mus. Vol. XII. Plate VII.

DICYNODON ALTICEPS, Broom & Havcuron.

DICYNODON TESTUDIROSTRIS, Broom & Havuauron.

(2%)

10.—Inwvestigations in South African Fossil Reptiles and Amphibia

(Parts 1-4).—By S. H. Haueurton, B.A., F.G.S., Asszstant

Director.

1. On a New Species of Trematosaurus (T’. sobeyt).

(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 ...........0..scce»-ases 517 mm.

ENG ALCSUROROAOGE! auemeniomen es sisiciaie ae Sawiieits steeeee ness 305m.

Tip of snout to occipital plate .................s26.0 ANN

Snout to plane of front of nostril..................068 65 _,,

48 Annals of the South African Museum.

Tengthiob nostril y ccgeacscossmsnsasaccoe seiiseeetreer 27 mm.

Internasal breadthywceoncm acces sseeueteeeetce eae Sl,

Preorbitalwlength) wasc-ccerec scoot +s sce tesereseniess PA es

Thength of Orbit) tec.cac-nacosneestees wreessiseedneuisere 24°5 ,,

Imterorbital widthie-eeessteeseee cc eects ce nese eeeee S00,

Length iobtirontal erences en ieesecancscesoe asker. 1GO

Lengthroimasaliay ween cceeacete cece ccennel anna 149 ,,

lhengthvol parietal (ac sascsateemee en cnteseoee eee ate Hale ee

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 Hryops, and it is known to occur in a number of small Stego-

cephalians. Unfortunately the type of Trematosaurus kannemeyert

does not show this region. If an interfrontal did occur in that

South African Fossil Reptiles and Anvphibia. 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 frontals

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 postorbital.

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 7’. kannemeyerr

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 Musewm.

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

Fic. 6.—Trematosaurus sobeyi, Haughton. Diagram of section of skull just

anterior to the orbits. x4.

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 mm.

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 New Dinocephalian from the Gowph.

(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 Rey. 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 :—

GreatostMen gb! eet aa-cncctesodscwssmec aeeemenee eee 580 mm.

Grentestiorentnemeeeecceer anda reer cereene ates 355 Ca,

Greatest helehipecrcnc: seser se toni: clsorsicweseaeeeees 230 ,,

Length of snout to back of internal nares......... 20000,

Breadth of snout at back of internal nares ...... 230s

Tene tiniol Orbitie cesarean ecco ether tcc siete 39153,

eich Gof Orbitinc. reese centre aatae nee sae COR:

Iheneth: of temporalfossar-.sns<2-05- 52 ease ayaa: SOs %;

Heightot, temporal tossaesccceste eet es eee nce 148 -,,

Interorbital width at front of orbit.................. dhol 0 Mies

Interorbital width at back of orbit .................. 295 ,,

Intertemporal width ............ UO ytd ak Rectory ee tee 130 ;;

Basal Tength. ia. 0scacac sets. aire aa aee neces eene ss ose Ola) 5

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

defined, 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 Therocephalians 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 Parezasaurus

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 frontals 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 Musewm.

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 mentum. 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 :—

Greutestenethyncucessdsee thoes ence t oe ceinnes secs 230 mm.

Distance from. snout to front of orbit............... eee

DiameterioMorbitwestnc sce. esse -eeecee cee rcenee eaccee 42

ingerorbitaliewidbhimas. dees ssc ececaescesee a tenes 46 _,,

Intertemporal width ?pss.:.-n.cssno-5-<. noes creeere 2305

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).

South African Fossil Reptiles and Amphibia. 57

TITANOSUCHUS 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 YT. cloetez 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.

HGS WMACISOL «os sccsienay scces se Imperfect 21 x 14 Imperfect

Interval between 7.1 and 7.2 10 10 9

HOCONG INCIGOL c:.c.scceceeess 21x lo 18 x 13:5 19 x 14:5

Interval between 7.2 and 7.3 10 151 4

PINT TA CIS OI cst cecee Se ce ve 20 x 12 20 x 14 20 x 15

Interval between 7.3 and 72.4 18 8 6

HMOUEGH INGISOM «<<< ¢siees =i 0% >< 33x17 18 x 14 16°5 x 13

Interval between 7.4 and ¢ ? i] 3)

GRANGE. 1k shoe seneeihasesaswee 50 x 35 30 x 26 47 x 20

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 TZ’. cloetei. The canine is narrower and

longer. In fact, whereas in 7’. cloetet 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 7’. cloetez the figures for

the same measurements are 67 mm. and 116 mm. respectively.

(58 )

4, On Some New Anomodonts.

(Plate XI.)

DICYNODON MUSTONIS, sp. 0.

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

Duictodon Kolbei, 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, 7.e. just below the

front of the orbit.

The following are the chief measurements of the skull :—

GE Remo Sule Mt Meas os tei nds are net wasincle te maswrlayienisasiaee 150 mm.

INA eare TATU GL bly eevee ala aiscctoa ernachswicien wee occ de Admob alone aOR

Maximum width of parietal region..............+... 24 ,,

Maximum width of frontal region ................0 195

Maximum width of nasal overhang.................. 29.

Minimum width across pterygoids .................. Loy.

Ibength of pineal foramen... g.cnsecccesrenc.oaeneecs. 6. |

Whidthvof pineal foramen yl 0.n.2:5.2m0sesinesiseos os Oe

Distance of pineal foramen behind postorbital

[CCAP EAN Reet hee) Giese ae era ae ae 10

Bacall etl se-wsase haceeacteeenmiatdeniasaerrecionee. IPAS es

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

Maximum depth of lower Jaw ..............seeceseees Dons,

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.)

DICYNODON BREVICEPS, sp. 0.

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 Murraysburg, 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 mm. 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 mentum descending

below the level of the bottom of the jaw.

LyYSTROSAURUS 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 Lystrosawrus 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 River, 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 ZL. 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 mm. 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. latirosiris, 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 pterygoid is of the usual

Lystrosaurus type, having a long articulation with 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 intertemporal widths, in the shape of

the orbit, and in other minor features. When viewed posteriorly

the skull is seen to differ from both LZ. declivis 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.

FIG.

1. Top view of skull. x 4 nearly.

2. Diagram showing arrangement of bones of top of skull. x 4 nearly.

PLATE IX.

2. TREMATOSAURUS SOBEYI, Sp. Nn.

1. Palatal view of skull. x 4 nearly.

2. Diagram showing arrangement of bones and teeth of palate. x 4 nearly.

PLATE X.

STRUTHIOCEPHALUS WHAITSI, g. et sp. n.

1. Side view of skull. x +4.

2. Palatal view of skull. x ¢.

3. Top view of skull. x +.

PLATE XI.

1. Top view of skull of Dicynodon mustonis, sp. n. x $.

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

breviceps, sp.n. xX 1.

. Top view of skull of Lystrosaurus oviceps, sp.n. X 5.

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

se)

EXPLANATION OF LETTERING.

F., 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.

no. S. Afr. Mus. Vol. XII. ea WL

TREMATOSAURUS SOBHYI, Havucuton. xX 4 NEARLY.

mmc. Afr. Mus. Vol. X11 Ree,

SASS x \

\ \ \ *

TREMATOSAURUS SOBEYI, Havucuron. xX 4, NEARLY.

Plate X.

ol. XIN.

mec, Air. Mus. V

HAUGHTON.

WHAITSI,

PHALUS

STRUTHIOCE

Ann. S. Afr. Mus. Vol. XII. Plate XI

1. DIGYNODON MUSTONIS, Havcuron. xX 4.

2. DICYNODON BREVICEPS, HavGHton. x le

3,4. LYSTROSAURUS OVICEPS, HatGHTON. X 3.

( 65 )

11.—Investigations in South African Fossil Reptilia and Amphibia

(Parts 5-9)—By 8. H. Haveuton, B.A., F.G.S., Assistant

Director.

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

Species.

THE genus Rhinesuchus 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 Ff. whaitsi was given. The genus was said then

to include also the species named by Lydekker Evyops africanus and

the European species called Macromerion Giimbeli by von Ammon,

At first the genus was thought to have no affinities with Hryops, 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 Fhinesuchus 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 Rhinesuchus as given by

Broom is as follows :—

“Skull probably like that of Capitosawrus 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

66 Annals of the South African Musewm.

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 Rhinesuchus

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 &. whaitst 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—Lhinesuchus, Broom—for which the following

short diagnosis may stand :—

‘‘Medium-sized to large temnospondylous 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

transpalatine with teeth present. Coronoid carries number of

denticles on upper surface.”’

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

Investigations im South African Fossil Reptilia and Amphibia. 67

in being more pointed, the skull of Rhinesuchus differs from that of

the American genus Hryops 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.

RHINESUCHUS WHAITSI, Broom.

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

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

An almost complete skull and lower jaw (S.A.M. Cat. No. 3009)

collected by the Rev. J. H. Whaits at Beaufort West (Hndothiodon

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 Hryops

megacephalus and Ehinesuchus senekalensis. 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 EHryops. The articular region extends well behind

the occiput. The following are some of the principal measurements

of the skull :—

AVOUT STA Mp estore sio.caeaiitewe's va eeeriseme saw veuiee 312 mm.

Heisman OMPMG GIA LING ee 5.00... eocecss ev ncesien s omer ns 265 ,,

MMieixtraa MMA WEN eee etc 4 s'ce's cceciea occsinetsisivcwss 258: ;;

PE MOR OMB NVC DU cee ee ow cee, cess scene avccesceescorcle 40 ,,

AT DOTENAG al WUE espe ee ce ccles ve assis sawlainsecdesine Anlee

Length from back of nostril to front of orbit...... 12555;

Length from plane of snout to front of orbit...... iF Sye yes

NCS MMOMOV DMP meee ct accetioss «scx \- 2sgcacle mcisteielsraee os se 33,

AV silts note OR Ditters cr ones sin) <jatee:sieis ies Ze\n'e Bisa ne 34,

Comparison with the type specimen of £2. 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 South 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 Hryops.

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 im South African Fossil Reptilia 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.

Fia. 7.

Sections across left ramus of lower jaw of Rhinesuchus whaitsi :

a, 85 mm., behind point of jaw.

3. 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-Meckelian

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 Hryops, as described by Broom and by Williston,

this ‘“‘coronoid”’ consists of three elements—the coronoid proper,

70 Annals of the South African Museum.

the intereoronoid, 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

allent 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 Hryops

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.

RHINESUCHUS SENEKALENSIS (v. Hoepen).

(Plate XII., figs. 1, 2.)

1911. Myriodon senekalensis, van Hoepen. Ann. Transvaal Mus.

i., 2, p, 103; pls. 1; 2:

1912. Rhinesuchus major, Broom. Trans. Geol. Soc. 8. Africa, xiv.,

p. 79; pl. 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 le 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 Reptilia 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 exampies come from

the same locality, I think there can be no doubt as to their specific

identity. Comparison 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 Hryops, 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

444mm. 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 well 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 Musewm.

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 Hryops. 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 Hryops. 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. Asin Hvryops, 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. Hach 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.

Memst hot areliye.sscacsets5cs eee eeieereeeer 45 mm. 52 mm.

length of dorsal spine)... 25..40<.-. seaeeea 36, Zils es

Height of spine above transverse process 31 ,, oo.

Width between extremities of processes... 88 ,, Ole

All the vertebrae carry ribs. The ribs from the 2nd to the 8th

have their distal thirds strongly expanded with a well-defined

Investigations 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 Hryops. 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 Hryops. 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, 50 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.

The distal end has a width of 24 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 Musewm.

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 Hryops. Only

four metacarpals and four digits are preserved on each side,

and the phalangeal formula as preserved is 2, 2, 3, 8 (?) or 4.

The following table gives the chief measurements :—

Length Proximal width

in mm. in mm.

difsepasateytNerhe ofd lls dhantrica acer eceescc hao. 23 15

PS tphalanx cr Boewesceha tea eee 15 14

Claw str, Rte seek Seren ee eet ala) al

iy MO CUACAI Dele. elena eee eee eee 29 14

LE SUR) O) OE Esha b: Carns oe 21 15

GH awe cat ce Mey et Soe Me ato 12 et

Srdemehacarmels ¢ 1 mew. es Re Notre -48essice 28 Lg

LG qohia lea vactes Sees aes ee awa 19 14

And phalanx asters asses Aye 10

Cla Wisi te ketenes 4 eee ee ? 2

Aten e ta Cag pall. .ja.c25e saqncs eee eee 23 15

Astuphalaax, ..ee. sesso eee 16 12

AN ogo elanx, waive. fea ee ee 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 Ophiacodon than of

Hryops 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 Reptilia 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.

lst metatarsal) “s52sscsesetoens2seveeeroaer 24-5 18

list phalanx © 25.2565 saeco. nore 15 18

Glaiweaeeewcceseeasecies merce ses reece: approx. 10 13

Drv mnetiatatsal cow tesueat-<ecoueneasarseees se 36 21

ligt balami- <dscu.cseatteneea.-keeee 26 19:5

Cla walie peccsnenesepememeeas te nteaneys approx. 11 12

Srdemotatarsal cs 4.5 scene ac eee ese ea 3¢ 19

istrphalanine Wise ecsaeeis cence 24-5 20

Ding pole ase iss Meee Oe poe eect ie 15

Claws acceapettce erate Sac eee 11 ity

Atha inG batAGAl) cecasctase te Wee creme enaua 37 19

Ist phalanx ........ shee ee nee ota 24 19

Piya bay 0) ak: NN ob. ee eer ERO eae ae 15 15

10 bg 0) 0k | 1a aa yee Soe tere aera ee 12 10

Clitwis com totect doa ereeents. Jatonate missing —

fan ales eo =1 se Vast: | Ma ge ORs ein AB oe 23 16

listo tial AM Rw fie oone vo anne ener 21 15

Didi lal lane et omeras ee ecee eee 16 12

CUAW. ects oscoretais Saskaneeiee ees 11 8

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. Hach 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 AFRICANUS (Lydekker).

1890. Eryops africanus, Lydekker. Quart. Journ. Geol. Soc., xlvi.,

p. 291; pl. xti., fig: 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 Lhinesuchus by Broom.

Investigations in South African Fossil Reptilia and Amphibia. 77

The species is imperfectly known. A crushed and incomplete skull

and lower jaw (S.A.M. Cat. No. 3010) collected by the Rev. J. H.

Whaits at Dunedin, Nieuweveld, Beaufort West (Cistecephalus zone),

probably belongs to the species. It shows a type somewhat smaller

than R. senekalensis, with a basal length of about 400 mm. The

lower jaw is more robust than that of R. 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 New 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 Parezasaurus 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 Tapinocephalus and Struthiocephalus. 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

14 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°55 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 215mm. The dentary covers almost the whole of

the anterior half of the jaw, the lower border being formed by the

Fic. 8.—Anterior view of vertebra of Moschosaurus 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 Reptilia 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 Ist, 3rd, and 5th of the series :—

Ist Vertebra. 3rd Vertebra. 5th Vertebra.

Mena ihiol Gembeunts..... ce -.02--2- 02° 30 mm. 33 mm. 32 mm.

Eleight of CentrwiMe is5..5.J62n 0202-027. OASh. ae Siliees,, Sills

Mo talline tg liieemaese saree sc csecea sa. isis ep jae Ae

Width across transverse processes 70 ,, GSe.. eee

Height of spine ............. SNe eee oO 5; See GS on

engi) Of Spine 2.52; sacaanecscncss-* Sng? Ree DIB og 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

croup that I propose to erect for it a new genus, and to name it

Moschosaurus longiceps, g. et sp. nov.

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

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

Horizon. ower Beaufort Beds. (Upper part of Pareiasaurus

zone.)

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 Paretasaurus 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 Hndothiodon and Cistecephalus zones. The antero-posterior

diameter of the orbit is 32 mm., and the interorbital width 30 mm.,

being equal to the intertemporal. 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

capensis.

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.

Inwestigations in South African Fossil Reptilia 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

Fic. 10.—Galesuchus gracilis.

Sketch of top of skull showing arrangement of bones. x #.

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 Scymnognathus. The dentary is very large, and its

posterior process rather more powerful than in the other Gor-

gonopsians. The angular is large.

Type. Incomplete weathered skull. (S.A.M. Cat. No. 2754.)

Locality. Abraham’s Kraal, Prince Albert Division.

Horizon. Beaufort Beds. (Parevasawrus zone.)

2. GORGONOGNATHUS LONGIFRONS, g. et sp. nov.

(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 14 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

305mm. ‘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 im South African Fossil Reptilia 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 75 cl m4. 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 frontals 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 Dimetrodon 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-temporal fossa, and covers part of

the back of the inner ramus of the squamosal.

Thus this form, occurring as it does in the Cistecephalus 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 Scymmnog-

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 Reptilia 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. Galesuchus gracilis

from the Pareiasaurus zone, a snout of Gorgonops sp. (S.A.M. Cat.

No. 3038) from the Hndothiodon zone of Beaufort West, and

Scymnognathus serratidens 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 2 3

Ere. 11.

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

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

3. Cross-section through vomer of Scymnognathus serratidens 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 110mm. In

88 Annals of the South 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. 14 miles E. of Homestead, Dunedin, Nieuweyeld,

Beaufort West.

Horizon. Beaufort Beds. (Cistecephalus zone.)

3. SCYMNOGNATHUS SERRATIDENS, sp. Novy.

(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 caleareous 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. tigriceps, S. angusticeps, and S. nuinor 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. tigriceps,

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 20 cl m4. 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 am South African Fossil Reptilia and Amphibia. 89

its length below the edge of the bone 25mm. The following gives

the length of the incisors: 21, 19 mm.; 22,21 mm.; 73 25 mm.;

74,24 mm.; 29,20 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 postfrontal. 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. whaitsi it is very

doubtful whether it plays any part in the border at all. In

Galesuchus and Gorgonognathus 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. tugriceps.

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.

Annals of the South African Musewm.

The following are the chief measurements of the type :—

Lip ot snout to front Of Orbit” ..2.+....s2es.---: 132 mm.

Tipwei snout to back of orbit <2.....:s22s000.+6 ae ee

materorbitalgwic these... sees scence cake acae sc: 66; =;

isstertenporal’ wadth we. ss-..02 sidca ce sce ae ese one 64 =a,

Depth—snout to mentum .................008 BO! | ee

Type. Incomplete skull. (S.A.M. Cat. No. 2672.)

Locality. Dunedin, Nieuweveld, Beaufort West.

Horizon. Beaufort Beds. (Cistecephalus zone.)

8. On a Skull of the Genus Kannemeyeria.

(Text-figs. 12-14.)

The genus Kannemeyeria was founded by Seeley in 1908 (Rep.

Brit. Ass., 1908, p. 713) on a skull of a large Anomodont 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. cvt.),

Kannemeyerta 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 ot 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 the South 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

Via 12.—Top view of skull of Kannemeyeria erithrea, x + 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.

Investigations in South African Fossil Reptilia and Amphibia. 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.

Fie, 13.—Side view of skull of Kannemeyeria erithrea. x 1 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 South African Musewm.

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, 11th,

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 les

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 hes

above the pterygoid and has a rugose surface which meets the

premaxilla. 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 :—

GreahestMlemotltece.wcsdos sctek sce scevnhsseecadcoscesd 450 mm.

(Caveenisisin lds tyelil el cts cesscoacSEBeceeeRceBA EEE Berrne 330 __,,

Minimum width of nasal overhang............ Gomes

Maximum width of nasal overhang............ JAIES)

Hiaberonoitellewadtileseeecee ss aevsaccecceSee dst ces ae

Mintertomporalewiditly s.5 + sc ecsiiesecccssinnceseees By hs

SuoUt to frontOb OLbIti see.60.le2-05.ccee0e0sse0s' WSC a

ene bi OMOr Dies. sc ccastacssces dacs daceesse st yates 82,

ac alplemet in wes. saa tetas tensa cache ca scesescccss. 302 =;

Minimum width across pterygoids

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 Kannemeyeria

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 Lystrosaurus 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 which

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,

Fie, 14.—Sketch of odontoid and axis of Kannemeyeria erithrea. x 4.

Vertebrae.—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 Reptilia 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 upperedge. 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.

Sie 3 4 #6 6 % 8 18? 172

Length of centrum ............... AVieeat oS) 39) 139) 39)) 36) “soya so

Width of centrum .................. 569) 65 51) 62 60 59 61 50 45

Motaleael platters. cconeeeccooress 128 142 161 ? 195 189 170 132 143

Height of spine above transverse

TROYES adoccnonantagsqodoendoaegion0o 66 71 80 ? 109 101 78 53 58

Width across transverse process 90 97 100 100 96 97 98 83 75

Width across prezygapophyses... 55 37 42 388 43 41 42 32 39

Width across postzygapophyses 32 35 30 48 38 385 27 32 32

[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.)

(98 )

9. A New Thecodont from the Stormberg 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 Red Beds of

Paballon, Mount Fletcher. It consists of an incomplete and some-

what crushed skull, with the cervical vertebrae attached, the two

Fic. 15.—Side view of skull of Sphenosuchus acutus. 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 Huparkeria, 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 Huparkeria or Ornithosuchus. The shape of the infratemporal

fossa is characteristic in that its anterior border, formed by the

Investigations in South African Fossil Reptilia and Amphibia. 99

jugal and postorbital, passes upwards and forwards instead of

upwards and backwards as in Huparkeria 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

Fie. 16.—Top view of skull of Sphenosuchus acutus. x 3.

(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 Hrpetosuchus the premaxilla is entire, and forms

an anterior border to the external nares, and Broom has considered

that the bone in Huparkeria 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 Huparkeria. 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 75 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 postfrontal. Hven 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 Reptilia and Amphibia. 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 parietals 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 Huparkeria 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 Huparkeria. The ascending process lies behind and superiorly

interior to the postorbital and is inclined slightly forwards. 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 parallel

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 Huparkeria,

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 Ornithosuchus 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 Musewm.

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, 7.e. half the outer and posterior

borders of the supratemporal 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 Ornithosuchus 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 exoceipital 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 Proterosuchus—two forms which von Huene groups together as

the Proterosuchia, 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 South African Fossil Reptilia and Amphibia. 108

The lower jaw is incomplete. About 100 mm. of the right ramus

is anchylosed to the anterior 40 mm. 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.—Sphenosuchus acutus.

. Outer view of right clavicle.

. Inner view of interclavicle.

. Inner view of left humerus.

. Outer view of right humerus.

All figures x 3.

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

symphysis. 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

well 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

Investigations of South African Fossil Reptilia and Amphibia. 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 humeri 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 25 mm.

Type. 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. Red Beds. (Stormberg series.)

Rowe

. Palatal view of skull of Rhinesuchus whaitst. 2

. Top view of skull of Gorgonognathus longifrons.

. Side view of skull of Gorgonognathus longifrons.

. Side view of skull of Scymnognathus serratidens.

( 106 )

EXPLANATION OF PLATES.

PLATE XII.

. Top view of skeleton of Rhinesuchus senekalensis.

. Top view of skull of Rhinesuchus senekalensis.

. Top view of skull of Rhinesuchus whaitst. 2,

x ia

x 16°

x 4.

xX 75:

PLATE XIII.

Top view of skull of Scymnognathus serratidens.

@ Afr. Mus. Vol. XII. Plate XII

1,2. RHINESUCHUS SENEKALENSIS (V. Horrrn).

3,4. RHINESUCHUS WHAITSI, Broom.

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

1,3. GORGONOGNATHUS LONGIFRONS, HauGHrTon.

294. SCYMNOGNATHUS SERRATIDENS, Havcuron.

Plate XIII.

( 107 )

12.— Foraminifera and Ostracoda from the Upper Cretaceous of Need’s

Camp, Buffalo River, Cape Province.—By Freperick CHAPMAN,

A.L.S., F.R.M:S., ete.

With two Plates, XIV, XV, and Text-figure.

Some little time since, I was favoured by Dr. L. Péringuey, 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.—Vhis 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, Bornemann sp.

Sections cut in various directions point to a comparison with this

species. It is a shallow water form, and has an e.censive geological

range.

Minronina 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 quinqueloculine plan. Foraminifera of this type are found in

all deposits ranging from the Lias to Recent.

108 Annals of the South African Musewm.

TEXTULARIA cf. 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* as in Spiroplecta, 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

water sands of Cretaceous and Tertiary times (see Text-figure 18).

F.C. Piolo.

Fre. 18.—Thin slice of limestone from the Upper Quarry, Need’s Camp,

Buffalo River, showing fine-grained calcitic matrix with Polyzoa and

Foraminifera (cf. Anomalina sp.). x 28 diameters.

PULVINULINA sp. (of the P. elegans group).

Vertical and tangential sections. The test shows redundant shell-

erowth, as in some Cretaceous forms.

THE MICROZOA FROM THE LIMESTONE OF THE

NEED’S CAMP LOWER QUARRY.

General Characters of the Rock.—The rock from this locality is a

white, friable limestone, almost chalky in parts. Amongst the finer

erushings 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.

Famity LITUOLIDAE.

Genus HAPLOPHRAGMIUM, Reuss.

HAPLOPHRAGMIUM NEOCOMIANUM, Chapman.

(Plate XIV, fig. 1.)

Haplophragmium neocomianum, Chapman, 1894, Quart. Journ. Geol.

Soc., vol. 1, p. 695, pl. xxxiv, figs. 2a, b.

Idem, 1904, Annals 8S. African Mus., vol. iv, pt. v, p. 223; pl. 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.

Famity TEXTULARIIDAE.

Genus SPIROPLECTA, Ehrenberg.

SPIROPLECTA ANCEPS, Reuss sp.

(Plate XIV, figs. 3, 4.)

Teatularia anceps, Reuss, 1845, Verstein. d. béhm. Kreideform., vol. 1,

p. 39, pl. vi, fig. 79; pl. xin, fig. 2.

Idem, 1860, Sitz. d. k. Akad. Wiss. Wien, vol. xl, p. 234, pl. xiii, figs.

2a, b.

Spiroplecta anceps, Rss. sp., Chapman, 1892, Journ. R. Micr. Soc.,

Dedols ple xi, tion 6:

This species is perhaps the most abundant foraminifer in the Need’s

Camp limestone. It is subject to great variation. Some of the short

and wide forms resemble S. 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 1s

partially unrolled and conspicuous, as in fig. 4

10§

110 Annals of the South African Museum.

S. anceps is a well-known Cretaceous species, occurring in the Chalk

of England, Westphalia, and Bohemia, and in the Gault of Folkestone.

SPIROPLECTA ANCEPS, Reuss 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, 1:175 mm.; length of primary test, 0°351 mm. ; width

of test at oral extremity, 0°675 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 S. praelonga, Reuss sp.*

that Gaudryina dispansa, Chapman { does to G. pupoides, VOrb.

Famity NODOSARIIDAE.

Genus NODOSARIA, Lamarck.

NoposaRIA ZIPPEI, Reuss.

(Plate XTV, figs. 6, 7.)

Nodosaria zippei, Reuss, 1845, Verstein. bohm. Kreideform., pt. 1,

p. 25, pl. vii, figs. 1-3.

N. zippei, Rss., Chapman, 1904, Annals 8. African Mus., vol. iv,

pt. v, p. 226, pl. 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, Reuss. Verstein. bohm. Kreideform., vol. i, 1845,

p. 89, pl. xii, fig. 14.

+ Journ. Roy. Mier. Soc., 1892, p. 753, pl. xi, figs. 10a, b.

Foraminifera and Ostracoda. 111

Chalk specimens; although Reuss figures one example more comparable

with the present in this respect.

N. zippei is recorded from the chalk of Bohemia, Westphalia, the

Upper Bavarian Alps, Maestricht, and the Isle of Riigen ; also from the

Gault (Albian) of France and England, and in the Cambridge Green-

sand (Albian in part).

Noposaria sutcaTa, Nilsson.

(Plate XIV, fig. 8.)

Nodosaria sulcata, Nilsson, 1825 (1826), K. Vet. Ak. Handl., p. 341.

Idem, 1827, Petrif. Suecana, p. 8, pl. ix, figs. la, A, B (error for 19).

N. suleata Nilsson, Hisinger, 1837, Lethaea Svecica, p. 33, pl. x, figs.

4a, b.

Reuss, 1845, Verstein. bdhm. Kreideform., pt. i, p. 26, pl. xiii, fig. 17,

Idem, 1855, Zeitschr. deutsch. geol. Gesellsch., vol. vii, p. 269, pl. viii.

fig. 140.

Sherborn and Chapman, 1889, Journ. Roy. Micr. Soc., p. 486, pl. xi,

fig. 24,

Egger, 1899, Abhandl. k. bayer. Akad. Wiss., Cl. ii, vol. xxi, Abth. i,

p. 79, pl. viii, fig. 24,

This is a form resembling the more extensively ranging Nodosaria

obliqua, L. 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, Riigen, 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, pl. vii, figs. 1, 2a, b.

Berthelin, 1880, Mém. Soe. Geol. France, ser. 3, vol. 1, Mém. v, p. 56.

Chapman, 1894, Quart. Journ. Geol. Soc., vol. 1, p. 712.

Idem, 1894, Journ. Roy. Mier. Soc., p. 647, pl. 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 (Aptian) of Surrey, also in the Gault (Albian) of

Folkestone, France and Germany.

112 Annals of the South African Museum.

CRISTELLARIA INTERMEDIA, Reuss.

(Plate XV, fig. 10.)

Cristellaria intermedia, Reuss, 1845, Verstein. bohm. Kreideform., pt. 1,

p. 33, pl. xiii, figs. 57, 58; pt. ii, p. 108, pl. xxiv, figs. 50, 51.

Alth, 1850, Haidinger’s Naturw., Abhandl. iii (41), p. 267, pl. Xi,

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 Reuss * from

the Greensand (Cenomanian) of New Jersey, and by Deecke + from

the Oxfordian of Montbeliard, France. In the former the test is

inflated and the segments higher as in Reuss’ C. planiuscula, whilst

the latter has the series of chambers more closely inrolled.

CRISTELLARIA SECANS, Reuss.

(Plate XV, figs. lla, 6.)

Cristellaria secans, Reuss, 1860, Sitzungsb. d. k. Ak. Wiss. Wien,

vol. xl, p. 214, pl. 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 Reuss’ C. secans, which that author

found in the Gault Clay of the Rhine. A variety of this species,

angulosa, has also occurred in the Folkestone Gault.

Famity ROTALIIDAH.

Genus DISCORBINA, Parker and Jones.

DiscoRBINA PILEOLUS, d’Orbigny sp.

(Plate XV, figs. 13a, b.)

Valvulina pileolus, d’Orbigny, 1839, Foram. Amér. Mérid., p. 47, pl. i,

figs. 15-17.

Discorbina pileolus, dOrb. sp., Parker and Jones, 1865, Phil. Trans.,

* Sitzungsb. d. k. Ak. Wiss. Wien, vol. xliv, 1861 (1862), p. 336, pl. viii,

figs. 2a, b.

+ Mén. Soc. d’Emulation de Montbeliard, vol. xvi, 1886, p. 380, pl. ii, figs. 19,

19a,

Foraminifera and Ostracoda. 113

vol. clv, p. 885; Brady, 1884, Rep. Chall., vol. ix, p. 649, pl.

Ixxxix, figs. 2-4; Chapman, 1894, Quart. Journ. Geol. Soc., vol. 1,

p- 719; Idem, 1896, Journ. Roy. Mier. Soc., p. 591, pl. 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, d’Orbigny.

TRUNCATULINA SCHLOENBACHI, Reuss sp.

(Plate XV, figs. 12a, b.)

Rosalina schloenbachi, Reuss, 1862, Sitzungsb. d. k. Ak. Wiss. Wien,

vol. xlvi, Abth. i, p. 87, pl. xi, figs. 5a-c.

Discorbina schloenbachi, Reuss sp., Egger, 1899, Abhandl. k. bayer.

Akad. Wiss., Cl. ii, vol. xxi, Abth. 1, p. 164, pl. xvii, figs. 19-21.

Chapman, 1904, Annals S. Afr. Mus., vol. iv, pt. v, p. 229, pl. 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 T'runcatulina on account

of its finely perforated or smooth test. It is a typical Cretaceous

form.

TRUNCATULINA UNGERIANA, d’Orbigny sp.

(Plate XV, figs. 16a, b.)

Rotalina ungeriana, d’Orbigny, 1846, Foram. Foss. Vienne, p. 15%,

pl. vii, figs. 16-18.

Truncatulina ungeriana, d’Orb. sp., Brady, 1884, Rep. Chall. vol. ix,

p. 664, pl. xciv, figs. 9a-c.

Egger, 1899, Abhandl. k. bayer. Ak. Wiss., Cl. ii, vol. xxi, Abth.

p- 150, pl. xix, figs. 4-6.

Chapman, 1912, Mem. Nat. Mus. Melbourne, No. 4, p. 43, pl. 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, Reuss sp.

(Plate XV, fig. 14.)

Rosalina ammonoides, Reuss, 1845, Verstein. bohm. Kreidef., pt. i

p- 36, pl. xi, fig. 66; pl. viii, fig. 53.

Anomalina ammonoides, Rss. sp., Perner, 1897, Foraminifery Vrstev

Belohorskych (Palaeontographica Bohemiae, No. iv), p. 72.

Chapman, 1898, Journ. Roy. Micr. Soc., p. 4, pl. 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.)

Rotalia karsteni, Reuss, 1855, Zeitschr. d. deutsch. geol. Gesellsch.,

vol. vil, p. 273, pl. ix, fig. 6.

Pulvinulina karsteni, 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, pl. xx, figs. 832-34.

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 BAIRDIIDAE.

Genus BATRDIA, McCoy.

BaArIrRDIA SUBDELTOIDEA, Minster sp.

Cythere subdeltoidea, Minster, 1830, Jahrb. fiir Min. etc., p. 64, No. 18;

1835, p. 446.

Foraminifera and Ostracoda. 115

Cytherina subdeltoidea, Miinst. sp., Romer, 1838, Jahrb. fiir Min. etc.,

p. oL¢; pl. vi, fig: 16.

Reuss, 1845, Verstein. béhm. Kreideform., pt. i, p. 36, pl. v, fig. 38;

pt. u, p. 104.

Buirdia subdeltoidea, Miinst. sp., Rupert Jones, 1889 (1890), Mon.

Cret. Entom., Suppl. (Pal. Soc.), p. 5, pl. 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 pl. 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, ete.

BarRpIA SUBDELTOIDHA, Minster sp. var. AEQUALIS, Var. Nov.

(Plate XV, figs. 17a, b.)

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,t a Miocene to

Recent species.

BartRDIA 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. 8. 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, pl. vii,

figs. 4-6. Id., Rep. Chall. Zool. vol. i, pt. iii, Ostracoda, 1880, p. 55, pl. viii,

figs. 8a—f, and figs. 2a-f.

+ Rep. Ghall. tom. cit., p. 54, pl. ix, figs. 5a-/; pl. x, figs. 2a-c,

116 Annals of the South African Museum.

Fam. CYTHERIDAE.

Genus CYTHERE, Miller.

CYTHERE POSTCULTRATA, Sp. NOV.

(Plate XV, figs. 18a, b.)

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. 8. 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 RESULTS.

Upper Quarry.

The meagre series of Foraminifera obtained from this locality does

not afford any data which can be used to decide the age of the deposit.

The genera found, viz. Miliolina, 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.

Tower Quarry.

The following list of species of Foraminifera and Ostracoda throws

a decided light on the age of this bed.

Foraminifera :

Haplophragmium neocomianum, Chapm.

Spiroplecta anceps, Reuss sp.

* Rep. Chall, vol. i, Zool., pt. iii, Ostracoda, 1880, p. 78, pl. xv, figs. 5a-d.

Foraminifera and Ostracoda. Lye

Spiroplecta anceps, var. infracta, nov.

ss deflata, sp. Nov.

Nodosaria zippei, Reuss.

Ms sulcata, Nilsson.

Cristellaria parallela, Reuss.

% intermedia, Reuss.

. secans, Reuss.

Discorbina pileolus, VOrb. sp.

Truncatulina schloenbachi, Reuss sp.

- ungeriana, V’Orb. sp.

Anomalina anmonoides, Reuss sp.

Pulvinulina karsteni, Reuss sp.

Ostracoda :

Bairdia subdeltoidea, Minster sp.

x, . var. aequalis, nov.

a africana, sp. NOV.

Cythere postcultrata, sp. nov.

Amongst the Foraminifera of generally Cretaceous aspect may be

noted Haplophragmium neocomianum and Spiroplecta anceps. The

species which lend to the deposit an Upper Cretaceous appearance

are Nodosaria zippei (a quite restricted form), N. sulcata (almost in-

variably Upper Cretaceous), Cristellaria parallela, C. 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

the 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.

Fig

(118 )

EXPLANATION OF PLATES.

(All figures magnified 36 diameters.)

PLATE XIV.

1.—Haplophragmium neocomianwm, Chapman.

2.—Spiroplecta deflata, sp. nov.

3.— 59 anceps, Reuss sp.

4.,— Be i. ,» An example with a well-developed aboral

extremity.

5.— 3 S53 » var. infracta, var. nov.

6.—Nodosaria zippei, Reuss.

7.— PA 33 _

8.— 55 sulcata, Nilsson.

9.—Cristellaria parallela, Reuss.

Puate XV.

. 10.—Cristellaria intermedia, Reuss.

11.— es secans, Reuss: a, lateral aspect; b, oral aspect.

12.—Truncatulina schloenbachi, Reuss sp.: a, superior aspect; 6, inferior

aspect.

13.—Discorbina pileolus, d’Orb. sp.: a, superior aspect ; b, inferior aspect.

14.—Anomalina ammonoides, Reuss sp.

15.—Pulvinulina karsteni, Reuss sp.: a, superior aspect; b, inferior aspect :

c, peripheral aspect.

16.—Truncatulina ungeriana, @Orb. sp.: a, superior aspect; 6, inferior

aspect.

17.—Bairdia subdeltoidea, Minster sp., var. aequalis, var. nov.: a, left valve,

lateral aspect; b, edge view, ventral aspect.

18.—Cythere postcultrata, sp. nov.: a, carapace from right side; b, edge

view.

19.—Bairdia africana, sp. nov.: a, carapace from right side; b, edge view,

ventral aspect ; c, end view of carapace.

Ann. 8S. Afr. Mus. Vol. XII. Plate XPV:

FORAMINIFERA. NEED’s Camp, Burrato R.

F.C. del. Adlard & West Newman.

Ann. 8. Afr.

Mus. Vol. XII.

Plate XoV.

12)

FoRAMINIFERA: Fias. 10-16.

F.C. del.

OSTRACODA :

Fies. 17-19.

Negp’s Camp, Burraro R.

Adlard & West Newman.

(119 )

13.—Some New Species of Anomodontia (Reptilia) —By R. Broom,

D.Sc., C.M.Z.S., and 8S. H. Haventon, B.A., F.G.S., Assistant

Director.

(With 6 Text-figures. )

Genus DICYNODON, Owen.

DicyNnopon CORSTORPHINEI, sp. nov.

A small skull and lower jaw collected by the Rev. J. H. Whaits at

Graaff Remet (S.A. Mus. Cat., No. 3337) seems to belong to an

undescribed 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 alone the postorbital

bar somewhat in the manner seen in Hocyclops 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 interparietal 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 . : : : . Alloy, dleK0) "5

Basal length . ; : : =) AS) a5

Interorbital width . : . . ; 2 Som ee

Intertemporal width : : : : 1 RsOhoe

Snout to front of orbit . ; : 1) AOL

Length of preparietal . : ; : goles

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 baini, 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

Fig. 19.—Dicynodon corstorphinei, Br. and Htn. Type, No. 3337. Tem-

poral and frontal regions. x O7.

Oudenodon baini 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.

3037).

Locality Heuning Nest Krantz, Graatt 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 New Species of Anomodontia (Reptilia). 121

Greatest length (oblique) —. , . 215 mm.

Greatest width ; . : 5 : alos 2.0 Oe

Interorbital width . , : ; ; Bee is Ws.

Intertemporal width — . . ; . Holes ye

Width across nasals . ; . aoe.

Width between nostrils . ; : ; abe one.

Width between canines . p h . i S80.

Basal leneth . ; . , ; : lice

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

Fra, 20.—Dicynodon cavifrons, Br. and Htn. 'l'ype, No. 747. Preparietal

region. x .

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 postfrontal 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.

DiIcYNODON ROGERSI, 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. kolbei. 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

JZ Ge | FA ex

/ FAN

( K WW \

Lf cg

vs SSS

Fic. 21.—Dicynodon rogersi, Br. and Htn. Type, No. 2356. x 3.

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) —. . ab. 290 mm.

Greatest width ; : : 5) 228! .,;

Interfrontal width ; : + NU) Pee

Intertemporal width —. ; ! e405;

Basal length . : : ; . ab. 2380 ,,

Width across nasals : ; 5 hel

Width between nostrils . . é Mp G28o

Fie. 22.—Preparietal region of same. x }.

Type.—Skull (S. Af. Mus. Cat., No. 2356).

Locality.—Thee Kloof, Nieuweveld, C.P.

Horizon.—Lower Beaufort Beds, bottom of Cistecephalus zone.

DICYNODON PYGMAEUS, Sp. Nov.

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

squamosals 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 Pa Ny : ; abs (oOhie

Interorbital width. ; : Fe oleae.

Intertemporal width (minimum). : evr lone.

SiS

Fig. 23.—Dicynodon pygmaeus, Br. and Htn. Type, No. 2664. Natural size.

Type.—Female skull (8S. Af. Mus. Cat., No. 2664).

Locality Dunedin, Beaufort West, C.P.

Horizon.—Lower Beaufort Beds, Cistecephalus zone.

Genus EMYDOPS, Broom.

EMYDOPS 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 Hmydops. It agrees, however, sufficiently closely with

the known species of Hmydops 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 Hmydops, 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 : : . ; : SS dane

Interorbital width . . 2 : ; Saeed (8 Rees

Intertemporal width — . : ; easton

Basal length . : ; : p WAS 55

Minimum width across pterygoids : oO

Width across palate between tusks . 125 mm.

Kia. 24.—Emydops platyceps, Br. and Htn. ‘Type, No. 2667. Natural size.

Type.—Skull (S. Af. Mus. Cat , No. 2667).

Locality.—Dunedin, Beaufort West, @ Ps

Horizon.—Lower Beaufort Beds, Cistecephalus zone.

INDEX.

PAGE | PAGE

eavifrons (Dieynodon) F . 120 | platyceps (Emydops) . ; . 124

corstorphinei (Dicynodon) . . 119 | pygmeus (Dicynodon) ‘ : 128

DICYNODON . : s . 119 | rogersi (Dicynodon) . F 5 Al

EMYDOPS . : : ; . 124

( 127 )

14.—Investigations in South African Fossil Reptiles and Amphibia

(Part 10).—By S. H. Haveurton, B.A., F.G.S., Assistant

Director.

10. Descriptive Catalogue of the Anomodontia, with especial reference

to the examples in the South African Museum (Part 1).

(With 3 Plates and 20 Text-figures.)

Tuts 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 Lystrosaurus

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 Hocyclops, Diictodon, Kanne-

meyeria, 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 Anomodont skulls, mainly

through the efforts of the Rev. 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 Museum.

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 Anomodont reptiles so that we may obtain, if possible,

some idea of the lines upon which variation has taken place. For

this purpose itis 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 prove 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 defined by the use of the species of

Dicynodon. 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

type, 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.

DicyNopoNn JOUBERTI, Broom.

1905. Broom. Rec. Albany Mus.,1, 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 postfrontal in any of the specimens in which the sutures

are displayed. If present, it must have been small.

There is no evidence of a, septomaxilla, which, if present, does not

appear on the face. The lachrymal is small.

Fig. 25.—Dicynodon jouberti, Br. Skull No. 577. Preparietal region.

Natural size.

The chief measurements of a typical skull are :

Greatest length . ; : ; : . 108 mm.

Greatest width ; : ‘ . LO:

Interfrontal width : : : : ; « 20a

Intertemporal width — . ; : pe Es

Basal length ; : ’ : ia BO

Width of palate between canines . Leisiyy a

The basioecipital 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 backwards. 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.— The Gouph.”

Horizon.—Lower Beaufort Beds. (Tapinocephalus zone.)

DicyNODON MEGALORHINUS (Broom).

1904. Broom, Oudenodon megalorhinus. Rec. Albany Mus., 1, p. 180.

The type skull is of interest as being the first Dicynodont in which

a postfrontal was clearly recognised.

Fic. 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-

frontals.

Of the occiput and palate but little is seen. It is interesting to

note, however, that the fenestra ovalis faces almost completely

laterally and but very little downwards; it is rather small. The

suture between the basisphenoid 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. 13]

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 le together in the

pit of the basisphenoid just behind the pterygoids.

Type.—Incomplete skull. (S.A. Mus. Cat. No. 640.)

Locality —Prince Albert Road Station, C.P.

Horizon.— Lower Beaufort Beds. (Tapinocephalus zone.)

DICYNODON CORSTORPHINEI, 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 notochordal pit in the condyle. The

post-temporal vacuity hes 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 fenestra 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, Graaff Reinet, C.P.

Horizon.—Lower Beaufort Beds. (‘Top (?) of Hndothiodon zone.)

DicyNopOoN CYCLOPS, Sp. Nov.

The specimen to be described, although somewhat reminiscent of

Dicynodon platyceps 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 . : SIE

Length from beak to front of cnt ; . .o2 2

Antero-posterior diam. of orbit. P 0 » S44

Interorbital width . ; : 3 Bourne

Intertemporal width —. ‘ , ; 340

Basal length . : : ey

In side view the skull agrees with that of D. pardiceps in the

coneavity at the preparietal region and the convexity of the parietal

132 Annals of the 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

Fia. 27.—Dicynodon cyclops, Htn. Type. No. 3447. x 4.

circular, and look more upwards than outwards. The snout is

moderately long, and the nostrils are midway between the beak and

Fie. 28.—Dicynodon cyclops, Htn. Type. Preparietal region. Natural

size.

the orbit, looking wholly laterally. The supranasal bosses are feeble,

as are the caniniform 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 Amphibia. 15

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. he 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 preparietal is small and forms half the border

of the pineal foramen. The postfrontal is short, but fairly broad

anteriorly.

Fie. 29.— Dicynodon cyclops, Htn. Type. Side view of snout. x 3.

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,

frontals, 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, Graaft Reinet, C.P.

Horizon.—Lower Beaufort Beds. (Hndothiodon zone.)

DicyNODON FELICEPS, Owen.

1876. Owen. Cat. Foss. Rept. 8.A., p. 45; pl. xlin.

»o

1889. Seeley. Phil. Trans. clxxx B; pl. x, fig. 3.

134 Annals of the South African Museum.

1890. Lydekker. Cat. Foss. Rept. & Amphuib. iv, p. 20.

1913. Broom. Bull. Amer. Mus. Nat. Hist., xxx, p. 445.

1914. Broom. Phil. Trans. B206; pl. 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. (Hndothiodon zone.)

DicyNopON GRACILIS (Broom).

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

Fic. 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 intertemporal 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 sutures 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. )

DicyNopon IctTrIpops, 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

Klipfontein, 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 condyle 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 condyle. 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, (OE

Horizon.—Lower Beaufort Beds. (Hndothiodon zone.)

136 Annals of the South African Museum.

DicyNopoN LUTRICEPS, Br.

1912. Broom. Proc. Zool. Soc., p. 870; pl. 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.

DicyNODON PARDICEPS, Owen.

1876. Owen. Cat. Foss. Rept. S. Afr., p. 42; pls. xxxvili, xxxix.

1889. Seeley. Phil. Trans., clxxxs, p. 244; fig. 2.

1890. Lydekker. Cat. Foss. Rept. & 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. (Hndothiodon zone (?).)

DicyNopvon psirracops, Br.

1912. Broom. Proc: Zool. Soc., p. 869 ;"pl. xen, fig. 17.

1915. Broom. Bull. Amer. Mus. Nat. Hist., xxv, 2, p. 189, 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

approach 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 (Hndothiodon 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 postfrontals are apparently small.

The chief measurements are :

Length of skull (oblique). ; : . 100 mm.

Greatest width : ; : : : < ''O4iqe2

Snout to plane of orbit in median line . 5 al ee

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, jouberti 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. (Hndothiodon zoue.)

DicyNODON RECURVIDENS, Owen.

1876. Owen. Cat. Foss. Rept. S. Afr., p. 46; pl. lxix, 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. (EHndothiodon zone (?) ).

DicyNODON TRIGONICEPS (Br.).

1901. Broom, Oudenodon gracilis (pars). Proc. Zool. Soc., p. 162.

1904. Broom, Oudenodon trigoniceps. Rec. Albany Mus., i, p. 73;

pl. 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. Shght supraorbital

ridge, and less distinct median ridge between the two frontals.

Narrow, fairly long preparietal.

Locality —Pearston, C.P.

Horizon.—Lower Beaufort Beds. (Probably top of Endothiodon

Zone. )

DicyNODON ALTICEPS, Br. & Htn.

1913. Broom & Haughton. Ann. S.A. Mus., xu; 1, p. 37; pl. vn,

figs. 1, 2.

Fig. 32.—Dicynodon alticeps, Br.and Htn. Type. No. 2347. Parietal

and frontal regions. x 3.

In the original description of this species a mistake was nade 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 postfrontal. 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. 2847.)

Locality.—-One mile E. of Oudeberg, Graaff Reinet, C.P.

Horizon.—Lower Beaufort Beds. (Bottom of Cistecephalus zone.)

. DicyNODON BOLORHINUS (Br.).

1911. Broom, Oudenodon bolorhinus. Proc. Zool. Soc., p. 1076; pl.

Ixiui, 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 Hndothiodon zone or

bottom of Cistecephalus zone.)

DiIcYNODON BREVICEPS, Htn.

1915. Haughton. Ann. S.A. Mus., xii, 2, p. 59; pl. 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 postfrontal 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.—V oetpad, Murraysburg, C.P.

Horizon.—Lower Beaufort Beds. (Cistecephalus zone.)

Dicynopon cavirrons, Br. & Htn.

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 processes curve inwards, so that the width between the

tusks is small.

The occipital condyle is tripartite and short. The foramen jugulare

les 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 le 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.)

DicyNODON GRANDIS, Sp. Nov.

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 clearly—the specimen being presumably an old individual.

The chief measurements are :

Greatest length . ‘ : ; . 553 mm.

Greatest breadth . ; : ; : a4 ounes

Interorbital width . : : : , . “123 8

Intertemporal width . : : : ode

Basal length . ; : . . 430

Width between caniniform eee ‘ Je tO ee

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 Hocyclops longus. In front of the pineal foramen, and

forming its anterior border, is a small preparietal, lozenge-shaped.

The interfrontal suture is seen along a median ridge, which terminates.

somewhat in front of the pineal foramen. The postfrontal 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

caniniform processes, which are placed below the front of the orbit, 1s

101 mm. The pterygoids are strong, the minimum width across them

being 48 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 and has a deep symphysis. The

splenial 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 1s 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.

DicyNopoN 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 interorbital width is equal to, or

slightly greater than, the intertemporal 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. Itisa slightly distorted skull and lower jaw (S.A.

Mus. Cat. No. 3414) collected from the Cistecephalus zone at Dunedin,

Beaufort West, C.P., by the Rev. J. H. Whaits. It shows the same

small face and general proportions as the type, and has apparently the

same small occipital condyle without tripartite division. A curious

feature is the height of the basicranium above the level of the

quadrates, which bones are also close together, lying directly below

the post-temporal vacuities. Im 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.—Knuil’s Poort, Beaufort West, C.P.

Horizon.—Lower Beaufort Beds. (Cistecephalus zone.)

DicyNnopon INGENS, Br.

1907. Broom. Ann. Natal Govt. Mus., i, p. 168; pl. 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.—Ennersdale, Natal.

Horizon.—Lower Beaufort Beds. (Probably top of Cistecephalus

zone.)

Dicynopon KOLBEI (Broom).

1911. Jaekel, Udenodon sp. Die Wirbeltiere, p. 192; figs. 210-212.

1912. Broom, Oudenodon kolbei. Ann. 8.A. Mus., vii, 5, p. 337;

figs. 1-5.

1913. Broom, Diictodon kolbei. Bull. Amer. Mus. Nat. Hist., xxxu,

p. 454.

Fra. 33.—Dicynodon kolbei (Broom). Type. No. 1886. Parietal and

frontal regions. x 2.

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.

DICYNODON LACERTICEPS, Owen.

1876. Owen. Cat. Foss. Rept. S. Afr., p. 30; pl. xxu.

1890, Lydekker. Cat. Foss. Rept. & Amphib., iv, p. 18.

Founded on a skull with lower jaw. Comparatively small. Greatest

length of type skull about 158 mm. 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.)

DicYNODON LATICEPS, Broom.

1912. Broom. Proc. Zool. Soc., p. 868; pl. xcii, figs. 12, 18.

1915. Broom. Bull. Amer. Mus. Nat. Hist., xxv, 2, p. 181; fig. 17.

Two skulls collected by the Rev. J. H. Whaits from the Graaff

Reinet 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

incomplete, 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 preparietal 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-

Fira. 34.—Dicynodon laticeps, Br. Skull, No. 3328 (Young). Preparietal

region. x 3.

frontal is a long, narrow bone, which in 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

sinking-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 Musewm.

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. )

DicyNODON LEONICEPS, Owen.

1876. Owen. Cat. Foss. Rept. 8. Afr., p. 32; pls. xxiv—xxvi.

1890. Lydekker. Cat. Foss. Rept. & 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 has the following approxi-

mate measurements :

Greatest leneth . é : : . 300 mm.

Length from snout to front of orbit — . ago an.

Interorbital width ; , : ; : 05 3

Intertemporal width —. . : : : By og,

Width across nasal overhang : : . (By 5

Orbital diameter . : ; : ‘ 5 est

Basal length . ; ; : 5 PHT) 4

Width between base of Rein : : By 4

The condition of the skullis such that sutures are auc 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 skullis its great

height, 7. e. the length of the downward portion of the squamosal.

Type in the British Museum.

Locality.—* Gats River, Sneeuwberg, Graaff Reinet,” C.P.

Horizon.—Lower Beaufort Beds. (Cistecephalus zone (?).)

DicyNODON LEONTOPS, Br.

1913. Broom. Bull. Amer. 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 Cistecephalus zone.)

Dicynopon uissops, Br.

1913. Broom. Bull. Amer. Mus. Nat. Hist., xxxiu, 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. Postfrontal very narrow.

Type in the American Museum of Natural History.

Locality.— Wilgebosch, New Bethesda, Graaff Reinet, C.P.

Horizon.—Lower Beaufort Beds. (Cistecephalus zone.)

DicyNoDON mMoscHopes, Br.

1913. Broom. Bull. Amer. Mus. Nat. Hist., xxxu, 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. Interparietal large, forming considerable

part of upper surface of skull.

Type in American Museum of Natural History.

Locality.—Oudeberg, Graaff Reinet, C.P.

Horizon.—Lower Beaufort Beds. (Cistecephalus zone.)

Dicynopon mustor, Htn.

1915. Haughton. Ann. S.A. Mus., xii, 2, p. 58; pl. 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. kolbei removes this point of

148 Annals of the South African Musewm.

difference, the two species can still be separated by the difference in

shape of the anterior part of the postfrontal, 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. kolbei the reverse is the case.

Fig. 35.—Dicynodon mustoi, Htn. Type. No. 2674. x 4d.

Type-—Skull. (S.A. Mus. Cat. No. 2674.)

Locality. Dunedin, Beaufort West, C.P.

Horizon.—Lower Beaufort Beds. (Cistecephalus zone.)

DicyNoDON PLANUS, Broom.

1913. Broom. Bull. Amer. Mus. Nat. Hist., xxxil. 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

condyle 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. (Cistecephalus zone.)

DicyNODON PLATYCEPs, Br.

1913. Broom. Bull. Amer. Mus. Nat. Hist., xxxii, p. 444; fig. 4.

1914. Broom. Phil. Trans. B 206, p. 44; pl. 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.)

DiIcYNODON PYGMAEUS, Br. & Htn.

1917. Broom & Haughton. 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 Cistecephalus 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 original 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 shghtly farther back along

the parietal 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. 2) so that the paroccipital

processes are deep and fairly short.

The only other small species known from the Cistecephalus zone is

D. testudirostris. The two species are readily distinguishable at a

elance by the nature 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 lewer 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 Le) 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 shght broadening of the

parietal crest.

Type.—Skull. (S.A. Mus. Cat. No. 2664.)

Locality.—Dunedin, Beaufort West, C.P.

Hovizon.—Lower Beaufort Beds. (Cistecephalus zone.)

DicyNnopon RoGERsI, Br. & Htn.

1917. Broom & Haughton. Ann. §.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; pl. vi, figs. 2, 3.

1876. Owen, Oudenodon (?) strigiceps. Cat. Foss. Rept. S.A., p. 61;

pl. xliv, fig. 4.

1890. Lydekker, Udenodon strigiceps. Cat. Foss. Rept. and Amphib.,

lv, 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. Rept. S.A., p. 45; pl. xliv, figs. 1-3.

1890. Lydekker. Cat. Foss. Rept. 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 River.”

DicyNopDoN TESTUDIROSTRIS, Br. & Htn.

1918. Broom & Haughton. Ann. 8.A. Mus., xii, 1, p. 36; pl. 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

152 Annals of the South 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

Fic. 36.—Dicynodon testudirostris, Br. & Htn. Type. No. 2854, 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 condyle. The fenestra ovalis is almost as low as the quadrate.

Between the tubera the basioccipital is deeply grooved. The basi-

sphenoid is short. Anterior to it the pterygoids form a broad, short

plate 11 mm. wide. Just behind the interpterygoid vacuity is a

median knob.

The quadrato-jugal is peculiar in that it is apparently not anchy-

losed to the quadrate. On the left side it is seen in position, lying on

the downward process of the squamosal 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 : : : ; 3 Mays 5

Snout to front of orbit . : : : Me RS Sess

Length of orbit : : : P Pei Oy ee

TInterorbital width . F : 5 d AliGh

Intertemporal width (min.) . ; a

Basal leneth . ; ; ; Set sae

Width between caniniform processes z b S2On As

Type.—Skull. (S.A. Mus. Cat. No. 2354.)

Locality.— Dunedin, Beaufort West, C.P.

Horizon.— Lower Beaufort Beds. (Cistecephalus zone.)

DiIcYNODON TIGRICEPS, Owen.

1855. Owen. Trans. Geol. Soc., vii, 2nd ser., p. 233.

1876. Owen. Cat. Foss. Rept. 8.A., p. 38.

1876, Owen, Dicynodon baini. Cat. Foss. Rept. S.A., p. 36; pls. xxx—

XXXV (NOM XXXVl, XXXVl1).

1889. Seeley. Phil. Trans., 1889 B, p. 286, pl. xiii.

1890. Lydekker. Cat. Foss. Rept. and Amphib., iv, p. 24.

This species is doubtfully represented in the South African Museum

collection by an incomplete skull (No. 749) from Pearston, 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 les at the level of the middle of the foramen

magnum,

Type.—Skull in the British Museum.

Locality.—Gonzia River, Kaffraria. (‘D. baini” is from Fort Beau-

fort, :C.P.)

Horizon.—Lower Beaufort Beds.

Dicynopon tTRUNcATUS (Br.).

1899. Broom, Oudenodon truncatus. Ann. S.A. Mus., 1, p. 405 ;

pl. 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 Musewm.

forms nearly the whole of the front of the roof of the mouth. No

septomaxilla showing on face. Nostrils well advanced, large. Canini-

form 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 les 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-

cephalus. )

DicyNODON TYLORHINUS, Br.

1913. Broom. Bull. Amer. 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, postfrontals entirely hidden.

Postorbitals meet behind the pineal foramen.

Type in American Museum of Natural History.

Locality.—Wilgebosch, New Bethesda, Graaff Reinet, C.P.

Horizon.—Lower Beaufort Beds. (Cistecephalus zone. )

DicyNnopon wuHaitTst, Br.

1913. Broom. Bull. Amer. Mus. Nat. Hist., xxxii, p. 443; fig. 5.

1915. Broom. Bull. Amer. Mus. Nat. Hist., xxv, 2, p. 142; fig. 33.

This species 1s one of the largest of the Dicynodons. 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 ramus 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 by the parietals.

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 Adendorp 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 Adendorp specimen (S.A.

Mus. Cat. No. 3714), however, the pineal foramen seems to be nearer

the postorbital arch than in the type. Otherwise the 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 Dunedin 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

Anomodonts 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 Dicynodonts found with the known skull types.

For example, the bones named by Owen as Platypodosaurus robustus

have been variously conjectured to be bones of Hocyclops magnus, E.

longus, and Dicynodon grandis, 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 acromion 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.

Humerus.—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 epicondylar 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 85 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.

Uina.—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

History.

Locality.—Uitspanfontein, Beaufort West, C.P.

Horizon.—Lower Beaufort Beds. (Cistecephalus 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 Rey. J. H. Whaits on the Graaff

Reinet Commonage (8.A. Mus. Coll. 3334).

Tt 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 3 ; f : » 730mm:

Intertemporal width —. : : A tesa” aoe

Basal length . : oe WS) es

Distance from snout to front of orbit . » 100)

Distance between nares and orbit . Se ee allies

Width between caniniform processes —. 00a

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 septomaxilla 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 septomaxilla.

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 shghtly 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 con-

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 no 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

* Qudenodon 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 8.A. Mus. collection (No. 2364),

which has hitherto been considered asa 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 witha

Fie. 37.—Chelyrhynchus lachrymalis, Htn. Type. No. 3384. 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 condyle 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,

importance among the many forms of Dicynodon and _ its allies P

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

Fig. 38.—Chelyrhynchus lachrymalis, Htn. Type. No. 3334. Side view

of skull. x ‘8.

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, 7.e. that forms which avree in all other characters

agree also in the possession, shape, and position of the preparietal.

In describing the following Anomodont species, Dicynodon platyceps,

Dicynodon ictidops, Emydops lougiceps, and Emydorhynchus 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 2864 and 3334 differ in that the former has a

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 lachrymalis, 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. (8.A. Mus. Cat. No. 3334.)

Locality.—Commonage, Graaff Reinet, C.P.

Horizon.—Lower Beaufort Beds. (Hndothiodon 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.

DitctopoN GALEOoPS, Br.

1913. Broom. Loe. 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. Postfrontal absent.

Type in American Museum of Natural History.

Locality.—Slachter’s Nek, C.P.

Horizon—Lower Beaufort Beds. (Probably top of Hndothiodon

ZONE. )

Genus EOCYCLOPS, Broom.

EocycLors Loneus, 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 Hocyclops longus, and which he said was closely allied with the

type of Owen’s form Oudenodon magnus. 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 by the Rev. J. H. Whaits, which agrees fairly

Fia. 39.—Eocyclops longus, Br. Skull No. 3425. x 4 nearly.

closely with Hocyclops, 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 interfrontal 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 surround 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 Jeast

part of the boss around the pineal foramen. In the type of Hocyclops

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-

parietal ? 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 Dicyuodonts 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 Hocyclops longus—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 Hocyclops

longus and Hocyclops magnus, as also in 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 antorbital

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 . ; , : ; . 602 mm.

Greatest breadth . . . ; 406

Interorbital width : ' . ne LOS

Intertemporal width —. : : : 7 LOonG.

Snout to front of orbit . ; ; 3 Sy ANG es

Basal length . : : .) A065.

The nostril is very large, the internasal 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 EH. 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 shghtly 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 EH.

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 condyle 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 Paardekraal), Beaufort West, C.P.

Horizon.—Lower Beaufort Beds. (Cistecephalus zone.)

Genus MYOSAURUS, gen. nov.

MyYosauRUS GRACILIS, 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 : . Sek) 5

Interorbital width . : : eee ae

Intertemporal width : ‘ ; » Gye

Length of orbit : . , Boge aren

Width of orbit ? , : a4 Ve,

Snout to front of orbit (oblique) — . ; paler

Width of palate , ’ ; 5 SL Or are

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

Fie. 41.—Myosaurus gracilis, Htn. Type. No. 3526. Natural size.

is very thin in the anterior half and the caniniform process extremely

feeble—almost non-existent.

The lachrymal and prefrontal are both fairly small triangular bones

whose bases form portions of the antorbital border.

The frontal is fairly broad and extends back to the level of the

parietal foramen, 7. 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 prolongation 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 pineal 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

Fie. 42.—Myosaurus gracilis, 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 hes the tabulare,

a long, 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 les

in advance of the exoccipital portions, so that the plane of the back of

the condyle 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 and Amphibia. 167

The foramen jugulare lies below the level of the foramen magnum 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 PROLYSTROSAURUS, 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 shghtly smaller and shehtly 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 asin 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 larger than in Lystrosawrus. ‘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 aes been shghtly

flattened laterally.

Type. Co-type.

Greatest length (front of snout to squamosal) . 110mm. 118 mm.

Greatest width . , : , : : i Bo. SM PM Oon ae

Interorbital width —. ; : : , PE BS 43 ,,

Intertemporal width . : 3 : ose eee Jor 3s

Length of orbit : ; ; : i : BO) gs Soule

Height of orbit : ; : : ; 74 ee Oy ee

Height of suborbital bovdor above edge of

maxilla : ; , : ; hon Aillegeess Abi

Width between tusks : ' va 3 i OO ae

Length from snout to front of piven oe amen. TAL a Vode

The chief features of the skull can be seen from the figures given.

14:

168 Annals of the South 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.

Fig. 43.—Prolystrosaurus natalensis, Htn. Type. No. 3715. x 4.

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

they are narrowed by the encroachment on the top of the snout of the

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 he 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.

\ Ti RY ee, i ze

i) Zags

bie -_— Z

Wits are

(( | \\\i on en =

ASN ayy Wi Yi CZ / ——

VW, >

oo es Z

A! ey Gee

Fie. 44.—Prolystrosaurus natalensis, Htn. Type. No. 3715. x #4.

Of the vertebral column there are 34 vertebrae preserved—25 cervical

and dorsal, 6 sacral, and 38 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 Anomodont type. The clavicles

are slightly curved and are 77 mm. 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 16 mm.

The radius is 54 mm. long, 18 mm. broad proximally, and 26 mm.

broad distally. The bone is twisted so that the longer diameter of

the distal end is not continuous with that of the distal end of the ulna,

but is bert at an angle to it passing inwards and backwards.

The ulna is 57 mm. long, 7. e. slightly longer in proportion to the

humerus than in Lystrosawrus. As in that genus there is no bony

olecranon process ; but on the ulnar 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 is9 mm. The lower arm is carried at right angles to the

upper arm.

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 ot Lystrosawrus 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 :

Length. wondth.

MetacarpalI . : ped ores 10°5

Phalanx I. : ae 9 : 11

wae vi Bhd : ; : ep Pale? 12

Metacarpal II . : A Bie 14

Phalanx I: ; Ba: 12

Sen i ee . 9 : tal

Metacarpal ITT ; L Pileiesy 5, 145

Phalanx TE ’ F ; me “56 : 1k

UB Oe : P Sk Tie Vel

+P]

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 and Amphibia. 171

Neither ischium is complete, but the bone is greatly produced back-

wards. The pubis is not seen.

The femur is 95 mm. long, has a narrow proximal width of 40 mm.,

and a distal width of 31 mm. The shaft has a minimum width of

16mm. The median condyle 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 fronto-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 Broom’s species in this new genus under the

name Prolystrosaurus strigops (Broom).

It is possible that the bending down of the snout and Jenethening

of the premaxilla and maxilla which must have taken place in the

evolution of Lystrosawrus from the earlier Anomodonts is foreshadowed

in such a form as Dicynodon testudirostris from the Cistecephalus 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.)

Co-type.—Skull, lower jaw, and nearly complete skeleton (3319).

Locality —Loskop, Natal.

Horizon.—Middle Beaufort Beds. (Lystrosaurus zone.)

PROLYSTROSAURUS STRIGOPS (Br.),

1913. Broom, Dicynodon strigops. Rec. Albany Mus., 11, 5, p. 400.

1915. Broom, Dicynodon strigops. Bull. Amer. Mus. Nat. Hist., xxv,

2p. 140; fig. 29.

Skull rather small. Skull unusually flat, orbits directed more

‘

Way Annals of the South African Museum.

upwards than outwards; tusks passing as much forwards as down-

wards. Beak elongated, upper surface at 45° with frontoparietal

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 opisthotice.

Type in the American Museum of Natural History.

Locality. —(?) Harrismith, O.F.S.

Horizon.—Middle Beaufort Beds. (Lystrosawrus zone.)

Genus KANNEMEYERIA, Seeley.

K ANNEMEYERIA ERITHREA, Htn.

1915. Haughton. Ann. §.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

grandis, Hocyclops, and similar large forms from the Cistecephalus

zone, and also some of the species of Hndothiodon. 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

pronouncedly convex. From Endothiodon it differs, of course, in the

lack of molar 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 grandis 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 basisphenoid are thick bones even in the middle line;

and the basioccipital tubera lie considerably below the level of the

South African Fossil Reptiles and Aimphibia. 175

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 (Proe. Zool. Soc., 1912, Pl. LVI). Between the small opening

for the 12th nerve and the larger one for the 9th—l1th nerves there

is a broad transverse ridge of bone which crosses the floor of the

cranial cavity. The opening for the 9th—1]th nerves is connected

with the upper end of the vestibule by a narrow groove. ‘The upper

end of the vestibule is in a pit in the floor and hes 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.—W wmnaarsbaaken, Albert, C.P.

Horizon.—Upper Beaufort Beds. (Cynognathus zone.)

KANNEMEYERIA LATIFRONS (Br.),

1899. Broom, Dicynodon latifrons. Ann. 8.A. Mus., 1, p. 452; pl. x,

figs. 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 postfrontal. 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.

alticeps (Dicynodon) .

bolorhinus (Dicynodon)

breviceps (Dicynodon)

cavifrons (Dicynodon)

CHELYRHYNCHUS.

corstorphinei (Dicynodon) .

cyclops (Dicynodon)

DICYNODON

DITCTODON

EKOCYCLOPS

erithrea (Kannemeyeria)

feliceps (Dicynodon)

galeops (Diictodon)

gracilis (Dicynodon)

gracilis (Myosaurus)

grandis (Dicynodon) .

halli (Dicynodon)

ictidops (Dieynodon) .

ingens (Dicynodon)

jouberti (Dicynodon) .

KANNEMEYERIA

kolbei (Dicynodon)

lacerticeps (Dicynodon)

lachrymalis (Chelyrhynchus)

laticeps (Dicynodon) .

INDEX.

PAGE PAGE

138 | latifrons (Kannemeyeria) . is

leoniceps (Dicynodon) : . 146

139 | leontops (Dicynodon) . : . 146

139 | lissops (Dicynodon) . : . 147

longus (Eocyclops) . : . 160

140 | lutriceps (Dicynodon) : : 136

156

131 | megalorhinus (Dicynodon) . . 180

131 | moschops (Dicynodon) ; . 147

mustoi (Dicynodon) . , oy AT

128 | MYOSAURUS . ; : . 164

160

natalensis (Prolystrosaurus) Loa

160

172 | pardiceps (Dicynodon) : . 136

planus (Dicynodon) . : . 148

133 | platyceps (Dicynodon) : . 149

PROLYSTROSAURUS . > 167

160 | psittacops (Dicynodon) : . 1386

134 | pygmaeus (Dicynodon) F . 149

164

140 | reeurvidens (Dicynodon) . 5 dlssz

rogersi (Dicynodon) . : . 150

142

strigiceps (Dicynodon) : a low

135 | strigops (Prolystrosaurus) . a abefal

142

testudiceps (Dicynodon) . = Loi

128 | testudirostris (Dicynodon) . > lol

tigriceps (Dicynodon). ; 2 ll58

172 | trigoniceps (Dicynodon) . . 138

143 | truncatus (Dicynodon) ; . 158

tylorhinus (Dicynodon) i . 154

144

156 | whaitsi (Dicynodon) . : . 154

144.

EXPLANATION OF PLATES XVI—XVIII.

PLATE XVI.

Dicynodon grandis, Htn.

Fig. 1—Upper view of type skull. x 1 nearly.

Fig. 2.—Side view of type skull. x 2.

Fig. 1—Upper view of skull. 8.A. Mus. No. 3425.

Fig. 2.—Side view of skull.

Upper view of skeleton of Prolystrosaurus natalensis, Htn. x +.

PAG ATR ReXeV Lie

Eocyclops longus, Br.

x 3 nearly.

S.A. Mus. No. 3425. x 4 nearly.

PA ATER XeVeTE

Ann. S.A. Mus., Vol. XII. Plate XVI.

: DICYNODON GRANDIS, Hauauron.

Adlard 4- Son 4 West Newman, Ltd:

Amn. so Ae Mus: Vol. Xai: Plate XVII.

EOCYCLOPS LONGUS, Broom.

Adlard 4 Son & West Newman, Ltd.

Ann. S.A. Mus., Vol. XII. Plate X VIII.

PROLYSTROSAURUS NATALENSIS, Havucuron.

Adlard & Son d&- West Newman, Ltd.

(175)

15.-- Investigations in South African Fossil Reptiles and Amphibia

(Part 11)—By S. H. Havenron, 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 mght

hand side and are bent; and the hind limbs are flexed at the

knee, the femora lying thrown forward.

Of the skull but httle remains. The head was obviously tri-

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

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

the palatine, forming the inner border of an oval suborbital 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 and Amphibia. 177

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 tor

articulation with the ilium. Only a very small portion of the

Fic. 45.—Impression of type. Macroscelesaurus jansent. x 0.48.

2nd sacral is preserved ; it was very much more slender than the

first, but articulated with the ilium. The 5rd 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

ptecoracoid 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

of 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 Galepus 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 Galepus 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 [cti-

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, Heleophilus, Broomia, 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 Therocephalha 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 Annals 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 7%).

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 angusticeps. 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-

ticeps 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. Nn.

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 16, and has a diameter

of 12 mm. Then follows a diastema of 8 mm., behind which are

4 small molars occupying 26mm. 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 Fossu Reptiles and Amphibia. 181

and the anterior ends approximate to one another. The whole

of the lower border is formed by the septomaxillary which also

forms 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.

Fic. 46.— Upper view of type skull of -llopecognathus minor. xX 3.

Fic. 47.—Ventral view of type skull of Alopecognathus minor as

developed. x i.

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 mm. behind the postorbita]

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

Scymnognathus. 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

been described it almost invariably extends down outside this

fossa to meet the paroccipital process; but the lower limit in

this form is indefinable. 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 watson. 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 eae type

specimen :—

Greatest length 2 t AP i .. 240 mm.

Greatest width ae sd 2g (about) 110 33

Snout to front of orbit A ae a2 ore. el “3

Interorbital width .. a - a Rete: 5 aan

Intertemporal width aff it me caw) ld ear ae

Length of temporal opening : 58 45

Type.—Skull and lower jaw (S.A.M. Cat. Aa. 3415).

Locality.—Klipbank, Beaufort West Division, Cape Province.

Horizon.--Lower Beaufort Beds (Tapinocephalus zone.)

184 “Annals of the South African Museum.

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 Cistecephalus 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 220mm. 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

Fre. 48.—Upper view of type skull of Whattsia plityceps. x 2.

Fic. 49.—Ventral surface of type skull of Whattsia platyceps. x

sto

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 Museum,

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-

noqnathus 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 about 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 by the basioccipital. the

basisphenoid lying below as in the Dicynodonts. Between 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 whaitsi 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 surface 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.

Fic. 50a.—Transverse section just behind ‘transverse bar’’ of

type.

Fic. 50b.—Section across distorted snout of co-type.

Fic. 50¢.—Section of prevomer just in front of “transverse bar’’

in type.

The palatine articulates with 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 hes

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

fairly 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,

by 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 what 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

Cynognathids, 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 watsoni (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 sclatert 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. Hach 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

* Jn a specimen consisting of the front halves of both dentaries from Kruger’s

Kraal, Graaff-Reinet, 3 incisors occur in front of the canine. This specimen

probably belongs to the species under discussion.

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 Scymnognathus. 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 Whaitsia platyceps named in

honour of the Rev. J. H. Whaits, to whose zeal, energy, and

knowledge as a collector South African paleontology 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. No. 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, Graafi-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 54mm. 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 15 c2 m7 or 16 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 Scaloposawrus

and Ictidognathus 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 22mm. 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 Ist

For. Jus

Fic. 51.—Palatal view of type skull of Akidnognathus parvus.

canine and the 5th incisor of 2mm.; and one of equal size between

the large canine and the Ist 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 Annals of the South African Museum.

The nostrils are large, close together, almost terminal, and

look upwards and forwards.

The bones of the skull surface are similar to those of Scalopo-

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 Scalopo-

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 very

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 dorsally

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

South African Fossil Reptiles and Amphibia. 195

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.

AKIDNOGNATHUS PARVUS n.g. et n. sp.

Fie. 52a.—Section across skull just behind internal nares.

Fic. 52b.—Cross-section at middle of interpterygoid vacuity.

Fig. 52c.—Section across basisphenoid at front of keel.

Fic. 52d.—Section across prevomers near anterior end.

Int Pr Vac

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.

6. The great distance between the posterior ramus of the

pterygoid and the paroccipital 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 (Cistecephalus 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 claystone

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 i4clin8—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 isctm;, 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 whaitsi. The skull was found at Weltevreden, in

the Sneeuwberg range, Graaff-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 14mm. They are simple

pointed conical teeth without serrations and curve slightly back-

wards. Kach tooth has a few longitudinal grooves which do not

Pr. Par. Foss.

Pa.

C : B.Spu.

CyNOSUCHUS WHAITSI Nn. sp.

Fic. 53a.—Palatal view of type skull. x 1.

Fic. 53b.—Canine and molars of right side

Fic. 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 5mm. which is followed by the large canine tooth.

This has a compressed oval cross-section with a longer—antero-

posterior—diameter of 8.5mm. It, too, is unserrated. Closely

following the canine are the 8 molars (only 6 are preserved on

the left side). These occcupy a space of 25mm. 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 Ist 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 5th 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

by a thin internasal septum formed by the premaxillae and the

nasals. The floor of the nostril is formed by a small septomayilla

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 Museum

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 XIIth 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 foramen, 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 basisphenoid 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, Ill, 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 line.

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 Cynosuchus 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

Cynosuchus

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 ;

the molars are only cusped in a simple manner ; and

. 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

Oo. Gr

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, Graaff-Reinet, C.P.

Horizon.—Lower Beaufort Beds—Cistecephalus zone (near top of).

NOTES UPON THE BRAIN-CASE IN CERTAIN THERAPSIDA.

DrnocepHattiaA.---Hitherto comparatively little has been known

about the detailed structure of the brain-case of the Dinocephalha.

Watson, in his paper on the Order, has given the fullest account

of the region and, ¢nter 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

Pir. Foss

Fic. 54.—Median section through the back part of skull of a large

Tapinocephaloid. (S. A. M. Cat. No. 750.) x } 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 [X-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, Watson said: “* The prootic, of course,

contributes to the fenestra ovalis, above which it is perforated

by the aquaeductus fallopii for the VIIth 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 Lumiasaurus showing the 1od-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 thé basioccipital

condyle—the structure approximates more to that of Lamiasaurus

than to that of Mormosaurus. The fenestra ovalis does not le

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 ofthe parietal

which overlaps on to the side of the occipital plate; the lower

portion with the tabulare. The post-temporal fossa is very small.

GorGonopstA.—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

Arctops and Scymnognathus 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 Scymnognathus 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 Annals 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

Ornithorhynchus, has a long suture with the parietal.”

SCYMNOGNATHUS TIGRICEPS ?

Fic. 55a.—Diagrammatised sagittal section through back part of

skull No. 4334.

Fic. 55b.—Section across vestibule of same. Both figures x 3.

In the skull of Scymnognathus mentioned above (S.A. Mus. Cat.

No. 43834) 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 les 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 les at about the level of the notochordal

South African Fossil Reptiles and Amyphibia. 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 XIIth 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 [X-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 VIIth nerve has not been seen.

208 Annals of the South African Museum.

The foramen for the exit of the VIth 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 fenestra

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 pesterior

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.

GORGONOGNATHUS LONGIFRONS Htn.

Fic. 56.—Back half of type skull sectioned vertically, seen from

right and showing some of the foramina. x 4 nearly.

Fic. 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

Wellwood, Graafi-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

Tapinocephalus 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. Saat

The posterior part of the basisphenoid 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 rounded 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.

Post CANAL

ve Vest

Far Oce

ALOPECOGNATHUS MINOR Htn.

Fre. 58a.—Median longitudinal section showing wall of brain-case

from within. ;

Fic. 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 [X-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 VIIth 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 VIth 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 VIth 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.

Pit Foss.

Fic. 59.—Irregular longitudinal section of Therocephalan 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 respect, 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 VIth 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

Scymnognathus ? 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 Custecephalus

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

AKIDNOGNATHUS .

ALOPECOGNATHUS

BAURID: .

capensis (Scylacops)

CYNOSUCHUS .

DICYNODON

DIMETRODON .

DINOCEPHALIA

GALECHIRUS

GALEPUS :

GOMPHOGNATHUS

GORGONOGNATHUS

GORGONOPSIA

ICTIDOGNATHUS

ICTIDOSUCHUS

janseni (Macroscelesaurus)

INDEX.

PAGE|

192 | LAMIASAURUS

180, 210 | longifrons (Gorgonognathus)

196 | MACROSCELESAURUS .

| minor (Alopecognathus)

208, 210

197, 215 |

| platyceps (Whaitsia) .

190

214

202

Annals of the South African Museum.

parvus (Akidnognathus)

SCALOPOSAURIDA&

SCALOPOSAURUS

selateri (Scylacosaurus)

SCYLACOPS

SCYLACOSAURUS

| SCYMNOGNATHUS .

| suppostus (Cynosuchus)

THERAPSIDA

tigriceps (Scymnognathus)

whaitsi (Cynosuchus) .

WHAITSIA .

180,

197, 2

16.—On Cretaceous Cephalopoda from Zululand.—By L. F. Sparu,

MeSe:, E:G-S:

(With a Sketch Map, 4: Text-figures and Plates XIX to XXVI.)

CONTENTS.

I. INTRODUCTION : , :

Il. THE UMKWELANE HILL FAUNA

DESCRIPTION OF SPECIES:

A. AMMONOIDEA:

Family DesmMocERATID# :

Gen. Parapuzosia, Nowak.

1. P. sp. nov. ? ind.

Gen. Parapachydiscus, Hyatt.

2. P. sp. nov. aff. colligatus, Binkhorst sp. .

3. P. cf. wittekindi, Schliiter sp. .

4 P. Spin.

Family PRIoNOTROPIDS :

Gen. Mortoniceras, Meek.

5. M. woodsi, sp. nov.

6. M. aff. wmkwelanense, Crick

7. M. sp. aff. soutoni, Baily sp.

Gen. Pseudoschloenbachia, nov.

8. P. umbulazi, Baily sp.

9. P. umbulazi (Baily) var. acuta, nov.

Gen. Diaziceras, nov..

10. D. tissotiaeforme, nov.

Family PLACENTICERATIDS :

Gen. Placenticeras, Meek.

ll. P. subkaffrariwm, sp. nov.

Family NosTocERATiIp= :

Gen. Nostoceras, Hyatt.

12. N. ? natalense, sp. nov.

13. N.? subangulatum, sp. nov.

Gen. Bostrychoceras, Hyatt.

14. B. ? sp. nov.

15. B. ? sp. ind. :

Gen. Diplomoceras, Hyatt.

16. D. ? indicum, Forbes sp. .

PAGE.

219

224.

224:

226

229

231

232

234:

235

236

240

241

242

245

247

256

218 Annals of the South African Museum.

Family Bacuirip# :

Gen. Baculites, Lamarck . : . 2 . : =) 20

17. B. capensis, Woods . d : ‘ : : . 257

18. B. sp. aff. capensis, Woods ; : : : . 258

19. B. ef. aspero-anceps, Lasswitz . : ; ; . 259

20. B. cf. brevicosta, Schliiter : : 3 : . ‘260

21. B. sp. cf. sulcatus, Baily . : : : : . 260

22. B. bailyi, Woods. ; : : : : . 261

B. NautiLoip#a :

Gen. Eutrephoceras, Hyatt.

23. E. aff. dekayi, Morton sp. : : : : . 262

24. E. cf. sublaevigatum (VOrbigny) var. indica

(Spengler) . . ; : : 4 : . 263

Gen. Cymatoceras, Hyatt.

25. C. ? sp. juv. aff. valudayurense, Blanford sp. . . 264

OBSERVATIONS ON THE UMKWELANE Hitt Fauna . : ; . 264

Til, THE MANUAN CREEK FAUNA . : ; heh wes : - 273

DESCRIPTION OF SPECIES:

A. ALBIAN :

1. AMMONOIDEA . ; 5 : : 4 : : . 273

Family PHYLLOCERATIDS :

Gen. Phylloceras, Suess.

1. P. velledae, Michelin sp. . ; : 3 . 2738

Family DesMocERATIDS :

Gen. Puzosia, Bayle.

2. P. cf. bhima, Stoliczka sp. ; : : . 274

Gen. Uhligella, Jacob. :

3. U. ? sp. nov. aff. stoliczkai, Kossmat sp. . . 275

Family DirpoLocERATIp@ :

Gen. Dipoloceras, Hyatt.

4, D. cristatum, Delue sp. 277

5. D. quadratum, sp. nov. . é ; F . 278

6. D. sp. nov. 5 2 : é ay fic . 280

Gen. Pseudophacoceras, nov.

7. P. manuanense, nov. : ; : 5 . 281

Gen. Subschloenbachia, nov.

8. S. prerostrata, nov. . : , : : . 284

9. S. ef. trinodosa, Bose sp. . : : : . 285

10. 8. bispinosa, nov... : : é ‘ . 285

Family LYELLICERATIDA:

Gen. Stoliczkaia, Neumayr.

11. S. sp. ind. : : 3 : : : - 286

Family Hamirips:

Gen. Torneutoceras, Hyatt.

12. @. sp. inde : : : : : : . 288

Family ANISOCERATIDS :

Gen. Anisoceras, Pictet.

13. A. sp. ind. ; : . : ; é . 288

On Cretaceous Cephalopoda from Zululand. 919

Family Turrimiripa :

Gen. Turrilites, Lamarck.

14. T. cf. gresslyi, Pictet & Campiche . ; . 289

2. NAUTILOIDEA :

Gen. Cymatoceras, Hyatt.

15. C. manuanense, G. C. Crick sp. 4 : . 290

16. C. cf. manuanense, G. C. Crick sp. . : - 291

17. C. cf. albense, d’Orbigny sp. . : c 3 292

18. C. sp. ind. : ; : : : : . 293

19. C. sp. nov. ? ind. ; : : < : - 293

20. C. cf. kossmati, Spengler sp. . : 5 . 294

21. C. ? cf. clementinwm, d’Orbigny sp.. c . 295

B. SENONIAN:

22. Peroniceras cf. dravidicwm, Kossmat sp. . . 295

23. Kossmaticeras (Madrasites) bhavani, Stoliczka

sp. : ; : : : : ‘ 5 2)

24. Placenticeras cf. subkaffrarium, sp. nov. . . 800

25. Cymatoceras ? sp. ef. justum, Blanford sp. . 301

OBSERVATIONS ON THE MANUAN CREEK Fauna : ; : . 802

IV. AMMONOIDEA FROM ISOLATED LOCALITIES 5 : 5 SiO

A. Norru-West SHORE oF FausE Bay (SENONIAN).

3 Mortoniceras vanuxemi, Morton sp. . : 5 : . 308

2. Bostrychoceras ? sp. . : : : : . 3808

Bt Pomme: s Camp, UPPER CATEMBE (eras) : : : - 3809

1. Aconeceras nisoides, Sarasin sp. . : : 5 : ~ oll

2. Cheloniceras gottschei, Kilian sp. : 5 . 312

3. Cheloniceras (Acanthoplites ?) delagoense, enrol sp. 7 ol6

I. INTRODUCTION.

Tue 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. multifida,’ Steinmann, 7. 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. Crickt+ had considered that they “ most probably represented a

somewhat higher horizon (possibly Senonian) than that indicated by

the False Bay fossils,” which latter were regarded to be Cenomanian.

Since Etheridget already had described a Dowvilleiceras and a Lyelli-

ceras ?§ from the neighbouring Umsinene River, 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. delaruei, @Orbigny sp. ; the latter, a typical Dipoloceras of the

cristatum-group (s./.) 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, Cenomanian and Senonian 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. Crick] stated that the occurrence (in the

Manuan Creek district) of Cretaceous beds of an age younger than

Cenomanian was somewhat doubtful. On the other hand, Mr. R. B.

Newton, in his paper on “The Cretaceous Gastropoda and Pelecypoda

from Zululand,’** discusses the evidence in favour of a Senonian

* Read before the Royal Society of Edinburgh, December 6th, 1920. (See

‘Nature,’ vol. evi, No. 2669, December 23, 1920, pp. 554-5.)

+ “Cretaceous Fossils of Natal,” pt. 11, No. 2: “ The Cephalopoda from the

Tributaries of the Manuan Creek, Zululand,” ‘ Third Rep. Geol. Surv. Nat. and

Zulul.,’ 1907, p. 249.

t Ibid., pt. ii, “The Umsinene River Deposit,” p. 87.

§ Gen. nov. (type, A. lyelli, Desh. in Leym.; d’Orbigny, ‘Pal. France. Ter.

Crét.,’ pl. Ixxiv, figs. 1 and 2) dealt with in the writer’s Angola paper.

|| Loc. cit., pp. 247-8.

{ “Cretaceous Rocks of Natal and Zululand,” ‘ Geol. Mag.,’ n.s., dec. v, vol. iv

(1907), p. 347.

** «Trans. Roy. Soe. 8. A.,’ vol. i, pt. 1 (1909), pp. 94-5.

On Cretaceous Cephalopoda from Zululand. HPAL

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, Manuan Creek” (19 specimens).

(b) Albian and Senonian Ammonitids from the “south side of

Manuan Creek Valley,” from “high ground on north side of United

Manuan Creek and Umsinene 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).

(2 specimens).

(dq) Aptian Ammonites from Powell's Camp, Upper Catembe

(5 specimens ).

(ce) A Senonian fauna from Umkwelane Hill. This includes over

forty specimens (half of which number are Baculites), 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 (= bon map),

and the Umfolozi 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.

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

puff-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 Report Geol. Surv. Nat. and Zulul.,’ 1904, pp. 69-93, pls. i-ill.

+ “The Cretaceous Fauna of Pondoland,” ‘ Ann. 5. Afr. Mus.,’ vol. iv, pt. vii,

No. 12 (1906).

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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 8.E. at 1° or 15°; 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 fzom one horizon,

and the beds and their fauna can therefore be regarded as a whole.”

It will be advisable to describe the Umkwelane Hill fauna separately

from that of the Manuan Creek and the other localities. There is a

large collection of Ammonites from the Umzamba (‘ Umtamvuna ”)

Beds of Pondoland in the British Museum, approximately corresponding

in age with the Umkwelane Hill fauna, as stated by Woods * and by

Crick | 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 Hoepen, 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 Pondoland 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 Campanian

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

Umkwelane Hill, of a much greater thickness, can be regarded as

belonging to ‘one horizon.”

The third part of this paper will deal with the Manuan Creek

* Loc. cit., p. 847.

t Loc. cit. (‘ Geol. Mag-’), p. 343.

t Woods, loc. cit. (1906), p. 3387. Kossmat, “ Die Bedeutung d. Siidind.-

Kreideform.,” ‘Jb. K.K. Geol. R.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 (¢) 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.

Bayzand, 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. Kitchin 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.

Il THE UMKWELANE HILL FAUNA.

DESCRIPTION OF SPECIES.

A. AMMONOIDEA.

Famitry: DESMOCERATIDA :

Gen. PARAPUZOSIA, Nowak.

1. PaRApPuzosia sp. nov. ? ind.

Pl DeLX ig P25" Pl Oe tiesn Ay io. Pl xe, figs)

2 1906. Desmoceras (Puzosia) gaudama (Forbes) Boule, Lemoine &

Thévenin, “ Pal. de Madagascar,’ III, “Céphal. Crét. Diego-

Suarez.” Ann. de Pal., vol. i, fasc. iv, p. 20, pl. 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 (Pl. XXTV, fig. 3) was forwarded, the specimen itself being

too bulky to be sent. The measurements, according to Mr. 8S. H.

Haughton’s note, accompanying the photographs, are as follows:

Diameter ; . 670 mm.

Height of last Thor . 325 [=48°5 per cent. of the diameter].

Thickness ,,_,, . 225 [= 33 be om se Ul

Umbilicus : 20 = 18 a - gel:

The specimen was alee clamtererised 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 Thévenin * non Forbes, the

* Loc. cit., fig. 4a of pl. iv, which, however, is more compressed.

On Cretaceous Cephalopoda from Zululand. DO

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-

phylla (Sharpe) and of certain Pachydiseus and Parapachydiscus, are

not apparent in the photographs of the form here described. P.

gaudama itself (B.M. Geol. 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. daubréei, Grossouvre sp.,t

though, owing to the absence of the inner half of the lateral areas, the

primary coste 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 Zululand form. On the other hand Nowak’s figure of the

suture line § apparently shows good agreement, as does that of P.

leptophylla, Sharpe sp.|| The fine example of P. dawbréei figured by

Miller and Wollemann™ has a larger umbilicus and very strong

primary costation.

Nowak: is inclined to unite these two Santonian species, and considers

P. tannenbergica, Fritsch and Schloenbach** to be closely related,

but he also quotes, as an example of Parapuzosia, Stoliczka’s A.

denisonianus, which is pre-Senonian, like P. austini, Sharpe sp.,t? a

form inuch nearer the ancestral Puzosia-type. P. stobae Nilsson,tt

* “Rech. s. 1. Craie Sup.,” II., Pal., “Les Amm. d. 1. Craie Sup.” ‘Mem.

Carte Géol. France,’ 1893 (1894), p. 174, pl. xxvii, figs. 1 a, b.

+ Ibid., p. 154, pl. xxviii. (“ Sonneratia,” in Grossouvre.)

t “Unters. ii. d. Ceph. d. Ob. Kreide Pol.,” iii, ‘ Bull. Ac. Sci. Cracovie

ser. B (1918), pl. xliii, fig. 32, p. 363.

§ Itid., pl. xliv, fig. 40.

|| ‘Moll. Chalk England, ITI, “ Cephal.” (1856), pl. xxi, fig. 2.

q “Moll. Fauna d. Unter-Senon v. Braunschweig,” II, “ Ceph.,” ‘ Abh. Preuss.

L.A.,’ N.F., Heft 47 (1906), p. 8, pl. v.

** ¢Cephal. d. Bohm. Kreideform.,’ Prague, 1872, pl. ix.

+t Loe. cit., II, 1854, p. 28, pl. xii, figs. 1a, b.

tt ‘ Petrif. Suec. form. cret.,’ p. i, London, 1827, p. 5, pl. i. Moberg, ‘ Ceph.

i Sverig. Kritsyst.,’ II, “ Artbeskrifn.,’ Sver. Geol. Unders., ser. C, No. 73, 1885,

p. 18, pl. ii, figs. 1-5. Nilsson’s figure is somewhat diagrammatic, but Moberg’s

example (1a) represents a form apparently similar to the specimen here

deseribed, 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 (oc. cit., p.56) thought perhaps belonged to P. stobae.

226 Annals of the South African Museum.

which is a pachydiscoid development (Parapachydiscus) later than

P. denisoniana, and which has Puzosia-like inner whorls, is also grouped

here by Nowak. But the forms that continue the Puzosia-type into

the Senonian (P. gaudama, Forbes, P. indopacifica, Kossmat, etc.), and

once more produce transitional forms to Kossmaticeras (P. darwint,

Philippi in Steinmann), cannot be classed in the same genus with

P. denisoniana which developed a ‘“* Pachydiscus-stage ” 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. daubréei is the genotype of Parapuzosia. The ornamentation of

the inner whorls shows that it is not a Parapachydiscus; and the

writer believes that it has nothing to do with the gigantic forms of

the type of P. seppenradensis, Landois,+ which Nowak, in the writer’s

opinion wrongly, also considers possibly to belong to Parapuzosia.

Locality.x— Railway cutting, Umfolozi. Coll. Dr. A. L. du Toit.

S.A.M. Cat. No. 5513.

Gen. PARAPACHYDISCUS, Hyatt.

2. ParapacHypiscus sp. nov. aff. connigatus, Binkhorst sp.§

(Plo Oi figs: Va, b.)

1861. Anmonites colligatus, Binkhorst. Mon. d. Gast. & Céph. de la

Craie Sup. d. Limbourg, ii, p. 25, pl. vill a only.

1894. Pachydiscus colligatus, de Grossouvre. Amm. Craie Supér., p. 202.

1908. Pachydiscus colligatus, de Grossouvre. Descr. d. Amm. d. Crét.

Sup. du Limbourg, ete. Mdcm. Mus. Roy. d’Hist. Nat. Belg.,

vol. iv (1908), p. 28, pls. 1v—viil.

* Forbes (‘Trans.,’ 2nd ser., vol. vii, 1846, 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. Soe.

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., pl. xiv, fig. 6), but there is no ornamentation

remaining at this pachydiscoid stage.

+ “ Die Riesen-Amm. v. Seppenrade,” 23. Jahresb., ‘Zool. Sect., Westfiil.

Prov.-Ver. f. Wiss. and K.’ (1895), pp. 99-108, pls. i and ii.

t Loe. 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 Zululand. 227

1915. Pachydiseus colligatus, Nowak. Unters. Cephal. ob. Kreide

Polen, iii, Bull. Ac. Sci. Cracovie, ser. B, p. 361, pl. xliii, fig. 30,

pl. xhv, fig. 39.

A completely septate specimen (No. 5489) of 145 mm. 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 beimg only

an internal cast; but there are about 25 costae on the outer whorl,

weakened at the grooved siphonal line 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-

discus as P. quiriquinae (Philippi), Stemmann sp.

The specimen shows close resemblance with the examples figured by

Binkhorst on pl. vili a, and which were selected as typical by

Grossouvre, though Pervinquitre * was of opinion that the large

form figured by Binkhorst on pl. vii, and which was renamed

P. van den broecki, should be taken as type of Binkhorst’s species. 7

The present example differs from the typical P. colligatus (pl. iv,

fig. 3, and pl. v, fig. 1, in Grossouvre) 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 imner whorls of

P. colligatus to a larger diameter. The costation, possibly, also is.a

httle too distant in the Zululand example.

One of Schliiter’s | examples of Parapachydiscus wittekindi 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 Schhiter’s

species from the form here described.

Parapachydiseus ? 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

* «Bt. de Pal. Tunis,” I, “Céph. d. Ter. Sec.,” ‘Carte Géol. d. 1. Tunisie’

(1907), p. 175 (footnote).

+ This “species” was withdrawn in 1908 by A. de Grossouvre, who then

figured Binkhorst’s original (in the Berlin Museum)—a poorly preserved example.

+ *Ceph. d. Ob. Deutsch. Kreide.,’ I, Palaecontogr., vol. xxi (1872), e. g. example

2 of p. 68 (A. robustus), with thickness = 57 per cent. of the diameter, pl. xxi,

figs. 5 and 6; pt. ii (1876), p. 160. Boule, Lemoine and Thévenin (Joc. cit., IL,

1907, p. 22) wrongly consider this species of Parapachydiscus to be the type of

Pachydiscus, Zittel.

§-“The Fossil . . .. Mollusca. . . Chalk of England,” II, * Ceph.,’

Pal. Soc. Mon., 1854, p. 30, pl. xiii, figs. 2, 3.

228 Annals of the South African Musewm.

two forms are closely comparable, though the adult portlocki

develops ventral tuberculation.

P. epiplectus, Redtenbacher,* included by Grossouvre 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. quiriquinae (Philippi), Steimmann sp.} resembles the Zululand

example in suture-line, and has a similar straight principal lobe, but

is too thin and too closely costate.

“A. newberryanus,’ Meek sp., as figured by Gabb has a

similar suture-line, but the costation 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 Parapachydiscus here

discussed and related to the iscwlensis-group 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. nawmanni, 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.R.A., vol. v (1873), p. 121,

pl. xxviii, fig. 1.

+ “ Beitr. z. Geol. and Pal. S. Amer.,” III, “ D. Alt. und d. Fauna d. Quiriquina

Sch. in Chile,” N. Jb. f. Min., ete., Beil. Bd. x (1895), p. 74, pl. vi, fig. 3, Text-

fig. 5 on p. 77.

i “Pal. of Calif.” vol. i (1864), p: 61, pl. xxvii, fig. 199,,and pli xxviii,

fig. 1994. 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. complewus ?, var. suciensis (‘U.S. Geol.

and Geogr. Surv.,’ Bull. No. IT, 1876, p. 369, pl. v, fig. 2).

§ ‘Mesoz. Foss., I, pt. ii. ‘“ Foss. Cret. Rocks Vancouver, etc.,’ ‘Geol. Surv.

Can.,’ 1879, p. 109, pl. xiv, figs. 1 and 1a.

|| Associated with P. otacodensis, Stoliezka (Whiteaves, loc. cit., I, V, 1903, p.

340, pl. xlvi, fig. 1, and ‘Trans. Roy. Soc. Can.,’ 2nd ser., vol. i, sect. iv, 1895

[1896], p. 181), and other species of Parapachydiscus labelled by Kossmat (see

‘Jb. K.K. Geol. R.A.,’ vol. xliv [1894], p. 472).

* « Beitr. z. Kenntn. d. Fauna d. Kreidef. v. Hokkaido,” ‘ Paleontol. Abh.

Dames & Kayser,’ vol. vi, pt. iii (1894), p. 30 (176), pl. iii, fig. 1.

** «“ Verstein. a. d. Japan Kreide,” ‘ Paleontograph.,’ vol. xxxvi (1890), p. 187,

plcxaiy fierell

On Cretaceous Cephalopoda from Zululand. 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 pl. v with fig. 3 a of Stoliezka’s pl. lxiii, 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

costation 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 ‘“ Pachydiscus,” including Jacobites

from other parts of the island.f Woods stated§ that Pachydiscus

was absent in Pondoland; but there are four specimens from the

Umtamvuna River in the British Museum,|| including two large

examples that may belong to forms of the colligatus-group. The

presence of “* Pachydiscus” both in Pondoland and in Zululand thus

forms a further point of resemblance with the Indo-Malgascan fauna.

Locality —Umkwelane Hill. Coll. Dr. A. L. du Toit.

3. Parapacuypiscus cf. wiTTEKrINpbI, Schliiter, sp.

(Pl. XXIV, fig. 1.)

1872. Am. robustus, Schliiter, “Ceph. d. Ob. Deutsch. Kreide,” Palaeon-

toer., vol. xxi, p. 67, pl. xxi, figs. 5 and 6, pl. xxii, figs. 1-3.

1876. Am. wittekindi, Schliiter (ibid.), vol. xxiv, p. 40 (160).

* In Blanford and Stoliczka, “ Foss. Ceph., Cret. Rocks of $8. India,’ ‘Mem.

Geol. Surv., India, Pal. Indica’ (1865), p. 126, pl. bxili, fig. 3 only.

+ Loc. cit. (1890), i, p. 186, pl. xxi.

t See Boule and hévenin, ‘ Bull. Soc. Géol. France, ser. iv, vol. iii, 1903,

p. 486; Boule, Lemoine and Thévenin, Joc. cit. (1907), pp. 28-25; Kilian and

Reboul: “Les Céph. Néocrét. d. Hes Seymour et Snow-Hill,’ ‘Wiss. Erg.

Schwed. 8. Pol. Exp. 1901-03,’ ITI, 6 (1909), p. 25.

§ Loc. cit. (1906), p. 346.

|| Two of these, named P. wmtafunensis by Crick (MS.), Nos. C19434-5, and

compared with P. tweenianus, Stoliezka (loc. cit., 1865, p. 107, pl. lv 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

Parapachydiscus. They are similar to certain South-American “ Pachydiscus ”

(Paulcke “Die Ceph. d. Ob. Kreide Siidpatagoniens,” ‘ Ber. Naturf. Ges. Freiburg

i. B.. vol. xv (1907), e. g. pl. xix [x]). The other two gigantic specimens, not

described by Crick, may belong to Parapachydiscus 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, pl. xlii).

230 Annals of 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: 4. #54) Sees me eh ee Dag) as

Umbilicus . : : eo ils; a i

The costation 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. colligatus, Binkhorst sp., by Grossouvre and Nowak.t P.

fresvillensis, Seunes,; and P. epiplectus, Redtenbacher,§ show a similar

change, but one of the examples of the former species, figured by

Seunes (pl. xii (111), 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. egertonianus (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,++ 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 costation at an earlier stage,

* « Kreidefauna d. Peterwardeiner Geb.,” ‘ Palaeontogr.,’ vol. lii (1906), p. 88,

Die wa ged:

+ Loc. cit. (1908), p. 29 and (1913) p. 361.

t “Contrib. A ’Et. d. Céphal. Crét. Sup. France,” ‘ Mem. Soe. Géol. Fr.,’ Pal.,,

vol. ii, fase. iii (1891), p. 3, pl. i, fig. 1.

§ Loc. cit., p. 121, pl. xxviii, fig. 1.

|| Loc. cit. (1894), p. 80 (176), pl. iii, fig. 1.

{ Loc. cit. (1865), p. 104, pl. liii, fig. 1.

**® Toc. cit. (1894), p. 29, pl. ii, fig. 2.

++ “Vancouver Cret. Foss.,” ‘Trans. Roy. Soc. Can.,’ 2nd ser., vol. i (1895),

sect. iv, 1896, p. 182, pl. iii, fig. 6.

tt Loe. cit. (1907), p. 230 (64), pl. 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 lewisiensis) by Zittel, retains its costation

to a very large diameter, but apparently belongs to the true Pachydiscus

of the perampius group, with comparatively simple suture-line.

P. leryi, Grossouvre sp.,* 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 pl. xvi.

The large forms of Parapachydiscus from Pondoland, referred to

under P. sp. nov. ef. colligatus, are more compressed than the present

example.

The suture-line of P. wittekindi is less complex than that of the

Zululand form, which resembles in this respect the suture-line of

P. quiriquinae (Philippi), Stemmann sp.,f 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

eroove on the periphery, marking the position of the siphuncle, as in

P. colligatus.

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. Ulineworth.

4, PARAPACHYDISCUS, sp. ind.

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. 5969) in the suture-line. Only the peripheral portions of two

camerae, including the greater part of theexternal 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 Riesen-Amm. v. Seppenrade,” 23. Jahresb., ‘ Westfil. Prov.-Ver. f.

Wiss. and Kunst.,’ 1895, p. 104, pls. i and ii.

+ Loc. cit. (1894), p. 178, pl. xxi.

f Loe. cit. (1895), text-fig. 5, p. 77.

§ Loc. cit. (1908), pl. vi, fig. la.

232 Annals of the South African Museum.

The shell is partly preserved and 5 mm. thick in places, but

less than 2 mm. in others. It is in two thick layers, fibrous and of

a white porcellanous aspect, like certain Inoceramus shells in the

chalk, with a thin inner and outer coating. The specimen may belong

to a fat form of Parapachydiscus, like the two examples previously

described, but is too incomplete for specific determination. The large

Parapachydiscus from Pondoland are far more compressed.

Locality.—Lake Itesa (Eteza), Umfolozi. Coll. W. J. Wybergh.

Famity: PRIONOTROPIDA.

GEN. MORTONICERAS, Meek.

). MoRTONICERAS WOODSI, sp. nov.

(Pil. XXI, figs. 1 a-d.)

The single specimen (No. 5451) upon which this species is based

has the following dimensions :

Diameter . . : . 80mm.

Height of the last hor : . 45 per cent. of the diameter.

Thickness 53 - . 2, 40° PA ps

Umbilicus . ; . ie) ae . ms

The important eharantanienics 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 Mortoniceras of the type of M. soutoni

(Baily)* from the Umtamvuna River, in the British Museum, there

are some transitional forms to the present species, showing decline

of tuberculation on the outer whorl (at a considerably larger diameter)

and a decrease in the size of the umbilicus. On the other hand, one

of the forms figured by Stuart Wellert as Mortoniceras delawarense,

* The example figured by Woods (loc. cit., 1906, p. 837, pl. xliui, fig. 1) repre-

sents a more evolute shell. Baily’s type in the British Museum (Geol. Soe.

Colln. 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.

+ “Report on Cret. Pal. of New Jersey,” vol. iv (Pal. Ser.), ‘Geol. Surv. of

New Jersey, 1907, p. 837, pl. civ, figs. 1-3 only. (See under Mortoniceras

Vanuwemi, 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 delawarense-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.

Pervinquitret 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 M. 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.t 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 4. marroti (Coquand), Grossouvre,§ is striking.

None of the species of Mortoniceras from the European Senonian

resemble the form here described. The suture-lines of Grossouvre’s

Coniacian species are considerably simpler than are those of the Cam-

panian delawarense-group. 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. texanwm (Romer) as figured by Schliiter|| 1s

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 Raritan Clays, etc.,” ‘Mon. U.S. Geol. Surv.,’ vol.

XViii (1892), p. 252, pl. xliil, figs. 1, 2 only.

+ “Etudes de Pal. Tunis,” I, ‘ Céph. d. Ter. Second,’ 1907, p. 243, pl. xi, figs. 21

and 22. Pervinquiére, in his note (1) on p. 244, somewhat misrepresents

Whitfield, for the remark in the latter author’s description of M. vanuwemi

(p. 254) refers to the compressed specimen he figures (pl. 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.

~ “Up. Cret. Dep. of Maryland,” ‘Maryland Geol. Surv. Baltimore,’ 1916,

1) BUI Fae rats waked Yc

§ Hoplites vari, Schliiter var. marroti in Grossouvre, loc. cit. (1894), p. 119,

pl. vi, fig. 3b = Hoplitoplacenticeras, Paulcke.

| “Cephal. d. Ob. Deutsch. Kreide,” ‘ Paleontogr.,’ vol. xxi, part 2 (1872),

p. 41, pl. xii, fig. 3.

234 Annals of the South African Museum.

example of MW. sowtoni (Baily), very close to 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. Morvroniceras aff. UMKWELANENSE, G. C. Crick.

(Text-fig. D 2, p. 297.)

1907. M. wmkwelanense, G. C. Crick, loc. cit. (Third Report), p. 228,

pl xvepligs.19s.0 a.

Crick briefly characterised this species, which he considered to be

related to M. soutoni 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. C18154) 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 . : : : F , ; . 250 mm.

Whorl height . : : . : : 5 sey

Whorl! thickness : 5 "OO aee

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 sight, 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. texanum (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. texanwm (Romer) and M. quinquenodosum

(Redtenbacher), the umbilical tubercle is near the edge, and M. cam-

paniense, de Grossouvre,* 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. wmkwelanense 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. MorvroniceERas sp. aff. souront, Baily sp.

A large but somewhat fragmentary example (No. 5492), of 180 mm.

diameter, agrees with the evolute specimen figured by Woods,t and

has fairly smooth inner whorls, with the lateral tuberculation only just

indicated. The ribs also are, then, quite indistinct. The five

tubercles become as conspicuous as they are 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. souwtoni ; 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. sowtoni 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. cit. (1894), pl. xiii, figs. 1 and 3.

+ Loc. cit. (1906), p. 337, pl. xliii, fig. 1a.

{ “Cephal. Fauna d. Gosau-Sch. i. d. N.O. Alpen.,” ‘ Abh. K.K.R.A.,’ vol. v

(No. 5), 18738, p. 108, pl. xxiv, fig. 3.

§ ‘Die Kreidebild. v. Texas, etc.,’ Bonn, 1852, p. 31, pl. iii, fig. 1b (and Le ?).

|| Loc. cit., p. 338.

236 Annals of the South African Museum.

Grossouvre, to be quite different from Baily’s species. On the other

hand, the specimen figured as M. texanwm 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. teaanust = 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’s M. campaniense, a form near to M. dela-

warense, with which, indeed, de Grossouvre|| subsequently united it.

M. wmkwelanense, Crick,4{ 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. sowtoni, and whereas in M. wmkwelanense 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.—Umkwelane Hill. Coll. Dr. A. L. du Toit.

Gren. PSEUDOSCHLOENBACHIA, nov.

Genotype: A. wmbulazi, Baily, 1855, pl. x1, fig. 4.

A. umbulazi, Baily, described as Schloenbachia by Woodstt and

Crick,{{ has been considered by Kossmat §§ to belong to the ‘‘narrowly-

* Loc. cit. (1894), pl. xvii, figs. la, b, non pl. xvi, figs. 2-4.

+ “Ceph. d. Gosau-Sch.,” ‘ Beitr. z. Pal. v. Osterr.,’ vol. i, 1858, pts lap lO}

pl. ii, figs. 4-6.

# oc. cit., p- 108; pl. xxiv, fos:

§ “Kreide-Amm. v. Texas,’ ‘Geol. und Pal. Abh. Koken,’ n.¥., vol. vi

(1904), p. 31.

|| ‘Recherches s. 1. Craie Supér.’ (1901), p. 379.

{ Loc. cit., p. 228, pl. xv, fig. 9.

** See above, p. 234.

tt Loc. cit. (1906), p. 336.

+t Loc. cit. (1907), p. 250.

§§ “ Die Bedeut. d. Siidind. Kreideform., etc.,” ‘Jb. K.K.R.A.,’ vol. xliv (1894),

p. 464.

On Cretaceous Cephalopoda from Zululand. 237

umbilicated Schloenbachia (Prionocyclus) forms that appear in the

Lower Senonian,’ and to be closest to A. paon, Redtenbacher, 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. wmbulazi in Muniericeras,

appears to have been much nearer the mark. The young A. wmbulazi

reproduces the ornament of Muniericeras lapparenti, Grossouvre,t and

has chevrons on the venter, but no keel (Fig. B2). But before this

stage, the young A. wmbulazi reproduces Puzosia, with constrictions

(Fig. B38), 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 t-Schloenbachia§

with certain contemporary Desmoceratids; also, possibly, Munieri-

ceras itself includes Prionotropid forms (Grossouvre’s species, M.

gosauicum, Hauer sp., M. dresdense, Petraschek) and Desmoceratid

species, e.g. the lower Senonian A. elypealis, Schliiter and Brauns,||

and “ Desmoceras” clypealoides, Leonhard,£ apparently connected

with the Turonian Puzosia hernensis, Schliiter sp. Thus the presence

of constrictions on the inner whorls of P. wmbulazi, and a suture-line

that resembles that of certain Desmoceratids, are not definite proof

that Pseudoschloenbachia be derived from Puzosia rather than from

Prionotropidae, vii 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 fourniert, Grossouvre

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 Hawericeras, a Desmoceratid

* « Hoss. Mungokreide,” ‘ Beitr. z. Geol. v. Kamerun, IT’ (1904), p. 205.

+ Loc. cit. (1894), p. 158, especially pl. xxxv, fig. 3.

t Gen. nov. for studeri group, Jacob em. Spath (genotype, A. renauxianus,

d’Orbigny, ‘ Pal. Frang. Ter. Crét.,’ pl. xxvii).

§ Varians-group only.

| de Grossouvre (loc. cit., p. 175) has pointed out that Brauns’ form is

different from Schliiter’s type.

q “Fauna d. Kreidef. Oberschles.,”’ ‘Paleontogr., xliv (1897), p. 97,

pl. vi, figs. 2a, 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-Hawericeras, not pre-Hauwericeras, speaking

phylogenetically, and irrespective of age—in other words, it would have

travelled farther away from the Desmoceratid stock than Hawericeras.

Now Nowak* proposed the new genus Prohauericeras for those

forms of Schloenbachia that showed a decided tendency in the direction

of Hauericeras. The derivation of the latter genus from Schloenbachia

(vid Prohauericeras) can on no account be admitted. Hauwericeras has

a truly Desmoceratid suture-line, as a comparison of its internal

sutures (Fig. A 7) with those of Puzosia planulata (Sowerby), Kossmat,t

JText-FiGc. A.—Hauericeras gardeni, Baily, sp. Umtamvuna River, Natal.

(B.M. No. C 18528 [1-7 and 9] and C 18531 [8]). 1. Suture-line at

diameter = 4 mm. si = siphuncle; tv = umbilicus. 2. Ditto at 6 mm.,

without external saddle. 38. 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-line 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 (vu). a.u. = antisiphonal lobe.

8. Whorl-fragment at D = 90 mm., k = hollow keel, on barely carinate

venter of cast. (Photo. by G. C. Crick.) 9. Outline whorl-section at D =

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 gardeni (see Fig. A 1—5)¢ is similarly developed in many

forms of Puzosia (e.g. subplanulata, Schliter, compressa, indopacifica,

* «Unters. Ceph. Ob. Kreide Polen., III,” ‘ Bull. Ac. Sci. Cracovie,’ June,

1913, p. 370.

+ Loc. cit. (1898), pl. xvi, fig. 4.

t+ The specimen (B.M. No. C18528) from which these figures were drawn,

agrees in all respects with Baily’s type (B.M. Geol. Soe. Coll., No. 113870),

and the larger of the two fragmentary co-types (No. 113871) 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 authors, in exactly the same manner.

On Cretaceous Cephalopoda from Zululand. 239

Kossmat, gaudama, Forbes, ete.). When ornament appears in

Hawericeras (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. gardeni at a diameter of 60 mm., which accounts for the fact that,

e. g., Pervinquitre,t who, unlike Nowak, clearly recognised the hollow

keel, found the casts of his small examples perfectly rounded.

The assemblage mentioned by Nowak includes Albian, Cenomanian,

Turonian and Senonian forms and is most heterogeneous. Sharpe's

A. goupilianus (non @Orbigny) is only a smooth variety of Schloen-

bachia varians, Sowerby sp. Whiteaves’ newly-created Schl. propinqua

has as little to do with Stoliczka’s earlier A. propinquus, as

Anderson’s Schl. propinquat (Stol.), possibly a Turonian Prionotropid,

is related to the Indian species. Such Albian forms as A. acuto-

carinatus, Shumard, belonging to the group of A. roissyanus,

@Orbigny,§ again, are as little related to the true Cenomanian

Schloenbachia or to ‘* Prohauericeras,’ as the Neocomian Oosterella

cultrataeformis, Uhlig sp., is to either.

Since Prohauericeras has heen proposed, it may be convenient to

retain it|| for A. gowpilianus, dOrbigny, which apparently was intended

to be the type, since Nowak mentions it first, and refers to it again in

connection with “ Schloenbachia ” fournieri, Grossouvre, and “8S.” obesa,

Stohezka sp. In suture-line, as well as in other characters, this restricted

Prohauericeras ditters both from the true Cenomanian Schloenbachia and

from its own (‘Turonian) contemporaries among the Prionotropidae. On

the other hand, Pseudoschloenbachia 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 Schloenbachia 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.” sugata, 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. d. W. Miinsterland.,” ‘Zeit. D. Geol. Ges.,’ vol. lvii

(1905), p. 208, pl. viii, figs. 1 « and b.

+ Loc. cit. (1907), p. 165.

t Loe. cit. (1902), p. 123, pl. ii, figs. 34-88. Anderson (p. 63) also compared

Whiteaves’ “ 8.” propinqua with the Utatur form.

§ See under Pseudophacoceras (p. 283).

|| The name is no more unsuitable than is, e. g., Frech’s Paralytoceras for a

Devonian Clymenid.

240 Annals of the South African Museum.

The hollow keel of Hawericeras, 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 Pseudoschloenbachia is not in favour

of a possible connection of this genus either with Schloenbachia or

with Prohanericeras. 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 (Hulophoceras 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 Psendoschloenbachia with those genera (Hulopho-

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 vid Muniericeras (and

like Gauthiericeras, to which probably S. bertrandi, S. fournieri, Gros-

souvre,* and the forms of the Syrian Senonian have to be added) is

derived from Prionotropidae.

Sand 9. PsEUDOSCHLOENBACHIA UMBULAZI, Baily sp.

(Pl. XX, figs. 2 and 3; Text-fig. B 2-7).

1906. Schloenbachia wmbulazi (Baily). Woods, “Cret. Fauna of

Pondoland,” Ann 8. 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)

(Pl. 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 ,, Be » 20 a * Ae

Umbilicus : : els ‘i is ”

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. 24

diameter, but develops the typical ornament, with distinct umbilical

tubercles, on the outer whor!. It has the following dimensions :

Diameter : , > 59 mim

Height of last whorl . 54 per cent. of the diameter

Thickness us Fe Of bs ie a

Umbilicus ; LS

” ” ”

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.

il ; 4a

Text-ric. B.—1. Pseudoschloenbachia umbulazi, Baily sp., Umtamvuna River,

Natal (B.M. No. €C19427). Sectional outline of inner whorls (at diameter

= 382 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. Psewdoschloenbachia wmbulazi

(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 3). 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 varians (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) (Pl. 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, Nov.

Mr. Woods stated that P. wmbulazi appeared to be confined to

Pondoland. 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 Pl. XX, fig. 3.

242 Annals of the South African Museum.

there is an immature, evolute specimen from the Umpenyati River,

Natal, recorded by Crick“; on the other hand, five specimens from

the Umtamvuna River, 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. papillata, G. C. Crick MS. sp., is not represented in the present

collection,* 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. DIAZICERAS, nov.

Genotype: D. tissotiaeforme, sp. nov., p. 245, Pl. XIX, figs. 1 a—k.

This genus is created for one form in the collection that cannot be

included in any of the known genera of keeled Sencnian Ammonites.

Its suture-line stamps it as being near to the genera Hulophoceras,

Hyatt, and Spheniscoceras, Crick MS.,f and in general outline this

* Loc. cit. (1907), p. 250.

+ 'I'wo additional specimens of this 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

S. tenue, G. C. Crick MS., which obviously are the “ other species of Hulopho-

ceras”’ referred to by Woods (loc. cit., p. 387). 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.1

with that of Eulophoceras natalense, Hyatt (‘Text-fig. C 2, after Woods, loc. cit.,

pl. xhi, fig. 3), and with Hyatt’s type (Pseudoceratites, 1903, p. 86, pl. 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, pl. xxvi, fig. 7),

or of certain Dipoloceras and Pseudophacoceras described in this paper (see, e. g.

Pl. XXV, fig. 1 d, and Pl. XXVI, fig. 5b), placed on a very thin and acute whorl.

This is an exaggeration of the feature shown in Hyatt’s fig. 6 a of pl. xi, but of

doubtful generic importance. Hauericeras rembda, Forbes, shows a similar

feature occasionally. In fact the writer is not convinced that Crick was right in

On Cretaceous Cephalopoda from Zululand. 2.43

suture-line agrees with the South American Lenticeras and Paralenti-

ceras included by Hyatt in the family Eulophoceratidae.* 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 Senonian in certain Pseudoschloenbachia

(P. papillata, G. C. Crick MS.), in Barroisiceras (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 pecuhar rounding

separating the forms he described from Eulophoceras. Eulophoceratidae may be

distantly related to the contemporary Sphenodiscidae (not the Turonian Coilo-

poceratidae, often confused with Sphenodiscidae), but there is no connection

TExT-FIG. C.—l. Spheniscoceras, G. C. Crick (MS.), Upper Senonian,

Umtamvuna River, Natal. (After drawings, of the natural size, by the

late G. C. Crick.) la. S. africanum, Crick MS. (genotype), B.M. No.

C19421. 1b. S. minor, Crick MS. (B.M. No. C19422). lc. S. tenue, Crick

MS. (B.M. No. C19428). 2. Eulophoceras natalense, Hyatt (after Woods),

Pondoland. For comparison.

whatever between Spheniscoceras or any other Eulophoceratid and the

Placenticeratidae.

* Loc. cit., 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, and the

beginning development of adventitious lobes.

Barroisiceras includes shells (e.g. B. desmoulinsi, Grossouvre sp.)

that are near the new genus in external shape, though less so in

suture-line, and 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 Pseudotissotidae 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 Psendoschloenbachia in

ornamentation, Spheniseoceras and Eulophoceras 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 Eulophocerasand Spheniscoceras (ef. Fig. C la) and less so with

Pseudoschloenbachia, the similarity of the suture-line of Diaziceras

with those of Lenticeras, Paralenticeras and other pre-Campanian

genera makes it doubtful whether in the present state of our know-

ledge a subdivision of the Senonian Prionotropidae can yet be

attempted. Moreover, an undescribed form, closely resembling

“ Barroisiceras haberfellneri”’ from Madagascar, as figured by Boule,

Lemoine and Thévenin,* but possibly nearer to the Upper Chico

“Schloenbachia” chicoensis, Trask sp.( Anderson? ) occurs in Pondoland,

associated with many Campanian species, but also with “ Puzosia”

sugata, Forbes sp., which oceurs in the Lower Chico Beds of California,

and in Lower and Upper Senonian beds in India and elsewhere.

The likeness of these presumed Campanian 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 Pseudoschloenbachia 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, pl. xi, fig. 3.

ft Loc. cit. (1902), p. 116, pl. ii, figs. 23-25.

On Cretaceous Cephalopoda from Zululand. 945

Pseudoschloenbachia and Gauthiericeras and Muniericeras on the

other.

The uncertainty as to the presence of pre-Campanian horizons in the

Senonian 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 andi, Gabb sp., and DL. baltai, Lisson,*

might be explained by the mechanical laws that govern the formation

of the Ammonite-septum and its edge,t yet it is curious that as

Lenticeras is associated with Mortoniceras texanum, so Diaziceras

oceurs together with the comparable M. wimkwelanense, whereas the

presence of Peroniceras in Zululand, and of “ Puzosia” sugata and

the above new form in Pondoland, suggests that the new genus may

be closer to Coniacian genera than is here assumed. Unfortunately

the writer has no comparable material for dissection.

10. DiazicERAS TISSOTIAEFORME, Nov.

(Pl. XIX, figs. 1 a-k.)

This species is based on a completely septate specimen (No. 5478)

having the following dimensions :

Diameter 5 : ; . 80mm.

Height of the last whorl . 00 per cent. of the diameter.

Thickness ,, " : Poo os 4 ‘3

Umbilicus : : : a 14 3 x

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 whor! 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), pls. xiii and xiv.

+ See the writer’s “ Notes on Ammonites,’ ‘Geol. Mag., 1919, January to

May numbers, and compare, e.g., the suture-lines of Pseudophacoceras (Pl. XXV,

fig. 1b and c), and Ozynoticeras (Pia, 1914, pls. viii-xi), or of Aconeceras

nisoides (Fig. B 9, p. 33) and Pseudoschloenbachia griesbachi (Fig. B 8).

24.6 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 Leuticeras andii, Gabb sp., L. baltai,

Lisson, and of Paralenticeras sieversi, Gerhardt sp.,* and of the two

close allies—EHulophoceras and Spheniscoceras, 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,t 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 Coniacian forms may belong to quite a different

stock from the presumably Campanian Diaziceras.

Pseudoschloenbachia 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. fowrnieri in

Grossouvre,{ and has much more prominent umbilical tubercles. But

in Pseudoschloenbachia the suture-line is quite complex and charac-

terised by a large and deep first lateral lobe. A comparison of

fig. B 4a (p. 241) with fig. 1% of Pl. XIX will show that whereas

Pseudoschloenbachia 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 Pseudoschloenbachia 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

“Geol. und Pal. v. 8. Amer.,” V,‘N. Jb. f. Min., etc.,’ Beil-Bd. xi, 1897, p. 82,

text-fig. 6, and p. 79, text-fig. 5.

+ Loe. cit. (1904), pp. 168 and 169, text-figs. 538 and 54.

+ Loc. cit. (1894), pl. xxix, fig. 6b, and pls. xxxv, figs. 1 b and 1 ¢.

On Cretaceous Cephalopoda from Zululand. 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.—Umkwelane Hill. Coll. Dr. A. L. du Toit.

Famity : PLACENTICERATID 4.

Gren. PLACENTICERAS, Meek.

11. PLACENTICERAS SUBKAFFRARIUM, Sp. Nov.

(Pl. XXI, figs. 2 a-d.)

This species is based on a fairly well-preserved specimen (No. 5106),

of the following dimensions :

Diameter : : . 85 mm.

Height of last whorl . 90 per cent. of the diameter

Thickness ,, Ps : oe ES 7 aN

Umbilicus 2 : : , WG o a ie

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. 2d 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. kaffrarium,

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 striw, both on the shell, as in P. tamulicum

(Blanford) Kossmat,t and on the cast of the body-chamber, as in

P, stantoni, var. bolli, Hyatt.t.

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, pl. ili, fig. 16.

+ Loc. cit. (1895), p. 174, pl. xxii (vill), figs. 1 a-c.

““Pgeudoceratites of the Cretaceous,” ‘Mon. U.S. Geol. Surv.,’ vol. xliv

(1903), pl. 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.g. the first adventitious lobe is

smaller than the second one in P. subkaffrariwm, whereas the reverse

proportions are noticeable in the Indian species.*

P. stantoni, var. bolli, Hyatt,t also is a close ally of the present

species, as is P. intermedium, Johnson,{ the latter only distinguished

from the Zululand form by a narrower ventral area and a wider

umbilicus.

Locality.—Umkwelane Hill, Umfolozi, Zululand. Coll. J. 8. Hedges.

Famity: NOSTOCERATIDA.

Gren. NOSTOCERAS, Hyatt.

12. NosTocERAS ? NATALENSE, sp. nov.

GP Xx, flesh a0)

A depressed turricone (No. 2746) with the apical portion missing,

like the somewhat similar Didymoceras ? newtoni, Whitfield sp., cannot

definitely be referred to either Didymoceras 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? sub-

angulatum, nov., 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.? tricostatum, Whitfield,|| which

Hyatt" considered to be the possible gerontie stage of D. ? newtoni.

* P. syrtale, Morton, var. tamulicum (Blanford), Kossmat in Boule, Lemoine

and Thévenin (loc. cit., II, 1907, p. 47, pl. xii, figs. 3 and 4), is very close to the

Zululand specimen, but apparently possesses the outer tubercle of P. syrtale.

+ Loc. cit., “ Pseudoceratites,” 1903, as above, also, e.g., pl. xliii, fig. 1.

t ‘The Geol. of the Cerrillos Hills, New Mexico,’ part ii, ‘ Palaeont.,’ School

of Mines Quarterly, vol. xxiv, no. 2, 19038, p. 206, pl. viii, figs. 27 a, b.

§ “Invert. Cret. and Tert. Foss.,” ‘U.S. Geol. 8. Territ.,” vol. ix (1876),

pl. xxii, fig. 5.

|| “ Pal. Black Hills, Dakota,” ‘U.S. Geol. 8S.’ (1880), pl. xv, fig. 7.

"| “Phylogeny of an Acquired Characteristic,’ ‘Proc. Am. Phil. Soce.,’

vol, xxxii (1894), p. 574.

On Cretaceous Cephalopoda from Zululand. 249

Only in the Zululand 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 pl. xxxv, figs. 1-4,as Heteroceras 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’s loosely coiled form,* and less rounded than that

of D.? newton, which has a larger umbilicus. That of the present

species is as small as the umbilicus of D. pawper, Whitfield sp., or that

of D. archiacianum, @Orbigny sp., which latter also shows a similar

forward sweep of the costation 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 Turrilites acutus, Passy, also

recorded from Zululand by Crick.f ‘ Turrilites” tridens, Schliter,t

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 under-surface of the whorls.§

If the form here described is a Nostoceras, it probably represents a

|; for the single costae

less specialised type than N. stantoni, Hyatt

with two lines of tubercles, so characteristic of Hxiteloceras, are found

in the young of Nostoceras stantoni and of N. helicinum, Shumard sp.%

The latter form has a very short spire like the Zululand form, with a

deeply impressed suture, but the costation is quite different.

Locality—Umfolozi 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, pl. 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).

+ Loc. cit. (1907), p. 176, pl. xi, figs. 3 and 4.

ft Loc. cit. (1872), pl. xxxv, fig. 9.

§ The trituberculate (?) Turrilites peramplus Lasswitz (loc. cit., p. 14, pl. ii,

fig. 1) also resembles the present form somewhat in shape, but may be a true

Turrilites, though Lasswitz compares it with Schliiter’s “ Turrilites” tridens.

|| Loc. cit. (1894), p. 570.

{ “Deser. of New Cret. Foss. from Texas,” ‘ Proc. Boston Soc, Nat, Hist.,’

yol. viii (1861), p. 190.

250 Annals of the South African Musewm.

13. NostocERAS ? SUBANGULATUM, Sp. Nov.

(Pl. XXII, figs. 3 a—c:)

A sinistrally coiled, fairly elevated but fragmentary turricone

(No. 27464), 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 Haiteloceras, and the fragment

of Hx. angulatum figured by Meek,+ 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 H'vite-

loceras,t 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.

stantoniand “ N.?”’ 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 ? stevensoni, Whitfield sp.|| 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, ete.,’ ‘Bull. Am.

Mus. Nat. Hist.,’ vol. xiv (1901), p. 219, pls. xxix and xxx.

+ Loc. cit., 1876, p. 484, pl. xxi, fig. 3 (perhaps a fragment of a Didymoceras ?).

+ Hyatt (loc. cit., p. 577) calls Hamites fremonti, Marcou (holotype in B.M.,

Geol. Soe. Coll., No. 12667), probably a gerontic stage of some species of

Ewiteloceras, which is doubtful, the former probably being of Albian age, and

close to “Anisoceras alternatum,’ Pict. & Camp. non Mantell (loc. cit., 1861,

pl.li). Helico. pariense, White, also, in the writer’s opinion, is not an Eviteloceras,

and with the so-called “ Crioceras ellipticum, Mantell,” of Schliiter and other

authors, and similar Turonian forms, belongs to a new, unnamed genus.

§ Loc. cit. (1894), pp. 568 and 571.

|| Loc. cit. (1880), pl. xiv, fig. 7, and loe. cit. (1901), pl. xxx. This form seems

to differ from the species here described only in the uncoiling of the body-

chamber.

On Cretaceous Cephalopoda from Zululand. 251

a reverse curve that is strongly convex forwards. A somewhat

similar costation is shown on the upper whorl-surface of D. pauper,

Whitfield sp.,* and of D. ? conradi, Morton sp., in Whitfield.

The elongated tubercles, situated at each side of the siphonal zone

(which latter becomes almost smooth towards the end of the shell) are

small on the septate portion, but 4 mm. high and very sharp, where

preserved on the last whorl. There is only one untuberculate inter-

mediate rib here and there, as on the last whorl of D.? stevensoni, but

the bituberculation is rather irregular as regards spacing.

The suture line shows good agreement with that of Didymoceras

tortum, Meek sp.,f except that the latter, taken at a larger diameter,

shows correspondingly greater complication. There is the same high

external lobe; only in the present form the siphonal line hes between

the two rows of tubercles, which are placed more centrally. The

principal lateral lobe shows a similar smaller outer and larger inner

branch ; the lateral saddle is equally bifid, the smaller second lateral

lobe is bifid in the two forms, and the arrangement of the dorsal lobe

and saddles is very similar. In D.? stevensoni, the very complex

suture-line, taken at a much larger diameter, shows a different deve-

lopment of the principal lobe, the outer branch being the larger.

The suture-line of D. pauper (Whitfield) also is very similar to that

here figured, and the whorl-section agrees, but the costation is

different.

D. hornbyense, Whiteaves sp.,§ a form somewhat resembling D. ?

binodosum, Hauer sp.,|| is much more finely costate than the specimen

here described, to judge by an example from Vancouver Island in the

British Museum,{{ and other specimens of Didymoceras in the same

collection from the Upper Missouri, etc., differ in the same respect.

* Loc. cit. (1892), pl. xlv, fig. 3.

+ Ibid., pl. xlv, fig. 10.

t Loc. cit. (1876), p. 481, pl. xxii, fig. 4c.

§ Loc. cit. (Mesoz. Foss. I), p. 332, pl. xlii, figs. 1-4.

\| “ Neue Ceph. a. d. Gosaugeb. d. Alp.,” ‘ Sitz. B. K, Akad. Wiss.,’ vol. liii

(1866), p. 8, pl. i, fig.6. This, however, may be a Bostrychoceras.

{ This was labelled by Kossmat “ Heteroceras sp. aff. cooperi, Meek,” whereas

another example of a Didymoceras, corresponding somewhat to Whiteaves’ A.

cooperi (Loc. cit., pl. xliii, fig. 1), but not with Meek’s specimen or Gabb’s fragment

(the latter compared with Emperoceras by Hyatt, loc. cit., p. 576), was wrongly

labelled by Kossmat “ Acanthoceras vancouvrensis, Meek.” In form and costa-

tion this second fragment recalls D. ? conradi (Morton), Whitfield sp., and

Emperoceras simplicostatum, Whitfield sp., but there is also a striking resem-

blance of the looped tubercles with the ornament of Jacobites anderssoni, Kilian

and Reboul (loc, cit., 1909, p. 35, pl. viii, fig. 3).

952 Annals of the South African Museum.

On the other hand, there is a superficial resemblance to certain Albian

Turrilites, e.g., T. circwmtaeniatus, Kossmat,* or T. catenatus

(VOrbigny),+ but not to Cenomanian species. The suture-line, however,

is placed differently in these true Twrrilites, and in the writer’s

opinion the Nostoceratidae cannot he considered to be descendants

of the earlier Twrrilitidae. The grouping of the uncoiled forms of

the Senonian may be provisional and more or less unsatisfactory in

the present state of our knowledge; but we must reject Nowak’s t

opinion that the Senonian “ Heteroceras”” (“ Helicoceras” is a strictly

Albian development), can, with the Aptian true Heteroceras, be con-

sidered to belong to one branch of uncoiled Parahoplitids.

Locality.—Umfolozi Valley, East of Railway. Coll. Mr. Illingworth.

Gren. BOSTRYCHOCERAS, Hyatt.

14. BostRYcHOCERAS ? sp. nov.

1906. Heteroceras sp. Woods: “ Cret. Fauna of Pondoland.” Ann.

S. Afr. Mus., vol. iv, part vil; No. -12, p. 339, pl. xlii, fig. 5a, b.

This form is represented in the collection by several fragments.

One of these (No. 5477) about 55 mm. in leneth, with almost circular

cross-section (long and short diameters 12 mm. and 11°5 mm. respec-

tively) corresponds with the figured example ; another larger fragment

of 16 mm. diameter and about 40 mm. length (No. 5476), like the

impression of a third and still larger example (No. 54774), appear to

have some of the ribs more pronounced than others, so that it would

seem as though, at a larger diameter, this form develops costation

like that of the Bostrychoceras sp. ind. next described and compared

with certain flared Japanese forms. The ribbing of the examples

under discussion, however, is of quite a peculiar character. The

inner shell, like the cast of the interior, only shows very indistinct

costation. The second layer forms a broad and flat septum at the

base of each rib and slight concavities in between these septa. The

* Toe. cit. (1895), p. 141, pl. xviii, figs. 4 and 5. Kossmat renamed Stoliczka’s

Turrilites brazoensis, since it does not agree with Roemer’s type, which Kossmat

considered to be Lower Senonian. Whether Kossmat’s example (p. 142 (46),

pl. xx, fig. 4), agrees with the quadrituberculate Texas species may be doubtful,

but Turrilites brazoensis is a true Cenomanian Turrilites, occurring in the Upper

Denison Beds (Grayson Formation), about 200-300 ft. above the horizon of

Subschloenbachia leonensis, probably of rostrata (s.1.) date. (R. 'T. Hill, “ Geogr.

and Geol. Black and Grand Prairies, Texas,” ‘21st Ann. Rep. U.S. Geol. 8,’

(1901), p. 247).

+ ‘ Pal. Frang., Ter. Crét.,’ vol. i, pl. exl, figs. 1-3,

+ Loc. cit. (1913), p. 379.

On Cretaceous Cephalopoda from Zululand. 253

third layer is very thin between the ribs, but apparently continuous

with the material deposited on the rib-bases and forming compara-

tively high and very sharp ridges, and the whole is covered by a

fourth outer layer of shell. Where the acute costation is worn off,

the septate rib-bases show as illustrated in Wood’s fig. 5 a.

Woods compared the form with Stoliczka’s Heteroceras indicum,

which is considered by various writers to be identical with, or closely

allied to, Bostrychoceras polyplocum, Romer sp., but which, like

Turrilites saxonicus, Schliiter, may be a Hyphantoceras, not a Bostry-

choceras. On the other hand, the small constricted specimen figured

as Helicoceras indicum ? by Anderson* and which is not identical with

the Indian species, shows open coiling similar to the South African

form, but it also may be a Hyphantoceras.

Of the many forms included in Bostrychoceras polyplocum (Romer)

by Schliter,+ that figured on pl. xxxv, fig. 8, or the evolute form

figured by Geinitz{ may belong to species allied to the South African

examples. B. ? declive, Gabb sp.,§ B. japonicum, Yabe sp.,|| further, the

whorl-fragments figured by Jimbo§] as ‘ T'urrilites sp.” and by Meek**

* <Cret. Depos. of the Pacific Coast,” ‘ Proc. Calif. Acad. Sci.,’ 3rd ser., Geol.

II, i, p. 91, pl. iii, figs. 96 and 97.

+ A specimen of a Bostrychoceras sp. n.,in the British Museum (No. 74042)

from Haldem in Westphalia, comparable with Schliiter’s pl. xxxiii, figs. 6 and

8, from the same locality (Heteroceras polyplocum (A. Romer) in Schliiter) has a

costate early portion, then a constriction, and a bituberculate final portion.

This form and, perhaps, more so Schliiter’s fig. 1 of pl. xxxiv, then Bostry-

choceras ? carlottensis, Whiteaves sp. (loc. cit., p. 271, pl. xxxiv, fig. 1) and

B.? oshimai, Yabe sp. (loc. cit., 1904, p. 12, pl. iii, figs. 5 and 6, which, however,

may be a Hyphantoceras), show close resemblance in ornamentation to certain

Didymoceras, e.g. D. nebrascense, Meek sp. (B.M. No. 83907 from Upper Missouri),

and D. hornbyense, Whiteaves (B.M., Hector Collection from Vancouver Island).

The separation of fragmentary examples of the two genera Bostrychoceras and

Didymoceras, based only on tuberculation and mode of coiling, seems somewhat

artificial, as a comparison of such typical forms of Didymoceras (in Hyatt) as

D. cochleatum and D. tortum, Meek sp., with some of the tuberculate varieties

of B. polyplocum in Schliiter will demoastrate. The suture-lines, also, are of a

similar pattern in the whole family Nostoceratidae.

ft “ Das Elbthal-Geb. i. Sachsen,” II, ‘ Paleontogr.,’ vol. xx (1872-5), p. 195,

pl. xxxvi, fig. 3 only. (“ Turrilites poiyplocus var. of Helicoceras type.”)

§ ‘ Pal. of California,’ vol. i, p. 73, pl. xxviii, figs. 200, 200 a. Upper Chico in

Anderson, loc. cit. (1902), p. 27, Maestrichtian in Haug, ‘ Traité, p. 1847.

|| “ Cret. Ceph. from the Hokkaido,” ‘J1. Coll. Sci. Imp. Univ. 'l'okyo, vol. xx

(1904), p. 17, pl. ii, fig. 8.

{| “ Beitr. z. Kenntn. d. Kreidef. v. Hokkaido,” ‘Pal. Abh.,’ vol. vi (1894),

p. 41, pl. i, fig. 8.

** Loc. cit. (1876), pl. xxi, fig. 4.

254 Annals of the South African Museum.

as ‘* Heteroceras ? sp. ind.,” agree with the specimens here described in

whorl-shape and coiling, but all have oblique costation.

A fragment of a “‘ Heteroceras sp.,” from the “ Iron Mines of Hok-

kaido, Japan” (B.M., No. C104108) resembles the South African

form in coiling and whorl-section, and being a cast, in the faint

ribbing, but it is considerably larger. Its suture-line is ‘“ lytoceratid,”

like that of Pravitoceras siqgmoidale, Yabe,* which it closely resembles,

and it is associated with fragments comparable with the inner, closely

coiled whorls of Pravitocerast and with the terminal portion of such

a species of Bostrychoceras or Didymoceras, as, e.g., the variety of

“ Heteroceras polyplocus” figured by Schliiter on pl. xxxiv, figs. 2 and 3,

or D.? cooperi, Gabb, in Whiteaves.{| The age of Pravitoceras

unfortunately is not known, but it is to be noted that the suture-line

of the South African form, figured by Woods, and that of the

Turoman Hyphantoceras, are very similar to that of Pravitoceras, and

the writer is of opinion that the resemblance of all these to the suture-

line of Lytoceratidae 1s a case of convergence, correlated with the

rounded whorl-shape. NipponitesS also represents a development

related to the loosely-coiled Bostrychoceras here discussed, but the

reference of the South African form to this genus cannot, of course, be

considered definite so long as only fragmentary specimens are avail-

able. The young of Hmperoceras|| show hamitid, helicoid coiling, and

the twisted form figured by Woods | as Hamites (Anisoceras), sp.,

seems to form a transition to such species as “ Ancyloceras”’ retrorsum,

Schliiter,and to the group of *‘ Anisoceras” referred to in the description

of Diplomoceras ? indicum, which groups stand in the same relation-

ship to Bostrychoceras as (ptychoceratid, hamitid or ancyloceratid)

Oxybeloceras** does to Exiteloceras. The similarity, however, of the

* «Note on Three Upper Cretaceous Ammonites etc.,” ‘Jl. Geol. Soc.

Tokyo,’ vol. ix (1902), No. 100, p. 3, pl. i, figs. 2-4.

+ Ibid., fig. 3.

t Loc. cit., p. 336, pl. xliu, fig. 1.

§ Yabe, loc. cit. (1904), p. 20, pl. iv, figs. 4-7, pl. vi, fig. 6.

|| Hyatt, loc. cit. (1894), p. 575, pl. xiv, figs. 15-17.

¥ Loc. cit., p. 340, pl. xliv, fig. 3.

** Unless new genera are introduced again for the different forms of coiling,

such species as “ Hamites” wernickei, Wollemann (“ Fauna d. Liineburger

Kreide,” ‘ Abh. K. Preuss, L A.,’ n.¥., Heft. 87 (1902), p. 95, pl. iv, fig. 4, and pl. v,

figs. 1 and 2—crushed ?), or “Ancyloceras” bipunctatum, Schliiter (loc. cit., p. 98,

pl. xxix, figs. 1-8), will have to be included in Ozxybeloceras, whereas “ Crioceras”’

plicatilis, Kner, non Sowerby (Lemberg, ‘Naturw. Abh.,’ ii, 1850, pt. 2, p. 9,

pl. ii, fig. 3) = Helicoceras schloenbachi, Favre (‘ Moll. Craie, Lemberg,’ 1869,

p. 80, pl. vii, figs. 5 a—-c), and Helicoceras hibernicum, Tate, possibly belong to

Exiteloceras. The presence of at least one other unnamed group is indicated by

On Cretaceous Cephalopoda from Zululand. 255

young of Emperoceras simplicostatum, Whitfield sp., to Oxybeloceras

shows that the interrelations of these genera are very complicated, and

that the determination of mere fragments is very difficult.

Locality—Umkwelane Hill. Coll. Dr. A. L. du Toit.

15. BosrrycHocEras? sp. ind.

CBE PRXTV, fig. 2.)

An impression (No. 54784) of a whorl-fragment of a form allied to

the Heteroceras sp., figured by Woods* but with only three, not four,

intermediate ribs and the flares much closer together, is doubtfully

referred to the genus Bostrychoceras. In a length of about 40 mm.

there are six of these flares, as against four in the Pondoland example.

This closeness of the costation approaches the form to ‘“ Helicoceras (?)”

venustum, Yabet and “ Helicoceras” scalare, Yabe,t but these have

more numerous intermediary ribs and may possibly belong to

Hyphantoceras. The peripheral portion of the impression here

described, and its whorl-section being unknown, comparison with

“ Helicoceras” breweri, Gabb,§ and “ Crioceras (?)” cingulatum,

Schliiter,|| is difficult. They also represent similar fragments with

flares, apparently connected with the typical Bostrychoceras by such

forms as B. japonicum, Yabe sp.)

Locality.—Umkwelane Hill. Coll. Dr. A. L. du Toit. (Impression

in matrix of specimen 5478 = Diaziceras tissotiaeforme, nov. )

such forms as ** Hamites” phaleratus, Griepenkerl (“ Verstein. d. Senon. Kreide

v. Kénigslutter,” ‘Pal. Abh.,’ vol. iv (1889), Heft 5, p. 104, pl. x1, fig. 8, and pl. xii,

figs. 3 and 4), “ Ancyloceras ” pseudo-armatum, Schliiter (loc. cit., p. 164, pl. xii,

figs. 8 and 9, ? 5-7), and “ Ancyloceras” kossmati, Simionescu (“ Fauna Cret. Sup.

d. 1. Urmis.,” ‘Acad. Romana,’ Publ. Fd. Vasilie Adamachi, No. 4 (1899), p. 21,

pl. i, figs. 6-8). A beautiful, but fragmentary, example of a new form of this

group, comparable with “ Hamites ” quadrinodosus, Jimbo sp., from the Umzamba

beds of Pondoland (Coll. Geol. Survey), was sent to the writer, after the com-

pletion of this paper, through the kindness of Mr. Henry Woods. The Durban

Museum Collection, already referred to, also includes Owybeloceras ? sp., cf.

interruptum, Schliiter, and wernickei, Wollemann sp. ; further, several gen. nov.

(Hyphantoceras ?) ef. spinigerum, Jimbo sp., all from the Pondoland Senonian.

* Loc. cit. (1906), p. 339, pl. xlii, fig. 4.

+ Loc. cit, (1904), p. 11, pl. iil, fig. 4.

{ Ibid., p. 9, pl. iu, figs. 2 and 3.

§ Loc. cit., vol. i, p. 72, pl. xiv, fig. 22 (Upper Chico in Anderson, p. 27).

|| Loe. cit., p. 101, pl. xxx, figs. 18 and 14.

Sip Locnctt.opelds plats fion8:

256 Annals of the South African Museum.

Gren. DIPLOMOCERAS, Hyatt.

16. DipLoMocERAS? INDICUM, Forbes sp.

(Pl, XL, ies oz)

1895. Hamites (Anisoceras) indicus, Forbes. Kossmat. Stidind. Kreidef.

Beitr. z. Pal. und Geol. Ost.-Ung. ete., vol. ix, p. 145, pl. xix,

fig. 4.

1906. Hamites (Anisoceras) indicus, Woods. Cret. Fauna of Pondoland,

p- 340, pl. xliv, fig. 2.

This form is represented in the collection by a fragment (No. 5465)

that corresponds with Kossmat’s fig. 4a; but the hooked portion

forms the smaller end, not the larger, as in Kossmat’s figured example.

The last few suture-lines, shown on the hooked portion of the shell,

are of the general outline of that of D.? indicum as figured by Kossmat,

but have a less minutely frilled edge. The costation is closer than

it is in the fragment figured by Woods, but not so close as in Kossmat’s

specimen or in D. ? rugatum (Forbes), Kossmat sp., the cross-section of

which latter species, also, is more elliptical. The costation is very

sharp and not septate, 7. e. the ribs are as acute on the cast as they are on

the shell. One of the ribs is higher than the others (about 15 mm.

high, measured from the concavity at each side, at a whorl-diameter

of 9mim.). Specimens of D. ? large-sulcatum, Forbes sp., and D. ? ruga-

tum, Forbes sp., 1 the British Museum show similar irregularities, as

does the Japanese form, figured as ‘ Hanvites sp.,” by Jimbo* and

compared with D. ? large-sulcatum by Kossmat.t The costation is too

distant, however, in the latter species, as it is in the small fragment

of a “ Hamites” from Umkwelane Hill figured by Etheridge.{ This

was compared with Griesbach’s Anisoceras rugatum, Forbes sp., from

the Umtamvuna Beds (a form that was included by Woods in the

synonymy of D. ? indicum), but probably is nearer to D. ? large-sulcatum,

Forbes sp.

Such species as D. obstrictum Jimbo sp.,§ and D. ellipticwm,

Anderson sp.,|

seem to form a connection with the typical gigantic

Diplomoceras of the cylindricum and notabile group, but with increased

knowledge of these forms it will probably be necessary to separate

from the highly specialised Diplomoceras the Indian ‘“ Anisoceras”

* Loc. cit. (1894), p. 40, pl. vii, fig. 7.

+ Loc. cit. (1895), p. 147.

Loc, cit. (Second Report, 1904), p. 90, pl. iii, fig. 23.

In Whiteaves, loc. cit., p. 334, pl. xliv, fig. 3.

| Loe. cit., 1902, p. 87, pl. iii, figs. 102-3,

On Cretaceous Cephalopoda from Zululand. 257

assemblage on the one hand, which connects directly with the loosely

coiled Bostrychoceras above described, and on the other the various

European “ Hamites” (“ H.” vroemeri, Geinitz), “ Toxoceras” (T.

aquisgranensis, Schliiter), “ Ancyloceras” (A. retrorsum, Schliter) of

slightly earlier date and simple suture-line. Whether these Cam-

panian “ Hamitids ” are ancestral to the Maestrichtian Diplomoceras

and what their relationship is to the contemporaneous “ Ptychoceras”

and Solenoceras it is for future investigation to determine. It may

be added that a form indistinguishable from the Maestrichtian

Diplomoceras cylindricum, @Orbigny sp., occurs in the Cambridge

Greensand (Uppermost Albian), but the writer is convinced that this

is only a case of convergence, and that even the Turonian “ Hamitids ”

should be separated generically from the Senonian forms. The last

true Hamites are comparatively rare in the Cenomanian.

Locality.— Umkwelane Ball. Coll. Dr. A. L. du Toit.

Faminy: BACULITIDA.

Gen. BACULITES, Lamarck.

17. Bacuunites capensis, H. Woods.

(Pl. XXIV, figs. 6 and 7.)

1906. Baculites capensis, Woods. ‘ Cret. Fauna of Pondoland,’ Aun.

S. Afr. Mus., vol. iv, part vu, No. 12, p. 342, pl. xliv, figs. 6 and 7.

21907. Baculites vagina, Forbes in Boule, Lemoine & Thévenin.

“ Céph. Crét. Diego-Suarez,” Ann. de Pal., vol. ii, p. 65, pl. xv,

fig. 5.

This is the commonest cephalopod at Umkwelane Hill, sixteen

examples being referred to this species, in addition to a number of

fragments in the matrix of other fossils. The young is merely striate,

like B. bailyi, and the nodes first appear where the long diameter is

about 8 mm.

Woods compares the species with B. asper as figured by Morton,

Romer and Stanton, the last probably of Turonian age. The less

coarsely nodate form figured by Meek,* and a specimen of this in the

British Museum from “ Mississippi” are very close to the South

African species in all characters but the suture-line.

Some of the larger examples (No. 54794, 5403) seem to develop

coarser striation on the siphonal side, much like the example here

compared with B. sulcatus, Baily. The suture-line differs rather

* Loc, cit. (1876), p. 404, pl. xxxix, fig. 10a only.

258 Annals of the South African Museum.

considerably from that of B. incwrvatus, Dujardin,* and is closer to

that of B. fairbanksi, Anderson,t characterised by broad and low

saddles and small lobes. B. vagina, Forbes, has an entirely different

suture-linet; and in that of the probably Turonian B. gracilis,

Shumard, in Stanton,$ which also is comparatively simple, the lateral

saddle is too high and too narrow, and the second lateral lobe too

deep. On the other hand, the suture-line figured by Boule, Lemoine

and Thévenin || as that of Cyrtocheilus baculoides, Mantell sp., agrees

very well with that of the South African species, and that of B.

bohemicus, Fritsch and Schloenbach,¥ also is very similar.**

The form figured by Boule, Lemoine and Thévenin tf as Baculites

vagina (Forbes), var. otacodensis Kossmat, seems to agree much

more with Woods’ species than with the Indian form.

Localities.—Nos. 5479, 5448, 5490, 5454, 5470, 5458, 5474, 5484

(pars), 5486, 5408, 5475, 54794 and 5405 from Umkwelane Hill. Coll.

Dr. A. L. du Toit. No. 4852 from Railway Cutting, Umfolozi. Coll.

W.J. Wybergh. No. 5108 from Umkwelane Hill. Coll. J. 8. Hedges.

18. BacuuitEs sp. aff. capensis, H. Woods.

A number of poorly preserved specimens (5484 (pars), 54798, 5509B)

are comparable with the fragment figured by Etheridge ff from Umkwe-

* In Schliter (after Geinitz), loc. cit. pl. x], fig. 3.

+ Loc. cit. (1902), fig. 194, pl. x, p. 92.

t The suture-line in Steinmann (Quiriquina, loc. cit., 1895, p. 91, text-fig. 8),

is different from that of Forbes’ type (B.M. No. R10488, Geol. Soc. Coll.) and

of Indian specimens in the writer’s collection.

§ Loe. cit. (Col. Form., 1893), p. 166, pl. xxxvi, fig. 2. See also Solger (loc. cit.,

«Kamerun, 1904), text-fig. 4, on p. 102.

| Loc. cit., II (1907), text-fig. 29, on p. 65.

| In Schliiter, loc. eit. 11, 1876, fig. 5 on pl. xxxix.

** The suture-lines of the varieties valognensis and leopoliensis of B. anceps

(Nowak, “* Untersuch. Poln. Kreide.,” I, Baculites,‘ Bull. Ac. Sci. Cracovie,’ 1908,

p. 331, text-figs. 1-4 and 5-10) are of the same type, but with a more complex

ventral lobe. ‘The suture-line (drawn by the late G. C. Crick) of a Pondoland

example (B.M., No C19420), represented in fig. 7 of Pl. XXIV, is characterised by

a wider inferior lateral lobe than that of the specimen 5486, here figured (PI.

XXIV, fig. 6). On the other hand, the very similar suture-line of B. oberholzeri,

Bohm (in Béhm and Heim, “ Senonbild. d. O. Schweiz. Alp.,”’ ‘ Abh. Schw. Pal.

Ges., vol. xxxvi (1909), p. 52, pl. i, fig. 9), varies in just the opposite direction.

The minute B. vn. sp.in Jahn (‘ Beitr. z. Kenntn. d. Bohm. Kreide. Jb. K.K.R.A.,’

vol. xlv [1895], p. 136, pl. vill, figs. 8 a-c) has a suture-line very similar to that

of the Pondoland example, but the lobes are not clearly bifid.

tt Loc. cit. (1906), p. 65, pl. xv, fig. 3.

t{ Loc. cit. (1904), p. 90, pl. iii, fig. 24.

On Cretaceous Cephalopoda from Zululand. 259

lane Hill. They probably belong to Woods’ species in spite of the

apparent absence of nodes, though this lack of ornament suggests

affinity with B. bailyi, Woods (which apparently is rare, and

characterised by a very distinct type of suture-line), or with other

smooth species of Baculites.

The examples that Crick * records from the South Branch of the

Manuan Creek, apparently related to B. capensis, Woods, are similarly

poorly preserved and more or less indeterminable specimens.

Specimen No. 5484, containing at least twenty to thirty examples,

has in addition to B. capensis, and forms close to B. bailyi, a number

of more or less unidentifiable fragments that had best be included

here. One example has the mouth border complete, but no initial

whorls were discovered in this block, the reason being, perhaps, that

this coarsely sandy, conglomeratic matrix (with pebbles of fossil wood)

was not suitable for the preservation of so delicate a structure. The

mode of life in the embryonic stage, also, possibly was different

(planctonic ¢) from the mud-boring existence of the adult shells.

Locality.—Umkwelane Hall. Coll. Dr. A. L. du Toit.

19. Bacunites cf. ASPERO-ANCEPS, Lasswitz.

(Pl. XXIV, figs. 4, 4a.)

1852. Baculites anceps, Lamarck. Romer, Kreidebild. v. Texas, etc.,

p. 36, pl. ul, figs. 8b and e only.

1904. Baculites aspero-anceps, Lasswitz. ‘ Kreide-Amm. v. Texas,”

Geol. und Pal. Abh., vol. x, 4, p. 16, pl. iii (xv), figs. la and b.

A small portion of a Baculites (No. 5480), 38 mm. in length and

forming part of the body-chamber, differs from the many examples of

B. capensis that occur in the same rock, merely in having the nodes

closer, there being six in the length represented, as against half as

many in Woods’ species. The nodes are rounded, as in B. capensis,

and the cross-section also, perhaps, resembles that of B. asper, Morton

(in Romer), and of B. capensis more than it does that of B. aspero-

anceps. The form here described probably is only a variety of B.

capensis, comparable to the Texas form in the closer spacing of the

nodes. It may be added that some of the examples included in B.

capensis (e.g. No. 5490, and No. 5470) have the nodes closer than the

(larger) Pondoland examples, and thus are transitional to the form

here described.

Locality Umkwelane Hill. Coll. Dr. A. L. du Toit.

* Toc, cit. (1907), p. 240.

Annals of the South African Museum.

op)

20. Bacunrres cf. BREvicosTA, Schliiter.

(Pl. XXIV, figs. 5, 5a.)

1876. Baculites brevicosta, Schliiter. “Cephal. d. Ob. Deutsch.

Kreide,” Palaeontogr., vol. xxiv, p. 141, pl. xxxix, figs. 9 and 10.

| Non 1885. Baculites brevicosta, Schliiter, in Moberg, loc. cit., p. 37,

pl. iv, figs. 5 and 6.]

One example (No. 5461), showing nine nodes in a length of about

30 mm., apparently agrees with Schliiter’s species, but it is not

definitely identified with the species of the Emscher marls, since it

probably only represents a variety of B. capensis; that is to say, its

exact agreement with Schliiter’s species* may be a case of hetero-

chronous homoeomorphy. The suture-line agrees with that of B.

capensis, and differs from that of B. anceps as figured by @Orbignyt in

having the two lateral lobes much narrower, and in having the

siphonal portion of the ventral saddle smaller than the internal

branch, an arrangement also seen in a specimen of B. incurvatus,

Dujardin, in the writer’s collection. The suture-line of B. anceps?

figured in Schliter{ also has a comparatively large second lateral

lobe. What Schliter states to be observable in the type of B. brevi-

costa, namely, “ saddles that are considerably broader than the lobes,

and inferior lateral lobes that lie almost completely on the anti-

siphonal side,” agrees with the characters of the suture-line of the

Zululand form. B. fairbanksi, Anderson,§ though differing in orna-

ment, has the same type of suture-line as B. capensis and B. cf. brevi-

costa, but with a more minutely frilled edge.

Locality —Umkwelane Hall. Coll. Dr. A. L. du Toit.

21. Bacuires sp. cf. suucatus, Baily.

1906. Baculites sulcatus, Baily. Woods, ‘** Cret. Fauna of Pondoland,”

Ann. 8. Afr. Mus., vol. iv, part vu, No. 12, p. 341, pl. xliv, fig. 4.

One terminal fragment of a larger specimen (No.5467), characterised

by the absence of nodes, is striated like Baily’s species, but the

* Wegener (in © Die Granulat. Kreide. d. Westl. Miinsterland.,” ‘ Zeit. Deutsch.

Geol. Ges.,’ vol. lvii (1905), pp. 207 and 228), who records this species from the

zone of Inoceramus cardissoides, states that the nodes (“ribs”) are crescent-

shaped, whereas in the present example they are rounded like those of B.

capensis.

+ ‘Pal. Frang. Ter. Crét.,’ I (1840), p. 565, pl. cxxxix, fig. 7.

ft Loc. cit. (1876), pl. xl, fig. 6.

§ Loc. cit., p. 92, pl. vii, figs. 152 and 153, and pl. x, fig. 194.

On Cretaceous Cephalopoda from Zululand. 261

folds are not so coarse as they are in Baily’s original fig. 5,

selected as type of the species by Woods. Also, whereas in Baily’s

holotype (No. 11573, Geol. Soc. Coll., British Museum), and still

more so in the co-type, the folds are coarser on the dorsal than on

the ventral sides, the reverse is noticeable in the present example, so

that the latter may only be a variety of B. capensis, Woods,

resembling some of the larger examples mentioned under the descrip-

tion of that species.

B. carinatus, Binkhorst,* has a somewhat similar ventral aspect,

but in the specimen here described, the whorl section is elliptical and

evenly rounded.

Locality—Umkwelane Hill. Coll. Dr. A. L. du Toit.

22. BacuLITES BAILYI, H. Woods.

1906. Baculites bailyi, Woods. ‘“ Cret. Fauna of Pondoland,’ Ann,

S. Afr. Mus., vol. iv, part vi, No. 12, p. 341.

Three fragments (No. 5463) agree with Baily’s original speci-

ment and have merely fine striation, not nodes, agreeing in this

respect with B. fawjasi, Lamarck, in Binkhorst = B. vertebralis,

Montfort (?), and with the large forms, B. ovatus, Say, and B. grandis,

Hall and Meek. B. chicoensis (Trask), Gabb,§ a number of speci-

mens of which from Vancouver Island are in the British Museum,

has only a slightly different suture-line and altogether seems very

close.

B. syriacus, Conrad, according to a number of more or less badly-

preserved specimens in the Egyptian Collection at the British Museum,

referred to below (and associated with B. cf. teres, Forbes, and B. cf.

leopoliensis, Nowak), probably also represents a similar smooth form

of this group.

Locality —Umkwelane Hill. Coll. Dr. A. L. du Toit.

* Toe. cit. (1861), Ceph., p. 43, pl. vd, fig. 2.

+ °O.J.G. S.,> vol. xi (1855), pl: xi, fig. 5a, b (non 5c), B.M. (Geol. Soe.

Coll.), No. 11872.

t Loe. cit. (1861), p. 40, pl. vd, fig. 1. The suture-line differs only in the

width of the dorsal saddle.

§ Loc. cit. (vol. i), p. 80, pl. xvii, fig. 27 a, and pl. xiv, fig. 276 (“commonest

form, having few or no ribs”). Meek (loc. cit., 1876, Bull. ii, p. 364), includes

in the synonymy of B. chicoensis, Trask, his own B. inornatus, and Whiteaves

(‘Mesoz. Foss.,’ 1908, p. 339) also includes in Trask’s species Meek’s B,

occidentalis,

262 Annals of the South African Museum.

B. NAUTILOIDEA.

Gren. EUTREPHOCERAS, Hyatt.

23. EUTREPHOCERAS aff. DEKAyI, Morton sp.

1907. Nautilus dekayi (Morton). Stuart-Weller, Report on Cret.

Pal. of New Jersey, vol. iv (Pal. Ser.), Geol. Surv., N.J., p. 817,

pl. ¢., figs. 2-5.

1910. Nautilus dekayi (Morton), Spengler. ‘ Untersuch. ti. d. Siidind.

Kreideform. Pt. iv: Die Nautil. und Bel. d. Trichinopoly

Distr.,” Beitr. z. Pal. und Geol. Ost.-Ung., vol. xxiii, pt. iii,

p. 157.

A small and poorly preserved specimen (No. 2751), of a little over

40 mm. in diameter, and of equal thickness, seems to agree with the

typical figures cited above, and with specimens of this form from the

Fort Pierre Shale, Black Hills, South Dakota, U.S.A., in the writer’s

collection. Since, however, the present example is somewhat crushed,

so that the original shape of the whorl-section cannot accurately be

determined, the specific identification must remain doubtful.

The position of the siphuncle (centran) corresponds with that shown

in Fig. 4 of the reference given above, and not with that of fig. 1 a in

Meek* (dorsocentran). Spengler gives the thickness as 90—100 per

cent. (typically 92 per cent.), which agrees with that of the specimen

here described. ‘There does not appear to be an annular lobe.

Eutrephoceras ovoideus, G. C. Crick sp.,f probably an older

(Cenomanian) species, has a more elevated, less depressed whorl

section. Nautilus | Cymatoceras ? | occlusus, G.C. Crick,f has a similar

globose, though less depressed, whorl-shape, but the position of the

siphuncle is centroventran, and there is an annular lobe—at least in

the young. Since the Senonian also occurs at the North-West end

of False Bay, it is impossible to state whether the six examples

described by Crick are all of the same (Cenomanian) age, or even

whether they are identical or co-generic, without breaking them up.

The Antarctic example of Nautilus blanfordianus, Kilian and Reboul,§

* «Report Invertebr. Cret. and Tert. Foss. Up. Missouri Country,” in

Hayden, ‘ U.S. Geol. Surv. of Territ.,’ vol. ix (1876), p. 496, pl. xxvii.

+ Loc. cit. (1907), p. 222, B.M., No. C18253-6.

t Ibid., p. 224, B.M., No. C18257-62.

§ “Les Céph. Néocrét. d. fles Seymour et Snow Hill,” ‘ Wiss. Ergeb. Schwed.,

Siidpol. Exped.,’ yol, iii, pt. vi (1909), p. 8, pl. i, figs. 1 and 2.

On Cretaceous Cephalopoda from Zululand. 263

represents a very similar form of Hutrephoceras to the specimen here

described, with the thickness about equal to the diameter, but the

siphuncle is ventrocentran in the Antarctic form, as it is in the far less

depressed N. hualeyanus, Blanford,

Locality.—Umfolozi Valley, East of Railway. Coll. J. L. Dling-

worth.

24, HUTREPHOCERAS cf. SUBLAEVIGATUM (d’Orbigny) var. INDICA

(Spengler),

1861. Nautilus bouchardianus, @Orbigny. Blanford, Cret. Fauna S.

India (Pal. Indica), vol. i, ‘“‘ Cephalop.,” pl. v, fig. 3.

1910. Nautilus sublaevigatus var. indica, Spengler. ‘ Untersuch. ii. d.

Siid-Ind. Kreideform. Pt. iv. Die Nautil. und Bel. d. Trichi-

nopoly Distr.,” Beitr. z. Pal. und Geol. Ost-Ung., vol. xxiii, pt. iii,

DD. Lor.

A nearly complete but slightly weathered example of a Nautilus

(No. 5509) agrees well in proportions with the large specimen figured

by Woods,* and at a diameter of 150 mm. has a thickness of

110-115 mm. The sides and ventral area, however, are more

flattened (weathered ?), giving the whorl a more quadrate shape. The

last half whorl of the specimen represents the body-chamber, with

indications of a mouth-border at the end. Several specimens of

Baculites are embedded in the matrix of this body-chamber.

Since the specimen was not broken up to reveal position of the

siphuncle, presence of an annular lobe, etc., the identification with the

Indian form of N. sublaevigatus, with which Woods also had compared

his Pondoland example, must remain doubtful. It is based on

similarity of whorl-shape and dimensions and general agreement with

Blanford’s fig. 3.

The probably Cenomanian Hutrephoceras ovoideum, G. C. Crick sp.,t

has a more elevated whorl-section, and Nautilus | Cymatoceras? |

ocelusus, G. C. Crick, is too inflated in the umbilical region.

Some of the Nautilus (Hutrephoceras) sp., described by Crickt from

the South Branch of the Manuan Creek, Zululand, may belong to the

present Senonian species.

Locality —Umkwelane Hill, Umfolozi, Zululand. Coll. Dr. A. I.

du Toit.

* Loc. cit. (1906), p. 330, text-fig. 1 on p. 331,

+ Loc. cit. (1907), p. 222.

t Loc. cit. (1907), p. 245, B.M., Nos, C18292-5.

264 Annals of the South African Museum.

Gren. CYMATOCERAS, Hyatt.

25. CYMATOCERAS? sp. juv. cf. VALUDAYURENSE, Blanford sp.

1861. Nautilus valudayurensis, Blanford. Cret. Fauna 8. India (Pal.

Indica), vol. i, “ Cephalop.,”’ p. 23, pl. xu, figs. 2-3.

1866. Nautilus valudayurensis, Stoliezka, ibid., p. 206.

A small and fragmentary specimen (No. 5469), consisting of the

casts of four camerae in a very good state of preservation, agrees with

Blanford’s figures, especially in the outline of the sectional view

(fig. 3), in position of the siphuncle (centrodorsan), and the presence

of a small annular lobe. The decussate ornament figured in 2 b also

is well shown in the dorsal impression, but on the outer whorl only

striae of growth are visible, not the coarse folds typical of Cymatoceras.

On the other hand, the presence of the linguiform annular lobe shows

the specimen to be distinct from Hutrephoceras,* some examples of

which (e. g. H. dekayi (Morton), var. montanaense, Meekt) agree with

it in whorl-section. Of the various forms included by Spenglert in

Nautilus | Cymatoceras| aft. atlas, Whiteaves, the small ammonite

figured by Blanford§ somewhat resembles the fragment here de-

scribed, but the siphuncle is ventrocentran in the Indian specimen, not

centrodorsan.

The small Nautilus sp. described by Crick) has a similar septal

surface, with annular lobe, but a more depressed section, and probably

is of Albian age, as also is Cymatoceras manuanense, Crick sp.| The

large examples of this species have no annular lobe, but agree in

section and position of siphuncle.

Locality —Umkwelane Hill, Umfolozi, Zululand. Coll. Dr. A. L.

du Toit.

OBSERVATIONS ON THE UMKWELANE Hitt Fauna.

To the Cephalopoda from Umkwelane Hill, described in the fore-

going pages, must be added three Ammonoids recorded by Etheridge,

namely :

Placenticeras kaffrarium, Etheridge.

a umkwelanense, Etheridge.

Diplomoceras ? cf. large-suleatum, Forbes sp.

* Hyatt, “Phylogeny of an Acquired Characteristic,” ‘Proc. Am. Philos.

Soc.,’ xxxii, No. 148, 1894, appendix, p. 555.

+ Loc. cit. (1876), p. 498, pl. xxvii, figs. 2-2 f.

t Loe. cit. (1910), p. 135.

§ Loe. cit. (1861), pl. viii, fig. 4.

|| Loc. cit. (1907), p. 248, B.M., No. C18310.

{ Ibid., p. 243, pl. xv, fig. 6, B.M., Nos, C18282-5,

On Cretaceous Cephalopoda from Zululand. 265

Further, the typical—

Mortoniceras umkwelanense, Crick,

so that up to the present twenty-nine species and varieties have been

described from this locality.* The form figured by Etheridge as

“ Oreniceras (?) sp. ind.” is not included, since its systematic position

is quite uncertain, and since it may not even be a cephalopod.

Woods and Newton considered the Umkwelane Hill fauna to be of

the same age (Campanian) as that of Pondoland. The occurrence, at

Umkwelane Hill, of a form (Mortoniceras woodsi, nov.) that is very

close to M. delawarense, confirms the presence of the Campanian, and

there certainly is no indication of any Cenomanian or “ Vraconnian ”

admixture in this fauna, as suggested by Lemoine.t On the other

hand, the Maestrichtian, or part of it, may also be represented in

South Africa. Parapachydiscus of the colligatus type are quoted both

from Campanian and Maestrichtiant deposits; and Placenticeras

umkwelanense, compared by Etheridge with P. placenta, also the two

Nostoceratids, recall Maestrichtian forms. According to Woods,

Pseudophyllites indra occurs in the basement bed of the Pondoland

deposit ; and Haug calls the ‘“Anisoceras” and Trigonoarca beds of

the Valudayur group (with Pseudophyllites indra) Maestrichtian, but

in the writer’s opinion, the many large Mortoniceras, characteristic of

South Africa, are pre-Maestrichtian.

The oceurrence, in the Pondoland Collections, of these Mortoniceras

in the same blocks with Hauericeras gardeni and with Pseudoschloen-

bachia, makes it probable that they are, indeed, Upper Senonian. De

Grossouvre’s contention that the Pondoland deposits are of Lower

Senonian age and somewhere near the limit of the Coniacian and

Santonian divisions has been questioned by Woods,§ who stated that

“the probability that one zone only is represented is supported by the

observations made by the Survey that most of the species range through-

out the deposit, as well as by the small thickness of that deposit.”

* Newton (loc. cit., p. 96) recorded the occurrence of a Baculites, closely

resembling B. bailyi, Woods, in the matrix of an Umkwelane Hill specimen.

+ «Etudes Géol. dans le Nord de Madagascar,’ Paris, 1906, p. 396. On this

page Lemoine puts part of the Umkwelane Hill beds (with Mortoniceras and

“ Anisoceras”’) as equivalent to the Utatur beds of Southern India, but on p. 403

he classes the fauna, described by Etheridge, as Senonian, whereas in the table

on p. 405 the former beds (with Mortoniceras and “ Anisoceras”) are, perhaps

through a slip, included in the Turonian.

t{ The specimen here described as P. n. sp. aff. colligatus shows very good

agreement with a typical French example (B.M., No. C524), except that it is

thicker.

§ Loc. cit. (1906), p. 346.

266 Annals of the South African Museum.

At Umkwelane Hill, as in Pondoland, it is chiefly the resemblance

of the Mortoniceras to M. tevanum, and of Pseudoschloenbachia to

Grossouvre’s Coniacian-Santonian forms that suggests the presence of

pre-Campanian horizons.

The additional evidence, however, is not very satisfactory so far as

exact dating of the beds within the Senonian is concerned ; and it is

hardly safe, from the evidence available, to assume the complete

absence of pre-Campanian horizons at Umkwelane Hill. Placenticeras

subkafirarium, nov., is close to Pl. tamulicum, Kossmat, which occurs

in the Upper Trichinopoly group of India; and at the Manuan Creek,

this new form, or a close ally, is associated with Kossmaticeras

(Madrasites) bhavani, Stoliczka sp., also common to the Upper

Trichinopoly and the Aryalur groups. Now the former group includes

Peroniceras dravidicum, Kossmat sp., which is here described from

the junction of the Manuan and Umsinene Rivers, whereas two other

forms of Peroniceras (P. cf. czirnigi, Redtenbacher sp., and P. cf.

rousseaual, Grossouvre) were included in the collection described by

the late G. C. Crick. The presence of the Coniacian in Zululand is

thus established.

Again, the new genus Diaziceras, the type-specimen of which is

associated in the same block with Bostrychoceras ? sp. ind., a form

that is comparable with certain Upper Senonian types of the Hokkaido

and of California, is here considered to be related to Pseuwdoschloen-

bachia papillata from Pondoland, but in suture-line the new genus is

very close to Lenticeras andii, Gabb sp., or L. baltai, Lisson, whereas

“ Barroisiceras’’ desmoulinsi, Grossouvre sp., resembles it very much

in external characters. Both these genera are of Lower Senonian

age; Barroisiceras occurs associated with Peroniceras in Madagascar

and the Cameroons; Lenticeras is associated with Mortoniceras

texanum in South America.

It may also be pointed out that the new collection of Pondoland

fossils belonging to the Durban Museum, and referred to in the intro-

ductory part of this paper, contains ‘‘ Puzosia” sugata, Forbes sp., and

an Ammonite resembling the Madagascar example of “B. haber-

fellneri (Hauer), figured by Boule, Lemoine and Thévenin.* This

new form may, perhaps, be more nearly related to certain Upper Chico

types, e.g. “ Schloenbachia”’ chicoensis (Trask), Anderson,+ though

similar “‘ Barroisiceras”’ also occur in the Lower Chico formation, but

are not satisfactorily separated from numerous forms that may be true

Prionocyclus. ‘‘ Puzosia” sugata, also, is recorded from the Lower Chico

* Toc. cit. (1907), ii, p. 43, pl. xi, fig. 3 only.

+ Loc. cit. (1902), p. 116, pl. ii, figs. 23-25.

On Cretaceous Cephalopoda from Zululand. 267

formation® ; but in India it occurs in the Lower Aryalur and Upper

Trichinopoly groups, and Haug? has it, both in the Santonian and in

the Maestrichtian, on the same page.

The difficulties of exact correlation were probably felt by Kilan and

Reboul,} who put the beds of Snow Hill and Seymour Islands, that

both contain Kossmaticeras (Madrasites) bhavani, into the Senonian

s.l. (= Santonian to Maestrichtian), placing the lower horizon as

equivalent to the Indian Upper Trichinopoly group, whereas the

upper beds show close affinity, not only with the Aryalur and

Valudayur groups, but also with the Campanian or Maestrichtian

deposits of Southern Patagonia. The fact that Placenticeras, similar to

the Zululand forms, occur in the Maestrichtian Fort Pierre Shale of

America, perhaps, is in favour cf the attribution of the fauna here

discussed, and of Pondoland, to this upper horizon of Antarctica ;

and it may be added that the two forms here described from the

north-west shore of False Bay (Mortoniceras vanuxemi, Morton Sp.

and Bostrychoceras’ sp.) also are of Campanian age; further, that

the isolated specimen of Peroniceras cf. dravidicui, representing a cast

in limonite, after pyrites(?), differs in mode of preservation from the

Ammonites of the Umkwelane Hill fauna as from the two forms of

Peroniceras described by Crick. If not all, at least the great majority

of the forms of the Umkwelane Hill and Pondoland Ammonite

faunas probably are of Campanian (and Maestrichtian ?) age, and in

his phylogenetic interpretation of the genera Psewdoschloenbachia and

Diaziceras, the writer assumed their Upper Senonian age. But the

occurrence, at about the limit between the Coniacian and Santonian—

at which level the Pondoland fauna had been placed by Grossouvre—

and associated with Mortoniceras texanum, of forms like Lenticeras,

Barroisiceras§ and what the writer considers to be Gauthiericeras

developments (bertrandi-fournieri group), attords the most striking

* Kihan and Reboul (loc. cit., p. 60) quote it as Upper Chico, but Anderson

(pp. 27 and 98) distinctly characterises it as a Lower Chico species.

¥ Loc. cit., II, ii, p. 1842.

t Table on p. 58, loc. cit., also p. 59. In this Antarctic fauna, also, the great

majority of forms are Upper Senonian. The little-known genera Grahamites

and Seymourites show a very striking resemblance to certain Canadian Fort

Pierre forms in the British Museum, including A. barnstoni, Meek (‘Sas-

katchewan Exploring Expedition: Geolog. Report,’ H. Y. Hind, Toronto,

1859, Chapter XIX (by F. B. Meek), p. 197, pl. ii, figs. 1-8).

§ E.g. B. dentuto-carinatum, Roemer (Hill), a form very near to “ Schloen-

bachia”’ siskiyouensis, Anderson (Lasswitz, p. 29, thought them identical), which

perhaps resembles the Pondoland form, referred to above, as much as does the

Upper Chico “ Schl.” chicoensis (Trask) Anderson.

268 Annals of the South African Museum.

parallel to the Mortoniceras-Diaziceras-Pseudoschloenbachia assemblage,

here recorded from Umkwelane Hill.

It may be added here that whereas, in East Africa, the succession

from the Bathonian up to the Aptian* is represented by generally

ammonitiferous deposits, in Zululand, as probably also in Mozam-

bique,t there is a fairly complete succession from the Aptian to the

Maestrichtian, with the exception of the Turonian, the presence of

which in Madagascar also has not been clearly demonstrated. It is

of interest to note that the Coniacian Peroniceras, mentioned above,

and which is almost indistinguishable from a Bohemian P. subtri-

carinatum, VOrbigny sp.,f is closely comparable with a type that

occurs in India and Madagascar and has also been recorded from the

Cameroons. Marine connection across Africa certainly did not exist,

and the writer thinks the evidence’ favours Lemoine’s§ contention

that the communication between the Indian and Mediterranean seas

did not, as Kossmat thought, take place vid the south of the African

continent. The genus Peroniceras also occurs in Tunis and in the

Egyptian-Syrian Coniacian, and the Turonian faunas (with Fagesia

and Neoptychites) of Tunis and India are closely allied. Unworked

Nigerian collections with Pseudotissotia, Vascoceras, etc. (Falconer,

Kitson, and Temple Colls., British Museum), show that during the

Turonian (as during the Albian) there was connection between the

Cameroons Bay and the great sea that covered the whole of

the Sahara || and extended across to India, but no further (‘Turonian)

extensions down the east or west coasts of Africa, can be traced by

ammonitiferous deposits, though a different facies may represent the

Turonian both in Angola and in Madagascar. Peroniceras dravi-

dicum, thus, probably came to the Cameroons by way of Tunis, and

not vid South Africa. The distribution of this form, therefore, is not

* See Zwierzycki, “Ceph. Faun. d. Tendaguru-Sch. i. Deutsch-Ostafrika,”

loc. cit. (1914), pp. 90-91 ; also Spath, “ Jurass. Amm. fr. E. Africa,’ ‘Geol. Mag.,’

vol. lvii (1920), pp. 311-20, 351-62.

+ The Dipoloceratidae of the Albian are poorly represented there, and

“ Mortoniceras cfr. candollei” in Choffat (‘Conducia,’ 1908, p. 24, pl. vi, figs. 3

and 4) cannot be definitely identified as a form of the Upper Albian candolli-

anus group.

t B.M. No. 88991, Coll. Dr. Fritsch.

§ ‘Et. Géol. Nord de Madagascar,’ Paris, 1906, p. 397.

|| A collection of 'Turonian Ammonites from Sinai (‘l’. Barron Coll.), described

in an unpublished paper by Crick, contains Hoplitoides ? and Vascoceras.

§ According to Kilian and Reboul (loc. cit., 1909, p. 64), this Turonian

Mediterranean, extending from Brazil to India, did not communicate directly

with Madagascar.

On Cretaceous Cephalopoda from Zululand. 269

of great significance, and, at any rate, there is no record from the West

Coast of Africa, of an ammonite indicative of the great Campanian

transgression which left the deposits of the Indo-Pacific type (with

Kossmaticeras and Lytoceratidae), and of the Atlantic type (with

Mortoniceras and Placenticeras) discussed by Kilian and Reboul.*

Of the twenty-nine species of Cephalopoda of the Umkwelane Hill

fauna, only one Ammonite (Pseuwdoschloenbachia wmbulazi) is identical

with a Pondoland species, in addition to a number of uncoiled and

straight forms (Bostrychoceras?, Diplomoceras’, Baculites), and to a

Nautilus. The assemblage of these genera suggests close affinity of this

South African fauna with that of the Egyptian Maestrichtian +; but the

place of Pseudoschloenbachia umbulazi, there, is taken by a new species

(doubtfully classed as a Cenomanian Schloenbachia by Blanckenhorn

(in Coll.|), whereas the plentiful Hutrephoceras desertorum, Zittel sp.,

replaces the form here described as FE. aff. dekayi, Morton sp. The

abundance of Bacuwlites and the frequent occurrence of uncoiled

forms, in the Egyptian as well as the South African deposits, are

further points of similarity ; and it may be noted here that Mortoni-

ceras of the delawarense group are common to Tunis and Zululand,

and that Parapachydiscus colligatus, also, has been recorded from

Tunis as well as from Madagascar.

The two genera Mortoniceras and Parapachydiscus, of course, are

other elements common to the two faunas, even if the species are

different, and it may be recalled here that Newton{ found “ very few

of the shells [from Manuan Creek] to occur in contiguous areas,

suchas . . . UmkwelaneHilland . . . Pondoland. a

The great abundance of large forms of Mortoniceras, with the equally

frequent occurrence of Hauericeras gardeni, and the presence of the

(less common) Pseudoschloenbachia, form the characteristic feature of

the Pondoland deposits, and distinguish them from the Egyptian and

Madagascar faunas. In the “new collection of Natal fossils, at the

Natural History Museum, far surpassing all collections hitherto

made” (Kossmat$), out of a total of 105 specimens, 43 are Mortoni-

ceras. But it is significant that Hawericeras gardeni, a typical Indo-

* Loc. cit. (1909), pp. 64-5. Through the kindness of Mr. Beeby Thompson, of

Northampton, the writer has lately (February, 1921), been able to study a new

collection from Angola, including Upper Senonian Ammonoids (Didymoceras):

+ In Haug (loc. cit., p. 1385), who considers the presence of the Campanian

to be doubtful, so that the great Upper Senonian transgression may here have

been of a slightly later date.

t Loe. cit. (1909), p. 95.

§ ‘Rec. Geol. Surv. India,’ vol. xxviii (1895), pt. ii, p. 43. See also Newton,

loc. cit. (1909), p. 14.

270 Annals of the South African Museum.

Pacific element, of which the latter collection includes no fewer than

thirty-seven specimens, is unrepresented at Umkwelane Hill, as are the

genera Gaudryceras, Tetragonites and Pseudophyllites, the last repre-

sented in the new collection by three specimens, one of which reaches

the diameter of 290 mm.

The gigantic Parapuzosia and the Placenticeras, also, are represented

only in the Umkwelane Hill fauna, and though oceurring in Mada-

gascar, are not known from Pondoland. The Zululand locality also has

the unique Diaziceras as a strictly local type, whereas in Pondoland

Bulophoceras and its close ally Spheniscoceras form special develop-

ments, not occurring elsewhere.

It is clear that this indicates a difference of facies, the stenothermal

Lytoceratidae being dependent on deeper water or warm currents.

Lithologically the difference is indicated by the absence of glauconite

in Zululand ; and what changes of facies may be observed in a distance

equal to that separating the Pondoland Umzamba beds from Umkwe-

lane Hill, is seen when comparing the deposits of the warm coralline

Gosau Sea with the contemporaneous beds left by the colder Chalk

Sea. These Lytoceratidae are absent also in Egypt and Baluchistan,

but occur in Madagascar and Southern India, and, as regards Pondo-

land, their distribution cannot be said to support the view put forward

by Grossouvre (and not accepted by Woods), that there was at least

as close a relationship to the fauna of the Chalk of Europe as to that

of Southern India. The fact that Mortoniceras, which forms one of

the most important elements of the South African faunas, is absent in

India, of course constitutes a striking difference, and might unduly

encourage comparison with corresponding European assemblages such

as those of Galicia or Poland, where Hawericeras gardeni and Koss-

maticeras (?), two Indo-Pacific elements, occur. Taking the South

African fauna as a whole, however, its affinity with the Indo-

Pacific fauna is undeniable, and, as has been mentioned before, the

“Atlantic”? type of deposit (with Mortoniceras and Placenticeras)

found in Zululand, and the Indo-Pacific type (with Lytoceratidae and

Kossmaticeras) occurring in Pondoland,* are connected by the presence

in both faunas of Pseudoschloenbachia, probably an active swimmer,

and of benthonic crawlers (Diplomoceras, Bostrychoceras) and mud-

boring Bacuwlites.

Of particular interest, perhaps, are the two turricones of the

* It is interesting to note that the additional Ammonites from Pondoland,

lately described by Dr. van Hoepen, and, with one or two exceptions, the

fauna sent by the Durban Museum, consist of such “ Indo-Pacific” types,

unknown at Umkwelane Hill.

On Cretaceous Cephalopoda from Zululand. 271

Umkwelane Hill fauna, partly because they represent benthonic types

of limited powers of migration, compared with the oxycone develop-

ments of the Upper Senonian, that might be thought to have been

active swimmers, but are often curiously restricted, just as other marine

organisms often may have a limited horizontal distribution.* Their

nearest allies, hitherto described, are European, Japanese and North

American forms, but in an unworked Egyptian Collection in the British

Museum there are, besides Bostrychoceras, which also occurs in Tunis,

Baluchistan, India and Madagascar, fragmentary Nostoceratids, com-

parable with the Zululand species and with Fort Pierre types from

the United States and from Canada.t These forms again point toa

direct connection with North Africa, as did Mortoniceras, and it appears

probable that ever since the breaking up of Suess’s Gondwanaland, or

at least from the time of the Aptian transgression, the Zululand

Cretaceous Sea was open not only to Antarctic-Pacific elements

coming from south-west, but was in direct communication, through

the Mozambique Channel, with the sea to the northt that led to the

Mediterranean on the one hand and to India on the other.

Boule, Lemoine and Thévenin§ stated that there was a gradually

diminishing number of forms common to the North African and the

Madagascar faunas, as the beds became higher in the Cretaceous

succession ; and they noted the absence of Tissotia, which constitutes

an important element in the North African fauna. Such a relation,

perhaps, is also to be observed in Zululand; but the figures, at any

rate, prove little, considering the differences in the facies of two

neighbouring areas such as, e.g., Pondoland and Zululand. In the

much more completely known Tunisian fauna, both the “ Atlantic”

and the ‘ Indo-Pacific” types (though the latter without Kossma-

ticeras) are represented. If, however, the Umkwelane Hill fauna be

compared with the neritic Egyptian or European faunas and the

Pondoland fauna with a corresponding bathyal|| assemblage of North

‘

* Mr. S. S. Buckman (“Jurassic Chronology: I, Lias,” Suppl. I, ‘Q.J.G.S.,

vol. lxxvi, pt. i, 1920, pp. 66-67) considers that analogy with modern organisms

does not hold, but his remarks are unconvincing, as a study of the distribution,

and dependence on facies, of the two fundamental stocks of Ammonites

(Lytoceratide and Phylloceratidx) will show.

Eziteloceras cf. angulatum (Meek), Didymoceras ?, sp. (cf. Heteroceras poly-

plocum, Romer sp. (pars), in Schliiter, 1872, p. 112, pl. xxxiv, fig. 1 only). Com-

parable forms have also been discovered in Angola (see footnote on p. 269).

+ In Krenkel’s sense (“ Unt. Kr. v. D.-Ostafr.,” ‘Beitr. Pal. Ost.-Ung.,’ vol.

xxiii, 1910, p. 249).

§ Loc. cit. (1907), p. 71.

|| The term “ bathyal” is misleading, for Lytoceratidae, e. g., may occasionally

occur in comparatively shallow-water deposits ( jwrense-zone).

272 Annals of the South African Museum.

Africa or Europe, the agreement may be found to be less close than it

was during Albian times.

Uhlig* thought that the aspect of the [Jurassic and Lower Creta-

ceous | fauna as a whole justified the establishment of an Ethiopian

province by Neumayr, later regarded by Dacqué and Krenkel as a

sub-area of the Indian Province. In the Upper Cretaceous, this

“Ethiopian Province”’ had lost its individuality, if it ever formed a

separate province ; for, e.g., Haug points to the presence of the peculiar

genus Bouleiceras in Madagascar as possibly indicating a separate

zoological province, but the writer has found a specimen of B. nitescens,

Thévenin, in a Domerian-Toarcian collection from Baluchistan.t At

any rate, in the Upper Cretaceous, the Indo-Malgascan fauna shows

the closest relations with those of the Pacific and Antarctic provinces ;

and the most characteristic element of this vast ‘ Indo-Pacific ”

province is the genus Kossmaticeras, as pointed out by Haug.

Through a slp, this author also stated$ that Kossmaticeras was not

known either in Madagascar or in South Africa, whereas on pp. 1544

and 1355 he quotes it from both localities.

Under the description of Kossmaticeras (Madrasites) bhavani,

Stoliczka sp., in the Manuan Creek fauna, the writer has referred to

various South African forms, and other hitherto unrecorded species of

Kossmaticeras and allied genera from New Zealand, and the presence

of this genus in South Africa shows the fauna to belong to this

great “ Indo-Pacific Province’’|| in spite of the number of “ Atlantic”

types introduced from the North vid Egypt.

* “Marine Reiche d. Jura and d. Unterkr.,” ‘ Mitt. Geol. Ges. Wien,” iv

(1911), 8, p. 406.

+ British Museum (Geol. Society Coll.), from Valley of Kelat, Baluchistan,

together with Phylloceras, Rhacophyllites, Lytoceras, Fuciniceras, Protogrammo-

ceras, Dactylioceras, ete.

t Loe. cit., vol. i1, 2; p. 1369.

§ Ibid., p. 1369.

|| The communication, to the West, with Graham Land, Southern Patagonia

and Chili, is perhaps even more certain, and in all these areas, as pointed out

below (p. 807), the deposits consist largely of glauconitic, calcareous sandstones ,

contain the same fossil assemblages, and apparently pass uninterruptedly into

the lower Eocene.

On Cretaceous Cephalopoda from Zululand. 273

Ill. THE MANUAN CREEK FAUNA.

DESCRIPTION OF SPECIES.

A. ALBIAN.

1. AMMONOIDEA.

Famity: PHYLLOCERATID AS.

Gren. PHYLLOCERAS, Suess.

1. PHYLLOCERAS VELLEDAE, Michelin sp.

1906. Boule, Lemoine & Thévenin: ‘Pal. de Madagascar, III,

Céph. d. Diego-Suarez,’ Ann. de Pal., vol. i, fase. 4, p. 7, pl. i,

fig. 11.

1907. Crick. Third Report, p. 286 (Phylloceras sp.), B.M., No.

C18264.

This well-known form is represented by a fragment (No. 4992)

completely septate, of a large example, agreeing both with the Zulu-

land and Madagascar specimens and with d’Orbigny’s* and Pictet’st

European types, in whorl-shape, ornamentation and suture-line. At

a diameter of 150 mm. the thickness is 47 mm., or 36 per cent. of the

diameter, which agrees with that of d’Orbigny’s type. The fragment,

being nearly half of the Ammonite, shows the inner whorls in section,

and at a diameter of 54 mm. the thickness still is 36 per cent.,

whereas the example figured by Crick as P. velledae, Stoliczka (non

Michelin ?), has very flat sides, considerably more compressed than

Stohezka’s large example.§ Crick’s figured specimen, which also

shows striation only on the periphery, though the sides are weathered,

probably also is of later (7. e. Cenomanian) age, and may be closer to

the variety figured by Boule, Lemoine and Thévenin in fig. 10.|| On

the other hand, the Phylloceras sp., from the South Branch of the

Manuan Creek, recorded by Crick, is identical with the form here

described.

Locality.— Middle Branch, Manuan Creek. Coll. W. J. Wybergh.

* Loc. cit. (1840), p. 280, pl. Ixxxii.

+ In Pictet and Roux, loc. cit. (1847), p. 30, pl. ii, fig. 1.

t Loc. cit. (Third Report, 1907), p. 166, pl. x, fig. 11 (B.M., No. C18137).

§ Loc. cit. (1865), p. 116, pl. lix, fig. 2.

|| Of pl. i. This is dated, in explanation of plate, as “Cénomanien

supérieur,” but in the text (p. 8) the horizon is given as “ Sénonien inférieur.”

274 Annals of the South African Museum.

Famity: DESMOCERATID AS.

GeN. PUZOSIA, Bayle.

2. Puzosta cf. BHiImA, Stolicezka sp.

Compare :

1865. Amm. bhima, Stoliczka. Cret. 8. Ind. vol. i, p. 157, pl.

Ixix, fig. 2.

1898. Puzosia bhima, Kossmat. Unters. S. Ind. Kreidef. pt. in,

Beitr. Pal. und Geol, Ost.-Ung., ete., vols xi; p19:

1907. Puzosia pinguis, Crick. Third Report. Geol. Surv. Nat. and

Zulul., p. 218.

A fragmentary example (No. 4907), originally of about 200 mm.

diameter, but still septate at the end, shows close agreement with

Stohezka’s large specimen and with the very similar False Bay form.

The dimensions probably were :

Height of the last whorl . 46 per cent. of the diameter

Thickness fe Bs 5 Bh) ; 99 ”

Umbilicus f : ean Pr i 5.

On the test, which is 1°5 mm. in thickness, the varices are not

shown, but there is a labial ridge across the periphery, corresponding

with a suleus on the cast, though this sulcus extends across the sides

as well. In addition to these ridges, which form a less acute sinus

on the venter than they do in P. bhima or P. pinguis, there is faint

and irregular striation on the test. The presumably Cenomanian P.

subtilis, Crick,* has very similar ornament, but is more compressed

(thickness = 31 per cent.) and more involute (umbilicus = 20 per

cent.). P. pinguis, Crick, with a thickness of 36 per cent., appears to

be merely an inflated form of the subtilis type; but since it may be

of Cenomanian age, and since the suture-line cannot be compared, its

exact relations to P. bhima must remain somewhat uncertain. The

larger fragment in the British Museum (No. C18243) is extremely

close to the specimen here described, and differs, apart from slightly

varying proportions, only in having more acutely linguiform ridges

on the periphery—a character in which it is nearer to the Indian

species than to the specimen here described.

P. insculpta, Kossmatt is more compressed, but the large examples

* Loc. cit. (Third Report), p. 217, pl. xiv, figs. 5, 5 a.

+ Loc. cit. (1898), p. 120, pl. xviii, fig. 5.

On Cretaceous Cephalopoda from Zululand. 275

from Madagascar figured by Boule, Lemoine and Thévenin* are very

close to the present specimen. On the other hand, the writer would

doubt whether Sharpe’s P. octosulcata,t included by Pervinquitre {

in P. mayoriana, VOrbigny sp., is as close to P. bhima as Kossmat

thought. P. compacta, Crick,§ from the Middle Tributary of the

Manuan Creek, and probably of the same age as the form here

described, i.e. Upper Albian, may be the young of a form of the

planulata group, comparable with P. octosulcata. P. concinna, Crick, ||

from the same locality, was stated to come nearest P. bhima, but has

only five constrictions, distinct costation at a small size, and a far

less conspicuous forward sweep of the sulci, so that this species also

has closer relations with the planulata group than with the insculpta-

bhima group.

Locality.—Middle Branch, Manuan Creek. Coll. W. J. Wybergh.

Gren. UHLIGELLA, Jacob.

3. UHLIGELLA? sp. nov. aff. sroLiczKa1, Kossmat sp.

Compare :

1865. Amm. beudanti, Stoliezka. Cret. S. Ind., vol. i, p. 142, pl. lxxi,

figs. 2-4, non fig. 1.

1898. Puzosia stoliczkai, Kossmat. ‘ Untersuch. 8S. Ind. Kreidef.,”

pt. ili, Beitr. Pal. und Geol. Ost.-Ung., xi, 3, p. 119, pl. xviii, fig. 6.

1907. Puzosia stoliezkai ; Crick. Third Report Geol. Surv. Nat. and

Zulul., p. 216.

1908. Puzosia (7) stoliczkai; Jacob. “Et. Pal. and Strat. Part. Moy.

Ter. Crét.,” Trav. Lab. Géol. Univ. Grenoble, vol. vin, p. 350.

The specimen (No. 4902) that is referred to this species is some-

what fragmentary, but permits of the dimensions being measured at

the following two Ciameters :

Diameter ‘ : : . 120 mm. 54 mm.

Height of last whorl : . 46 per cent. 41 per cent.

Thickness _,, = : ogee SOY ots:

Umbilicus : ; : . 24 30

” 9

* Loc. cit., I (1906), p. 19, pl. iii, figs. 2-4. These authors state that P.

stoliczkai has a notably wider umbilicus than P. inscu/pta, whereas Kossmat’s

figures are 22 per cent. and 25 per cent. respectively for the umbilical widths

in these two species.

t+ Loe. cit, (1856), p. 42, pl. xix, fig. 3.

t Loe. cit. (1907), p. 157; but see the writer’s “ Cret. Amm. from Angola,

to be published shortly.

§ Loe. cit. (1907), p. 246, pl. xv, fig. 7 (B.M., No. C18307).

|| Loe. cit. (1907), p. 245.

276 Annals of the South African Museum.

These figures indicate that with increase in size, the umbilicus

becomes narrower, whereas the Indian examples appear to be more

involute in the young. The inner whorls of the present example are

smooth, as are Stoliczka’s specimens; but on the periphery of the

outer whorl costation appears, between the numerous sulci, as in

P. compressa, Kossmat.* A further point of difference is the slight

flattening of the sides in the Zululand example, combined with a

wider venter, making the whorl-section more rectangular than that of

Stoliczka’s larger example (fig. 3 a), i.e. more like that of his fig. 2a.

The umbilical slope, also, is inclined, not perpendicular, or even

overhanging, as in Stoliczka’s species, and agrees with that of Crick’s

specimen.

The constrictions, however, are very similar to those of Kossmat’s

species, and it may be recalled that Crick already had described his

example as being slightly more compressed than Stoliczka’s fig. 3 a.

The False Bay example also has, at a diameter of 82 mm., practically

the same dimensions as the larger specimen here described.

Etheridge? figured a very large “ Desmoceras sp.” from the Albian

of the Umsinene River, in the neighbourhood of the Manuan Creek,

and thought it possibly allied to the ‘ Ootatur form of D. beudanti

(Brongniart) ” = P. stoliczkai, Kossmat, 1898. The presence of cos-

tation might suggest that it is a large example of the form here

described, in which ribbing appears near the end ; but the constrictions

appear to be quite different, and it is probable that Etheridge’s form

is related to the forms of the planulata-group, with straight constric-

tions, found in Angola.

Puzosia of the insculpta-bhima group, referred to above, have a

different course of the constrictions, and, the costate planulata group

is too evolute. The reference of the present form to Uhligella, after

Jacob, the author of the genus, appears somewhat doubtful.

Locality.—Middle Branch, Manuan Creek. Coll. W. J. Wybergh.

* = 4. durga, Stoliezka, non Forbes, loc. cit., pl. 1xxi, fig. 7.

+ A specimen, not noticed by Crick in his paper, but labelled by him ? Beudan-

ticeras beudanti, Brongniart sp. (B.M., No. C18303), is identical with the form

here described, and Hauericeras sp. (recorded on p. 243), together with a second

Hauericeras ? sp., not mentioned in the paper (B.M., Nos. C18276-77);

probably also belong here.

+ “Cret. Foss. Natal,’ II, ‘Third Report Geol. Surv. Nat. and Zulul.,

1907, p. 88, pl. vi.

On Cretaceous Cephalopoda from Zululand. 2774

Famity: DIPOLOCERATIDA.

Gren. DIPOLOCERAS, Hyatt.

4, DipoLOCERAS cristaTuM, Deluc sp.

(Pl. RXV, fig: 2; Pl XOX tigsiGs)

1822. Amm. cristatus, Delue in Brongniart. Environs de Paris,

pp. 95 and 395, pl. vu, fig. 9.

1907. Mortoniceras (/) cristatum (Deluc), Pervinquitre. Et. d. Pal.

Tunis. Céph. Ter. Second, p. 239.

1908. Mortoniceras (2) cristatum (Deluc), Jacob. “ Et. Pal. and Strat

Part. Moy. Ter. Crét.,” Trav. Lab. Géol. Univ. Grenoble, vol. viii,

pp. 826 and 384.

This well-known species is represented in the collection by two

examples of the following dimensions :

No. 2728. No. 2727.

Diameter , : . 144 mm. és mm.

Height of last ee ; . 939 per cent. 39 per cent.

Thickness _,, », (at promi-

nent costae) : ; apt ge: | aes 44,

Umbilicus 3 SOON ts, 40 ,,

There is particularly eal agreement with the coarse form figured

by Brongniart. The keel is narrowed at its base, like that of the form

described below as D. sp. nov. or of Ooster’s A. roissyanus (VOrbigny )

varietas.* The extremely pronounced forward sweep of the costation,

near the end of the large example (see fig. 2, Pl. XXV), is particularly

striking. This probably represents the mouth border with its rostrum,

but unfortunately the latter is not perfectly preserved.t The length

of the body-chamber is well over half a whorl; the last few (approxi-

mate) suture-lines are very simple but could not be exposed sufficiently

well for complete delineation. What can be seen differs from the

suture-line figured by Pictet{ (fig. 5c) in having shorter and wider

elements and from that of fig. 2c in having simpler outlines. The

internal portion shows part of the dorsal saddle, but the antisiphonal

lobe is hidden by matrix that did not permit of further preparation.

* «Catal. Céph. Foss., etc.,’ 1860, iv, p. 144, pl. xxvi, figs. 6 and 7

+ @Orbigny’s figure (/oc. cit., pl. Ixxxviii, fig. 1) probably is restored and

composite, for according to the evidence of the example here described, the

curve of the costae near the end is quite different, and all tuberculation has

disappeared.

t In Pictet & Roux, ‘ Moll. Foss. Grés. Verts.,’ 1847, pl. viii.

278 Annals of the South African Museum.

The smaller example is distinguished from equal-sized European

specimens of the swberistatus type, as figured by d’Orbigny and

Sowerby (in Fitton), in having straighter and more rigid costation.

This is reminiscent of D. cornutum, and some fine specimens of the

latter species, much larger than Pictet’s type, from the Astier Collec-

tion in the British Museum (e.g. Nos. 37610 [88]) show comparable

ornamentation near the end, after decline of the tuberculation has set

in. Only the thickness is far greater in Pictet’s species than in

D. cristatum, and the prominences on the inner whorls are blunter.

The Schloenbachia sp., recorded by Crick* from the Middle Tributary

of the Manuan Creek, and compared with d’Orbigny’s A. delaruei and

Pictet’s A. cornutus, is a badly preserved and immature example of a

Dipoloceras ; but a larger fragment (No. C18301) of a similar form,

close to both the large D. cornutum from Escragnolles, mentioned

above, and to the typical D. cristatum in Brongniart, was worked out

of the matrix of one of the Nautili described on p. 244 by Mr. Crick,

but was not referred to in the paper.

The large example, in peripheral view of the final, costate portion,

somewhat resembles Marcou’s A. shumardit; but that species has

no “ flares,’ has weak outer and strong inner tubercles, and is tran-

sitional from Dipoloceras to Subschloenbachia.

Locality.-—Manuan Creek. Coll. Resident Magistrate, Ubombo.

5. DIPOLOCERAS QUADRATUM, sp. Nov.

(Pl. XXV, figs. 3 a—c.)

Cf. 1847. A. bouchardianus (d’Orbigny) Pictet. In Pictet & Roux,

Moll. Foss. Gres. Verts., p. 350, pl. viii, fig. 9.

This species is based on a specimen (No. 4955) of the following

dimensions :

Diameter ; ‘ : . 43 mm.

Height of last whorl. . 40 per cent. of the diameter.

Thickness ,, E : . PADS “a e. 5

Umbilicus 2 BD a x 2

Like the closely comparable specimen figured by Pictet, this

* Loe. cit. (1907), p. 247, B.M., No. C18308.

+ ‘Geology of N. America’ (1858), p. 33, pl. i, fig. 1 (misspelt schumardi on

plate), holotype in B.M. (Geol. Soc. Coll.) No. 12662. The geological position

of this form, in the Upper Duck Creek Formation, below the Fort Worth Beds

with Subschloenbachia leonensis, seems to correspond with that of the European

Dipoloceras cristatum.

On Cretaceous Cephalopoda from Zululand. 279

Ammonite is distinguished from the true D. bouchardianum, a’ Orbigny,*

by its evolute and square whorls, with a wide, carinati-sulcate ventral

area. It may be considered to form a transition towards the swb-

cristatum and subinflatum groups, as dV Orbigny’s compressed type has

leanings towards Pseudophacoceras roissyanum, VOrbigny sp. The

resemblance of Pictet’s example to D. subcristatum consists chiefly of

the peculiar trifurcation of some of the costae; whereas in the smaller

Zululand specimen, comparable with the inner whorls of Pictet’s

form, this resemblance is due to the slightly greater prominence of one

or two of the costae, though there are no conspicuous flares, such as

are characteristic of the cristatum group.

The present example, on the other hand, does not show the umbilical

tuberculation that is so notable a feature in, Pictet’s much larger

specimen, but, as in d’Orbigny’s species, a pair of costae may be

thickened where they meet at the umbilicus. The example, perhaps,

might be thought to represent only the inner whorls of a large

specimen, such as Boule, Lemoine and Thévenin’s “ Schloenbachia ef.

bouchardiana.’ + It appears probable, however, that the last half-

whorl, at least, of the present example belongs to the body-chamber,

though the suture-line, unfortunately, cannot be made out.

The Madagascar specimen, referred to above, which is larger than

any European form of this group, and therefore difticult to compare,

apparently does not agree either with @’Orbigny’s species or with the

species here discussed, and may represent a new type. In D. sub-

inflatum, Pictet sp.,f and in D. rowxianum, Pictet sp.,§ the umbilical

tuberculation is far too pronounced.

Forms comparable with the present species, but not with d’Orbigny’s

form, are found in bed VIII at Folkestone, but in the succeeding

zones, forms belonging to the group of Brancoceras symmetricum,

Sowerby sp., are common, and often confused with D. bouchar-

dianum.|| Other transitional forms of Dipoloceras from the cristatus

zone of Folkestone differ from the present species in having the point

of bifurcation of the costae moved farther away from the umbilicus.

Locality. —Middle Branch, Manuan Creek. Coll. W. J. Wybergh.

* Loc. eit. (1840), p. 301, pl. Ixxxviii, figs. 6-8.

+ Loe. etés (1907, 11), p: 39; pl: ix, fie. 11.

¢ In Pictet and Roux, loc. cit. (1847), p. 104, pl. x, fig. 1. A specimen in the

British Museum, also from Mt. Saxonet (No. C. 10399) forms a transition to

Brancoceras symmetricum (Sow.).

§ Lbid., p. 99, pl. ix, figs. 2 a, d.

|| See Price, “On the Gault of Folkestone,” ‘Q.J.G.S., vol. xxx (1874), table

on p. 362; and Jukes-Browne and Hill, ‘Cret. Rocks Britain, I, “Gault and

Up. Gr. Sd. of Engl.,” tables on pp. 82 and 459.

280 Annals of the South African Museum.

DIPOLOCERAS sp. Nov.

(Pl. MOVE, figs oa,b))

Cf. 1910. Schloenbachia n. sp. Bose, “ Mon. Geol. & Pal. d. Cerro

de Muleros,”’ Bol. Inst. Geol. Mexico, No. 25, p. 74, pl. vin, fig. 6.

The fragment (No. 4903) to be described, unfortunately, is too

incomplete to justify the creation of a new species, but the characters

of the body-chamber, as well as of what is preserved of the inner

whorls, clearly distinguish it from the previously described forms of

this group. The measurements, based on the restoration of the

complete shell, shown in fig. 5 a, are:

Diameter : py mm,

Height of the last w shoe . 46 per cent. of the diameter

Thickness ,, e . 39 * es s

Umbilicus ,, > 22 3 4 >

Except for the smaller arabicus of the present rues these

measurements agree with those of D. sergipense, White sp.* There is

a similar strong outer tubercle and high keel, but the latter, im the

Zululand specimen, is of the shape of that of A. roissyanus (d’Orbigny)

varietas, Ooster,t or of many specimens of Dipoloceras cristatum, that

are well enough preserved—that is to say, it is thinner at its base than

at its middle height, and becomes thin and sharp again at the edge.

But the present species has a second small tubercle half way between

the much more prominent outer tubercle and the umbilical suture.

Similar bitubereulation is shown in some varieties of D. (Mojsisoviesia /)

delaruei, V’ Orbigny sp., e.g. the form figured by Parona and Bonarelli,f

only in the present, species it is more developed, and altogether the

new form, like Mojsisovicsia ventanillensis, Gabb sp., shows a decided

resemblance to the later Subschloenbachia. Of the bituberculate forms

included by Bose in “ Schloenbachia n. sp.,’ his figs. 6-8, pl. viii,

represent a closely comparable form ; but the inner tubercle is hardly

indicated, whereas in figs. 4 and 5 it is the outer tubercle that is not

prominent enough. Besides, both these examples are represented as

having an acute periphery instead of a high keel on a flat periphery,

such as is characteristic of D. sergipense and the delaruei group.

* «Contrib. Pal. Brazil,’ ‘Arch. Mus. Nac. Rio de Janeiro,’ vol. vii (1887),

p. 221, pl. xxiv, figs. 1 and 2. A specimen intermediate between this species

and D. (Mojsisovicsia ?) delaruei (d’Orbigny) in the British Museum (No, C4255),

from Velez, Colombia, with a smaller umbilicus, is still nearer to the present

example, and also shows the steep forward edge of the costae very well.

+ ‘Catal. Céph. Foss.,’ ete., 1860, iv, p. 144, pl. xxvi, figs. 6 and 7.

+ “Foss. Alb. d’Escragnolles,” ‘ Pal. Ital.,’ vol. 11 (1896), p. 88, pl. xi, fig. 9 only.

On Cretaceous Cephalopoda from Zululand. 281

D. colladoni, Pictet sp.,* is too compressed and too evolute; and, in

the sectional view, the outer tubercle is not prominent enough; but

the character of the ribs, with an occasional rib that does not reach to

the umbilical border, is very similar. There apparently is no imner

tubercle in this small species, but the high umbilical border, shown in

Pictet’s fig. 1 b indicates how the line of minute inner tubercles (of

some of the costae) of the specimen here described first arises. On

the other hand, in such a form of Subschloenbachia ? as Quenstedt’s

evolute “A. varicosus,’+ with the Dipoloceras keel still retained, the

inner tubercle has become strongly developed, and the ribbing has lost

its flexiradiate and peculiar wedge-shaped character.

Locality.—Middle Branch, Manuan Creek. Coll. W. J. Wybergh.

GEN. PSEUDOPHACOCERAS, nov.{

Genotype: A. roissyanus, A Orbigny, ‘Pal. Franc. Ter. Crét.,’ p. 302,

pl. Ixxxix.

7. PSEUDOPHACOCERAS MANUANENSE, Nov.

(Pl. XXV, figs. 1 a-d.)

1907. Schloenbachia sp., Crick. ‘Cret. Foss. Natal (III),” Third

Report, Geol. Surv. Nat. and Zulul., p. 240.

This species is represented by two specimens, the larger of which

is taken as type. Their dimensions are as follows :

No. 2725. No. 2726.

Diameter. : . 235 mm. (at) 80 mm.

Height of last whorl 45 per cent. 52 per cent. of the diameter.

iwhickness) ,, ' 3 20 "4; 2s 99 99

Umbilicus : a 3 Smee, 29 %9

At a diameter of 170 mm. the umbilicus of the large example is

only 16 per cent., and the uncoiling that leads to an excentrumbilicate

(scaphitoid) shell is confined to the last half-whorl, belonging to the

body-chamber. At a diameter of 80 mm. there are 20-22 primary

ribs per whorl ; they bifureate at varying distances from the umbilical

border, and one branch bifurcates again higher up, so that generally

* In Pictet & Roux, loc. cit. (1847), p. 89, pl. viii, figs. 1 a, 6. Pictet &

Campiche (Joe. cit. [1859], p. 175) wrongly united this species with J. de C.

Sowerby’s Brancoceras symmetricum.

+ Non Sowerby (‘Cephalop.,’ 1849, pl. xvii, fig. 2).

+t Dealt with in the writer’s Angola paper, above referred to.

282 Annals of the South African Museum.

there are three secondaries to each primary rib.“ On the outer whorl

the ornament is considerably weakened. The umbilical border is

gently rounded and the slope slightly concave, except on the innermost

whorls. The keel is very prominent and narrowed at its base, as in

the form of Enea one described by Ooster and in many

Dipoloceras (see ante, pp. 277 and 280).

The smaller example, which has portions of the outer whorl

preserved (not shown in fig. le of Pl. XXV), has its last suture-line

at a diameter of about 115 mm., when, here also, excentrumbilication

sets in. Unfortunately only the umbilical portion of the outer whorl

here is preserved. The (last) suture-line of the larger example, taken

at a diameter of 160 mm., apparently differs from that of the smaller

specimen, the latter (in fig. 1¢) being nearly half a whorl away from

the beginning of the body-chamber, at a diameter of 80 mm. But

though comparison at the same (relative) size is impossible without

breaking up the type, the dissimilarity, in the writer’s opinion, is.

accounted for by the fairly frequently observed simplification of the

last few suture-lines, in Ammonites generally, often accompanied by

equalisation of the elements. This is a phenomenon of individual

erowth and cannot be applied to phylogeny; and it seems to the

writer that the suture-line of the smaller specimen (fig. 1 ¢) indicates

that descendants of this group would show the Sphenodiscus type of

suture-line, with adventitious elements.

The suture-line of the present species differs from that of the less.

compressed P. multifidum, Steinmann sp.,t its nearest relative, chiefly

in the greater depth of the ventral lobe and the presence of a larger

number of auxiliaries (corresponding to its greater involution)—a

character that also is not shown in the more evolute form figured by

Lasswitz.{ The suture-line of d’Orbigny’s A. roissyanus§ has this

deep ventral lobe, but differs in the auxiliaries. On the other hand,

in a specimen of P. aff. roissyanum (from Escragnolles, in the British

* This type of ornament is somewhat reminiscent of that of Neoharpoceras

fgen. nov.| ef. hugardianum (VOrbigny) in Pictet (/oc. cit., Pictet & Roux,

1847, pl. x, fig. 3), but the suture-lines are very different in the two stocks.

(Genotype = A. hugardianus, d’Orbigny, ‘ Pal. Frang. Ter. Crét., pl. Ixxxvi,

figs. 1 and 2.)

+ “Wb. Tithon and Kreide i. d. Peruan. And.,” “N. Jb. f: Min., etc. ID,

1881, p. 139, pl. vii, fig. 1 [ Schloenbachia acuto-carinata (Shum. sp.) Marcou).

+ “ Kreide-Amm. v. Texas,” ‘Geol. and Pal. Abh.,’ N.F., vol. vi, Heft 4, 1904,

p- 22, pl. v, fig. 2. Schlagintweit (see below) questions Lasswitz’s identifications,

but the present examples indicate that in the case of the secondaries it occa-

sionally is impossible to say whether they result from dichotomous Pee

or are simply intercalated.

§ Loe. cit. (1840), p. 302, pl. lxxxix, figs. 1-3.

On Cretaceous Cephalopoda from Zululand. 283

Museum, No. 50065), transitional to P. mirapelianum, d’Orbigny sp.

there is only one auxiliary lobe less than in the present specimen, and

the suture-line describes a similar curve, but the lateral lobe is shghtly

deeper than the ventral lobe. :

Shumard’s original P. acuto-carinatum* is more evolute and much

more distantly costate than the present species; but the form figured

as ‘ Sonneratia acuto-carinata, Typus,” by Lasswitzt forms a closer

approach to P. multifidum and to P. manuanense than to Shumard’s

type or to @Orbigny’s species.f The specimen figured by Bose§ as

Schloenbachia aff. acuto-carinata is similarly closely costate, but

differs from the Zululand species in the whorl-section, and in the very

pronounced forward projection of the peripheral portion of the costae.

A specimen of P. cf. peruvianum, v. Buch, from Velez, Colombia, in the

British Museum,|| shows similar peripheral projection ; but the ribs,

all of which are single, have the pecular perpendicular forward edge,

reminiscent of such large examples of P. roissyanum, d’Orbigny sp.,

as that figured by Parona and Bonarelli.. The suture-line of the

evolute Mexican example, on the other hand, shows great resemblance

to that of the large specimen here figured, except that there are two

more auxiliary lobes in the latter.

One of the specimens included by White** in his A. buarquianus

seems close to the present species, but the type (White’s figs. 5 and 4)

has distant and single costae.

The fragments recorded by Crick, since mounted in _ plaster,

probably belong to a large specimen of this form. Sphenodiseus sp.,

* In Marcy, “Explor. Red River Louisiana” (1853), Appendix E, ‘ Pal.,’

p- 209, pl. iii, fig. 1. The thickness of this form appears to be only about

20 per cent. of the diameter.

+ Loc. cit. (1904), p. 21, pl. xvii (v), fig. 1, = A. peruvianus, Marcou, V, 1a, b

only, non v. Buch (B.M., No. 12718, Geol. Soe. Coll.).

t Schlagintweit (“D. Fauna d. Vracon und Cenoman in Peru,” ‘N. Jb. f.

Min., ete., Beil. Bd. xxxiii (1912), pp. 64 and ff., includes in “ Seh/oenbachia”’

roissyana, @Orbigny sp., not only P. acuto-carinatum (already united with

d’Orbigny’s species by Lasswitz, 1904, and Douvillé, 1906), P. multifidum,

P. mirapelianum (as varieties), further P. buarquianum (White), P. peruvianum

(v. Buch), P. carbonarium (Gabb), P. belknapi (Marcou) [holotype in B.M., No.

12663, Geol. Soc. Coll.], and other comparable forms, but, probably quite

wrongly, also Lea’s A. americanus, apparently a Pulchellia.

§ Loe. cit. (1910), p. 65, pl. ii, figs. 1-3.

|| No. C4266, comparable with fig. 6 of pl. iin v. Buch (‘ Petrif. Recueill. en

Amer. p. M. A. Humboldt et M. Ch. Degenhardt,’ Berlin, 1839, p. 5), and more

closely costate than A. peruwvianus, Marcou, V, 1, non v. Buch (B.M., No. 12664

Geol. Soc. Coll. and writer’s coll.).

{ Loe. cit. (1896), p. 88 (36), pl. ii, fig. 8.

** Toe. cit. (1887), p. 222, pl. xxiv, figs. 5 and 6 only.

284 Annals of the South African Museum.

figured by Boule, Lemoine and Thévenin* from the “ Lower Ceno-

manian” of Madagascar, may represent an allied form of this genus,

distinguished by more numerous auxiliaries.

Locality —Manuan Creek. Coll. Resident Magistrate, Ubombo.

GreN. SUBSCHLOENBACHIA, nov.

Genotype: A. rostratus, J. Sowerby, ‘Min. Conch.,’ pl. clxxiii (Oxford

University Museum).

8. SUBSCHLOENBACHIA PREROSTRATA, NOv.

(Pl. XXIV, fig. 10.)

A fragmentary example (No. 4970), consisting of portions of two

whorls (completely septate) of a specimen of 160-170 mm. diameter,

belongs to a type of Subschluenbachia that is found in Southern Europe,

and to which probably the large example figured by Boule, Lemoine

and Thévenint is related, though the latter acquires single costation,

resembling that of S. stoliczkai, n. nov.,{ at an earlier stage.

The whorl-section at first is wider than high,§ but becomes quadrate at

a later stage. S. picteti,n.nov.,|| has a less depressed whorl-section, and

shows less coarse ornamentation; on the other hand, S. orbignyi, nu. nov.,§|

or at least the French examples that the writer would consider typical of

d’Orbigny’s form, agree in the more robust ornamentation, but are com-

pressed. On the shell there is very distinct spiral striation, but the east is

almost smooth. The suture-line has deeper lateral lobes than that figured

by @Orbigny. A peculiar feature is a distinct groove on the umbilical

slope (Pl. XXIV, fig. 10, marked by arrows) of the cast, near the end.

S. stoliczkai, nov., which has a depressed whorl-section,** shows

the ornamentation of S. aequatorialis, Kossmat sp., on its inner

whorls; and its near ally, S. pachys, Seeley sp., is too closely costate

and less tuberculate. The depressed examples of the quadrituber-

culate perinflata group (n. nov. = A. inflatus, pars, Pictet & Campiche,

pl. xxii, fig. 3 only), and others described in the Angola paper already

referred to, are distinguished by their double row of outer tubercles.

Locality— Middle Branch, Manuan Creek. Coll. W. J. Wybergh.

* Loc. cit. (II, 1907), p. 50, pl. xi, fig. 5.

+ Loc. cit. (II, 1907), p. 40, pl. ix, fig. 7.

+t = A. inflatus (Sow.) Stoliczka, loc. cit., pls. xxvii and xxix, fig. 2.

§ In fig. 10 of Pl. XXIV, the height of the smaller whorl should have been

only 27 mm.

|| =A. inflatus, Pictet, in Pictet & Roux, 1847, pl. ix, fig. 6, e.g. B.M., No.

C3822 (holotype of species) and No. 62116.

{ =A inflatus, VOrbigny, 1840, pl. xc, e.g. B.M., No. C901 (holotype of

species ; thinner, more distantly, coarsely [and recte-] costate, than S. picteti).

** Stoliezka, loc. cit., pl. xxix, fig. 2.

On Cretaceous Cephalopoda from Zululand. 285

9, SUBSCHLOENBACHIA ef. TRINODOSA, Bose sp.

(Pl. XXV, fig. 4.)

1910. Sehloenbachia trinodosa, Bose. ‘ Mon. Geol. & Pal. Cerro de

Muleros,”’ Bol. Inst. Geol. Mexico, No. 25, p. 78, pl. x. figs. 2—4.

A whorl-fragment (No. 4972), about 100 mm. long, is comparable

with the smaller specimen figured by Bose, but has a slightly more

depressed whorl-section. The spiral ornamentation, also, is very well

preserved on this (body-chamber?) fragment, whereas the Mexican

original, a septate cast, does not show it. The present example is

distinguished from the form above described (S. prerostrata) by its

whorl-section, which difference results in very dissimilar ventral

aspects of the two species. The trituberculation also is a distinctive

feature of the form here described, quite different from the ornamen-

tation of the earlier group, with the high umbilical tubercle almost at

the middle of the side.

Forms similar to this species occur in the Cambridge Greensand

and in the Red Chalk, and there exist (at Blackdown) transitions to

S. aequatorialis, Kossmat sp., on the one hand, and to the bituber-

culate species, referred to below, on the other.

Locality—South side of Manuan Creek Valley. Coll. W. J.

Wybergh.

10. SUBSCHLOENBACHIA BISPINOSA, NOV.

(Pl. XXIV, fig. 9.)

This species is based on a somewhat fragmentary specimen

(No. 4993) that is weathered on one side, so as to expose, ina natural

section, the distant septa of the inner whorls and the more approxi-

mate septa of the last quarter of a whorl, which shows the beginning

of the body-chamber. The dimensions are as follows :

Diameter ; : ; . 1448 mm.

Height of the last whorl . 32 per cent. of the diameter.

Thickness ‘ s 4) BD - - 95

Umbilicus 44. ee 5 9

The peculiar whorl-section (Pl. XXIV, fig. 9), with its depressed outer

tubercle, shows the distinctive features of this form. The inner whorls

agree in ornamentation with those of the Bellegarde variety of S. pre-

rostrata (B.M., No. C10547, less depressed than fig. 10, Pl. XXIV),

and show spiral striation ; the outer whorl, with only a peripheral and

an umbilical tubercle (on the ribs that reach to the umbilicus) is

286 Annals of the South African Museum.

comparable with that of one of Etheridge’s Queensland specimens* and

of a common Blackdown form (e. g. B.M., No. 52045). This latter is

interesting, since it is connected by quite a series of transitions with S.

orbigny?t on the one hand, and with compressed forms converging towards

Prohysterocerast goodhalli on the other. The, Blackdown form stands

in the same relationship to S. bispinosa as S. orbignyi to S. picteti—t hat

is to say, the Zululand species here described, like 8. picteti, represents

the less coarsely ornamented type. Near the end of the shell the costae

become single and equally bituberculate, and on the cast show no trace

of spiral striation. The Angola S. cycloceratoides, Spath, has similar

lateral ornament, but is compressed and has no distinct keel, but

merely an angular periphery, like certain Acanthopleuroceras.

S. leoennsis, Conrad sp.,t includes a group of bispinous forms that

have a certain resemblance to the specimen here described. Whether,

however, such forms as, e.g., Hill’s S. leonensis,§ are identical with

Conrad’s type seems doubtful. Lasswitz|| probably has misinter-

preted the species altogether, putting it in the Senonian.

A worn fragment (No. 4971) of the body-chamber of a large

specimen, in which the costation has become single, in whorl-section

and peripheral aspect agrees with the species here described. In

somewhat similar body-chamber fragments of S. stoliczkai from

Angola, the peripheral tubercle is more prominent, elongated longitu-

dinally and projected upwards.

Locality —Middle Branch, Manuan Creek. Coll. W. J. Wybergh.

Famity: LYELLICERATIDA.

Gen. STOLICZKAIA, Neumayr.

11. SroniezKata sp. ind.

1888. Stoliczkaia dispar (d’Orbigny) Choffat. In Choffat & de

Loriol, “Matériaux . . . Angola,’ Mém. Soc. Phys. and

d’Hist. Natur. Genéve, t. xxx, No. 2, p. 69, pl. ii, fig. 6 ?.

1894. St. clavigera (Neumayr) Kossmat. ‘‘ D. Bedeut. d. Siidind. Kr.

F.,” Jb. K.K.R.A., vol. xliv, p. 465.

* « Low. Cret. Foss., ete.,’ Il, Ceph. ii, ‘ Records Austral. Mus.,’ vii, 4 (1909),

pl. xvi, fig. 1 only.

+ Gen. nov. Genotype = P. wordiei, nov., a new Angola species of the candolli-

anum-goodhalli group.

t “ Deser. of Cret. and Tert. Foss.,” ‘Geol. Rep. Mexican Boundary * (1857 ?),

p. 160, pl. xvi, figs. 2 a, b.

§ Loe. cit., 1901, pl. xxxvi, figs. 1, la (as 4. (Schloenbachia) leonensis, Romer).

|| Loe. cit., p. 28.

On Cretaceous Cephalopoda from Zululand. 287

A whorl-fragment (No. 4939) of a specimen of about 35 mm.

diameter, with the umbilical portion imperfectly preserved, appears to

belong to a flexicostate form of this genus, like some of the varieties

of S. dispar, @Orbigny sp., figured by Choffat and by Pictet &

Campiche.* The square whorl-section agrees with that of S. clavigera,

Neumayr,+ but in this species, as in the still more quadrate-whorled

S. tetragona, Neumayr,{ the costation is too straight. There are

nine ribs on the fragment, very thick on the periphery, and of

varying lengths.

The specimen of S. claviyera from Madagasear, figured by Boule,

Lemoine and 'Thévenin,$ has the inner whorls too poorly preserved for

comparison with the much smaller fragment here described.

It should be mentioned that there is no indication of tuberculation

on the periphery, such as is shown in Choffat’s figs. 5 and 7, in Pictet

and Campiche’s fig. 1b, and in Stoliczka’s fig. 3—a feature that becomes

permanent in the later Mantelliceras, e.g. M. hoplitoides, Lasswitz,||

and M. martimpreyi (Coquand), Pervinquitre.{ = Mantelliceras is

known to occur in Zululand,** but the presence of a Stoliczkaia in the

Manuan Creek Fauna is of interest in view of the occurrence of this

genus in India and Madagascar, on the one hand, and in Angola and

North Africa on the other.

Locality—South side of Manuan Creek Valley. Coll. W. J.

W vbergh.

* Loc, cit. (1860), pl. xxxvili. 8. dispar, dOrbigny sp., and S. notha, Seeley

sp., are too compressed and too rounded ventrally.

+ In Stoliezka, Joc. cit. (1865), pl. xlv, fig. 1 (vefigured in Lasswitz, ‘ Kreide

A. v. Texas,’ loc. cit., pl. iv, fig. 2), and fig. 3 (refigured Kossmat, loc: cit.,

pl. xxiv [x], fig. 2). Kossmat included the Angola specimens in S. clavigera

Neumayr, but they belong to several distinct varieties.

t In Stoliezka, loc. cit., pl. xlv, fig. 2.

§ Loe. cit. (1907), pl. ix, fig. 1, p. 83 (apparently reduced by half, though

stated to be natural size).

| Loe. cit. (1904), p. 19, pl. iii (xv), fig. 3. The suture-line of this form

(text-fig. 4, p. 19) is little advanced from that of Lyelliceratide, so that Man-

telliceras is not a direct descendant of Stoliczkaia, but more or less a parallel

development with Neophlycticeras, nov. (genotype = A. brottianus, d’Orbigny,

‘Pal. Franc. Ter. Crét.,’ pl. Ixxxv, figs. 8-10), and Stoliczkaia, of Lyelliceratide.

(See, however, Nowak, “ Unters. Cephal. Ob. Kreide Pol.,” II. Skaphiten, ‘Bull.

Acad. Sci. Cracovie, July, 1911, p. 554, text-figs. 1 and 2.)

| Loc. cit. (1907), pp. 289 and 389.

** KE. eo. M. choffati, Kossmat sp.

288 Annals of the South African Museum.

Famity: HAMITIDAL.

Gen. TORNEUTOCERAS, Hyatt.

12. ToRNEUTOCERAS sp. ind.

1861. Hamites virgulatus, Pictet & Campiche (non d’Orbigny). ‘‘ Foss.

Ter. Crét. de Ste. Croix,” Mat. Pal. Suisse, IIT, pt. 2, p. 85, pl. liv,

figs. 7 a—d only.

A fragment (No. 4967), about 20 mm. in length, and slightly curved,

has seven thick and round ribs that are very prominent on the ventral

side but disappear on the dorsum, so that the concave side of the

shell appears quite smooth. This agrees with the ornamentation

common to several species of “ Hamites”’; but the cireular cross-

section and very slight obliquity of the ribs suggest comparison with

the above form. The example, however, is only preserved as a very

poor cast in a brownish, friable, sandy matrix, so that its identification

must remain doubtful.

Specimens of 7. virgulatum from Angola (B.M. No. C20180-1)

show very similar ornament, but a more compressed section, like the

types of Brongniart and d’Orbigny, and Pictet & Campiche’s fig. 6.

Some of the examples figured by Pictet & Campiche on pl. li as

Anisoceras alternatum (Mantell) show some resemblance to the frag-

ment here described (e. g. fig. 6), and it should be mentioned that there

isa specimen of a comparable form of Anisoceras in the British Museum

(No. C18300) from the South Branch of the Manuan Creek, pre-

served in a similar matrix, but not referred to in Crick’s paper.

The tuberculation of alternate ribs, however, is very distinct in this

fragment.

Locality—South side of Manuan Creek Valley. Coll. W.

Wybergh.

Famity: ANISOCERATIDA.

Gen. ANISOCERAS, Pictet.

13. ANISOCERAS sp. ind.

(PES eValy is: 7.)

Cf. 1861. Helicoceras thurmanni, Pictet & Campiche. “ Foss. Ter.

Crét. Ste. Croix,” Mat. Pal. Suisse, III, pt. 2, p. 118, pl. lvi, fig. 5.

On Cretaceous Cephalopoda from Zululand. 289

A small fragment (No. 4982) of a completely septate, free and

unsymmetrical whorl resembles the above form in size, coiling and

quadrituberculation, but the costae are broader and all equal. The

tubercles are less prominent, agreeing in this respect with those of

Anisoceras pseudopunctatum, Pictet & Campiche,* and the costae

are slightly weakened between the ventral tubercles, but flattened,

after the manner of those of A. perarmatum, Pictet & Campiche,t

between the ventral and lateral tubercles.

“ Helicoceras”’ astierianum, VOrbigny in Parona and Bonarelli,t

shows only slightly closer coiling than the present form, and other

comparable forms, e.g. Twurrilites elegans, Pictet & Campiche, pars,

non VOrbigny,§ have somewhat similar ornamentation. It appears

most probable, however, that the example here recorded represents

part of the helicoid initial whorls of an Anisoceras of the typical

group of A. saussureanum, Pictet,|) and A. oldhamianum, Stoliezka.§,

The suture-line differs from that of the typical Anisoceras in the wide

external saddle, and from those equally simple ones of certain Hamitids

(Torneutoceras) in the unsymmetrical first lateral lobe (Pl. XX VI,

fio. 7).

Locality.—Low Ridge, about three miles east of foot of L~eebombo

Mountains, north of M’Kusi River, due east of Ubombo. Coll.

W.J. Wybergh. This is the only specimen in the present collection

from this locality, further north than that of any of the other (post-

Aptian) Cephalopoda ; but the facies appears to be the same as that

of the Albian Manuan Creek fauna.

Faminy. UR REELED AG.

Gen. TURRILITES, Lamarck.

14. Turriires cf. GREssiy1, Pictet & Campiche.

1907. Turrilites gresslyi. Boule, Lemoine & Thevenin: “ Céph. d.

Diego-Suarez,”’ Ann. d. Pal., vol. ii, p. 57, pl. ii, fig. 2.

A small portion of a Turrilites (No. 4954), with five rows of tubercles,

agrees with the example from Madagascar, in having the upper two

tubercles very close together. Since these coincide with the upper

* Loc. cit., p. 74, pl. lii, figs. 1-3.

+ Ibid., pl. xlix, figs. 2, 4a.

+ Loc. cit. (1896), p. 102 (50), pl. v, fig. 13.

§ Loc. cit., e.g. pl. lvi, fig. 104.

|| In Pictet and Roux (1847), p. 118, pl. xiii (Hamites).

{ Loc. cit. (1866), p. 175, pl. Ixxxiii, figs. 1-4.

290 Annals of the South African Musewm.

suture and the lowest (indistinct) row of tubercles with the lower

suture, only the two median rows are clearly visible at the middle of

the sides, as pointed out by Boule, Lemoine and Thévenin. In the

Ste. Croix specimens, however, the three rows of lateral tubereles

appear to be more equal-sized, and the upper two rows are not so close

together.*

Specimens of J’. cenomanensis, Schliiter,+ from Wiltshire and

Wissant, France, of undoubted Cenomanian age (zone of Schloenbachia

varians), Show the closest agreement with the present fragment in the

arrangement of the tubercles ; but shape and coiling of the whorls are

slightly different in the Cenomanian species. The Twrrilites recorded

by Crick{ from the Cenomanian of False Bay, Zululand, belong to

different species.

Locality.—South side of Manuan Creek Valley. Coll. W. J.

Wybergh.

2. NAUTILOIDEA.

Gren. CYMATOCERAS, Hyatt.

15. CyMATOCERAS MANUANENSE, G. C. Crick sp.

1907. Nautilus manuanensis, G. C. Crick. Loc. cit. (Third Report),

p. 243, pl. xv, figs. 6, 6 a.

This species was founded for a “‘ less tumid, more finely ornamented

and more narrowly umbilicated shell than Nautilus pseudo-elegans,

VOrbigny.” Crick stated that it was ‘‘numerously represented in

Mr. Anderson’s collection from the Manuan Creek,’ but it is possible

that the specimens do not all belong to the same species, and that

some fragments (e.g. B.M., No. C18283) belong to such a Senonian

Cymatoceras as Nautilus elegans (Sowerby) in Boule, Lemoine and

Thévenin;§ for Albian and Senonian forms occur together at the

south branch of the Manuan Creek, and there are slight differences

in the matrices. Taking the figured example (B.M., No. C18282) as

type of Crick’s species, the differences from d’Orbigny’s N. pseudo-

* Pictet et Campiche, ‘Ter. Crét. de Ste. Croix,’ 2nd ser. (1861), p. 132,

pl. lvii, figs. 11-13.

+ Loc. cit. (1876), p. 11 (181), pl. ii (xxxvii), figs. 6-8. (Cf. Turrilites

tuberculatus (pars) Sharpe, loc. cit., III, 1856, pl. xxv, fig. 3 only.) (Writer’s Coll.)

t Lee. cit. (1907), pp. 173-8.

§ Loe. cit. (1907), p. 66, pl. xv, fig. 4.

On Cretaceous Cephalopoda from Zululand. 29]

elegans* seem to be a less broad periphery and less coarse, if equally

distant, costation. The proportions of diameter to thickness (3 : 2)

are about the same in the two forms, but on account of its broad

periphery d’Orbigny’s figure appears considerably more tumid than

that of N. manuanensis, as does the figure of the type-specimen given

by Foord.+ On the other hand, C. pseudo-elegans is of Barremian

age, whereas among Albian forms the example figured as

N. albensis, VOrbigny, by Pictet & Campichet appears to be almost

indistinguishable from Crick’s species. The differences are the

slightly greater thickness of the European species and the less pro-

nounced peripheral sinus in the costation, possibly also the absence

of bifurcation. These may not be characters of specific importance,

for sexual dimorphism is proved in the recent Nautilus, and among

some fifteen presumably Albian Cymatoceras from the Manuan Creek

there are no two alike.

One example in the collection (No. 4991) of the following

dimensions—

Diameter. : ; : . 185 mm.

Thickness . , . : : ay AO) nis)

ditters from the holotype only in showing fine costation at the end of

the shell, which, then, is still septate, whereas on the body-chamber of

Crick’s figured specimen, in the one place where the test is preserved,

the ornament does not show this tendency to become finer and closer.

C. virgatum, Spengler sp.,§ has coarser ornamentation than C. manua-

nense; and C. carlottense, Whiteaves sp.,

shallow umbilicus.

| differs in its small and

C. crebricostatum, Blantord sp.,4 differs from the form here described

chiefly in its more sinuous septa, and in a less linguiform peripheral

sinus of the costae.

Locality.—Middle Branch, Manuan Creek. Coll. W. J. Wybergh.

16. Cymarocrras cf. MANUANENSE, G. C. Crick sp.

One example (No. 4990) agrees with Crick’s holotype of N. manna-

nensis, but the coarse plication only appears on the shell of the ventral

* «Pal. France. Ter Crét.’ (1), 1840, p. 70, pls. viii and ix, fig. 1. The ratio of

diameter to thickness is 3 : 2 according to the text.

t+ ‘Catalogue Foss. Ceph. Brit. Mus.’ (IT), 1891, p. 253, fig. 59 on p. 255.

t Loe. cit. (Ste. Croix, I) (1859), p. 134, pl. xvii, figs. 1 @ and 6 only.

§ Loe. cit. (1910), p. 180, pl. xi, fig. 3.

|| ‘Mesozoic Fossils,” I, ‘Geol. Surv. Canada,’ pt. iv, 1900, p. 269, pl. XXi.

" «Pal. Indica,’ I, “ Foss. Ceph. Cret. Rocks 8. India,’ 1861, p. 36, pl. xxi,

fig. 3, and pl. xxi.

292 Annals of the South African Museum.

area at a late stage, and up to a diameter of about 90 mm. there are

only striae of growth. This is reminiscent of the Barremian N.

pseudo-elegans, d’Orbigny, the genotype of Cymatoceras, and the

measurements also agree with those given by the author of this

species. These dimensions are, in the specimen here described :

Diameter : : . 180 mm.

Thickness. : . 115 ,, (64 per cent. of the diameter)

Height of the last whorl

(in siphonal plane) . 65 ,,

Another apparently Albian but imperfect specimen of a Cymato-

ceras, 1n the British Museum, No. C18298, not referred to in Crick’s

paper, shows a similarly late appearance of costation. This and

another example (No. C18299) of the same imperfect set of Nauwtili,

not mentioned in Crick’s Report, are interesting as showing approxi-

mation of the last few septa (see Crick, ‘ Proc. Geol. Soc.,’ No. 979,

p. 3, November 11th, 1915).

Foord* describes, from the Gault, a “species which, on the whole,

agrees very closely with Pictet and Campiche’s description and figures

of N. albensis, but differs from it in respect that up to a certain

” The present

example thus would seem to stand in a similar relation to C.

[variable | stage of growth the test is smooth.

manuanense.

One of the imperfect specimens mentioned by Crick, on p. 245

(B.M., No. C18295) as “ probably referable to Hutrephoceras”’ may

belong to the same species as the form here described, and at any rate

is a Cymatoceras. Another of the examples (C18294) is referable to

Cymatoceras kayeanum, Blanford sp., or a similar compressed species,

whereas the remaining two specimens (Nos. C18292 and 93), preserved

in limonite, may be of Senonian age.t

Locality.—Middle Branch, Manuan Creek. Coll. W. J. Wybergh.

17. CymMaroceras cf. ALBENSE, d’Orbigny sp.

1859. Nautilus albensis, @Orbigny. In Pictet & Campiche, Pal.

Suisse, 2nd ser., p. 134, pl. xvu, figs. 1 a and 6 only.

1891. Nautilus albensis, Foord. Catal. Foss. Ceph. Brit. Mus., i,

p- 258.

One specimen (No. 4989) differs from the holotype of C. manuanense,

Crick sp., and from the examples above described in being slightly

* Loe. cit. (1891), p. 259.

+ Loe. cit. (1907), Third Report, pt. ii.

~ See under Cymatoceras? sp. ef. justum, Blanford sp., p. 301.

On Cretaceous Cephalopoda from Zululand. 2938

thicker, having a more obtuse sinus on the periphery, and a coarser

costation. Its dimensions are :

Diameter. . 135 mm.

Thickness. . 100 ,, (74 per cent. of the diameter).

The whorl-shape agrees with that of the type-figure im Pictet and

Campiche, and is more rounded than that of N. pseudo-elegans,

d’Orbigny, or those of the three forms of Cymatoceras described below.

Locality — Middle Branch, Manuan Creek. Coll. W. J. Wybergh.

18. CyMATOCERAS, sp. ind.

One example of Cymatoceras (No. 4987) differs from the above

forms of the manuanense-albense group in having a flattened periphery,

but it agrees with some of the fragmentary co-types of C. manuanense,

Crick sp. These differences in whorl-shape are not important; and

since the present example is somewhat fragmentary, a definite identi-

fication is impossible. The last septum is shown at a diameter of

about 90 mm., followed by a portion of the body-chamber.

The smooth earlier whorls, in addition to the flattened periphery,

separate it from the holotype of C. manuanense, and the finer,

dichotomous costation also from C. albense. The example agrees with

C. pseudo-elegans, dOrbigny sp., in ornamentation, dorsocentran

position of the siphuncle and flattened periphery, but is not quite so

depressed. C. kossmati, Spengler sp.,* has a narrower whorl-section.

Locality.—Middle Branch, Manuan Creek. Coll. W. J. Wybergh.

19. CymaTocERAS, sp. nov? ind.

1907. Nautilus sp., Crick, loc. cit., p. 244.

Among five specimens of Cymatoceras, referred to by Crick as repre-

senting ‘‘a large species which, in external characters, differs from

N. manuanensis chiefly in having flatter sides and a more broadly-

rounded periphery,’ four are comparable with the various forms

of the manuwanense group described above, but are not well enough

preserved for definite identification. On the other hand, one example

(No. C18291) agrees in its squarish whorl-shape and fine ornamenta-

tion with a large specimen in the present collection (No. 2589) that

has the following dimensions :

Diameter oO) man

Thickness . . 125. ,, (66 per cent. of the diameter).

The ratio of diameter to thickness, about 3:2, corresponds with

* Loe. cit. (1910), p. 129 (= N. pseudo-elegans, Blanford, non d’Orbigny).

294 Annals of the South African Museum.

that of C. pseudo-elegans and C. manuanense, but the costation is much

closer in the present example, and in the specimen described by Crick,

than it is in either of the two species mentioned, and the flat, sub-

parallel sides and broader periphery separate them especially from

C. manuanense.

C. virgatum, Spengler sp.,* and C. striaticostatum, Crick sp.,t have the

squarish whorl-shape of the form here described, but their costation

is much coarser or more distant than that of any of the Zululand

examples of the genus Cymatoceras here dealt with.

In closeness of costation this species approaches to the Cenomanian

Cymatoceras elegans, J. Sowerby sp.,f though its whorl-shape is more

like that of the older C. pseudo-elegans, VOrbigny sp.

Locality.—Manuan Creek. Coll. Resident Magistrate, Ubombo.

20. CyMATOCERAS cf. KossMAtI, Spengler sp.

1910. Nautilus (Cymatoceras) kossmati, Spengler. “ Untersuch. ti.

d. Siidind. Kreidef.,” IV, loc. cit., p. 129.

One specimen (No. 2588), representing a wholly septate cast, has a

portion of the shell preserved at the diameter of 80 mm. and shows

only striae of growth at that stage, but obscure folds on the ventral

area of the cast of the following portion. Its dimensions are:

Diameter . 120 mm.

Thickness . @5 ,, (63 per. cent. of the diameter).

The specimen is a more compressed example of Cymatoceras than

those described above, and resembles in general appearance C. koss-

mati, but this species is strongly costate already at a considerably

smaller diameter. The siphuncle in the present example is nearer

the ventral side and not in the median plane.

C. kayeanum, Blanford sp.,§ is a still more compressed species, and

is more distinctly costate on the ventral area than the specimen here

described.

The Cenomanian C. imbricatum, Crick sp.,|| also is too compressed.

Locality Manuan Creek. Coll. Resident Magistrate, Ubombo.

* Loe. cit. (1910), p. 130, pl. xi, fig. 3.

+ Loe. cit. (1907), p. 221, pl. xiv, fig. 7.

t Non Nautilus elegans (Sowerby), in Boule, Lemoine and Thévenin, Joc. cit.

(1907), p. 66, pl. xv, figs. 4 and 5.

§ Loc. cit. (1861), p. 41, pl. xviii, fig. 1 (type) ; Spengler, loc. cit. (1910), p. 127,

pl. xi, figs. 1 and 2, pl. xii, figs. 2 and 7 a.

|| Loc. cit. (1907), p. 220, pl. xiv, fig. 6.

On Cretaceous Cephalopoda from Zululand. 295

21. Cymaroceras? cf. CLEMENTINUM, d’Orbigny sp.

1840. Nautilus clementinus, d’Orbigny. Pal. France. Ter. Crét., vol. 1,

ps 77>. pla xii bas:

1861. Nautilus splendens, Blanford. Loc. cit. (Foss. Ceph. Cret. Rocks

S. India), p.-21, pl. ix, fig 5, pl. x, Teele

1866. Nautilus splendens, Stoliczka, ibid., p. 205.

1910. Nautilus cf. clementinus (d’Orbigny) Spengler. Loe. cit. p. 143.

A small specimen (No. 4988), slightly worn, and showing the striae of

growth on the test in one or two places, agrees with d’Orbigny’s type

in whorl-shape and general appearance. At a diameter of 70 mm.

the thickness = 45 mm., which is less than the thickness of d’ Orbigny’s

example, but slightly more than that of N. splendens, Blanford, the

type of which, however, is crushed. Spengler united the two species,

which may be open to objections, but since the specimen here

described is not well preserved, it also is included in d’Orbigny’s well-

known Gault species.

Foord * described the species in detail, and was inclined to separate

the Indian from the European forms on account of their greater

whorl-thickness. Spengler + has since created the var. indica of N.

clementinus for the later Indian form, but the present example is

much closer to the European Gault form than to the Trichinopoly

species.

Locality— Middle Branch, Manuan Creek. Coll. W. J. Wybergh.

B. SENONIAN.

AMMONOIDEA.

Gren. PERONICERAS, Grossouvre.

22. Prroniceras cf. DRAVIDICUM, Kossmat sp.

(Pl. XXIII, figs. 1 a-d.)

1865. Am. subtricarinatus, VOrbigny. Stoliezka, ‘* Cret. 8. India,” I,

p. 54, pl. xxxi, figs. 3, 3 a-c.

1895. Schloenbachia dravidica, Kossmat. ‘‘ Unters. S. Ind. Kreidef.,”’

Beitr. Pal. und Geol. Ost.-Ung., vol. ix, Heft 3 and 4, p. 190

(94), pl. xxiii (ix), figs. 3 a-d.

* Loc. cit. (1891), p. 285 (see there for synonymy).

+ Loc. cit. (1910), p. 143.

296 Annals of the South African Museum.

1904. Peroniceras dravidicum, Kossmat. Solger, “ Foss. d. Mungo-

Kreide,”’ Geol. v. Kamerun, II, p. 181, text-fig. 71, p. 182, and 72,

p. 183.

A fragment (No, 4950), 72 mm. in length and belonging to a shell

just a little larger than the inner whorls of this species, refigured by

Kossmat, 7.e., of about 85 mm. diameter, has a whorl-height and

thickness of 24mm. _ It differs from the type of this species only in

having the inner tubercle nearer the umbilicus and more prominent,

i.e. projecting laterally and representing the region of greatest thick-

ness—a character that would approach the present example more to

P. subtricarinatum. The suture-line, however, has no independent

second lateral saddle, hike that of P. subtricarinatum, but, as in P.

dravidicum, this second lateral saddle forms only the internal branch

of the first lateral saddle, and there is a large umbilical lobe. The

principal lobe, though, is deeper in the specimen here described, and

the umbilical lobe shorter, than they are in Kossmat’s figure. It may

also be added that the internal portion of the suture-line is very

variable ; the stem of the umbilical branch of the dorsal saddle may

be much broader than drawn, so that the details of the umbilical lobe

and the size of its lower internal and higher external branch may differ

considerably in consecutive suture-lines. No similar suture-line appears

to have been observed in any other species of Peroniceras, e.g. P.

schneeblii, Boule, Lemoine and Thévenin sp.,* which may be a related

form, occurring in Madagascar, does not show this type of umbilical

lobe, nor do the two examples of Peroniceras recorded by Crick + from

Zululand. One of these, sp. « (B.M., No. C18245), compared with

P. tridorsatum, Schliter sp., resembles P. czirnigi, Redtenbacher sp.,{

but not the “variety” figured by Grossouvre,§ and has a thinner

whorl-section than the present example. Since Crick only figured the

external half of the suture-line, the internal portion is here given for

comparison (PI. XXIII, fig. 2). The other example, sp. 8 (No. C18246),

is much fatter, and in section and crescent-shaped costation resembles

P. rousseauaxi, Grossouvre,|| but owing to the large size of these

specimens comparison with the small European types is difficult.

The tricarination is less distinct in this second specimen, and the

suture-line also is less well preserved, though very similar in the

character of the auxiliary lobes.

* Loe. cit. (ii, 1907), p. 37, fig. 20, pl. xii, figs. 2, 2a.

+ Loc. cit. (Third Report, 1907), pp. 226-7.

~ Loe. cit. (18738), p. 105, pl. xxiii, fig. 4.

§ Loc. cit. (1893), pl. xi, fig. 2.

|| Zoid. pl. xi, fig. 5a, b, p. 102.

On Cretaceous Cephalopoda from Zululand. 297

P. westphalicum, Schliiter sp.,* and particularly the specimen figured

by Grossouvre,f are close to the present example in ornamentation

and general appearance, but the tubercles are coarser; on the other

hand, in P. tridorsatum, Schliiter sp.,f and P. mouwreti, Grossouvre,§

costation is too fine and close.

Fic. D.—1. Mortoniceras stangeri, Baily sp. Upper Senonian, Umtam-

vuna River, Natal. la. Specimen No. C19444, British Museum, 1b.

No. C19448, comparable with Baily’s co-type No. 11368a (Geol. Soc. Coll.).

1c. No, C19440 (penultimate septum of an example 340 mm. in diameter).

Figs. 1 a-e are after drawings by the late G. C. Crick. 2. Mortoniceras

aff. wmkwelanense, Crick. Upper Senonian, Umkwelane Hill, Zululand.

(Specimen No, 5491.) (P. 234). All reduced to 3.

Since Mortoniceras stangeri, Baily sp., has tricarinate inner whorls,

and has, indeed, been included in Peroniceras by some writers’ (e. 9.

Kossmat), the external suture-lines of three examples, drawn by the

late G. C. Crick, and the internal portion of another example, showing

* Loc. cit. (1872), p. 45, pl. xiii, figs. 5 and 6.

+ Loe. cit. (1894), p. 98, pl. xii, figs. 1 and 4, a, 5.

t Loe. cit. (1876), pl. xli, figs. 3-5, ‘Jiingst. Amm.’ (1867), p. 26, pl. v, fig. 1.

§ Loe. cit. (1894), p. 100, pl. xi, figs. 3 and 4.

24,

298 Annals of the South African Museum.

good agreement with Baily’s type and one of Baily’s co-types

(11368a),* are here given for comparison (Pl. XXIII, fig. 3). The

distinct second lateral lobe and independent second lateral saddle (on

the umbilical wall) differ greatly from those of the specimen here

described, though the whorl-section, at a diameter of about 25 mm., is

that of a Peroniceras. At a stage corresponding to that of Peroni-

ceras sp. ind. (in Redtenbacher: A. sp. indet. cfr. A. tridorsatus,

Schliiter +), the costae of M. stangeri continue across the ventral area

with its three faint keels, something after the style of the ventral area

in Pseudotropites ultraliasicus, Canavari in Wahner.f If Baily’s

species is really of Upper Senonian age, as would appear from its

association with Hauwericeras gardeni and Pseudophyllites indra,s it

forms a striking case of convergence of a late Mortoniceras, charac-

terised by its suture-line, towards the Lower Senonian Peroniceras.

P. dravidicum occurs, in India, in the Middle Trichinopoly group,

which is considered to be of Coniacian (Lower Senonian) age. In

Europe, e.g. the North of France,|| Peroniceras subtricarinatum, west-

phalicum and moureti occur in the zone of Micraster cor-testudinarium.

Grossouvre | has all the species of Peroniceras, the exact horizon of

which is known, in the Lower and Middle Coniacian. Boule, Lemoine

and Thévenin** record what appears to be the Indian species from

Madagascar; Solger describes it from the Cameroons.

Locality.— High ground on north side of United Manuan Creek and

Umsinene River, almost opposite Junction. Coll. W. J. Wybergh.

The mode of preservation of this specimen (limonite, after pyrites ?)

is different from that of any other Zululand Ammonite that the writer

has examined.

* Baily’s type of A. sfangeri in the British Museum (Geol. Soe. Coll.,

No. 11366), a gigantic specimen of 325 mm. diameter, has the inner whorls

corroded ; the three co-types (11367, 11368 and 113684) represent three slightly

differing varieties, but the large series in the British Museum includes many

transitions and shows the great variability of the species.

+ Loc. cit. (1878), p. 125, pl. xxx, fig. 3.

t “Beitr. Kenntn. Tief. Zon. d. Unt. Lias N.O. Alp.,” part vii, ‘ Beitr. Pal.

Ost.-Ung.,’ vol. ix (1894), pl. iii, fig. 1 ¢ only.

§ According to Woods, loc. cit., pp. 346-7. Two examples of Pseudoschloenbachia

umbulazi, in the British Museum, from the Umtamvuna River, Pondoland,

have impressions of large Mortoniceras of stangeri affinity, in the same pieces

of matrix.

| Pruvost, “ Les Ammon. Sénon. d. Nord.,’ * Ann. Soe. Géol. Nord., vol. xxxix

(1910), pp. 365-8.

{ Loc. cit. (1894), p. 106.

** Toc. cit. (1907), p. 42 (as “ Schloenbachia (Peroniceras) subtricarinatum

d’Orb.”). See also Pervinquiére, loc. cit. (1907), p. 250.

On Cretaceous Cephalopoda from Zululand. 299

Gren. KOSSMATICERAS, de Grossouvre.

Sus-Gen. MADRASITES, Kilian & Reboul.

23. KossMATICERAS (MaprasITEs) BHAVANI, Stoliezka sp.

(BI EXE ches)

1865. Am. bhavani, Stoliczka. “Cret. Fauna 8. India” (Pal. Indica),

I. “Cephalopoda,” p. 138, pl. Ixix, figs. 4-7.

1897. Holcodiscus bhavani, Kossmat. ‘“ Unters. Siidind. Kreidef.,”

Beitr. Pal. und Geol. Ost.-Ung., vol. xi, p. 38 (145), pl. vill (xix),

figs. 5 and 6.

One example in the collection (No. 4909) agrees with the largest

form figured by Stoliczka (fig. 6), and has the following dimensions :

Diameter : ; : : . 65 mm.

Height of the Jast whorl . 40 per cent. of the diameter.

Umbilicus . ‘ ; , 29 * Ee

Thickness of the last whorl . . 80 es » wWhorl-height.

Stoliezka’s example has a whorl-height of 41 per cent., an umbilicus

of 26 per cent. and a thickness of 77 per cent., so that it differs only

in having a slightly narrower umbilicus; the umbilicus of the present

example, in width, agrees more with that of the smaller form figured

by Stoliczka. The inner whorls, shown in the umbilicus, are more

distinctly costate than are those of either Stoliczka’s or Kossmat’s

examples—possibly a matter of preservation.

The variety densicostata, Kilian and Reboul,* is much more finely

ornamented than the form here described, as are, to a lesser extent,

the slightly tuberculate var. seymowriana, Kilian and Reboult, K. (M. ?)

cumshewaense, Whiteaves, { and the variety of the latter form figured

as Holcodiscus cf. H. theobaldianus, Stoliczka, by Anderson.§ ‘The

writer has lately recognised the presence, in the Upper Senonian of

New Zealand, of Kossmaticeras (Madrasites) bhavani, Stoliczka sp.,

and K. (M.) cumshewaense (? Whiteaves) Kilian and Reboul,|| as well

as of K. (Gunnarites) aff. bhavaniforme, Kilian and Reboul, K.

* Loe. cit. (1909), p. 30, pl. xviii, fig. 1, pl. xv, fig. 4.

+ Lbid., p. 29, pls. xiv, Xv, xix.

t Loe. cit. (‘ Mesoz. Foss., 1884), p. 208, pl. xxiv, fig. 1.

§ Loe. cit. (1902), p. 101, pl. v, figs. 126-7. This form, however, is of

Horsetown age, and Whiteaves’ species also may be earlier.

|| Possibly close to the incompletely known 4K’. (Madrasites) meKayi, Hector

sp. (‘Catal. New Zealand Court.’ 1886, p. 57, text-fig. 19a, No. 4) (misspelt

mac Coyi in Haug, “'Traité,” II, ii, p. 1845), that Steinmann (Joc. cit., 1895, p. 28)

considered to belong probably to the group of K. (JZ) aemilianum, Stoliczka sp.

300 Annals of the South African Museum.

(Grossouvrites) gemmatum, Huppé sp., and of Pseudophyllites (Tetra-

gonites ?) sp. juv.*

The small K. (Madrasites) cf. madrasinum, Stoliezka sp., figured as

Holcodiscus sp. by Woods,t from Pondoland, is distinguished by its

pronounced umbilical tuberculation. The Madagascar form of Koss-

maticeras (Madrasites) figured by Boule, Lemoine and Thévenin{ as

“* Holcodiscus theobaldinus, Stoliczka sp. var.,” differs from the form

here described in having a larger umbilicus.

Crick described (in MS.) two species of Kossmaticeras from Pondo-

land as Holcodiscus natalensis and H. acuticostatus. He did not

compare them with Woods’ form, which is more involute, but stated

that their nearest ally appeared to be H. buddhaicus, Kossmat—a

statement with which the writer agrees. These two forms (B.M.,

No. C19432-3) differ little from each other, but are distinguished

from the species here described by being evolute (U = 34 per cent.

and 37 per cent. respectively) and by having umbilical tubercles

throughout. K. (Madrasites) fakwu and K. (M.) africanum, Hoepen

sp., are closely similar forms§$ represented in the collection from the

Durban Museum.

Kilian has recorded K. bhavani also from New Caledonia,|| and

related forms occur throughout the Indo-Pacific province, whereas the

European forms recorded as Kossmaticeras belong to other groups.

than the finely-ribbed South African Madrasites.

Locality.—South side of Manuan Creek Valley. Coll. W. J.

Wybergh.

Grn. PLACENTICERAS, Meek.

24. PLACENTICERAS cf. SUBKAFFRARIUM, Sp. NOv.

Two whorl-fragments (Nos. 4957 and 4958), possibly belonging

to the same individual, appear to be identical with the species described

* In collections kindly sent by Mr. Henry Woods, F.R.S., and by Dr.

Trechmann (see ‘ Geol. Mag.,’ n.s., dec. vi, vol. iv [1917], p. 338.)

+ Loe. cit., p. 336, pl. xli, fig. 2. Kilian and Reboul (Joc. cit., p. 62) compare

this form with the Antarctic K. (Jacobites) anderssoni. It may be near K. (1.)

africanum, v. Hoepen sp.

t Loe. cit. (1906), p. 26 (vol. ii), pl. vii, fig. 3 (vol. i).

§ “ Deser. of some Cret. Amm. from Pondoland,” ‘ Ann. Transvaal Museum,’

vol. vii, pt. ii (1920), p. 144, pl. xxv, figs. 3 and 4, and pl. xxvi, figs. 1 and 2;

p. 146, pl. xxvi, figs. 3-5.

|| There is a specimen of a Kossmaticeras (Madrasites) bhavani (Stoliczka),

var. densicostata, Kilian and Reboul, in the British Museum (No. C1536) from

St. Vincent, West Coast (Bourail), New Caledonia.

On Cretaceous Cephalopoda from Zululand. 301

from Umkwelane Hill (see p. 247), and the slight differences between

the fragments themselves and the figured specimen (Pl. XXI, fig. 2)

may be due to weathering. The ornament thus appears perhaps more

distinct, after the style of P. intercalare, Meek, and the periphery may

be just a trifle narrower. Like the type, these two examples differ

from P. tamulicum, Blanford sp., chiefly in being fatter in the umbilical

region. The suture-line is worn, but its plan agrees with that of the

Indian species.*

Locality.—South side of Manuan Creek Valley. Coll. W. J.

Wybergh.

NAUTILOIDEA.

Gen. CYMATOCERAS, Hyatt.

25. CyYMATOCERAS ? sp. cf. sustum, Blanford sp.

1861. Nautilus justus, Blanford. “Cret. Fauna 8. India” (Pal. Indica),

vol. 1, “‘ Cephalop.,” p. 22, pl. x, figs. 2-3.

? 1861. Nautilus bouchardianus (VOrbigny) Blanford, pars. Ibid.,

pl. iv, fig. 3 only.

1866. Nautilus justus (Blanford) Stoliezka. Ibid., p. 206, pl. xeiu, fig. 2.

1910. Nautilus justus (Blanford) Spengler, “‘ Untersuch. ii. d. Siidind.

Kreideform., pt. iv, Die Nautil. und Bel. d. Trichinopoly Distr.,”

Beitr. z. Pal. und Geol. Ost.-Ung., vol. xxiii, pt. iii, p. 142, pl. xiv,

A small specimen (No. 4935) agrees with one of the examples

deseribed by Crick? as Nautilus (2 Eutrephoceras) sp., both in whorl-

shape and elegant ornamentation, as in the mode of preservation

(brown limonite coating), but on account of its annular lobe and

elegans-like (if fine) striation is referred to Cymatoceras. At a diameter

of 40 mm. the whorl-thickness is 30 mm.; the siphuncle is centran

and the umbilicus not quite closed.

Spengler thought Crick’s Nautilus | Cymatoceras?| occlusus to be

close to the species here discussed, but the latter is less globose.

Cymatoceras kossmati, Spengler sp.,{ agrees in whorl-shape, but the

costation is much coarser, as it also is in the Utatur species C. kayeanum,

Blanford sp.

* Fig. 1c of pl. xxii, in Kossmat, loc. cit., 1895.

+ Loe. cit. (1907), p. 245, B.M., No. C18293.

ft Loc. cit, (1910), p. 129 (N. pseudo-elegans, VOrbigny m Blanford, loc. cit.,

1861, p. 33, pl. xviii, fig. 3).

302 Annals of the South African Museum.

This specimen is the only one in the collection from “ High Ground,

South Side of South Branch of Manuan Creek, just below Wagon-

Drift.” Coll. W. J. Wybergh. Its assumed Senonian age thus is

not supported by Ammonite evidence, and its mode of preservation

is distinct from that of any of the other specimens in this collection.

OBSERVATIONS ON THE MANUAN CREEK FAUNA.

The relations of the Manuan Creek fauna have been discussed in

detail by Crick and Newton, but it has been mentioned in the intro-

ductory part of this paper that the presence of Albian, Cenomanian

and Lower and Upper Senonian forms was not clearly recognised, so

that a revision of the faunas has become necessary. With regard to

the Ammonoids, the few Senonian forms here described have already

been referred to in the observations on the Umkwelane Hill fauna.

They include the Coniacian—

Peroniceras cf. dravidicum, Kossmat sp.,

to which have to be added the two Peroniceras recorded by Crick, with

the Cenomanian False Bay fauna, and here referred to on p.296, namely:

Peroniceras ct. czirnigi, Redtenbacher, sp.

Peroniceras cf. rousseauxi, de Grossouvre.

To the presumably Campanian Ammonites described in this paper,

namely,

Kossmaticeras (Madrasites) bhavani, Stoliczka sp.

Placenticeras cf. subkaffrarium, nov.

must be added the following forms described by Crick from the South

Branch of the Manuan Creek :

Gaudryceras pulchrum, Crick.*

Gaudryceras cf. kayei, Forbes sp. | Gaudryceras sp. in Crick,

p. 238. ]

Diplomoceras ? sp. | Anisoceras sp. in Crick, p. 239. |

Baculites ef. capensis, Woods | Baculites sp. in Crick, p. 240. |

Hauericeras sp.¥

The Upper Senonian fauna of the Manuan Creek District thus

shows a greater resemblance to the fauna of the Umzamba Group of

Pondoland (so-called Umtamvuna Beds) than to the Umkwelane Hill

fauna, which is much nearer, geographically, but which represents a

different facies.

* Gaudryceras sp. (p. 239 in Crick) probably is a badly weathered fragment

of this species.

+ P. 242, pl. xv, fig. 5, the inner whorls erroneously being represented as

having anacute venter. For Hauericeras sp. (p. 243), see above, under Uhligella

sp. n. ef. stoliczkai, Kossmat sp. (p. 276, footnote f).

On Cretaceous Cephalopoda from Zululand. 303

Acanthoceras sp. (in Crick, p. 241),

comparable with) A. datum, Crick, and A. quadratum, Crick, is of

Cenomanian age, like the “ False Bay” fauna described by Crick. The

resemblance of this fauna to that of Northern Africa on the one

hand and India on the other was noticed by Pervinquitre,* who

thought it quite evident that inter-communication between these areas

must have been easy.

There is, then, evidence in the Manuan Creek district of the presence

of deposits of Albian, Cenomanian, Lower Senonian (Coniacian) and

Upper Senonian (Campanian, incl. Maestrichtian?) age. The first

of these formations, perhaps, is the most important, and of Crick’s

Manuan Creek Ammonites the following probably belong to it:

Phylloceras sp. (p. 236) [identical with Ph. velledae, Michelin

sp., described in this paper].

Lytoceras crenulatum, Crick (p. 256).

Schloenbachia sp. (p. 240) [probably Pseudophacoceras manu-

anense, nov. |.

Desmoceras sp. (p. 241) [Latidorsella ? sp. ind. ].

‘“ Hauericeras” sp. (p. 243) [probably Uhligella sp. n. cf.

stoliczkai, Kossmat sp. |.

A second specimen of a “ Hauericeras”? sp. and a “? Beudanti-

ceras beudanti” also probably belong to this last form, and other,

unrecorded, specimens from the South Branch of the Manuan Creek

in Crick’s collection are :

Anisoceras sp.

Douvilleiceras sp.

* Schloenbachia aff. delaruei, V@Orbigny sp.” | Dipoloceras sp. |}.

All the forms recorded by Crick from the Middle Tributary of the

Manuan Creek, also, probably, are of Albian age, namely :

Puzosia concinna, Crick (p. 245).

Puzosia compacta, Crick (p. 246).

Schloenbachia sp. (p. 247) | Dipoloceras sp. |}.

Hysteroceras sp. (p. 248) | Brancoceras sp. |.

These Albian forms allow of more exact correlation. Including

those described by Etheridge from the Umsinene River (the types of

which, however, the writer has not examined), the list of Albian

Ammonoids, then, is as shown in Table I.

The forms mentioned in this list indicate that probably the Middle

Albian (mammillatum, delaruei, and cristatum zones), and the Upper

Albian, up to the upper rostrata zone above, are represented, the latter

zone apparently transitional to the Lower Cenomanian, exposed near

_ *«Amm. d. Crét. Algér.,” ‘Mém. Soc. Géol. France,’ Pal., vol. xvii, No. 42

(1910), p. 81.

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On Cretaceous Cephalopada from Zululand. 305

the junction of the Umsinene River and the Manuan Creek. These

horizons of the Upper and Upper Middle Albian, according to the

succession tentatively suggested by the writer in his description of the

Albian fauna of Angola, appear to he:

Upper rostrata horizon : . “‘post-rostrata” (XIII, XIT).*

Lower _,, ms, - : >» “rostrata s.s..” (XL):

Upper varicosum horizon . » <“prerostrata”’ (X, Xb).

Lower = * : . “bouchardianum’’ (1X a).

Upper cristatun . Pals Bg Sesser (VALI)

Lower - ie ; ‘cornutum” (VII, VI).

It is to be hoped that further wallesiine will be done, with a view to

tracing the succession of the Manuan Creek Albian in detail and eluci-

dating the somewhat uncertain relations of, e. g., the outcrop of the

‘‘Umsinene deposit,” exposed chiefly in the bed of the southernmost

tributary of the Manuan Creekt+ and apparently comprising Middle

Albian and Campanian forms, with the neighbouring Coniacian and

Cenomanian exposure near the junction of the Umsinene River and the

Manuan Creek. It may be added that the forms of the highest Albian

(upper rostrata zone, with Stoliczkaia) all come from one locality, namely

the “south side of the Manuan Creek Valley” (V in the list), and

that none of the other localities apparently have yielded examples

that are referable to this highest Albian. In the case of the Lytoceras

and the Desmoceratids (2, 3 and 7 in the list), the comparison with

Indian species might suggest a high horizon in the Albian, only the top

of this formation, apparently, being found in India; but the Ammonites

probably are not specifically identical. On the other hand, it is just

these forms that connect the Zululand fauna with the types special to

the Indo-Malgascan Province, for nearly all the remaining Ammonites

correspond with well-known European types, with the exception of

Pseudophacoceras manuanense and of Dipoloceras sp. nov., which are

more closely comparable with American} species, and which, e.g. in

Mexico, occur several hundred feet below the equivalent of the rostrata

zone. ‘The writer, in another place, when discussing the relations of the

Angola fauna, referred to the probable immigration of these elements

into the African region from South America along the southern edge of

* These numbers refer to the beds at Folkestone.

+ Anderson, loc. cit. (Third Report, 1907), p. 58.

t Lemoine (‘ Pal. Nord. Madagascar,’ 1906, pp. 204-5) recorded “ Schloenbachia

roissyt (= Sehl. acuto-carinata, Shumard)” and “ 8. mirapeliana, d’Orbigny sp.

(= 8. buarquiana, White sp.),” but in the following year, Boule, Lemoine and

Thévenin described a 8S. (Mortoniceras) ctf. inflatiformis, Szajnocha, which

probably is a Subschloenbachia, whereas their “ Lower Cenomanian Spheno-

discus”? may be a Pseudophacoceras, allied to P. manuanense and P. buarquianum.,

306 Annals of the South African Museum.

the Africano-Brazilian Continent (“ Brasilia”), whereas undoubtedly

there also was free and direct communication through the Channel

of Mozambique and the Sea to the North with the Mediterranean.

That a larger number of forms are common to the European and the

Zululand Albian, than to the latter, and e.g. the fauna of Tunis,

probably is partly due to the fact that this formation has been much

more thoroughly explored in Europe, and the Middle and Lower Upper

Albian of North Africa, are still incompletely known.

It is interesting to note that the South Queensland forms of

Subschloenbachia, described by Etheridge, are more closely allied to

Zululand than to Indian forms; but since similar types occur again in

Europe and in Mexico, perhaps no significance may be attached to this.

Moreover, some of the Queensland forms appear to belong to the preros-

trata horizon, which, probably, is not represented in Southern India,

the lowest Utatur Beds, according to Kossmat, including Stoliczkaia

and Mantelliceras, in addition to Subschloenbachia, and such forms as

Prohysteroceras propinquum and Neokentroceras (gen. nov.*) gracil-

limum, Kossmat sp. On the other hand, Dipoloceras of the cristatum

eroup, and Subschloenbachia have now been found in Nigeria, and the

genera Douvilleiceras, Subschloenbachia, Brancoceras (7), Stoliczkara,

also Desmoceratidae and Hamitidae occur in Angola; yet, it may be

held with Boule, Lemoine and Thévenint that “the resemblances

with West Africa (Angola) are feeble, and it seems . . . that

West Africa formed part of another zoological province.” This

latter is characterised by the special genus E/obiceras} and the group

of Subschloenbachia evoluta, nov., found only in Nigeria, the Elobi

Islands, and Angola, possibly also in Tunis, whereas, as has been

mentioned, the large Lytoceras, found again in Pacific regions, and

certain Desmoceratids, connect the Zululand fauna with the special

developments of the Indo-Malgascan Province. On the other hand,

easy communication with the Mediterranean facilitated extensive

faunal equalisation during the Aptian and Albian, extending to South

America along the Northern§ and Southern shores of the Africano-

* A post-Subschloenbachia development (Genotype = N. curvicornu, nov. from

Angola, allied to N. tectoriwm, White sp.).

+ Loc. cit. (1907), p. 72.

t Gen. nov. (Genotype = Schloenbachia elobiensis, Szajnocha, 1885, pl. iv,

fio. 1). Described fully in the writer’s Angola paper.

§ The westward extension of the Tethys is indicated by the close corre-

spondence with European developments shown by the Upper Jurassic of Mexico,

the Lower and Middle Cretaceous of Venezuela and Colombia, the “Gosau

facies” of Jamaica and Mexico, ete. (see Suess, ‘ Face de la Terre,’ vol. ili, pt iv,

p. 1680). An arm of this sea reached south as far as Angola.

On Cretaceous Cephalopoda from Zululand. 307

Brazilian Continent.* That the latter was still in existence in

Upper Senonian times appears to be indicated by the extremely close

resemblance of the Kossmaticeras-beds of Pondoland with those of

Antarctica, Southern Patagonia, Chili and New Zealand, all of which

are largely made up of glauconitic, calcareous sandstones, and

apparently pass uninterruptedly into the lowest tertiaries.+

The lower Albian beds (Clansayes horizon) that bridge over

the gap between the Aptian fauna of Powell’s Camp and the mam-

millatwm-zone of the ‘** Umsinene River deposit,” if present at all, have

not yet yielded Ammonites.

IV. AMMONOIDEA FROM ISOLATED LOCALITIBS.

A. Nortu-West SHore or Fatse Bay.

The present collection only includes two specimens from this locality

(Coll. W. J. Wybergh) namely :

Mortoniceras vanuxemi, Morton sp.

Bostrychoceras ? sp.

They are preserved in a brownish, marly sandstone, very friable, and

thus different from the matrix of Crick’s Cenomanian fauna, but this

difference may partly be due to weathering. On the other hand, M.

vanueemi can definitely be dated as Campanian (zone of M. delawa-

rense),, and the other specimen, as well, is comparable with a Pondoland

form, considered by Woods to be of Campanian age. The outcrops of

these beds along the north-western edge of False Bay are referred to by

Mr. W. Anderson in the Second§ and Third Reports|| of the Geological

Survey of Natal and Zululand. Crick*{ assumed that this was the

locality from which the fossils were obtained that he described under

the title of “The Cephalopoda from the Deposit at the North End of

False Bay, Zululand.” According to Mr. W. Anderson, however,**

these Cenomanian Ammonites came from the river-bank near the

* Engler (“Ub. Florist. Verwandsch. zw. d. Trop. Afr. und Am., ete.,” ‘ Sitz.

K. Preuss. Ak. Wiss. Berlin,’ 1905, i, p. 229) deduced the existence of large

islands or a continent connecting Brazil with Africa from a study of the

existing flora.

+ Seein O. Wilckens, “ Die Kreideform. v. Neu-Seeland,” ‘ Geol. Rundschau,’

vol. xi (1920), pp. 189-91.

t Haug, ‘ Traité de Géologie,’ IT, ii, p. 1170.

§ 1904, p. 48.

|| 1907, p. 57.

" Ibid., p. 164,

** 1907, p. 60, See also Crick (‘ Geol. Mag.,’ August, 1907), p. 344.

308 Annals of the South African Museum.

junction of the Manuan and Umsinene Rivers, whereas from the

western bank of False Bay he only ‘‘ obtained some fragmentary fossils,

very badly preserved, which Mr. Etheridge was inclined to think

belonged rather to the Tertiary than to the Cretaceous System.” *

Gen. MORTONICERAS, Meek.

1. MorronicERAS VANUXEMI, Morton sp.

(Pl. XXIII, figs. 4a, 0.)

1892. Mortoniceras vanuxzemi (Morton), Whitfield. ‘ Gast. and Ceph.

Raritan Clays,’ Mon. U.S. Geol. Surv., vol. xvi, p. 252, pl. xlu,

figs. 5-4.

1907. Mortoniceras delawarense (Morton), Stuart Weller. Rep. Cret.

Pal. New Jersey, vol. iv, Pal. Ser., p.887, pl. civ, figs. 4-5 only,

non pl. cil.

A fragmentary specimen (No.4947), showing interlocking suture-lnes

and an impression of portions of the inner whorls, agrees with the

above figures and undoubtedly belongs to this species, which is

distinguished from its close ally M. delawarense, Morton sp., by being

more compressed and less coarsely ornamented. ‘I'he suture-line is of

the same type as those of M. woodsi (Pl. X XI, fig. le) and M. soutoni

(Baily) (Pl. XX, fig. 4) figured in this paper, but differs in the terminal

branches of the lateral lobe, which is perhaps due to the interlocking

of the (last few ?) suture-lines.

M. woodsi, nov., is an extreme development of M. canwerenii, with

overhanging wmbilical edge and fine and close ornament.

GEN. BOSTRYCHOCERAS, Hyatt.

2. BostTRYCHOCERAS, ? sp.

1906. Heteroceras sp., Woods. ‘Cret. Fauna of Pondoland.” Ann.

S. Afr. Mus., vol. iv, pt. vii, No. xii, p. 339, pl. xlu, fig. 4.

A small fragment (No. 4952), about 28 mm. in length, belongs to

the form described by Woods and has four intermediate ribs between

the flares, but it is too fragmentary to determine whor]l-shape and

coiling. The reasons for referring the form to Bostrychoceras are

discussed under B.? sp. nov. (p. 252) of the Umkwelane Hill fauna.

Hamites sp., described and figured by Jimbo,t seems to belong to a

* Hbid.. ps Ol.

+ Loe. cit., p. 40, pl. ix, fig. 1 only.

On Cretaceous Cephalopoda from Zululand. 309

similar form. Woods noted the resemblance of this form to Hyphanto-

ceras reussianum, d’Orbigny sp.; H. fleznosum, Schliiter sp.,* and

H. sp. nov. (= Heteroceras sp. in Woods),? are other species of

the genus Hyphantoceras that show superficial resemblance, but are

of Turonian age.

The example is distinguished from the B. ? sp. ind., found at Umkwe-

lane Hill (p. 255), by having coarser costation and four, not three,

intermediate ribs, so that at a similar whorl-height the flares are

about twice as far apart in the present form.

B. POWELL’S CAMP, UPPER CATEMBE, PORTUGUESE

EAST AFRICA.t

The three Ammonites from this locality (Nos. 5117-19) are pre-

served, as casts, in a light yellowish-grey, caleareous sandstone, and

two of them show portions of the test. This matrix, stained yellow

with rust in places, is not unlike the lighter-coloured portions of the

calcareous sandstones of Umkwelane Hill, and apparently is similar

to that of the Aptian fauna of Delagoa Bay, recorded by Kilan§ and

described by Krenkel. ||

The three forms described below are—

Aconeceras nisoides, Sarasin sp.

Cheloniceras gottschei, Kilian sp.

Cheloniceras (Acanthoplites ?) delagoense, Krenkel sp.

The assemblage, thus, is the same as that recorded from Delagoa

Bay, of the same sandy, littoral facies, and undoubtedly of Aptian

age. Dr. Kitchin] has already referred to this Aptian fauna from

Delagoa Bay in his important memoir on the Uitenhage Beds.

Krenkel was of opinion that the Bedoulian (Lower Aptian) as well

* Loc. cit., p. 108, pl. xxxii, figs. 10-12.

+ ‘Q.J.G.S., vol. lii (1896), p. 75, pl. ii, figs. 7 and 8.

t When writing this account, the author was under the impression that

Powell’s Camp was in Zululand, but it now appears that though its exact posi-

tion and the meaning of “ Upper Catembe” are unknown, it may be in the

neighbourhood of Delagoa Bay, Catembe being marked on Jeppe’s map as being

on the right bank of the Tembe river, some five miles from the mouth.

_ § “Ub. Aptian in S. Afr.,” ‘Centralbl. f. Min., August, 1902, p. 465; also

‘Bull. Soc. Géol. France’ (4), II, 1902, p. 358; and ‘Comptes Rendus,’ exxxv,.

No. 1, (July 1902), pp. 68-71.

|| “D. Aptfossil. d. Delagoa Bai,’ ‘N. Jb.f. Min., ete.,’ 1910 (I), pp. 142-168,

pl. xvii.

| “The Invertebr. Fauna and Pal. Rel. of the Uitenhage Ss.,” ‘Ann. 8. Afr.

Mus.,’ VII, pt. 11, No. 3 (1908), p. 57.

310 Annals of the South African Museum.

as the Gargasian (Upper Aptian) were represented in the Delagoa Bay

fauna, and he considered “ Oppelia” nisus (VOrbigny) to be a typical

representative of the latter division. Haug* has a zone of “Oppelia”’

nisus above the zone of “ Parahoplites” deshayesi and Ancyloceras

matheronianum, Which ineludes the whole of the Bedoulian, but

v. Koenent records ‘* Oppelia” nisoides, indistinguishable from the

example here described, from the zone of “ Parahoplites” weissi, which

constitutes his lowest Aptian, below the deshayesi zone. Kiliant{ con-

sidered that the occurrence of Aconeceras niswim in the Upper Aptian

(Gargasian) of the South of France indicated a migration of this

genus from north to south, but it seems to the writer that the range

in the Aptian of this form and of its close allies, like A. nisoides, is

not yet known.§

It has to be noted in this connection that the suture-lines

of the two forms of Cheloniceras agree with that of Ch. cornue-

lianum (WVOrbigny) much more than with those of the <Acan-

thoplites of the bigoureti group; and Ch. cornuelianum is put into

the Lower Aptian both by Kilian|| and by R. Douvillé,{{ whereas,

according to Haug,** at la Bedoule. Ch. cornuelianum occurs both

in the lower Aptian, 7.e. in the true Bedoulian, and in the higher

division (Gargasian), and it is associated with ‘* Oppelia”’ nisus also

at other localities.+*

The relations of the Delagoa Bay fauna with those of other parts of

Africa, of Madagascar, India, ete, were ably discussed by Krenkel in

the paper quoted above, and in another memoir on the Lower Creta-

ceous of East Africa.tt Zwietzycki§§ later described a fauna, from

doubtful localities in Kast Africa, that included Acanthoplites (?) rauff,

Zwietzycki sp., and Diadochoceras nodosocostatumn, d’Orbigny sp. The

former is compared with Cheloniceras cornuelianum, and only differs

from the example here described as Ch. gottschei in retaining bitubercu-

lation to a larger diameter ; but it probably is an Acanthoplites of the

* Traite, II, ii, p. 1170.

+ “Amm. d. Nordd. Neoc.,” ‘Abh. K. Pr. Geol. L.A.,’ N.F., Heft. 24 (1902), p. 51.

t In Lethaea Geogn. II, Mesoz. 3, Kreide, I, 3, 1913, p. 338.

§ In his latest paper (‘ Trav. Lab. Géol. Univ. Grenoble, vol. xii [1919],

p. 94), Kilian has A. nisoides as Gargasian.

|| Loc. cit. (1910), p. 281.

G ‘Pal. Univers.,’ 1911, No. 209.

** Loc. cit., p. 1198.

Tip gs ps 1183, 1191, loc. cit. (Haug).

tt ‘ Beitr. Pal. Ost.-Ung.,’ vol. xxiii (1910), pp. 230-250.

§§ “D. Ceph. d. Tendaguru-Sch. i. Deutsch. O. Afr.,” ‘Wiss. Erg. d. Tendag.

Exp., 1909-12,’ pt. iii, ‘Arch. f. Biontol., 1914, vol. iii, pt. 4.

On Cretaceous Cephalopoda from Zululand. dll

lower Clansayes horizon = Zone III in Jacob.* Kilian also puts this

horizon as the base of the Albian, but Haug? has the Clansayes Beds

(= “zone of D. nodosocostatum”’) as the uppermost Aptian. At any

rate, this East African fauna described by Zwietzycki includes higher

horizons than do the Aptian faunas of Delagoa Bay and of the present

locality.

If the latter come from one horizon, then it is probable that this

corresponds with what is called the furcatus-zone (?) in the table

below. Stolley§, probably wrongly, quoted Ch. cornuelianum from his

“ Middle Gault”’ zone 1, which misled the writer when drawing up

the correlation notes, kindly inserted by Mr. L. Dudley Stamp in the

“Report of Excursion to Tilburstow Hill and Nutfield.”|| The corre-

lation with Stolley’s horizons of the Upper Aptian, therefore, has here

been corrected :

Upper Albian . Hor. IX-XIII (Foikestone) . Hor.7 (Stolley) VI (Jacob).

(Upper Gault)

(Hor. I-VIII (Folkestone) : sO . V

Middle Albian . / (Lower Gault)

(« mammillatum bed’

[ tardefureata Z { ed ee subz., Sal IV

Lower Albian .- imilletian. ,, » 4)

Lean _ (jacobi 2 \

| odosocostat. z. +" ; en

{ nolant lj

Upper Aptian . {subnodosocostatum z. 5 IIb

(Gargasian) .|fureatus z. (2) . ed Ila

Lower Aptian . ( ee a 5 ; a | I

(Bedoulian) sss) 05 zs {°

{ bodei z. 1

Gen. ACONECERAS, Hyatt, 1903.

1. ACONECERAS NISOIDES, Sarasin sp.

(Pl. XXVI, figs. 4a, b, text-fig. B 9, p. 241.)

1893. Oppelia nisoides, Sarasin. ‘“ Etude s. 1. Oppelia, ete.,” Bull. Soe.

Géol. France (3), vol. xxi, p. 155, pls. iv—vi, figs. 10 a—e, text-figs.

3 and 5 (p. 154).

1902. Oppelia nisoides, Sarasin. V. Koenen, ‘“*Amm. d. Nordd.

Neocom.,” Abh. K. Pr. Geol. L. A., n.F., Heft 24, p. 51, pl. xlv,

figs. 2 and 3.

* Loc. cit. (1907), pp. 296-806.

+ Loc. cit. (1913), p. 341.

tf <Traite, p: 1199:

§ “Die Glied. d. Nordd. Unt. Kreide,” ‘ Centralblatt f. Min.,’ 1908, p. 242.

|| ‘Proc. Geol. Assoc.,’ 1921, vol. xxxii, pt. 1, pp. 30-32.

312 Annals of the South African Museum.

1913. Adolphia nisoides, Kilian. ‘ Lethaea Geognostica. IT, Mesoz.,”

3, I, ‘ Unterkreide,’ fase. 3, pp. 337-8.

A completely septate specimen (No. 5119), with the suture-lines

well shown, has the following dimensions :

Diameter : 39°5 mm.

Height of last whorl. . Od per cent. of the diameter

Thickness: =n: : Sa We a BS

Umbilicus . 1 Bees s i

Being a cast and slightly worn, the specimen has the fine ornament,

which distinguishes the species from A. niswm, d’Orbigny sp., only

preserved in one or two places, and near the umbilicus, but there is

very good agreement in all characters with Sarasin’s type. Krenkel’s*

Delagoa Bay specimen (Oppelia nisus) agrees with the present example

in smoothness and in whorl-section.

The hollow keel is the important distinguishing feature of this form

from the flat and smooth examples of Pseudoschloenbachia, e. &.

P. griesbachi, Crick (M.8.) sp.t which have a very similar suture-line

(compare figs. B8 and BY, p. 241), but less parallel sides, and which

are of Senonian age.

Hyatt’s genus Aconecerast has priority before Stolley’s Adolphia,§

and Hyatt may have been right in assigning this genus to Desmocera-

tide. It probably has nothing to do with Oppelide.

Gren. CHELONICERAS, Hyatt, 1903.

2. CHELONICERAS GOTTSCHEI, Kilian sp.

(Pl. XXVI, figs. 1 a-d.)

1902. Acanthoceras (Parahoplites) martini, d’Orbigny sp., var. gottscher.

Kilian, “‘ Apt. i. Stidafr.,” Centralbl. f. Min., etc., p. 465.

1910. Douvilleiceras martini, var. gottschei, Kilian. Krenkel, “ Apt-

foss. d. Delagoa Bai,’ N. Jb. f. Min., etc. (1), p. 144, pl. xvii,

figs. 4, 5, 8, 9.

* Loe. cit., p. 142, pl. xvii, figs. 1 @ and 4, and p. 164.

+ = A. wmbolazi, Griesbach (‘Q.J.G.S.,’ vol. xxvii [1871], p. 63, pl. iii, fig. 1),

non A. umbulazi Baily (B.M., No. C19428), a smooth form of 103 mm. diameter,

with the greatest whorl thickness at the middle of the side, not near the umbilical

border, as in A. umbulazi, typus; with umbilical tubercles, but no costation,

and with faint constrictions, like P. papillata, G. C. Crick (M.S.) sp.

t ‘ Pseudoceratites,’ 1903, p. 100.

§ «Ub. Nordd. Aquival. Clansayes Fauna, ete.,’ ‘Centralbl. f. Min., 1907,

p. 269 (foot-note).

On Cretaceous Cephalopoda from Zululand. 313

A specimen (No. 5117) of 72 mm. diameter, with only the body-

chamber portion (occupying three-quarters of a whorl) well preserved,

and with the inner whorls largely replaced by crystalline calcite, has

the following proportions :

Height of the last whorl . 938 per cent. of the diameter

Thickness x 5 . 40 x %» 9

Umbilicus —. : . 8d 3 : -

The coronatiform innermost whorls (diameter = 4 mm.) have a

depressed whorl-section with prominent lateral spines and constrictions,

like the ammonite next described, and like Cheloniceras royerianum,

d’Orbigny sp. The following stage, represented by Krenkel’s figures,

and comparable with Sinzow’s examples of Ch. seminodosum (Sinzow)

and Ch. meyendorfi (d’Orbigny),* is badly preserved. Only the first

prominent rib, at the beginning of the well-preserved body-chamber

portion, has a lateral tubercle, in addition to the umbilical one, so that

there is loss of the characteristic bituberculation, as stated by Krenkel,

and as shown in the specimen of “ D. cornuelianuwm” figured by Kihan.t

The latter, however, has lost the umbilical tubercle, as well as the

lateral one, whereas in the form here described this umbilical tubercle

remains distinct on some of the longer ribs, which feature approaches

the present example to Ch. albrechti-austriae (Hohenegger), Sinzow

sp.,t to Ch. tschernyschewi, var. laticosta, Sinzow sp.,§ and to other

forms of the cornwelianum group.||

On account of the imperfect condition of the present example,%

identification with Kilian’s form, or with any of the other species of

the cornuelianum group, is, perhaps, impossible, but though Ch.

gottschei was only figured in small examples, it appears probable that

the present specimen represents the adult stage of that form, showing

“decline” of the costation, as does Ch. meyendorfi (VOrbigny) in

Sinzow.** The present example, at any rate, is closer to the

* “Beschr. einiger Douvilleiceras-Arten a. d. Ob. Neocom. Russl.,” ‘ Verh.

Russ. Kais. Min. Ges.’ (ser. 2), vol. xliv (1906), pl. i, figs. 6 and 8.

+ Loc. cit. (1913), p. 339, pl. ix, fig. 3.

+ Loe. cit. (1906), pl. iv, figs. 1 and 2, and Kilian, loc. cit. (1913), pl. viii, fig. 2.

§ Lbid., p. 187, pl. iii, fig. 1, and Kilian, Joc. ctt. (19138), p. 340, pl. ix, fig. 5.

|| A specimen in the British Museum (No. 46590), from the Isle of Wight, is

almost indistinguisable from the present form in ornament and suture-line,

but the whorl-section increases more rapidly in width.

€ The restoration of the inner whorl, given in fig. 1 ¢, is somewhat problem-

atical. Only two or three ribs are shown (on one side only), and of these one

has the inner, the other only the outer, tubercle preserved.

#* Loc. eit. (1906), pli, fic: 8:

314. Annals of the South African Museum.

Delagoa Bay form than is Kilan’s “ D. cornuelhianum,” cited above, to

the specimens of the same species figured by Sinzow.*

Kilian t put “ D.”’ martini, var. gottschei, into the group of “ D.”

martini, but if the present specimen is correctly identified with

Krenkel’s Ammonites, the character of the outer whorl and the

suture-line approach the South African form more to the cornuelianum

group. The writer, therefore, has raised this variety to an inde-

pendent species. 'I'here is no differentiation of the peripheral area as

there is in the martini group, e.g. in “ D.” martini, var. orientalis,

Jacob.

Ch. kiliani, v. Koenen sp.,§ differs in suture-line, but is close to

the South African form as regards costation. The lateral tubercle,

however, persists on some of the ribs in y. Koenen’s species, and the

whorl-section is less depressed.

The suture-line (only the last one or two are visible, in addition

to the immature ones at a diameter of 4 mm.) is characterised by a

high external saddle and a very wide lateral saddle. The leaflets

subdividing the equally wide lateral lobe are unusually large, and

there is good agreement with the suture-lines of Ch. cornuelianum

and Ch. meyendorfi (VOrbigny) in Sinzow.|| This type of suture-

line distinguishes the form here described from the somewhat. similar

Acanthoplites of the aschiltaensis-bigoureti-bergeroni group 4 of a higher

horizon, but it is interesting to note that in a still later group, namely

that of Acanthoceras giltairei, Pervinquicre** and A. euomphalus

(Sharpe),+¥ this wide lateral lobe with its prominent, median saddle tft

* Tbid., pl. i, figs. 1 and 2.

+ Loe. cit. (1913), p. 340.

t In Jacob and Tobler, “ Gault de la Vallée de ?Engelberger Aa,” ‘Mém.

Soc. Pal. Suisse, vol. xxxiii (1906), pl. i, figs. 1-3.

§ Loe. cit. (1902), p. 406, pl. xxxiii, figs. la and b.

|| Loe. cit., text-figs. 1 and 2, pp. 160 and 164.

§ See Seunes, “Amm. du Gault,” ‘ Bull. Soc. Géol. France’ (3), xv (1887),

pls. xii-xiv; and Anthula, “ Kreidefoss. d. Kaukasus,” ‘ Beitr. Pal. Ost.-Ung.,’

xii (1899), pls. ix—xiii.

** Cf. loc. cit. (1907), text-fig. 108 on p. 286.

+t “ Foss. Moll. Chalk,” ‘ Mon. Pal. Soe., II (1854), p. 31, pl. xiii, figs. 4a-c.

tt See Crick, “Note on A. ewomphalus,” ‘Geol. Mag.,’ n.s., dee. iv, vol. vi

(1899), pp. 252-8. A specimen of this rare species in the writer’s collection

has the lateral saddle rather larger than the saddle subdividing the lateral

lobe, which, in Crick’s figures, might be mistaken for the former. Sharpe’s

fig. 4c represents the relative sizes more accurately. This type of suture-line is

a development of that of Acanthoceras cunningtoni, Sharpe sp., and of A.

sussexiense, Mantell sp. (Sharpe, loc. cit., pl. xv, figs. 1d and 2c), and the resem-

blance to Douwvilleiceratide is a case of convergence.

On Cretaceous Cephalopoda from Zululand. 315

is found again. The inner whorls of the present example, however,

show that the resemblance with certain Acanthoceratids, notably

Calycoceras naviculare, Mantell sp.* is quite superficial.

The Persian example of “ Acanthoceras cornueli” recorded by H.

Douvillé+ is much more coarsely costate than the specimen here

described.

The very large Douvilleiceras, referred to on pp. 220 and 303 as

coming from the South Branch of the Manuan Creek, belongs to a

different group of forms. It somewhat resembles the large (and more

rapidly increasing) “ Pachydiscus’’ Waageni, Anthula,} im its closely

costate outer whorl, but appears to be a development of the Albian

mammillatum group (as far as can be judged by the poorly preserved

younger whorls), and possibly is a very large example of the form

figured by Etheridge.§ It does not appear to have anything to do

with the Aptian form here described.

* Also recorded from Madagascar, though the two forms figured by Boule,

Lemoine and Thévenin (Joc. cit., 1907, p. 30, pl. viii, figs. 1 and 2) are very

doubtful. Fig. 2 may be a Mantelliceras, with smooth ventral area on the

inner whorls, whereas Calycoceras, which is a post-Metlacanthoplites stock, has a

median row of tubercles in the young. In Calycoceras gentoni (Brongniart =

Sharpe’s figs. 3 and 5, pl. xviii) all the ventral tubercles disappear at about the

same time (‘ Pal. Univ.,’ 1911, No. 223) ; in C. naviculare, Mantell sp. (lectotype,

Sharpe’s figs. 1 and 8, pl. xviii (B.M. No. 36834), Mantell’s original being useless),

the two ventro-lateral rows persist longer than the median row. Mantell’s type,

refigured in a posthumous paper by Crick (“ 4. zavicularis, Mantell,” * Proc.

Mal. Soc.,’ vol. xiii, 1919, pp. 154-160, pl. iv) is too worn and scraped about to

show any tubercles, and the writer believes that what Crick (p.157) had

considered as differences of specific value cannot be relied on, some of the ribs

being artificially carved, or at least scraped, and the original shape of the whorl

is quite unrecognisable. ‘The resemblance to the Indian and Portuguese forms,

which are very tumid-whorled, and to d’Orbigny’s figure may not be so great

as appears from a comparison of the figures, and in India, as in the English

Chalk, a number of undescribed forms of Calycoceras occur; for Stoliczka

(p. 74) states that “there are specimens which have scarcely any trace of either

lateral or dorsal tubercles, even in the youngest stages,” 7.e. forms near to

C. baylei, Pervinquiére (= A. sarthacense, Bayle). Peron and Pervinquiére had

drawn attention to the similarity between Calycoceras of the Cenomanian and

the Turonian Fagesia. Crick (in coll.) had labelled Mantell’s type “ Fagesia

navicularis,’ but since he did not refer to this genus in his last paper, he

probably came to the conclusion that the specimen, after all, was a Cenomanian

* 4eanthoceras,” as he did in his earlier work on the False Bay fauna (p. 205).

+ “Mission Scientif. Perse, Morgan,” vol. iii, “‘ Et. Geol.,” pt. iv, ‘ Pal.,’ 1904,

p. 231, pl. xxviii, figs. 1 a, b.

t Loc. cit., p. 106, pl. ix, figs. la, 6; Sinzow (loc. cit., 1906), p. 164, pl. 1,

fig. 10 (as Douvilleiceras meyendorffi, var. waageni).

§ Loc. cit. (Third Report), pl. v, fig. 1. There also is a resemblance to

Choftat’s Acanthoceras marques-costai (‘ Conducia, 1903, p. 27, pl. vii, fig. 2),

but this form is compared with Cenomanian Ammonites.

316 Annals of the South African Musewni.

Hyatt’s genus Cheloniceras is adopted for these Aptian forms since

the genus Dowvilleiceras should be restricted to the Albian mammallatum

group. According to Kilan* the group of ‘ Douvilleiceras” royeri-

anum, @Orbigny sp. (= type of Cheloniceras), is closely connected

with the martini and albrechti-austriae groups (to which last the

form here described belongs), and not with the group of “ D.” bigoureti,

which “ leads to Acanthoplites.” The relations of these Aptian forms

with the Barremian Paraspiticeras have yet to be worked out. Sinzowt

would include the type of Paraspiticeras (P. percevali, Uhlig sp.)

with the Aptian Ch. meyendorffi, VOrbigny sp.,f but in the writer’s

opinion the two developments are distinct.

3. CHELONICERAS (ACANTHOPLITES ?) DELAGOENSE, Krenkel sp.

(Pl. XXVI, figs. 2 a—d.)

1910. Dowvilleiceras delagoense, Krenkel. ‘ Aptfossil. d. Delagoa

Bai,” N. Jb. f. Min., ete. (i), p. 147, pl’ xvii, figs. 6 and 7.

A small example (No. 5118), showing very good agreement with

Krenkel’s species, by its suture-line belongs to the same group of

forms as the specimen last described, but in loss of tubereulation, and

in the costation of the outer whorl, one-third of which already belongs

to the body-chamber, it has a superficial resemblance to the genus

Acanthoplites. The dimensions are as follows :

Diameter 4 : . 22 mm.

Height of the last whorl. 57 per cent. of the diameter.

Thickness _,, Ms 48 “ f 5

Umbilicus : : 40 7 zs si

At a diameter of 7°5 mm., at which the suture-line represented in

fig. 2c was taken, the whorl-section is depressed, and there are con-

strictions as in Cheloniceras royerianum, VOrbigny sp., and in the

specimen last described. The point of bifurcation of the ribs is

marked by a tubercle at this stage, as in Cheloniceras seminodosum,

Sinzow sp.,§ or in Acanthoplites Bigoureti (Seunes), Jacob,|| and the

* Loc. cit. (1913), p. 340.

+ Loc. cit. (1906), p. 163. ‘

t See also ‘ Pal. Univ.,’ 1911, pl. ccix.

§ “ Beschr. einiger Douvilleiceras-Arten a. d. Ob. Neocom. Russl.,” ‘ Verh,

Russ. Kais. Mineral. Ges.’ (2), vol. xliv (1906), p. 165, pl. i, fig. 3 only.

|| “Gisement de Clansayes,” ‘Bull. Soc. Géol. France’ (4), v, pl. xiii,

figs. 6 a, bd.

On Cretaceous Cephalopoda from Zululand. 317

intermediate ribs also may have a tubercle at this point, even when

single, giving the inner whorls a coronate appearance. After a

diameter of 10 mm., however, all tuberculation is lost, whereas

Krenkel’s type shows the last tubercle only at a diameter of 15 mm.

This loss of tuberculation is a feature found in certain varieties of

“ D.” tschernyschewi, Sinzow,* but much more typically shown in certain

Acanthoplites. A. bigoti (Seunes), Sinzow sp.,t e.g., has a very similar

outer whorl; and the suture-lines of this form, and of A. bigowreti,

Seunes sp., as figured by Jacob, are not very different from that of the

present specimen. Seunes’s type,§ however, and the form figured by

Pervinquiére,|| are less closely comparable. On the other hand, the

small example of Ch. seminodosum, the lateral and peripheral views of

which, as figured by Sinzow, greatly resemble the (somewhat less

depressed) South African form, instead of losing the original tuberele,

takes on a second one, that is to say, it becomes a more closely costate

type of Ch. cornuelianum.

The suture-line corresponds with that found in certain Acan-

thoplites, i.e. the bigoureti-bergeroni group, and with that of the

cornuelianum-meyendorfi group, referred to above; also with that of

Ch. hambrovii, Forbes sp.4/ Those of the typical Acanthoplites (aschil-

taensis group) ** and of the true Parahoplites (melchioris-group)t¥ are

different.

There is no close resemblance to any of the various forms of the

South American Aptian (e.g. A. roseanus, treffryanus, Karsten sp.tt

of which there is a large series in the British Museum, and which

(with forms like @Orbigny’s A. crassicostatus,$§ with A. peltoceroides,

* Loe. cit., e.g. pl. iii, fig, 4.

+ “ Unters. einiger Ami. a. d. Unt. Gault Mangyschlaks und d. Kaukasus,”

‘Verh. Russ. Kais. Min. Ges.,’ ser. 2, vol. xlv (1907), pl. iv, fig. 18.

t Loc. cit. (Clansayes), p. 412, fig. 5; p. 415, fig. 7.

§ Loc. cit. (1887), pl. xii, fig. 2, p. 568.

|| Loc. cit. (1907), pl. vii, figs. 36 a, bd.

"| See fig. 3, pl. vili, taken at a diameter of 10 mm., from an Isle of Wight

specimen in the writer’s collection.

** In Sinzow, loc. cit. (1907), pl. v, e.g. figs. land 3; also Anthula, lve. cit.

(1899) Splashes sro:

++ In Sinzow, Joe. cit. (1907), pl. ii, figs. 1-4; also Anthula, Joe. cit. (1899),

pl. viii, fig. 4c.

+t “Ub. d. Geogn. Verh. d. Westl. Columb., etc.,” ‘Amtl. Ber. 32. Vers.

Deutsch. Nat. F., etc.” Vienna, 1858, e.g. pl. ii, fig. 4; also Lea, “* Notice of Ool.

Form. i. Am.,” ‘Trans. Am. Phil. Soc.,’ Philad. (vii), n.s., 1840-1, pl. viii,

figs. 4, 5. ;

§§ Loe. cit. (1840), pl. lix, figs. 1-4.

318 Annals of the South African Museum.

Kilian non Pavlow sp.,* A. tobleri, Jacob sp.,*t and Sinzow’s various

Caucasian Acanthoplitest) show the peculiar ventral flattening of the

costation that is only just indicated in Krenkel’s species. The

Acanthoplites of the milletianus-jacobi group found in North

Germany (Schrammen, Collet, etc., Colls., British Museum) belong

to a different group again. On the other hand, there is a superficial

resemblance to certain Calycoceras of the Cenomanian, notably to

C. baylei, Pervinquiére, or to the form figured by Vilanova.§

Kilian || first named this species Acanthoceras (Parahoplites) abichi

(Anthula) var. africana, but Krenkel drew attention to the differences

between the African and the Caucasian forms. The suture-line

confirms this separation of Ch. delagoense from Acanthoplites abichi,

which latter belongs to the bergeroni-bigowreti group, and was, indeed,

united with the last species by Pervinquitre.4]

The genus Puaraspiticeras, which shows a change to costation,

following on tuberculation, does not have the slightly sigmoidal

ribbing which approaches the present example so much to the inner

whorls of the Acanthoplites bigoti figured by Sinzow (1907) on pl. iv,

fig. 18.

EXPLANATION OF PLATES XIX—XXVI.

PLATE XIX.

FIG.

1. Diaziceras tissotiaeforme, gen. and sp. nov. Upper Senonian, Umkwelane

Hill. Specimen No. 5478, p. 245. a, Side view of holotype. 6. Peri-

pheral view, not quite central. c. Sectional outline, at diameter =

50 mm. d. Inner whorls, at diameter = 3 mm., x 15. The keel

appears at a diameter of about 25 mm. e. Adult suture-line, x 2

external portion. (f. Internal portion of same. g-k. Development of

suture-line, at diameters of 1:5 mm. (g); 2mm. (f/); 8 mm. (7); 5°5 mm.

(7); 8mm. (#). All greatly magnified.

2. Parapuzosia sp. nov. ? ind. Upper (?) Senonian, Railway Cutting, Umfolozi.

Specimen No. 5513, p. 224. Reduced about +. After a photograph sent

* Pavlow’s form (‘Argiles de Speeton,’ 1892, p. 152, pl. xi. figs. 20 and 21),

according to specimens in the British Museum and information kindly given

by Mr. Lamplugh, does not belong to this family and formation.

+ Loc. cit. (‘ Engelberger Aa,’ 1906),"p. 11, pl. ii, figs. 4-6 (Parahoplites).

+t Loe. cit. (1907), pl. v.

§ ‘Mem. Geogn. d. Castellon,’ pl. ii, fig. 5 (as 4A. mantelli). The A. cor-

nuelianus (ibid., fig. 11) of this author is considered by Kilian (1918, p. 341)

to belong, probably, to Ch. albrechti-austriae, Hohenegger sp.

|| Loc. cit. (1902), p. 465.

Loc. cit. (1907), p. 195.

FIG.

iw)

On Cretaceous Cephalopoda from Zululand. 319

PLATE XX.

Parapuzosia, sp. nov ? ind. (Specimen figured Pl. XIX, fig. 2.) Reduced ?.

Photograph by Dr. A. L. du Toit. With restored outline-section (1a),

p. 224.

_ Pseudoschloenbachia umbulazi, Baily sp. Upper Senonian, Umkwelane Hill.

Specimen No. 5494, p. 240. (Genotype.)

. Pseudoschloenbachia umbulazi (Baily) var. acuta, nov. Same locality and

formation. Specimen No. 5450, p. 241. 3a. Peripheral view.

. Mortoniceras soutoni (Baily), Umtamvuna River, Natal. B.M., No. C19441.

Suture-line, after a drawing by the late G. C. Crick. (See p. 234.)

PLATE XXI.

. Mortoniceras woodsi. sp. nov. Upper Senonian, Umkwelane Hill. Specimen

N. 5451, p. 232. a. Side view. 4. Peripheral view, not quite central.

ec, Suture-line (restored), x 2. Ventral lobe on left, antisiphonal lobe

on right. d. Sectional outline.

. Placenticeras subkaffrarium, sp. nov. Same locality. Senonian. No. 5106,

p. 247. a. Side view. 4. Peripheral view, not quite central. ¢. Sectional

outline. d. Portions of suture-line, x 3, showing ventral lobe (arrow

on right) and umbilical tubercle (left).

PLATE XXII.

. Parapachydiscus sp. n. aff. colligatus, Binkhorst sp., x 4. Upper Senonian,

Umkwelane Hill. No. 5489, p. 226. Side and peripheral views.

. Nostoceras ? natalense,sp.nov. No. 2746. Upper Senonian, Umfolozi Valley,

East of Railway, p. 248. a. Side view. 6. 'Top view.

_ Nostoceras ? subangulatum, sp.nov. No. 2746A. Same locality and formation,

p. 250. a. Side view. 5. Septal surface at * in fig. 3a, x 2. D.= dorsal,

J”. = ventral side, 7.7. = impressed zone. ec. Suture-line, x 4 (restored),

at * in fig. 3a.

PLATE XXIII.

. Peroniceras cf. dravidicum, Kossmat sp. Lower Senonian. High ground on

north side of United Manuan Creek and Umsinene River, almost opposite

junction. No. 4950, p. 295. a. Side view. 6. Peripheral view. e. Sec-

tional outline. d. Suture-line, x 2.

_ Peroniceras cf. ezérnigi, Redtenbacher sp. (= Peroniceras a of Crick, 1907,

p. 226, text-fig. on p. 226). B.M., No. C18245. Lower Senonian, Zulu-

land. (See p. 296.) Internal (dorsal) portion of suture-line showing

considerable asymmetry.

_ Mortoniceras stangeri, Baily sp. Senonian, Umtamvuna River, Natal. (See

p- 297.) a. Sectional outline of No. C19459 (British Museum), x 2, at

diameter = 25 mm. 8%. Portion of suture-line of same specimen, from

lateral saddle (left) to antisiphonal line (right). U. = umbilical suture.

(At diameter = 125 mm.) c. Internal suture (at diameter = 60 mm.) of

Baily’s co-type, 11368A (Geol. Soe. Coll.).

320 Annals of the South African Museum.

FIG.

10.

Mortoniceras vanuxemi, Morton (Whitfield) sp. Upper Senonian, North-

west shore of False Bay. No. 4947, p. 308. @. Sectional outline. 6.

Suture-line, x 3. The dotted line on the right of the fig. 4a denotes

the position of the antisiphonal lobe.

. Diplomoceras ? indicum, Forbes sp. Upper Senonian, Umkwelane Hill.

No. 5465, p. 256.

PLATE XXIV.

. Parapachydiseus cf. wittekindi, Schliiter sp. Upper Senonian, Umfolozi

Valley, East of Railway. No. 3969, p. 229, x §.

Bostrychoceras ? sp. ind. Upper Senonian, Umkwelane Hill. No. 5478A,

p. 255. (Squeeze of impression in matrix of Diaziceras tissotiaeforme.)

Parapuzosia, sp. nov. ? ind. Senonian, Railway Cutting, Umfolozi. Speci-

men figured, Pl. XIX, fig. 2, Pl. XX, fig. 1, p. 224. Cast of dorsal impression

in outer whorl, showing ornamentation of missing inner whorls, x 9.

Baculites cf. aspero-anceps, Lasswitz. Upper Senonian, Umkwelane Hill.

No. 5480, p. 259. 4a. Sectional outline.

Baculites cf. brevicosta, Schliiter. Same locality and formation. No. 5461,

p. 260. 5a. Sectional outline.

Baculites capensis, Woods. Same locality and formation. Suture-line of

specimen No, 5486, p. 257, x 6.

. Baculites capensis, Woods. Upper Senonian, Umtamvuna River, Pondoland.

B.M., C19420. After a drawing by the late G. C. Crick (see p. 258), x 2.

. Kossmaticeras (Madrasites) bhavani, Stoliczka sp. Senonian, south side of

Manuan Creek Valley. No. 4909, p. 299.

Subschloenbachia bispinosa, nov. Upper Albian, Middle Branch, Manuan

Creek. No. 4998, p. 285. (Sectional outline, restored.)

Subschloenbachia prerostrata, nov. Upper Albian, Middle Branch, Manuan

Creek. No. 4970, p. 284. (Sectional outline.) The arrows indicate the

spiral grooves of the cast, not visible on the shell.

PLATE XXV.

. Pseudophacoceras manuanense, nov. Albian, Manuan Creek (p. 281). a. Side

view of holotype, No. 2725, x 3. 4. Suture-line of same, at diameter

= 160mm. ec. Side view of specimen No. 2726. d. Sectional outline of

same.

. Dipoloceras ecristatum, Deluc sp. Albian, Manuan Creek. No. 2728, p. 277.

Mouth-border of very large example, taken from a cast of impression in

matrix.

Dipoloceras quadratum, nov. Albian, Middle Branch, Manuan Creek. No.

4955, p. 278. a,b. Side and peripheral views. c. Sectional outline.

_ Subschloenbachia cf. trinodosa, Bose sp. Uppermost Albian, south side of

Manuan Creek Valley. No. 4972, p. 285. Sectional outline.

On Cretaceous Cephalopoda from Zululand. 32]

PLATE XXVI.

FIG,

1. Cheloniceras gottschei, Kilian sp. Aptian, Powell’s Camp, Upper Catembe.

Specimen No. 5117, p. 312. a. Side view of body-chamber portion. 6.

Peripheral view of same. c. Restored sectional outline of inner whorl.

d. Suture-line, restored from last two, at beginning of portion figured in

la (at *), x &.

2. Cheloniceras (Acanthoplites ?) delagoense, Krenkel sp. Same formation and

locality. No. 5118, p. 316. a,b. Side and peripheral views. c. Suture-

line, at diameter = 7°5 mm. (where whorl is coronatiform and con-

stricted), x 6. d. Same at15 mm. /, = antisiphonal line, x about 6.

3. Cheloniceras hambrovii, Forbes sp. Aptian, Atherfield Clay, Isle of Wight.

(Writer’s Coll.) Suture-line, at diameter = 10 mm., x 6. (See p. 317.)

4, Aconeceras nisoides, Sarasin sp. Aptian, Powell’s Camp. No. 5119, p. 311.

Side view (4) and sectional outline (44).

5. Dipoloceras sp. nov.? Albian, Middle Branch, Manuan Creek. No. 4908,

p- 280. a. Side view, with restored outline of complete shell. 4. Sec-

tional outline, showing peculiar keel.

6. Dipoloceras cristatum, Deluc sp. Albian, Manuan Creek. Portion of suture-

line, x 2, of specimen 2728, p. 277. /. = antisiphonal line, with keel of

previous whorl (impossible to remove) concealing internal lobe.

7. Anisoceras sp.ind. Albian? Low Ridge, about three miles east of foot of

Lebombo Mountains, north of M’Kusi River, due east of Ubombo. No.

4982, p. 288. Suture-line, x 8.

Ann. 8. Afr. Mus. Vol. XII. Plate XIX.

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( 323 )

17. — The fauna and Stratigraphy of the Stormberg Series. — By

S. H. Haucuton, B.A., D.Sc., F.G.S., Hon. Curator of the

Palaeontological Collections, formerly Assistant Director, S. A.

Museum, (With 55 Text-figures.)

INTRO CT LONe

In the following pages an attempt is made to bring together facts,

previously scattered through a number of publications, bearing upon

the stratigraphy and palaeontology of the Stormberg Series and_ to

incorporate among them the results of three collecting expeditions

made by the author mainly in the districts of Herschel and Wode-

house. The whole of the fossil collection of the South African

Museum has been reviewed — the animals by the author, the plants

by Dr. A. L. du Toit. The work of the latter is not yet ready for

publication; but Dr. du Toit has kindly forwarded a provisional list

of his identifications, which is incorporated below. To him I am

also indebted for constant critical advice and, as will be seen, I have

drawn largely upon his published descriptions of the stratigraphy

of the beds.

Mr. Macgregor, of the Rhodesian Geological Survey, has been good

enough to supply me with notes and specimens from Southern

Rhodesia. Rock-sections have been cut in the Geological Department

of the University of Cape Town, where Professor A. Young has

critically examined the petrological work, besides discussing the

many interesting points of palaeogeography which arose. Thanks

are due to the Royal Society of South Africa for a grant which

resulted in the discovery of the skeleton of Massospondylus harriesi ;

and, among others, [ am indebted for help to Dr. L. Pérmguey for

examination of the insect-remains and to Mr. K. H. Barnard for

inspection of the crustacea.

FAUNA AND FLORA.

The following list shows the forms found in the Stormberg Series.

PLANTS.

(List supplied by Dr. A. L. du Toit.)

Neocalamites (Schizoneura) Carrere, (Zeill.).

Neocalamites sp. Sew.

324

Annals of the South African Museum.

Danaeopsis hughes, (Feist.).

Thinnfeldia odontopteroides, (Morr.).

Thinnfeldia lancifolia, (Morr.).

Thinnfeldia Feistmanteli, Johnston.

Thinnfeldia aquilina, Shirley.

Thinnfeldia trilobita, Johnston.

Taeniopteris Carruthersi, (Ten.— Woods).

Taeniopteris Tenison— Woodsi, Eth. jun.

Taeniopteris crassinervis, (Feist.).

Chiropteris Zeilleri, Sew.

Chiropteris cuneata, (Carr.).

Chiropteris copiapensis, Stein. et Solms.

Gingkoites (Ginkgo) digitata, (Brongn.).

Ginkgoites antarctica (Sap.).

Ginkgoites, nov. sp.

Baiera Schenki, Feist.

Baiera stormbergensis, Sew.

Baiera moltenensis, Sew.

Sagenopteris longicaulis, du Toit.

Cladophlebis (Todites) Roesserti, (Presl.).

Cladophlebis nebbensis, (Brongn.).

Callipteridium stormbergense, Sew.

Stenopteris elongata, (Carr.).

Stenopteris rigida, Dun.

Stormbergia Gardneri, Sew.

Sphenopteris alata, Brongn.

Sphenopteris lobifolia, Morr.

Pterophyllum natalense, du Toit.

Pterophyllum cf. Footeanum, Feist.

Phoenicopsis (Desmiophyllum) elongata, (Morr.).

Pachypteris acuta, du Toit.

Pachypteris lanceolata, Brongn.

Zamites sp.

Maraitiopsis munsteri, (Goepp.).

Glossopteris browniana, Brongn.

Glossopterts conspicua, Feist.

Stachopitys sp.

Strobilites sp.

Rhexoxylon sp.

ANIMALS.

CRUSTACEA.

Cyzicus (Euestheria) draperi (Jones & Woodw.).

Cyzicus sp. Leriche.

Lepidurus stormbergensis, sp. nOv.

Unnamed Ostracod.

Fauna and Stratigraphy of the Stormberg Series. 325

INSECTA.

Coleopteron. Gen. et sp. indet.

Phthartus africanus, sp. nov.

Striatotegmen africanum, gen. et sp. nov.

Unnamed Blattid.

Archaegryllodes stormbergensis, gen. et sp. nov.

Eggs of Orthopteron?

PISCKS.

Semionotus capensis, Sm.—Woodw.

Helichthys (2) sp.

REPTILIA.

Pachygenelus monus, Watson.

Tritheledon riconoi, Br.

Lycorhinus angustidens, gen. et sp. nov.

Sphenosuchus acutus, Htn.

Notochampsa istedana, Br.

Pedeticosaurus leviseurt, v. Hoep.

Erythrochampsa longipes, (Br.).

Thecodontosaurus browni, (Seeley).

Thecodontosaurus skirtopodus, (Seeley).

Thecodontosaurus minor, Htn.

Gyposaurus capensis, Br.

Aristosaurus erectus, v. Hoep.

Massospondylus carinatus, Owen.

Massospondylus harriesi, Br.

Massospondylus schwarzi, sp. nov.

Aetonyx palustris, Br.

Dromicosaurus gracilis, v. Hoep.

Plateosaurus stormbergensis, Br.

Plateosaurus cullingworthi, sp. nov.

Gryponyx africanus, Br.

Gryponyx transvaalensis, Br.

Gryponyx taylori, sp. nov. .

Euskelesaurus browni, Huxley.

Euskelesaurus capensis, (Lyd.).

Euskelesaurus africanus, sp. nov.

Gigantoscelus molengraffi, v. Hoep.

Eucnemesaurus fortis, v. Hoep.

Melanorosaurus readi, sp. nov.

Geranosaurus atavus, Br.

( 326 )

jeyeU ani Be IIe

DESCRIPTION OF SPECIKS.

CRUSTACEA.

ENTOMOSTRACA.

Famity CYZICIDAE.

CYZICUS, Audouin.

Cyzicus (KUESTHERIA) DRAPERI (Jones & Woodw.)

1894. Estheria Drapert and E. stowiana, Jones & Woodward,

Geol. Mag., N.S. Dec. IV, vol. I, p. 289. Pl. IX figs. 1a—e, 2.

The original description runs as follows: — ‘Size: Length of valve,

16 mm. Length of hinge-line, 14 mm. Height 10!), mm. Valves

suboblong, straight above, slightly curved below, rounded at the

ends; anterior margin higher and less convex than the posterior.

The imterspaces on the surface are ornamented with coarse shallow

pits, making an obscure reticulation. The umbo is just in front of

the middle of the hinge-line.”

I have examined a number of specimens from the shale-band at

Siberia, Wodehouse, C. P. in the Cave Sandstone which belong to

this species. None of them attain the dimensions recorded by Jones &

Woodward; the largest valve seen by me is 12 mm. long, 8 mm,

high and has a hinge-line of 8'/, mm. The corresponding dimen-

sions on a number of the specimens are 7!/, mm., 5 mm. and 5 mm.,

while some have a length of not more than 2!/,; mm.

Several of the specimens show the details of the external markings.

The shape is somewhat variable, the smaller specimens approx-

imating more to that figured by Jones & Woodward as E£. stowiana.

All agree, however, in the possession of the concentric ridges and,

as far as can be seen, in the nature of the intercostal ornamentation.

Fauna and Stratigraphy of the Stormberg Series. 327

In a valve having a height of 4 mm. there are 13-14 regularly

spaced well-marked fine costae with broad interspaces. As the valves

erow older and larger the costae increase rapidly in numbers and

are crowded together near the ventral border in the larger specimens

as in C. mangaliensis.

The intercostal ornamentation consists of shallow pits when viewed

from without and pustules when viewed from within the valve.

These are arranged in irregular rows and are set closely together so

that there are 7 or 8 rows in each interspace. The pits are rounded

rather than polygonal in outline. The older portion of the valve

(that around the umbo) is smooth save for the concentric ridges.

The pits are more prominent near the ventral, anterior and posterior

borders of the valves. The abscence of pits near the umbo on the

larger specimens is presumably due to wear or some other cause,

as the small specimens occasionally show ornamentation over the

whole valve.

Some of the valves show a considerably greater degree of ribbing

than the more typical specimens. This variation in sculpture is

paralleled in C. maurchisonae in different valves of which Rupert

Jones recorded variable ribbing.

The similarity of ornamentation between the valves of all sizes

found at Siberia helps to prove the assumption here made that

C. stowiana and C. draperi are but growth-stages of the one species.

This possibility was mentioned by Jones & Woodward who had not,

however, material for observing the intercostal ornamentation on the

smaller forms nor intermediate valves showing the gradual changes

in size. The variation in shape between specimens of the same size

does not seem sufficient warrant for separating the specimens into

several species. The species figured by Rupert Jones in his ‘Monograph

of Fossil Estheriae” are variable in shape-characters; and the varia-

tions seen in C. draperi are no greater than those figured for his

Estheria ovata or E. minuta.

In general form C. draperi seems to approach fairly closely to

C. mangaliensis from the Rhaetic of India and from the Argentine.

Type. In the British Museum.

Locality. Harrismith, O. F.S.

Horizon. Shale-band in the Cave Sandstone.

CyZICUS sp.

1920. Leriche. Estheria sp. Rev. Zool. Afric. VIII p.78. Pl. II, fig. 4.

Founded on specimens from the Lubilash Beds at Kitari, canon

of the Inzia, Belgian Congo.

328 Annals of the South African Museum.

“The valves are sub-oval; the beak is anterior and not prominent.

The dorsal border is almost straight. The anterior border is rounded

and passes insensibly into the ventral border, with which it forms

a very regular curve. The ventral border is regularly convex; it

joins the posterior border, which is oblique to the dorsal border, by

a very obtuse angle. The valves carry about fourteen concentric

ribs. The state of preservation does not permit of the presence of

an intercostal ornamentation being stated.”

Leriche states that these fossils belong to a group of “‘E'stheria”

characterised by a regular concentric ornamentation formed of pro-

minent spaced ribs. This group has been called Euestheria by Deperet

and Mazeran. The Congo form approaches most closely EH. manga-

lensis but is distinguished by the shghtly more elongate shape and

the sharper demarcation of the posterior and ventral borders. E. greyi

has a similar ornamentation but has a more oval and more inequi-

lateral form.

Specific identification is not possible without a knowledge of the

intercostal ornamentation.

Fam. APODIDAE.

LEPIDURUS, Leach.

LEPIDURUS STORMBERGENSIS sp. nov.

This form is represented in the South African Museum collection

by a number of impressions on shale. Some represent isolated

carapaces of immature animals; others more or less complete mature

individuals. _No specimen is perfect, and therefore several have been

examined in the preparation of the following description. In some

cases Impressions of the appendages are fairly well-preserved, but

detailed description of these is postponed.

The carapace is almost circular, slightly longer than broad, and

deeply emarginate behind, the emarginate portion having an untuvothed

border. There is a strong median keel extending from the hinder

border to a short distance behind the eyes.

Measurements of four carapaces of varying size are as follows:

Median length Maximum length Maximum breadth

9 mm. 10°3 mm. 10 mm.

9 mm. 44. mm. 10 mm.

44 oinma: 13 mm. 12 mm.

49 mm. 17-5 mim.

Fauna and Stratigraphy of the Stormberg Series. 329

The smaller carapaces are isolated; the larger are attached in each

case to the body.

The first antenna is short, single, possibly divided into two joints.

As seen in specimen 5752 and possibly in 5751 it is similar in

appearance to that of the recent ZL. viridis. The antenna is best

seen in specimen 5758, :

The second antenna cannot be distinguished.

The mandible is apparently an elongate plate (breadth greater than

length) with a denticulate biting edge having 5 or 6 teeth. An

impression is seen in no. 5758 In no, 5752 the maxilla is seen

in juxtaposition with its neighbour of the other side.

The segments under the carapace are all provided with setose

Phyllopod appendages.

In one specimen there are about 17 segments protruding beyond

the carapace; of these at least 7 are longer than the others and may

be regarded as true abdominal segments, although none of the seg-

ments are seen to possess feet. Each segment bears an encircling

row of sharp spines which diminish in number posteriorly. The

spines arise from near the middle of each segment and project well

back behind the posterior border. Excluding the telson the body is

about half the length of the carapace. (In another specimen the

body is shorter, and the number of abdominal segments apparently

six. This is possibly a female).

The telson is moderately short, spatulate, with a fairly well

rounded posterior border and a pronounced longitudinal median keel.

It carries no spies, and the edge is entire. In specimen 5763,

however, the telson, instead of being spatulate, is somewhat pointed

at the end, like a spear-head, and the lateral edges seem to be

either serrate or provided with short spines. This may represent a

variety; but the specimen is otherwise too incomplete for lengthy

description.

The caudal filaments are fairly broad at the base, taper fairly

rapidly and are covered with long fine hairs.

Specimen no. 5752 shows an oval body 2°5 mm. long situated in

the neighbourhood of the 15th or 16th legs under the carapace,

behind the mid-line of the carapace and to the side. The body is

empty, but was probably an ovisac.

The form has been placed in the genus Lepidurus on account of

the characteristic telson which distinguishes it at once from Apus.

As far as can be discovered Lepidurus has not hitherto been des-

cribed from Triassic deposits, although an allied form <Apudites (or

Apus) antiquus has been named by Schimper from the Lower Trias

330 Annals of the South African Museum.

(Voltzia Sandstone) of the Vosges. Unfortunately it has not been

possible to consult the description of this form, which has been

placed in the genus Apus by Ph. C. Bill (14914). The genus Lepi-

durus is not a member of the modern South African fauna although

several species of Apus occur in the South African vleis, which

habitually suffer periodic dessication.

Co-types. S. Af. Mus. Coll. Cat. Nos. 5754-5754, 5759-5761.

Locality. Siberia, Wodehouse, C. P.

Horizon. Shale-band near base of Cave Sandstone.

Cxiass INSECTA.

OrpDER COLEOPTERA.

Text fig. 1.

A piece of shale in the collection of the South African Museum

contains the impression of what seems part of a coleopterous elytron.

The impression is that of the posterior half of a right elytron, the

anterior portion being overlain by another impression which, accor-

ding to Dr. L. Péringuey, cannot be coleopterous. The impression

Fig. 1. Coleopterous elytron.

shows 8 main sub-parallel veins and a fairly broad fold. The sixth

and seventh of the veins coalesce posteriorly. In addition there is

a series of pustules (left in the impression as punctures) forming a

fine line between the two outer veins, and another series Westen

the next pair of veins.

The fragment (Cat. No, 5635) is too incomplete to justify naming.

Triassic coleoptera have been described from Australia (families

Hydrophilidae’?, Elateridae?, Tenebrionidae? and Malacodermidae)

and from the Keuper and Rhaetic of Europe. No Permian forms

have hitherto been described.

Locality. Siberia, Wodehouse, C. P.

Horizon, Shale-band near base of Cave Sandstone.

Fauna and Stratigraphy of the Stormberg Series. 334

OrpER PLECTOPTERA.

Fam. EPHEMERIDAE.

PHTHARTUS AFRICANUS, sp. nov.

Text fig. 2:

This is represented by several imperfect larvae of which the

specimen figured (Cat. No. 5732) is nearly complete, lacking the

head and lateral appendages. The cercal setae are faintly indi-

cated — and have been drawn in on the evidence of a second spec-

imen (No. 5733) consisting of the hinder portion of an abdomen in

which the long setae are well-marked.

As preserved, the body without the cerci is 145 mm. long and

3D mm. broad in the middle.

Fig. 2. Phthartus africanus, Htn.

The earliest Ephemerid larvae belong to the genus Phihartus

Handl. from the Permian of Russia. The present form seems fairly

similar to that genus, but the cerci are shorter and stouter. It may

be grouped provisionally as Phthartus africanus sp. nov.

Type. Impression on shale — 8. Afr. Mus. Cat. No. 5732.

Locality. Road-cutting, Siberia, Wodehouse, C. P.

Horizon. Shale-band near base of Cave Sandstone.

OrpER ORTHOPTERA.

Fam. MESOBLATTINIDAE, Handl.

STRIATOTEGMEN AFRICANUM, gen. et sp. nov,

Text fig. 3.

Tillyard’s emendation of the characters of this family runs as

follows: — “Subcosta much reduced. Radial area extensive and

strongly developed, sending numerous branches towards the anterior

3o2 Annals of the South African Museum.

border, and reaching nearly to the apex. Media free, dividing into

a variable large number of branches directed towards the apical

border. Cubitus also free, branching variably, the branches directed

towards the posterior border. Anal field large, well-defined, usually

somewhat cultriform, with more or less curved veins running chiefly

towards the posterior border. Intercalated veins and_ reticulation

or cross-venation may or may not be present. Mostly small to medium-

sized forms.” (Proc. Linn. Soc. N.S. Wales, XLIV, 2, 1919, p. 366).

The fossil S. A. M. Cat. No. 5634 consists of a piece of shale and

a partial counter-piece showing a nearly complete insect. One

tegmen is folded back on the body; the other is spread out at right

angles to the body and is preserved without the apex as an impres-

sion on slab 5634a.

The width of the tegmen is just over one-third of the probable length

(which is about 9 mm.). The humeral area is narrow and distinctly

less in leneth, from base to apex, than the anal area, and is bounded

distally by a slightly double-curved sub-costa which is apparently

unbranched. The humeral area is sharply pointed at the apex.

The radius is strongly double-curved. It gives off eight primary

branches to the anterior border, excluding the forked distal end; of

these, the first six are simple veins, the seventh and eighth forked,

The median has four branches which apparently extend to the

border below the apex. The cubitus has at least six branches.

In the area covered by R, M, and Cu strong raised ridges sepa-

rate the veins. These are preserved as strong channels; but that

they are not true veins is shown by the fact that they do not unite

with one another basally, whereas the true veins do. There are

faint indications of crossvenation on the proximal portions of the

area covered by M and Cu.

The anal area has the vena dividens strongly marked. The first

anal vein rises from the vena dividens but ends at the wing border

just below the apex of the area. The third vein is forked at some

distance from the border. The fourth vein is waved, as is the fifth —

the latter being forked near its distal end. The sixth vein is also

waved. In the anal area, there are indications of faint ridges

between the veins. There is also a certain amount of irregular

cross-venation.

Comparison of this wing with the forms from the Ipswich Beds

of Australia gives the following results: — From Triassoblattina it

differs in the narrower humeral area, the reduced nature of the

subcosta, the stronger double-curving of the radius, and in the fact

that none of the anal veins end distally on the vena dividens. There

Fauna and Stratigraphy of the Stormberg Series. 333

are differences, too, from the genotype J. typica in the number of

branches of R, M, and Cu.

From Samaroblatta it differs in the shorter and narrower humeral

area, in that the distal portion of R probably reached the apex.

In other points it approaches fairly closely to this genus, especially

in the possession of intercalated ridges and a cross-venation. The

diflerence in branching of R, M, and Cu is not of great moment;

Tillyard has pointed out that not only different individuals of the

same species of Cockroach, but also even the right and left tegmina

of the same individual, show considerable differences in this respect.

Fig. 3. Striatotegmen africanum, Htn.

Venation of tegmen.

From Austroblatta, with which it agrees in size, it differs in the

smaller humeral area, the pronounced bending of R, and the pre-

sence of intercalated veins.

In general characters it agrees closely with Mesoblattula from the

Lias of Mecklenburg. It shows the same reduced subcosta; the hu-

meral area shorter than the anal, the same strongly double-curved

radius, and the same general features of M and Cu As in Meso-

blattula there are intercalated ridges, and evidence of occasional

cross-venation (cf. M. geinitziana). Unfortunately the anal area is

unknown in the genus Mesoblattula. Our form differs from this genus

in that the anterior branch of the medial is apparently not forked;

in this it differs also from Mesoblattopsis, Liadoblattina, Rhipidoblat-

tina and Caloblattina, and apparently agrees with Mesoblattina. It

agrees also with Mesoblattina in the reduced humeral area and the

simple subcosta; but the area covered by the radius is larger in the

South African form, the radius is more strongly curved, and there

are fewer primary branches. The tegmen, too, is somewhat broader.

Two genera of this family have been described from the ‘‘Rhaetic”

of Tonkin — Rhaetcblattina Handl. and Hongaya Handl. Both of

these are larger than our form. In each the humeral area is short,

304. Annals of the South African Museum.

the subcosta reduced to a single vein, the radius strongly double-

bent so that it touches at its Jowest point the middle line of the

tegmen. Hongaya agrees with the South African form in that the

upper branch of the median is not forked, in the general nature of

the cubitus and median, in the fact that the first anal vein rises

from the vena subdividens and ends distally on the wing border, and

in the presence of intercalated ridges; but it differs in the number

and nature of the radial branches and in the smaller size of the

anal area. Rhaetoblattina has a larger anal area, comparable with

that of our form, a small humeral area, and intercalated veins: but

the median and cubitus are not so similar to our form as those of

Hongaya. In neither of the Tonkin genera are cross-veins described ;

and Handlirsch was unable to discern them.

As was to be expected, this form shows close relationships with

the Triassic, Rhaetic and Liassic forms of the Mesoblattinidae, and

is more advanced than the Carboniferous and Permian members of

the family. It does not seem to fall readily into any one of the

described genera and for that reason the new generic name Sfriato-

tegmen is proposed for it, founded upon the following generic cha-

racters: —

Striatotegmen gen. nov. Tegmen small (under 40 mm. in length).

Width just over one-third of the length. Humeral area narrow,

and distinctly shorter than the anal area. Sub-costa slightly double-

curved, unbranched. Radius strongly double-curved, almost reaching

middle line of wing, with eight primary branches, of which the

anterior six are simple. Upper branch of median unforked; upper

branch of cubitus unforked. Vena dividens strong. First anal vein

arising from vena dividens, but not ending distally on it. Intercal-

ated ridges present, especially strong in radial, medial, and cubital

areas. Cross-veins faintly indicated.

For this form the specific name Striatotegmen africanum sp. nov.

is proposed,

Type. Insect on shale, and counter-impression. S. Af. Mus. Cat.

No. 5634,

Locality. Road-cutting, Siberia, Wodehouse, C. P.

Horizon. Shale-band near base of Cave Sandstone.

Gen. et sp. indet.

Text fig. 4.

A fragment from Siberia (S. Af. Mus. Cat. No. 2340) shows por-

tions of two Blattoid tegmina. Parts of the radius, median, and

Fauna and Stratigraphy of the Stormberg Series. 335

cubitus of each tegmen are seen, but nothing of the subcosta nor

of the anal area. Both ends of each tegmen are missing. Inter-

calated ridges are seen between the branches of the cubitus and

median, but none in the area covered by the portion of the radius.

Nothing of the subcosta is seen, and but little of the radius. The

distal branches of the radius are forked.

Fig. 4. Blattoidea incert sed.

Most of the distal portion of the median is preserved, The appea-

rances of the two tegmina are slightly different. Each shows the

primary branch forked near the extreme end, and, in addition, three

other branches which arise by dichotomous division of a branch

from the main stem. Intercalated ridges occur on one tegmen; on

this side there appears to be a second primary unforked branch of

336 Annals of the South African Museum.

the median which is connected to the first fork of the first primary

branch by a diagonal cross-vein. :

The cubitus occupies a larger area than the median. It is dichot-

omously forked, and intercalated ridges are well-marked.

There is no sign of a strong vena dividens, nor any indication of

the anal area.

Locality. Road-cutting, Siberia, Wodehouse, C. P.

Horizon. Shale-band near base of Cave Sandstone.

Fam. GRYLLIDAE.

ARCHAEGRYLLODES STORMBERGENSIS, gen. et sp. nov.

Text fig. 5.

This is founded on portion of the tegmen of a male Gryllid, about

11 mm. long as preserved and 5 mm. broad.

Fig. 5. <Archaegryllodes stormbergensis, Htn. X 12.

The general arrangement of the main veins is similar to that in

the recent form Brachythripus and is distinct from the hitherto

earliest-described Gryllid — Protogrylius from the Upper Lias of

Mecklenburg.

The subcosta is long and has a number of curved oblique branches

in its proximal half. The radius hes close behind the subcosta.

Fauna and Stratigraphy of the Stormberg Series. 337

The median is weak and joins the radius about half way along the

latter forming a space with an acute distal angle. Distal to this <

branch of the median again reaches up to the radius. Cross-veins

connect the proximal portions of the radius and median.

There are indications of a drum between the branches of the

cubitus, but the veining is not distinct. There are cross-veins between

the second branch of the cubital and the first anal.

This is the earliest Gryllid described; and that fact, coupled with

its lack of similarity with Protogryllus seem to justify the erection

of a new genus which can be called Archuegryllodes n. gen., the

species represented by the specimen under discussion being termed

A, stormbergensis.

Type. Partial tegmen on shale. 8. Af. Mus. Cat. No. 2344.

Locality. Road-cutting, Siberia, Wodehouse, C. P.

Horizon. Shale-band, near base of Cave Sandstone.

ORTHOPTEROUS EGGS.

Text fig. 6.

On the same slab of shale as that which contains the type of

Archaegryllodes stormbergensis (No. 2341) are from ten to twelve

Fig. 6. Eggs of Orthopteron? x 8.

elongate oval bodies, which seem to be eggs of an Orthopteron.

Each is about 1 mm. long by 0:4 or 0-5 mm. broad., and the specimens

lie together in one mass,

There is a rough approximation to the form of a double row of

eges placed transversely, although one or two have been displaced

from their original position; but the general appearance seems to

indicate that the eggs were originally in an ootheca similar to that

of some Orthopterous insects, but that the cover has disappeared.

338 Annals of the South African Museum.

PISCES.

TELEOSTOMI.

Gren. SEMIONOTUS, Ag.

SEMIONOTUS CAPENSIS, Sm.—Woodw.

1888. Smith—Woodward. Quart. Journ. Geol. Soc. XLIV, p. 438.

1901. Schellwien. Schrift. Physik.-oekonom. Gesell. Konigsberg.

1909. Broom. Ann. S. Afric. Museum VII, part 3, p. 262.

Since Broom’s description, the South African Museum has received

no further examples of this well-known fish, and it will suffice to

quote the main points of Broom’s analysis of the form.

“The majority of specimens measure from 160 to 210 mm. in length.

In the example which is 210 mm. long, the body is 42 mm. in

depth at the deepest part, and the head measures 48 mm. to the

back of the operculum.

Schellwien has recently described a number of specimens and has

shown the more important features of the skull structure. The

specimens I have examined confirm most of his observations, but in

one or two points I am inclined to differ from him.

Almost every detail of the skull is now known except the basi-

cranial region. The frontals are large, and extend from the nasal

region to behind the plane passing through the back of the orbit.

The back part of the bone is about twice as wide as the middle

portion. Behind it is a large oblong parietal. Below the parietal is

a slightly narrower squamosal. My specimens do not satisfactorily

show the supratemporal region, but Schellwien finds a narrow

supratemporal and a post-temporal. ;

The opercular bones are very like those of Lepidotus. The oper-

culum differs in being relatively considerably wider in its lower

half. Inferiorly it joins the subopercular in a manner very similar

to that in the better known genus. The subopercular in Semionotus

is only about one-third the size of the operculum instead of half as

large as in Lepidotus, while the interopercular is less than half the

size of that im Lepidotus. In front of these three opercular bones

is a narrow curved preopercular, along which there runs a mucous

canal. ;

In Schellwien’s diagrammatic restoration the postorbital seems to

Fauna and Stratigraphy of the Stormberg Series. 339

me to be rather too small, while the interopercular is much too

Jarge. Above the anterior end of the long preopercular is an elon-

gated suborbital smaller in size than the postorbital. The portion

of the figure dealing with this region is, im my opinion, erroneous.

The lower jaw has an elongated triangular dentary and a power-

ful angular.

The palato-pterygo-quadrate arch is fully ossified, but the exact

limits of the different elements cannot be made out with certainty.

There is a long narrow bone below the quadrate stretching from

the articular region to the lower end of the hyomandibular. This

would seem to be the symplectic. The hyomandibular is a power-

ful bone and fairly similar to that of the ordinary Teleosteans. In

addition to supporting the opercular bones and the quadrate arch,

it supports the hyoid arch. There is a large quadrangular epihyal

and an elongate triangular ceratohyal. The interhyal has probably

been cartilaginous, as has also probably been the hypohyal and the

urohval. Under the subopercular are six branchiostegals.

The clavicular arch consists of the clavicle, supraclavicle, post-

clavicle and post-temporal, but there seems to be no trace of an

infraclavicle. A mucous canal crosses the supraclavicle obliquely as

in the Palaeoniscias. There is a small ossification which possibly

may be the coracoid as is thought by Schellwien,

The pectoral fin consists of 14 rays with 5 or 6 fulcra in front.

The rays are much flattened distally, but apparently not branched.

The pelvic fin consists of 7 rays which are branched distally.

The fulera are powerful.

The dorsal fin begins exactly in the middle of the back of the

fish, and consists of 13 rays, of which the last 3 are small. All the

rays are branched distally and articulated. In front are a row of

very powerful fulcra, 9 in number. The anal fin consists of 9 rays

with 9 powerful fulcra in front.

The caudal fin consists of 16 rays, all of which are branched and

articulated. Below and in front of the first ray are 14 fulera, and

14 fulera also lie above the tail, gradually passing into the dorsal

scales in front. The rays of the dorsal, anal, and caudal fins are

double. Though the tail is in a sense brevi-heterocercal the upper

portion is really continued as a long, slender process bearing small

rhombic scales about 20 mm. beyond the end of the middle of

the tail.”

Type. In British Museum.

Locality. Ficksburg, O. F. S.

Horizon. Cave Sandstone.

340 Annals of the South African Museum.

HELICHTHYS (?) sp.

A piece of shale from the deposit at Siberia, C. P., shows portion

of the body of a small fish including the proximal portions of the

anal and pelvic fins.

The body must have been fairly long and shallow. The pelvic

fin is not entire, but the preserved portion consists of 13 rays,

spread out into something of a fan-shaped body. They show no

evidence of bifurcation. The anal fin is preserved in its proximal

half; it is composed of 35 or 36 rays, of which the first seven at

least increase rapidly in size from the first. There is no evidence

of the presence of fulcra. The dorsal fin is not preserved.

The flank-scales are small, rhomboidal, without external ornament.

There seems to be evidence of a single longitudinal groove on the

inner surface.

It is possible that the form is allied to that described by Broom

as Dictyopyge formosa. It is doubtful, however, whether that species

can be ranked in the genus Dictyopyge as defined by Egerton. Ex-

cept in the pectoral fin no fulcra are seen; while there is no evi-

dence of bifurcation of the fin rays in the specimens of formosa

which we possess. Rather does the form seem to be generically

identical with Helichthys, agreeing in the comparative scarcity of

fulcral bones, in the simple unbifurcated rays, and in general cha-

racters. Fulcra are not well-marked, but are present.

It is considered, therefore, that the type specimen of Dictyopyge

formosa (S. A. Mus. Cat. No. 2761) must be known as Helichthys

formosa.

Type. Incomplete small fish. S. A. Mus. Cat. No. 5130.

Locality. Siberia, Wodehouse, C. P.

Horizon. Shale-band near base of Cave Sandstone.

Crass REPTILIA.

SuPER-ORDER ANOMODONTIA, Owen.

OrpER THERIODONTIA, Owen.

PACHYGENELUS MoNUS, Watson.

1913. Watson. Geol. Mag. N. S. Dec. V. Vol. X., p. 145, figs 4, 2.

This form is known only from the anterior portion of a small

dentary. There are only two incisors, of which the first is much

bigger than the second. The canine is large, of oval section. There

is a long diastema between the canine and the first molar. ‘The

Fauna and Stratigraphy of the Stormberg Series. 341

(molar) teeth are small, single-rooted, and narrow from side to side;

the root is deep and closely fits its alveolus.” ‘The crown is of an

irregular oval shape, widest in front, where it is about three-quarters

of its length. There are four cusps arranged longitudinally and for-

ming the outer side of the tooth;.... the first was much the largest,

and they gradually decline in size and height to the fourth. On the

inner side is a strong cingulum. This shows a very faint crimping”.

Type. Imperfect dentary.

Locality. Witkop, Albert, C. P.

Horizon. Red Beds.

TRITHELEDON RICONOI, Broom.

Text fig. 7.

14912. Broom. Ann. S. Afric. Museum VII, 5, p. 334, Pl. XXII,

figs, 30-36.

The type of this form is a ‘portion of a left maxilla with the

roots of 7 teeth and two imperfect immature teeth, portion of the

left jugal, and a fragment of the left palatine” from Paballong, Mount

Fletcher District, Griqualand East, which is in the collection of the

South African Museum.

In 1914, the discoverer of this fragment, the late Dr. M. Ricono,

forwarded to the Museum a small parcel containing a portion of a

skull and eleven dissociated vertebrae from Paballong, all of which

undoubtedly belong to the same animal as the fragment described

by Dr. Broom. Dr. du Toit, who has since visited the locality,

informs me that the fossil came from the Red Beds. It was found

within a quarter of a mile of the spot from which the type of

Sphenosuchus acutus came, and from an horizon within 40 or 12 feet of it.

This new skull fragment consists of the greater part of the right

maxilla including the palatal portion, the base of the jugal arch,

part of the left maxilla, and a portion of the right palatine. The

first seven molars are present. They agree with the description

given by Dr. Broom as far as can be seen from the crowns of

the 4th and 6th teeth, which are the only teeth preserved entire.

The 7 teeth occupy a length of 15°2 mm.

In front of the 1st. molar there is a markedly concave diastema

of which a length of 9 mm. is preserved. There is no remnant of

the canine present, and if the base of the canine was on the same

level as the base of the molars, the diastema would probably have

been about 12 mm. long. It has a maximum height of 4 mm. above

the base of the teeth.

342 Annals of the South African Museum.

The maxillary part of the palate is complete on the right side.

It is 48 mm. long, and narrows slightly anteriorly. Between the

bases of the dst. molars the palate, when complete, had a width of

26-5 mm., while at the pomt of junction of the maxilla with the

premaxilla the width was 22 mm. The alveolar part of the bone

which contains the roots of the molars is fairly massive, but the

palatal portion of the bone has a thickness of but 2 mm.

In front of the maxilla a small palatal fragment of the premaxiila

is preserved, while posteriorly a section of the right palatine is seen

forming a vertical wall to the narial passages and curving downward

and inward to form part of the secondary palate, articulating on the

palatal surface with the maxilla. The vault of the palate is low,

being a flattened concavity.

Fig. 7. Tritheledon ricono, Br.

Palatal view of right maxilla.

By placing the type specimen in its correct position with respect

to this fragment it is seen that in all probability the first molar

tooth on the type is in reality the second molar of the animal.

It would thus appear that this form has 10 molars; for it is im-

probable that any are missing from the posterior part of the type,

the alveolar border of the maxilla thinning away rapidly behind the

last of the series displayed.

All the vertebrae preserved are incomplete. The largest — pro-

bably from the mid-dorsal region — has a centrum 34 mm. long,

whose ends are 22 mm. high and 20 mm. broad. The ends of the

centra are slightly concave. The centra are regularly constricted,

the median width being 16 mm., and along the ventral surface is

a well-defined median longitudinal grove, which broadens at the two

ends. The centra are relatively longer than those of Cynognathus.

Fauna and Stratigraphy of the Stormberg Series, 343

The neural canal is well-defined. The transverse processes are

placed midway along the vertebra, and arise at the level of the top

of the centrum. The base of the process occupies at least two-

thirds of the entire length of the centrum. No zygapophyses are

preserved.

Type. Skull-fragment S.A.M. Cat. No. 1885.

Locality. Paballong, Mt. Fletcher, Cape Province.

Horizon. Red Beds.

LYCORHINUS ANGUSTIDENS, gen. et sp. nov.

Text fig. 8.

The type of this new form consists of a portion of the left ramus

of the lower jaw of a Cynodont from Paballong, Mount Fletcher,

presented to the Museum by the late Dr. M. Ricono. The fragment

shows the canine, 7 molars and the impressions of 4 others, making

14 in all. The ramus lacks the lower border, and is broken off

anteriorly at the front of the canine and posteriorly at the ninth

molar.

The canine is long and powerful, with a long root, and curves

Fig. 8. Lycorhinus angustidens, Htn.

Type dentary.

slightly backwards. The point is absent and the inner anterior side

of the upper sixth is bevelled off by rubbing against the canine of

the upper jaw. In section the tooth is oval with the longer dia-

meter along the jaw. The anterior border is ridged and serrated

for the greater part of its length; the posterior border is also ridged,

but is unserrated.

Close behind the canine, separated from it by a diastema mea-

suring at the most 1:5 mm, come the molars. They are all of the

same pattern, increasing gradually in size to the fourth, and decreasing

344 Annals of the South African Museum.

from the eighth. The roots are long and nearly circular in cross-

section. Half-way up the crown on the inner side is a cingulum

above which the crowns are laterally compressed, so that seen

from above they are considerably longer than broad. Above the

cingulum on the inner side the crown slopes upwards and outwards

and the thinning of the crown thus produced is also accentuated

apparently by a somewhat inwardly-directed slope of the outer

face. Each molar from the 4th onwards is provided with two

cusps, a large anterior cusp occupying two-thirds of the grinding

surface, and a much smaller, somewhat lower posterior cusp. On

the inner face there is a groove running between the ridges which

descend from the cusps to the cingulum. The second and third

molars are provided each with an additional cusp in advance of the

large one which has been called the anterior cusp; and to a lesser

extent this feature is also seen on the first and fourth molars. The

crowns are but scarcely worn. The outer side is unfortunately not

seen, but its general characteristics can be obtained from the im-

pressions of the 8th, 9th, and 10th molars.

The total length of the fragment as preserved is 446 mm. The

crown of the canine is 19°5 mm. high, and 7 mm. long at the base.

The first ten molars together occupy a length of 33 mm. At the

canine the depth of the dentary was about 17 mm.

Type. Portion of lower jaw with teeth. S.A.M. Cat. No, 3606.

Locality. Paballong, Mount Fletcher, C.P.

Horizon. Red Beds.

SUPER-ORDER ARCHOSAURIA.

Fam. SPHENOSUCHIDAE nov.

SPHENOSUCHUS AcUTUs, Htn.

Text figs. J—16.

1915. Haughton. Ann. S. Afr. Mus. XII, 3, p. 98.

Since the original description of this form was published a cer-

tain amount of additional development has been done upon the type

specimen, which necessitates slight additions and emendations. It

has been thought best, therefore, to recast the description.

The skull is somewhat crushed but nearly whole, and shows all

the external details. It is larger than that of Huparkeria, and is

comparatively more pointed, longer, and narrower. The orbits are

rounded, and wholly in the posterior half of the skull. The supra-

Fauna and Stratigraphy of the Stormberg Series. JAD

temporal fossa is elongate, oval in shape, and larger than that of

Euparkeria or Ornithosuchus. The shape of the infratemporal fossa

is characteristic in that its anterior border, formed by the jugal and

postorbital, passes upwards and forwards instead of upwards and

backwards as in Huparkeria and Ornithosuchus; so that the superior

length of the opening is about equal to the inferior length.

The snout is characterised by the fact that the premaxilla does

not form an anterior border to the nostrils, these being quite ter-

minal. Further, there is no trace of a median septum 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

SEL pr

Fig. 9. Side view of skull of Sphenosuchus acutus Htn. x }.

(Shghtly restored.)

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.

There is no septomaxilla 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 anterior and lower borders is formed by the

maxilla, which extends back only as 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 car-

nivorous Dinosaurs. The teeth are flattened laterally, and vary

considerably in size. The first maxillary tooth has an antero-posterior

346 Annals of the South African Museum.

diameter of just over 1 mm.; the probable 6th, which is the largest,

has a diameter of 75 mm. The teeth do not increase nor 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

Fig. 10. Top view of skull of Sphenosuchus acutus Htn. X }.

(Shghtly restored.)

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

Fauna and Stratigraphy of the Stormberg Series. 3AT

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 postfrontal. 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

postorbital from the parietal. The interorbital region has a median

elevation, broadened at the level of the postorbital bars, and nar-

rowing 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

a . \\k 7

HA \\\\“S__

Fig. 11. Sphenosuchus acutus, Htn.

Occipital view of skull. x 1.

the supraorbital border. Its greatest length is in the middle line,

the sutures with the parietals 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 Hupa.keria and the allied

forms in that its upper end is in advance of the lower. The des-

cending portion of the postorbital is thus inclined backward instead

of forward, lying in front of the ascending process of the jugal.

The postorbital forms most of the outer border of the upper tem-

poral fossa and a small portion of the upper border of the lower

opening, anteriorly articulating with the frontal, and posteriorly over-

lying a part of the squamosal. Nowhere does it meet the parietal.

348° Annals of the South African Museum.

The jugal is a tripartite bone. Its anterior process forms the

inferior border of the orbit, and articulates with the lachrymal and

the maxilla Jt does not pass up in front of the orbit as in Eupar-

keria. ‘The ascending process lies behind and superiorly internal to

the postorbital and is inclined slightly forward. The posterior pro-

cess forms most of the zygomatic arch, lying outside the quadrato-

jugal.

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 both Euparkeria and Ornithosuchus the quadrato-jugal is a

fair-sized bone whose articulation with the jugal passes downward

and forward, and which passes up in front of the quadrate to meet

the squamosal. In this form, however, the quadrato-jugal is a com-

paratively small flat bone which lies in the lower posterior corner

of the lower fossa and whose articulation with the jugal passes

downward and backward. 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

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, 7.e. half the outer and poste-

rior borders of the supratemporal fossa. It passes over on to the

occipital plate and takes part in the border of the small posttem-

poral foramen.

The parietal has a strong median crest which divides posteriorly

and forms the upper border of the occipital plate. The bone passes

over this lateral crest to form part of the occipital plate. It articu-

lates on the plate on the inner side with the supraoccipital, on the

outer with the squamosal, and below with the exoccipital, forming

a smal] part of the border of the posttemporal fossa. Anteriorly the

bone articulates with the frontal, and below with the opisthotic and

alisphenoid. As in Ornithosuchus there is no interparietal,

The foramen magnum is an oval opening 12 mm. high and 10

mm. broad, lying high in the skull but well in the lower half of

the occipital plate.

The supraoccipital forms a very small portion of the upper border

Fauna and Stratigraphy of the Stormberg Series. 349

of the foramen magnum. It is a triangular bone with a narrow

short shaft passing down to meet the foramen magnum, Its apex

lies under the bifurcating ridge of the parietal. The surface is

slightly concave.

The suture between the exoccipital and paroccipital process is not

to be seen. The latter is fairly high and fairly thin; its outer cor-

ner is bent almost horizontally and rests on the backwardly-directed

part of the squamosal.

The basicranial region differs from that of the Theropodous Dino-

saurs as exemplified by Platevsawrus erlenbergensis.

The basioccipital forms the majority of the rounded condyle,

which lies wholly below the foramen magnum and is slightly hol-

A VIN Kfen. ov.

‘= j ») p>: g i 1X ~XxI

ie Ba

- for mag.

ys li

WZ St /

ff pp OH <= xi

ea EO:

B38. B.0.

int. car,

Fig. 12. Sphenosuchus acutus, Htn.

A. Side view of basicranium and brain-case.

B. Ventral view of basicranium.

lowed out in the middle of its hinder surface by a small notochordal

pit. The bone also forms the base of the foramen magnum. Ante-

riorly it thins considerably and then thickens again; seen from the

side its lower border is very strongly concave. The basioccipital

tubera are strongly projecting and between them the transverse ridge

is deeply hollowed out. The tubera are formed wholly of the basi-

occipital, in strong contra-distinction to those of Plateosaurus and

are thus somewhat reminiscent of the tubera in the Anomodontia.

This likeness is further mtensified by the fact that the basisphenoid

sends back a process on either side to support the anterior wall

of each tuber. Between these basisphenoidal processes and anterior

350 Annals of the South African Museum.

to the transverse ridge the basioccipital rises considerably so that

seen from below it has a deep central pit. This pit is triangular

in shape with its base behind and its apex in front. The base is

formed wholly by the basioccipital. The upper portion of the sides

is also formed by the basioccipital, but the larger, lower portion is

formed by the basisphenoid. Superiorly this pit divides into two

canals separated by a rod of bone which passes upwards from the

transverse basioccipital ridge and then turns forwards. The openings

to the canals are elongate vertically. Superiorly the basioccipital

meets the exoccipital. On the left hand side of the skull the suture

can be seen running from just below the top of the condyle — the

exoccipital forming a very small portion of the condyle — forwards

and slightly upwards to a point above the basioccipital tuber, and

then passing downwards so that the exoccipital forms part of the

lateral ridge running down to the tuber and meets the basisphenoid.

The exoccipital forms all the lateral border of the foramen magnum,

and the two almost meet above the foramen. The foramen for the

XIIth nerve (hypoglossus) is single and small.

The basisphenoid sends back two processes as supports for the

tubera, as mentioned above. In front of these the bone narrows

somewhat and then sends forward the two pterygoid apophyses.

At its narrowest the bone is deeply grooved in the median line.

The lower border of the pterygoid apophysis bends down sharply.

Between the exoccipital and the basisphenoid on the side is a

small foramen, presumably that of the carotis interna. Between the

exoccipital and a dumb-bell shaped piece of bone forming part of

the side wall of the brain and seen in oblique section is another

foramen. The bone is probably a portion of the prootic, and the

foramen, which lies above and slightly in advance of the supposed

foramen for the carotis interna, is probably the foramen lacerum for

the [Xth-XIth nerves. On the dorsal side of this bridge of bone

is a large irregular shaped opening which I take to be the fenestra

ovalis and the opening for the VIIIth (auditory) nerve. Anterior to

this and looking downwards and forwards is the large circular

foramen ovale for the Vth (trigeminal) nerve.

Anterior to the foramen magnum the brain case heightens rapidly.

Posteriorly it is considerably constricted and above the foramen

ovalis the bone of the side-wall is pierced by a small circular opening,

probably for a blood-vessel. Anterior to this the brain-case expands

laterally and is bounded by the parietal above and the alisphenoid

on the side — the two bones being divided by a straight horizontal

suture,

Fauna and Stratigraphy of the Stormberg Series. 301

The condition of the basicranium is somewhat similar to that seen

in Thecodontosaurus antiquus, but differs in that the basisphenoidal

tubera are placed further back and more ventrally in this new form,

so that the lower surface of the basioccipital instead of passing

almost entirely forward has a very concave profile when viewed from

the side. The basisphenoid, too, differs in that it is longer in com-

parison with its width, neither the outward ridges of the bone which

lead to the tubera nor the forwardly directed pterygoid apophyses

diverging at so great an angle from each other as in the higher form.

The postpalatal vacuity lies directly below the orbits, and has a

length of 26 mm., and a width of 9 mm. Almost the whole of its

inner border, its anterior border, and part of its outer border are

formed by the palatine. Its hinder border, and the remainder of

the outer border is formed by the transpalatine.

The internal nares are not seen, but must have been considerably

in advance of the postpalatal vacuity.

The anterior prolongation of the pterygoid passes far forward as

a narrow bone with a rounded keel, separated from its neighbour

by a well-marked groove. At the level of the front of the postpalatal

vacuity it begins to broaden out and passes back as a gradually

broadening plate articulating with, and lying below, the palatine.

Posteriorly the mesial rounded keel becomes less pronounced and

the platelike portion of the bone has a slight median depression.

The palatine is not wholly displayed. It articulates with the

pterygoid internally, with the maxilla externally, and with the trans-

palatine posteriorly, forming the anterior and inner borders of the

postpalatal vacuity. The front part of the bone carries a pronounced

ridge which runs zig-zag across the bone with deep hollows in front

and behind, There is no evidence that this ridge carried teeth.

The transpalatine is a stout bar with a strong expanded articu-

lation with the maxilla externally. It separates the postpalatal

vacuity from the large pterygoid vacuity; internally it is broken off,

but the impression of the upper surface of the bone on the matrix

remains in part, and it seems to have met both the palatine and

the pterygoid.

Of the whole palate the premaxilla, maxilla, and prevomer form

but a small part: in this, as pointed out in an earlier paper, the

form agrees with Erpetosuchus and differs from Ornithosuchus.

The lower jaw is incomplete. About 100 mm. of the right ramus

is anchylosed to the anterior 40 mm. 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

302 Annals of the South African Museum.

present. 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

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

symphysis. 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 part of the angular

lying within the splenial and dentary. There is a_postarticular

process passing behind the extremity of the quadrate.

Fig. 13. Sphenosuchus acutus, Htn.

Anterior caudal vertebra. x 1.

Vertebrae. The anterior cervical 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 preserved. 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 3rd cervical, slightly expanded at the crest, with a shallow

groove running down the posterior border. The anterior ribs are

double-headed.

There is an anterior caudal preserved (Text fig. 143). It is 16 mm.

long and has a total height of 37 mm. The body is somewhat con-

stricted in the middle, having a minimum width of 8 mm. The

ventral surface is broadly rounded. The ventral border is concave

with the posterior end lower than the anterior, The anterior surface

Fauna and Stratigraphy of the Stormberg Series. 300

is concave. The posterior surface is concave in the middle, but its

border slopes away to the edges of the surface so that its outer half

is convex. There is thus a large surface for articulation with the

haemapophysis below the concave portion of the posterior face of

the centrum. The neural canal is circular in cross-section. The

transverse processes rise high up and pass outwards, slightly upwards

and slightly backwards. They are long, narrow, and thin. The

zygapophyses diverge from each other strongly. The prezygapophyses

have flat upper faces which look inwards. The neural spine is high

Fig. 14.. Sphenosuchus acutus, Htn.

Internal view of left scapula. x 1.

and thin, narrower and higher than is usual in the Dinosauria or

Crocodilia, Its base is broader posteriorly than anteriorly, and lies

wholly in the hinder two-thirds of the neural arch. The width

across the transverse processes is 30 mm.; that across the prezyga-

pophyses is 125 mm.; the length of the upper border of the dorsal

spine is 8 mm.

Shoulder girdle. The shoulder girdle is preserved entire, and con-

sists of two scapulae, two coracoids and an interclavicle.

The scapula is 81 mm. long and expanded both at its proximal

304 Annals of the South African Museum.

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

45 mm. in width. There is no acromion process.

The bone figured and described in the original paper as a clavicle

is in reality a coracoid. The left coracoid is perfect save for the

extreme distal point. The coracoid is a long, thin bone, whose shape

can best be understood from the figure. The ventral margin is

slightly convex and thin. Above this thin border the bone swells

Fig. 15. Sphenosuchus acutus, Htn.

Inner view of left coracoid. x 1.

and on the outer side there is a very slightly roughened elongated

surface apparently for the insertion of the coraco-brachialis muscle.

Above the interclavicular portion the anterior border is concave to

a point directly below the coracoid notch, and then is again some-

what concave, the two curves being separated by a_ well-defined

obtuse angle. The upper border of the post-glenoid prolongation is

slightly concave. Anterior to what is apparently the scapular arti-

cular surface there is a small coracoid notch measuring only 2°50 mm.

in diameter, and beyond this is a thin spatulate pre-glenoid prolon-

gation with a length along its upper border of 14 mm. The greatest

length of the bone is 77 mm. The width at the coracoid notch is

13 mm.; that at the broadest part of the postglenoid prolongation

Fauna and Stratigraphy of the Stormberg Series. 399

is 144 mm. The length of the postglenoid process is 45 mm. while

that of the interclavicular articular surface is 37 mm.

This coracoid differs from that of any known forms. In the Phyto-

sauria the coracoid is rounded in form and has a large coracoid

notch, and the pre-glenoid portion is much longer than the post-

glenoid. In the Pseudosuchia the form is variable. Ornithosuchus

has a rounded coracoid with a short posterior ventral process, and

a supracoracoid foramen. Huparkeria has a large rounded coracoid.

Schleromochlus has a long, flattened, rod-like coracoid like those of

the Birds and Pterosauria. In the Crocodilia the coracoid approxi-

mates somewhat to that of Sphenosuchus. It also is composed of two

expanded ends joined by a relatively narrow neck. But in the

Crocodilia the bone is much higher and relatively shorter, and there

is a foramen instead of a coracoid notch. This latter feature, however,

is variable in the Crocodilia; for Andrews has figured two species of

Metriorhynchus from the Oxford Clay, one of which has a coracoid

notch and the other a coracoid foramen. In none of the Triassic

Coelurosauria is the coracoid known; but in Aetonyx and_ other

Triassic Saurischia it is a large rounded bone with a supracoracoid

foramen, somewhat similar to that of Kuparkeria. The South African

form which most closely approximates to Sphenosuchus in this respect

is Notochampsa.

Humerus. The humerus is 415 mm. long. The proximal end is

broad; in the inner view it is seen to be hollowed out between

the delto-pectoral crest and a well-marked ridge which runs down

on the inner side from the proximal condyle. The delto-pectoral

crest lies considerably below the level of the head of the bone, so

that the proximal expanded portion occupies two-fifths of the total

length. The shaft is slightly curved, and oval in cross-section, the

distal end being more strongly bent than the proximal. In the

narrowest part the shaft has a diameter of 10 mm. The distal end

is but slightly expanded and shows two distinct rounded condyles,

of which the inner is much the larger. Seen in lateral aspect, the

bone presents somewhat the appearance of that of Ornithosuchus,

but it is longer, more slender, and the distal condyles are more

rounded,

Tibia and Fibula. There is also preserved a tibia and the distal

third of a fibula from the same Jimb. The tibia is a long, thin

bone slightly curved, with an expanded proximal end, and a robust

distal end. Its greatest length is 120 mm.; the distal end is 15 mm.

wide, the shaft 9 mm., and the proximal end 29 mm. The tubero-

sitas tibiae is the highest point of the bone. From it the articular

356 Annals of the South African Museum.

surface slopes downwards, gradually being rounded off into the

posterior face of the bone. The inner edge of the proximal surface

is evenly rounded; on the anterior half of the outer edge there is

a prominent lateral condyle from which a short, prominent ridge

passes down on to the shaft. This condyle lies well below the level

of the head of the bone. At the distal end there is a differentiation

into two processes, one of which lies higher than the other, as in

the Triassic Saurischia; but the step from one to the other is not

so deep as in, say, Thecudontosaurus.

Fig. 16. Sphenosuchus acutus, Htn.

A. Inner view of left humerus.

B. Outer view of right humerus.

Both figures x 4.

The distal portion of the fibula shows the bone to have been more

slender than the tibia. The distal end is slightly expanded. The bone

is oval in cross-section.

Type. Skull and part of lower jaw with bones of the shoulder

girdle, part of fore limbs, and part of hind limb, with vertebrae and

rib fragments. (S.A.Mus. Cat. No. 3014.)

Locality. Paballong, Mount Fletcher, Cape Colony.

Horizon. Red Beds.

Affinities. In the general form of the skull Sphenosuchus agrees

with most of the members of the super-order Archosauria, i.e. in

the possession of supra-, infra-, and post-temporal arcades and vacuities.

Fauna and Stratigraphy of the Stormberg Series. 307

In some Dinosaurs the post-temporal vacuity is closed. In Sphenosuchus

it is very small, much smaller than in the Phytosauria or genera

like Huparkeria.

The skull has a smaller quadrato-jugal than EHuparkeria, but shows

an advance on- that form in the loss of the interparietal and the

reduction of the lachrymal.

Chief interest lies, however, in the post-cranial part of the skeleton

and in the lack of armour. This latter feature is noteworthy as the

larger number of the genera comprising the order Thecodontia, to

which this form was originally assigned, possess armour in the form

of scutes which are sometimes numerous and heavy. No armour

has, however, yet been seen in T'hecodontosaurus, Massospondylus,

and the genera of the Saurischia.

The shape of the coracoid (erroneously described in the original

description as the clavicle) is unique among reptiles, and strongly

resembles that of many Birds. Among Reptiles, it is approximated

most closely by the bones in Notochampsa and the Crocodilia. Hupar-

keria has a rounded coracoid, and so has Massospondylus, so that

Sphenosuchus cannot be ancestral to the latter nor to the Saurischia.

Its humerus and tibia, however, are more like those of the Saurischia

and Massospondylus and its allies than other Archosauria; and the

shape of the distal end of the tibia shows that the astragalus was

probably fairly immovably fixed to the tibia. Further, the possible

lack of clavicles is important. The other bones of the scapular arch

are so well preserved and so nearly in place that it is scarcely

possible to suppose that bony clavicles, if present in the living animal,

were not preserved. Lying anterior to the arch and dissociated from

the scapula and coracoid is a thin elongate bone — not complete —

which might be a clavicle or a neck rib or even a long bone of the

hyoid arch similar to the ceratobranchial described by Broom in

Euparkeria. The absence of clavicles would remove the form alto-

gether from the order Thecodontia. Further, it could only be put

with difficulty imto the Saurischia on account of its peculiarly spe-

cialised coracoid, which must be taken as a proof of a certain simi-

larity in musculature between this form and the Birds. On the other

hand, the coracoid is more elongate and birdlike than that of

Archaeopteryx, which has already acquired feathers; and it is probable

that this peculiarity in the coracoid of Sphenosuchus is merely a

specialisation which was not a step on an advance from the Reptile

to the Bird. The absence of clavicles would prevent Sphenosuchus

from being on the direct line of evolution of the class Aves.

Taking all the facts into consideration, it is necessary to found

358 Annals of the South African Museum.

for the genus a new family, which may be called the Sphenosuchidae,

and which on account of the uncertainty as to the occurrence or

absence of clavicles, cannot be satisfactorily included in any of the

defined orders of the Archosauria.

Fam. NOTOCHAMPSIDAE, nov.

NoOTOCHAMPSA ISTEDANA, Broom.

Texto As:

1904. Broom. Geol. Mag. Dec. V, Vol. I, p. 502.

The only hitherto-published description of Notochampsa is that by

Broom who gave a very brief notice of the two forms from the

Stormberg Beds which he included in the genus. At the time of

writing that paper, the remains of the larger animal (N. istedana)

had not been completely developed, but Broom considered that enough

was shown to place the forms in the Crocodilia (Mesosuchia of Huxley).

A short time ago Watson suggested that the larger form was possibly

not a crocodile at all, but was related to Stegomus. I have therefore

further developed the fossil in the hope that it would be possible

to settle its systematic position. There are now displayed .the im-

pressions of most of the bones of the top of the skull, most of the right

ramus of the lower jaw with the whole symphysis, the shoulder girdle

(right scapula, two coracoids, and an interclavicle), the proximal end

of the right humerus, the humerus, radius, ulna, carpus, and meta-

carpus of the left side, part of the femur, tibia, and fibula of the

left side, most of the dorsal armour, and some ribs. These remains

merit a somewhat fuller description than that already given.

Skull. Unfortunately, although further development has shown

a little more of the top of the skull, nothing can be seen of the

structure of the palate. ;

In the middle line, as preserved, the skull has a length of 101°5 mm.,

while its greatest length is 147 mm. The length of the lower jaw

from the symphysis to the articular surface is only 106 mm. Little,

therefore, can be missing from the front of the snout although the

anterior borders of the nares are not seen. The nares must, in con-

sequence, have been almost terminal. In his restoration of the skull

Broom figures the nares as being typically crocodilian, 1.e., coalesced

into one central opening in the premaxilla, and thus adds about

14 mm. to the length of skull as preserved. This, I think, is a

misinterpretation of the structure because, in the first place, the

disparity in length between the mandible and skull becomes too

Fauna and Stratigraphy of the Stormberg Series. 309

ereat and, secondly, there seems to be sufficient evidence in the nasal

region itself to prove that the nostrils were separate and somewhat

lateral. The whole structure of the remaining portion of the snout

seems to point to this conclusion, On the righthand side the im-

Fig. 17. Notochampsa istedana, Br.

Top view of skull. x 1.

pression of the bone is excavated as if it were the posterior border

of the nostril. This portion of the snout seen in profile is distinctly

raised, whereas the remainder of the snout is slightly concave along

the median line, and between the nostril and the centre line is a

shallow concavity. If this reading of the structure is correct, then

360 Annals of the South African Museum.

the nostrils were somewhat laterally placed and separate as in the

Pseudosuchia. It demands an additional length of about 5 mm. to

the tip of the snout, thus making the upper and lower jaws more

commensurate than does the earlier restoration.

The snout is fairly long, and tapers slightly anteriorly. Most of

its upper surface is formed by the nasals, which are separated by

a long median suture, and terminate behind in a digitating suture

with the frontals.

The maxilla bears a number of teeth, about 8 or 9 in all probability.

The first two preserved on the right maxilla are large teeth with long

roots each set in its own socket, and with backwardly-curved poimted

crowns somewhat oval in section. There were no prominent anterior

or posterior edges. The posterior teeth were smaller than the others,

but there is no regular diminution in size.

The snout is broken off anterior to the orbits, and the whole of

the orbital region is missing. Just anterior to the hiatus the snout

swells laterally and becomes raised on the two sides, the median

depression remaining constant. The lateral prominences look to be

each the impression of a separate bone, although sutures are extremely

difficult to delineate. Each contains a portion of the bone figured by

Broom as lachrymal, and each is probably wholly a prefrontal. If

this be so, the lachrymal was wholly on the side of the skull.

Only the impressions of the lower surface of the bones surroun-

ding the vacuity are present.

The upper temporal vacuities are present. Their anterior borders

are formed by the postorbitals. Broom figures the squamosal as

forming the whole of the bar between the upper and lower temporal

fossae: but on the right hand side there is a well-marked separation

between the bone forming the inner part and that forming the outer

part of this bar, and the separation is seen also in part on the left

hand side of the skull. The inner of the bones I take to be the

upper portion of the quadrate, which just meets the postorbital

anteriorly, and has the relations of the quadrate in the recent Croco-

dilia. Externally and posteriorly it articulates with the squamosal.

The squamosal also passes outwards, backwards, and downwards

to form the posterior border of the lower temporal vacuity and _ to.

meet the quadrate. On the inner side it meets the supraoccipital

and exoccipital.

The parietal bar is very broad.

The quadrate is partially displayed. The form of its articular

surface is not seen. The inner edge of the bone is bent upwards

to meet the squamosal; and its other relations have been stated above.

Fauna and Stratigraphy of the Stormberg Series. 361

The lateral temporal opening was long and fairly low. In it can

be seen a portion of the quadrato-jugal, lacking the outer edge, and

articulating with the quadrate.

The supraoccipital articulates above with the parietal in a fairly

lone suture. Ventrally it narrows rapidly and articulates on its

oblique edges with the exoccipitals, which are apparently large bones.

The weathering away of the bone shows a canal opening into the

auditory region between the quadrate and the parietal on the posterior

border of the upper temporal opening, and this is continued into the

post-temporal vacuity, bounded by the squamosal and parietal above

and the exoccipital and supraoccipital below.

Mandible. Part of the lower jaw is preserved. The symphysis is

fairly short, and formed apparently wholly by the dentaries. The

lower jaw is slender, its deepest part being just behind the Meckelian

cavity.

Of the right ramus, the dentary is wholly preserved. It meets its

neighbour in a symphysis 12 mm. long. The alveolar border is

straight and carries about 12 teeth of which 9 can be seen. The

first two are in the position of incisors and point strongly forwards.

The third is considerably larger than any of the others. Posterior

to it the teeth seem to be of approximately equal size to one another.

At the level of the last three or four teeth the dentary is extremely

shallow, while it broadens again towards the Meckelian cavity, of

which it forms the anterior half of the border. The ventral face of

the symphysial region is convex from side to side, and pitted.

The splenials are not preserved; but the shape of the imner

surface of the right dentary shows their approximate positions. The

bone did not enter the symphysis. For over one-half its length it

lay along the inside of the dentary. For a distance of 18 mm. it

was separated from the dentary by the angular, lying along the

inner surface of that bone almost to the level of the outer fossa.

The angular has relations with the other bones, as far as can be

seen, similar to those in the Crocodilia. It forms the posterior border

of the outer fossa, and passes on the ventral side below the dentary,

separating it from the splenial. None of the other bones are seen.

Scapula. The right scapula is displayed in full, as well as the

glenoid end of the left scapula. The bone is extraordinarily expanded

at its upper end, and somewhat swollen below. The posterior border

from the narrowest part of the bone upwards is almost straight,

save at the upper extremity; but the anterior edge is strongly

bent owing to the anterior prolongation of the upper part of the

bone. The lower part of its anterior border is swollen and convex,

362 Annals of the South African Museum,

while the articulation with the coracoidal element is almost straight.

Approximately half the glenoid cavity is formed by the scapula.

From the narrowest part upwards the bone is very thin. From the

upper posterior corner to the middle of the glenoid cavity the scapula

measures 43 mm. At its narrowest it Is 45 mm. wide; at its upper

border it is 50 mm, wide. The articulation with the coracoid measures

8:5 mm.

Coracoid. Both coracoids are seen. The bone is smaller than the

scapula, strongly compressed and expanding considerably at each end.

Fig. 18. Notochampsa istedana, Br.

A. Scapula and coracoid.

B. Proximal half of humerus.

It forms the lower half of the glenoid cavity. Anterior to the

glenoid cavity and just below the articulation with the scapula the

bone is pierced by the coracoidal foramen. The distal expansion

is very broad, terminates in a convex border and is prolonged more

posteriorly than anteriorly.

The coracoid has a length of 25 mm. in the middle line. Its

lower border measures 15 mm., while its narrowest part has a width

of only 3 mm.

The Interclavicle is a narrow elongated strip of bone, not wholly

preserved.

Humerus. The humerus has a long, thin, flattened shaft with a

considerably expanded upper end. The lower end is not preserved

on either side of the body. The total length of the bone was probably

Fauna and Stratigraphy of the Stormberg Series. 363

about 60 mm. The maximum width at the proximal end is 14 mm.,

while the shaft is 4 mm. broad and 1:5 mm. thick as preserved.

The head is rounded, and there is no very prominent deltoid crest

although some distance below the head of the bone a slight crest

was present. The anterior face of the expanded portion is concave.

The radius and ulna are both long, slender bones but slightly

expanded at their ends. The bone lying as the outer of the two on

the left-hand side of the body is slightly longer than the other, and

this I take to be the ulna. The left forearm is lying parallel to the

body with the elbow behind and the hand in front, bent almost at

an angle of 150° to the plane of the humerus.

Radius. The radius has a length of 52 mm., while the middle

of the shaft has a greater diameter of 25 mm. The bone is com-

pressed and has its end expanded to a width of about 4mm. Save

at the lower end, where it is slightly curved away from the ulna, it

is straight throughout its length.

Ulna. The ulna is a slightly curved bone 54 mm. long, apparently

somewhat more slender than the radius, but with similar expanded

ends. Just below the surface for the articulation with the humerus

the radial border is markedly concave.

Carpus. The carpus of the left side is partially displayed. Two

bones of the proximal row can be seen, one below the ulna and one

below the radius. These I regard respectively as the ulnare and

the radiale.

The whnare is an elongated bone with a narrow shaft and expanded

ends. In cross-section it is apparently oval. Its length is 10 mm.,

the proximal width 4 mm., and the distal width 5 mm., while the

shaft of the bone has a width in its narrowest part of less than

2mm. The proximal articular surface has a small outer convexity,

the remainder being hollowed out. The distal surface is cushion-shaped.

Attached to the lower end of the bone is a small bone, oval in

section as seen, 2 mm. high and 4 mm. broad, which may be the

fused 3rd, 4th, and 5th carpals. The radiale is a similar hour-glass

shaped bone 11°5 mm. long with its ends somewhat less expanded

than those of the ulmare. The narrowest part of the bone is 2

mm. wide.

Metacarpus. Remains of all five metacarpals are seen. The third

and fourth articulate with the bone lying below the ulnare, but not

the fifth. Hach metacarpal is an elongated bone with expanded ends.

The first is 9°7 mm. long, the second 10 mm., the third 95 mm.

and the fourth 83 mm., while the fifth is not fully displayed.

Femur, The distal half of the femur is slightly curved and has

364 Annals of the South African Museum.

an expanded end, which was probably slightly cartilaginous. The

preserved portion of the bone is 48 mm. long, and the distal end

measures 114°5 mm. in its greatest diameter. The shaft at its narrowest

is between 5 and 6 mm. thick. The outer condyle is larger than

the inner and on the anterior surface between the two is a shallow,

broad depression.

The tibia and fibula are both long, slender bones, longer and more

robust, however, than the bones of the lower part of the fore-leg.

The tébia has a length of 62 mm. Its shaft is long and straight;

the proximal end of the bone is swollen to a thickness of 65 mim.,

and rounded. The distal end is somewhat crushed, but it is wider

than the proximal end and has a protuberance on the fibular side.

The fibula is a more slender bone, 60 mm. long, with slightly

expanded ends.

Armour. The back was covered with a series of paired dorsal

scutes, one of which was figured by Broom. There were apparently

neither lateral nor ventral ossifications

at least, none are preserved

on this specimen. Each pair of scutes covered one vertebra. Remains

of 20 pairs are preserved, almost wholly as impressions of the under

surface of the bone. Each scute was roughly rectangular, but towards

the outer end of the posterior border there was a projecting process.

From the twelfth pair onwards the width gradually decreases. The

first scute is short, the second shghtly longer. The 3rd is a little

longer than the 4th or Sth. From this onwards the length is

constant. The 41th. scute is 20 mm. broad and 9°5 mm. long at

its inner edge. The 19th. has a breadth of 1445 mm. and a length

of 9 mm.

Affinities. In the elongation of the rostrum the form agrees with

the Crocodilia and Phytosauria; but in the possession of paired and

lateral outer nares it diflers from the former, and in the short

premaxillary and forward position of the nostrils from the latter.

The elongation of the rostrum is a secondary character,

I am not able to determine with absolute satisfaction whether or

not a preorbital vacuity was present. The orbit itself is absent, but

both prefrontals can be seen. On the right hand side below the

prefrontal on the side of the skull there is a distinct depression with

superior, anterior, and inferior borders of bone. This may — and

probably does — represent a preorbital vacuity, looking wholly

outwards and placed below the orbit and but slightly in advance of

it, a condition paralleled to a certain extent in the Phytosaurians

Mystriosuchus and Rhytidodon.

The scapula is without parallel among Archosauria in the extreme

Fauna and Stratigraphy of the Stormberg Series. 365

broadening of its dorsal half. The coracoid has the general shape

seen in the Crodilia; but an even greater departure from the rounded

bone of the Pseudosuchia and Phytosauria is presented by the

coracoid of Sphenosuchus, described in this paper, which is undoubt-

edly a Pseudosuchian of rather advanced type. In the possession of

an interclavicle and the absence of clavicles Notochampsa agrees with

the Crocodilia and differs from the Dinosauria, Phytosauria, and

earlier Pseudosuchia.

The fore-limb is not Crocodilian in structure. The humerus in

its expanded proximal portion with concave anterior surface, its

deltopectoral crest well below the level of the head of the bone, and

its narrow shaft is reminiscent of that of Stagonolepis which, however,

departs from the normal Phytosaurian form and approximates to

that of the Pseudosuchia and Saurischia.

The long and slender limbs show affinities with such forms as

Stegomus and Schleromochlus, although the front limb approximates

in length to that of the hind limb much more nearly than in the

latter form. The genus herein differs greatly from the Jurassic Crocodilia.

The armour is peculiar in that it apparently consists only of two

rows of scutes along the back. These are broader than long as in

Stegomus, Aetosaurus and Dyoplaxr. Possibly, however, ventral armour

was also present.

With the form described as Pedeticosaurus leviseuri by van Hoepen

Notochampsa possesses several characters in common. Both have the

snout somewhat elongate, with the nostrils separate, lateral, and

nearly terminal. Both have thecodont teeth, of which the two just

behind the premaxilla seem to be the longest and strongest. In

Pedeticosaurus there is a small antorbital vacuity on the side of the

face — smaller than in other Pseudosuchia -— an arrangement which

may, as stated above, be present in Notochampsa. ‘There is also a

general agreement between the two forms in the broadened distal

end of the scapula, the long and slender bones of the upper arm

and fore-arm, the elongate carpals, the comparatively short ribs, and

the double row of plates down the back. The bone called by van

Hoepen the left scapula appears to be more likely to be a coracoid.

If that be so, it will agree with the coracoids of Notochampsa in

its general shape, having two expanded ends connected by a narrower

shaft. The hind leg in Pedeticosaurus is relatively somewhat longer

than in Notochampsa.

The points of agreement are, I think, sufficiently numerous

provisionally to place the two genera Notochampsa and Pedeticosaurus

in the same family which must be called the Notochampsidae. This

366 Annals of the South African Museum.

displays affinities both with the Pseudosuchia and the Crocodilia.

By von Huene the Crocodilia are thought to have been derived

directly from the Aetosauria. Such a descent implies, among other

things, elongation of the snout, the loss of preorbital vacuity, the

loss of the clavicles, and the production of a typically Crocodilian

coracoid. These changes are partly brought about and partly fore-

shadowed in the Notochampsidae: and although the limbs are not

moditied in the Crocodilian manner save in the lengthening of the

carpals, it may be concluded with some degree of truth that the

family occupies an intermediate position between the Aetosauria and

the Crocodilia, possibly on the direct line of descent. It is unfortunate

that nothing is known of the palate in either of the two members

of the family. :

Type. Skull and portion of skeleton and armour. (S. A. Mus.

Cat. No. 4013).

Locality. Funnystone, Barkly East, C.P.

Horizon. Cave Sandstone.

PEDETICOSAURUS LEVISEURI v. Hoepen.

1915. van Hoepen. Ann. Transv. Mus. V, p. 83. Pls. XIDI-XIV.

The type consists ‘fof a nearly complete individual on two slabs

of matrix. The fossil is an impression of the right side of the skull

and limbs, and besides this, consists of some vertebrae, a few ribs,

a great part of the tail and of dermal ossifications”.

The skull is only seen in lateral view. It has a total length of

abouth 90 mm. The snout is low and fairly long, the front of the

orbit lying a little in advance of the middle of the skull. The orbit

is large and rounded, the antorbital vacuity small with a straight

lower border. The nostrils are almost terminal. The bar separating

the upper and lower temporal vacuities is not clearly displayed.

The lower border of the lower opening is formed mainly by a

shallow jugal. Posteriorly this meets a bone, which, as preserved,

has a free upper anterior end, and occupies the same position as

the bone which I have called quadratojugal in Nolochampsa and

Sphenosuchus. Behind this is a portion of the quadrate with which

the lower jaw still articulates.

The premaxilla carries three small teeth; the maxilla probably

carries 16 or 17 teeth, of which the anterior two or three are large.

All are pointed, backwardly curved, aud apparently unserrate.

The bone which van Hoepen has called a ‘‘hyoid” is apparently

a double-headed anterior cervical rib.

Fauna and Stratigraphy of the Stormberg Series. 367

The account given by van Hoepen of the post-cranial skeleton is

correct save that the bone which he describes as the less-completely

preserved scapula I take to be a coracoid, elongate in shape some-

what similar to that seen in the Crocodilia and in Notochampsa.

It is also possible that the small fragment lying above the supposed

coracoid and below the humerus of the other side is the impression

of part of a small straight interclavicle. The assumption of the

presence of a coracoid is justified by the long straight distal articular

surface of the scapula lying anterior to the glenoid cavity; this is

equal in length to the articular surface of the supposed coracoid,

which has a comparatively narrow middle portion with strongly

expanded ends.

If a coracoid be truly present, then an interesting comparison can

be drawn between Pedeticosaurus and Notochampsa, as I have pointed

out in the re-description of the latter form. Pedeticosaurus is a

slightly, smaller animal having, however, its hind legs longer and

more strongly developed than those of Notochampsa. The resem-

blances are close enough to suggest a family connection, and I have

therefore placed both genera in the same family.

| have to thank Mr. M. Leviseur and the Bloemfontein Museum

authorities for permission to examine the type.

Type. Incomplete skeleton’ on two slabs in the Orange Free State

Museum.

Locality. Rosendal, Senekal Distr., Orange Free State.

Horizon, Cave Sandstone.

OrDER CROCODILIA (?),

ERYTHROCHAMPSA nov.

ERYTHROCHAMPSA LONGIPES (Broom).

1904. Broom. Notochampsa lonyipes. Geol. Mag. N.S. Dec. V.

Vol. I. p. 582, figs. 2 and 4,

Pelvis. In his original description of this form, Broom says ‘‘the

pelvis is typically Crocodilian in that the pubis does not enter the

acetabulum. The ilium is of small size.”

The pelvis as preserved is seen partly on the main slab and partly

on a small piece of rock which was broken off in the development

of the fossil for the purpose of examining the pelvis. This latter

shows the two ischia, most of the two prepubes, and a portion of

the left ilium (partly in bone and partly as a mould). The main

slab shows moulds of these bones with small pieces of bone adhering.

The ilium is incomplete. It was a stout bone, nearly as large as

368 Annals of the South African Museum.

the ischium, with a long dorsal border. There is a long pre-

acetabular portion with a bluntly-pointed extremity.

The ischium is of the usual Crocodilian form. Ventrally it forms

a long union with its fellow by means of a straight suture 145 mm.

long. The ventral portion of the bone is a triangular plate with an

acute posterior angle and a more obtuse anterior angle. The dorsal

portion of the bone is bifurcated. with a larger posterior portion for

articulation with the illum, and a prominent anterior protuberance

directed upwards and forwards for articulation with the unossified pubis.

On neither side is the prepubis complete. Each is a slender bone

with a spatulate anterior portion. The two bones do not meet

ventrally and must have been connected by cartilage. The proximal

end is shghtly expanded and lies at present some distance in front

of the ischium and wholly disconnected from it. The length was

23-5 mm., and the greatest breadth at the spatulate end 45 mm,,

while at its narrowest the shaft is Just under 145 mm. broad. The

shaft is nearly circular in cross-section.

Femur. The right femur is partially preserved. It consists of a

slightly bent shaft with somewhat swollen ends. The ends of the

bone are spongy while the shaft is hollow

a condition which is

seen in the other bones of the hind limb and the pelvis. The length

is about 48 mm. The head is 43 mm. broad, the shaft 3°8 mm.

In cross-section the shaft is a somewhat flattened oval. The distal

end has two condyles separated by a well-marked intercondylar groove.

The outer condyle is larger than the inner. The width of the distal

end is 8 mm.

Tibia. The tibia is 4 or 5 mm. shorter than the femur. It is a

very slightly curved shaft with an expanded proximal end flattened

antero-posteriorly. The fibular side of the shaft is somewhat flattened,

while the inner side is regularly curved. Distally the bone widens

out. The breadth of the proximal end is 6 mm., while the width

of the shaft is 3-5 mm.

Fibula. The fibula is approximately of the same length as the

tibia, but is a much more slender bone. Between the shaft and the

proximal end there is a narrow neck. The shaft has a longitudinal

ridge on the inner anterior border. The distal end is not seen.

The shaft is 2 mm. broad.

Tarsus. The structure of the tarsus is somewhat difficult of deter-

mination. The cleavage of the rock has broken the bones across

the middle so that neither the dorsal nor palmar surfaces are seen,

The astragalus is much larger than the calcanewm, which is a small

bone with a small posterior tuber joined to the main body of the

Fauna and Stratigraphy of the Stormberg Series. 369

bone by a narrow neck. There are two other bones of which the

one articulating with the calcaneum may be the cuboid and the

other the reduced fifth metatarsal or the fused 4th and 5th tarsalia,

Metatarsus. Metatarsals | — IV are similarly elongated narrow

cylindrical bones with slightly expanded ends. The 2nd and 3rd

are larger than the others, while the 4th is the shortest. The lengths

from the 1st to the 4th are 22 mm., 23 mm., 23 mm.. and 20 mm.

respectively.

One phalanx of the 2nd. digit and two phalanges of the third are

preserved. Hach is a fairly long slender bone with concave proximal

and convex distal articulation.

Armour, Portions of the dorsal and ventral armour are preserved.

One of the dorsal plates has been figured by Dr. Broom. It has

a width of 15 mm., and a length of 8 mm. in the middle line.

Its dorsal surface is pitted save for the portion overlapped by the

anterior plate. At the outer sixth the plate is bent downwards,

the bend being strengthened by a dorsal ridge. The total width of

the body was probably about 25 mm. The ventral plates are paired,

but are smaller and more numerous than the dorsal plates. In the

abdominal region each was about 9 mm. wide and 4 mm. long,

while in the caudal region they were smaller.

The two forms described by Dr. Broom as Notochampsa were

included in the one genus on account of resemblances between the

dorsal plates. Further investigation shows that while the pelvis in

“N. longipes” is typically Crocodilian the skull of N. istedana shows

features which remove it from the true crocodiles. The question

then arises, are the two forms congeneric? Dorsal plates are known,

not only im the Crocodilia, but in the Pseudosuchia such as Schlero-

mochlus, Aetosaurus, and Kuparkeria; and it is doubtful whether they

can be considered of generic importance. Unfortunately the pelvis

is absent in the type of N. istedane and also in the allied type

Pedeticosaurus. If Broom’s N. longipes is to be kept in the genus

Nolochampsa then the genus must be considered to be characterised

by the possession of a skull differing from that of a true Crocodile

and of a typically Crocodilian pelvis. This is not impossible; but

until more is known of. these forms it would seem best to separate

the two forms from one another, classing istedana as one of the

higher Pseudosuchians and erecting, as above, a new genus Evythro-

champsa for the more truly Crocodilian Erythrochampsa cvongipes.

Type. S. Af. Mus. N® 445 f.

Locality. Eagles Crag, Barkly East, C. P.

Horizon. WRed Beds.

S&S

Annals of the South African Museum.

Fam. THECODONTOSAURIDAE von HueEne.

4905. von Huene. Zeitschr. d. dtsch. geol. Ges. LVII, p. 345.

THECODONTOSAURUS BROWNI (Seeley).

1895. Seeley. Massospondylus (?) browni, Ann. Mag. Nat. Hist.,

Ser. VI, vol. 15, pp. 102-132.

4906. von Huene. Geol. u. Pal. Abh. XII, 2, p. 45, figs. 82-85

Pl. XII (XIX), figs. 7-8.

49114. Broom. Ann. S. Afric. Mus. VII, 4, p. 293.

The type consists of two femora, 2 cervical vertebrae, a dorsal

vertebra, 3 caudal vertebrae, and some foot-bones from the Telle

River, separating Herschel from Basutoland. There is some doubt

as to whether all the remains are from the same animal, and von

Huene in his re-description has classed the vertebrae with Theco-

dontosaurus skirtopodus. From von Huene’s account, the following

are taken as the salient points of the form.

The femur is small, markedly S-shaped, with a compressed distal

end and a sagittal groove which divides the articular surface. The

bone is strongly compressed laterally. The fourth trochanter lies

wholly in the upper half of the bone. The head is rounded and

very thick. The length of the type is 24 cm.

Von Huene notes that the femur is as large as that of Thecodon-

tosaurus cylindrodon, but at the distal end is smaller and has higher

and smaller condyles than J. skirtopodus. The proximal end_ is

smaller and thicker than in 7. antiguas and T. cylindrodon.

Van Hoepen has described under the name of Massospondylus

browni a fairly complete skeleton in the Transvaal Museum. I be-

lieve this to be a specimen of Massospondylus harriesi and have

discussed it in my description of that form.

Type. Isolated bones in the British Museum.

Locality. Telle River, Herschel, C. P.

Horizon. Red Beds.

THECODONTOSAURUS SKIRTOPODUS (Seeley).

Text figs. 19, 20.

4894. Seeley. Hortalotarsus skirtopodus. Ann. Mag. Nat. Hist. (6).

Vol. XIV, p. 444-419.

1906. von Huene. Thecodontosaurus skirtopodus. Geol. u. Palaeont.

Abh. N. F. Bd. VI1l, Hft. 2) p: 424 otigs) 72-78. Pls) XT Xai.

The type is a portion of a hind limb from Barkly East. Von

Fauna and Stratigraphy of the Stormberg Series. 371

Huene has also described some vertebrae, humeri, a femur, and a

tibia from “the Karroo formation” — now in the Vienna Museum —

as members of this species.

Discussing the generic position of the form von Huene says ‘The

form of the tibia is entirely characteristic of Thecodontosaurus; the

long, laterally-directed projection at the proximal end, the broad

lateral condyle at the proximal end, and the nature of the expan-

sion of the proximal and distal ends are elsewhere only seen in

Thecodontosaurus and: Anchisaurus. The astragalus, also, corresponds

closely with that of Thecodontosawus from Bristol. [I can see no

eround for separating Hortalotarsus from Thecodontosaurus.”

While collecting at Foutanie, Fouriesburg, O. F. S., Mr. A. R.

Fig. 19. Thecodontosaurus skirtopodus (Seeley).

Right ulna. X 4.

Walker obtained some bones from the top of the Red Beds. They

comprise (S.A.M., Cat. No. 3429) some vertebrae, a scapula, the

distal end of a humerus, an ulna, an ilium, part of an ischium, a

femur and a tibia which belong to an animal somewhat smaller

than, but closely comparable with, the type of J’. skirtopodus. The

bones are in good condition and worthy of a short description,

especially as some portions of the skeleton were hitherto unknown.

Scapula. The right scapula is almost complete. Its length as

preserved is 125 mm., but it lacks the distal end. It is small and

slender, the minimum width across the bone bemg 21 mm. _ Distally

the shaft expands very slightly, but proximally it has a greatest

width of 47 mm. The proximal portion is very like that of Massos-

pondylus carinatus. On the lateral face the supracoracoidal surface

is concave. The whole bone is curved in its length, convex out-

372 Annals of the South African Museum.

wards. At the glenoid cavity the bone has a thickness of 17 mm.

Both the anterior and posterior borders of the blade are fairly thin

and keeled, but proximally the anterior border becomes thicker and

more rounded.

Ulna. The right ulna is complete. Its length is 87 mm. The

proximal articular surface is triangular with an acute rounded anterior

angle, and an obtuse lateral angle. The posterior end stands higher

than the anterior. Below the proximal end the bone narrows rapidly.

In its proximal half the medial face of the bone has a shallow broad

longitudinal groove. Distally the bone is twisted and flattened so

that the broad faces of the distal end look directly forwards and

backwards. The distal end is 19 mm. broad. The medial end of

the articular surface is pointed, the lateral end truncate. The

Fig. 20. Thecodontosaurus skirtopodus (Seeley).

Right ilium.

maximum thickness at this surface is 11 mm. At its thinnest the

shaft is 12 mm. broad and 7 mm. thick.

Ilium. The right ilium is typically Thecodontosaurian with its

short anterior spine and its elongate posterior spine. The length of

the upper border is 105 mm.; the border is slightly curved, more

so anteriorly than posteriorly. The posterior spine is more pointed

than in JT. antiquus. On its inner surface the inner crest is strongly

developed. The acetabulum is 60 mm. broad and 30 mm. high.

The supra-acetabular crest is strongly developed above the anterior

part, but disappears posteriorly. The height from the postacetabular

process to the upper border of the bone is76mm. The preacetabular

process is 48 mm. long.

Ischium. The distal end of a left ischium is 147 mm. long. The

shaft is slender, strongly expanded distally. The anterior edge is

straight and sharp, the posterior border regularly concave and flattened.

Fauna and Stratigraphy of the Stormberg Series. 373

The bone was in contact with its fellow for a length of about 80

mm. <At the bottom of the groove on the posterior surface the bone

is 17 mm. thick from back to front. At the distal end the maximum

thickness is 29 mm.

Femur. The distal portion of a right femur is complete from the

trochanter quartus downwards. The distal end agrees closely with

that figured by von Huene in its general contours, its compression

sharply marking it off from Massospondylus carinatus. The length

from the bottom of the fourth trochanter to the distal end is 105 mm.

Both condyles are high; at the lateral condyle the distal surface is

at its broadest. There is a deep, narrow groove between the condyles

on the posterior surface. The minimum breadth of the shaft is

24 mm. The length of the distal articular surface is 39 mm., its

breadth at the lateral condyle 34 mm., and at the medial condyle

29 mm.

Tibia. The tibia is complete save for the proximal articular surface.

The length is 174 mm. The bone seems slightly crushed, so that

the medial edge of tbe proximal end is not very convex. The medial

condyle lies nearer the posterior end of the surface. Behind the

anterior point the lateral border is somewhat concave. The shaft

is flattened laterally and has a minimum thickness of 35 mm., and

a minimum breadth of 11 mm. At the distal end the bone is broader

in front than behind, the breadths bemg 341 mm. and 47 mm.

respectively. The anterior condyle lies 13 mm. above the posterior.

Discussing the relations of this species with European members

of the genus von Huene says Thecodontosaurns skirtopodus has some-

what shorter dorsal vertebrae than TY. antiquus, the cross-section

being the same. The humerus is of similar size to J. antiquus,

but the processus lateralis reaches much deeper here than there.

A bone, which is doubtless the end of the ischium, agrees with a

similar bone in 7’. antiquus and differs considerably from 7’, polyzelus.

The distal end of the femur is smaller and thicker than in 7’. antiquus

and T. cylindrodon, and the condyles are higher than in J. polyzelus.

The length of the tibia agrees with that of 7’. MacGillivrayi; the

proximal end is tolerably small, the lower half of the shaft and

distal end thicker than in TJ. antiquus. The astragalus is as in

T. antiquus. Metatarsal 1V corresponds best with 7. polyzelus.

Type. Portion of hind limb in British Museum.

Locality. Barkly East Division, C.P.

Horizon. Cave Santstone.

374 Annals of the South African Museum.

THECODONTOSAURUS MINOR Htn.

Dextchos2de

4918. Haughton. Ann. Mag. Nat. Hist. IX, II, p. 468.

“The specimens forming the type of this new form were presented

to the South African Museum by the late Dr. M. Ricono. They

consist of a left tibia, a cervical vertebra, and a portion of the

left ilium.

Left tibia. The tibia is 109 mm. long. The proximal articular

surface is 31 mm. long, and 18 mm. broad. This surface for the

most part slopes obliquely backwards and laterally, the inner border

being convex from front to back and higher in front than behind.

The tuberositas tibiae is almost the highest point of the bone; it is

prolonged anteriorly and turned slightly outwards. The lateral con-

dyle is strongly developed. Below the head the shaft thins rapidly

until at its middle it has an anteroposterior thickness of 12 mm.,

and a width of 10 mm. Thence it thickens towards the distal end.

The anterior face is flat with a prominent edge on the lateral side

and a rounded edge medially. The outer sharp edge is continued

down to the anterior distal process. The posterior border of the

shaft is rounded.

The distal surface is trapezoidal in form. The inner anterior

border is 20.5 mm. long, the posterior outer border 16 mm. long,

while the posterior inner border is 12 mm. long. The anterior

process les 7 mm. above the posterior process. Between the two

on the outer surface of the bone is a shallow groove.

Cervical vertebra. The length of the body is 31 mm. The anterior

articular surface is slightly larger than the posterior. Both are con-

siderably higher than broad. The body is pronouncedly amphicoelous.

There is a prominent median ventral keel, sharper in its anterior half.

The whole body is strongly compressed laterally, having a width at

the middle of 5 mm.. and the anterior end of 8 mm. The canal

has a height and breadth anteriorly each of 5 mm. The ends of

the zygapophyses are missing. The dorsal spme was low and fairly

long with a somewhat convex upper border.

Ischium. A portion of what is probably the left ischium is pres-

erved, including the proximal articular surface. The bone is bent

strongly backwards, more so than in Thecodontosaurus antiquus as

figured by von Huene, so that the ischium must have been directed

very strongly backwards. At the broken distal end the bone is

12 mm. thick, and 65 mm. broad. The inner border of the proxi-

mal surface is straight, the lateral border has a prominent outward

Fauna and Stratigraphy of the Stormberg Series. 379

Fig. 21. Thecodontosaurus minor, Htn.

la. Left tibia. 1b. Left proximal end. le. Left distal end.

2. Cervical vertebra. 3. Ischium. All natural size.

376 Annals of the South African Museum.

projection — the maximum width of the surface being 9 mm.

The nature of the tibia and the ischium mark these remains off

from the Plateosauridae and place them among the T'hecodontosau-

ridae. They indicate a member of this family smaller than any

hitherto described from South Africa and which cannot be exactly

identified with any European species. I propose therefore to give

it a new specific name Thecodontosaurus minor.

Type. S. A.M. Cat. No. 3451.

Locality. Pitsing, Maclear, C. P. Cutting in road to Naude’s Nek.

Horizon. Red Beds.”

There is in the Bloemfontein Museum a slab of Cave Sandstone

from Ladybrand, O. F.S., which I was kindly permitted to examine.

The slab contains the cast of a right femur, tibia, the distal end of

the fibula, metatarsals I-[V, and the first three digits of the pes of

a Dinosaur; also part of the right ischium and two fragments of

jaw with teeth. The form seems to correspond pretty closely with

the type of Yhecodontosaurus minor, and I refer it to that species.

It differs from Gyposaurus in being smaller and in having the meta-

tarsals proportionately shorter.

One fragment of jaw is 50 mm. long and 9 mm. deep at the

mentum. 44 mm. from the front it carries a single tooth 6°5 mm.

long and 1:5 mm. in diameter. At the back of the fragment there

are five teeth closely set together, the five occupying a space of

12 mm. and gradually decreasing in size from the first backwards.

The largest is 6 mm. long. The teeth are widest some distance

above the jaw and are serrated, at least on their posterior borders,

in their upper halves. 3 mm. in advance of the largest is a slightly

more slender tooth.

The other jaw fragment is 25 mm. long and carries 6 teeth in a

distance of 18 mm. The longest tooth is 7 mm. long. All the teeth

are serrated coarsely on their posterior and anterior borders.

The femur as preserved is a slightly curved shaft with very

slightly expanded ends. The ends are, however, incomplete. The

greatest length im a straight line is 155 mm., and along the front

curve is 1454 mm. The proximal width is 52 mm., the distal width

30 mm., and the width at the narrowest part of the shaft is 25°5 mm,

The tibia is 1417 mm. long, has a proximal width of 33 mm., and

a distal width of about 25 mm. The following give the measurements

of the metatarsus and pes.

Fauna and Stratigraphy of the Stormberg Series. 3717

Length prox.. width distal width

Metatarsal I 4Q

dst phalanx 28 14 10

claw 36 15

Metatarsal II 56 17 14

dst phalanx 26 13

2nd_ phalanx 19 14 15

claw 29 12

Metatarsal IIT 64 ? 16

dst phalanx 26 18 ?

2nd_ phalanx 21 14 12

3rd phalanx 16 11 11

claw 21 10

The ischium as preserved has a greatest length of 100 mm. Near

its Junction with the pubis and ihum it has a width of 45 mm,,

while at its narrowest part the width was probably not more than

15 mim.

THECODONTOSAURUS DUBIUS sp. nov.

Two slabs of Cave Sandstone from Ladybrand, O.F.S., presented

to the South African Museum by Mr. van Niekerk contain the larger

part of the skeleton of a Thecodontosaurus which seems to be specific-

ally distinct from any yet described from South Africa. A slab and

counter slab from the Cave Sandstone of Rosendal, O.F.S., now in

the Bloemfontein Museum, which | was kindly permitted to examine,

contain remains of a somewhat smaller animal apparently of the

same species,

As usual with the Cave Sandstone fossils the bones are badly

preserved, and it has only been possible to expose one side of the

type specimen. On the larger slab the dorsal and sacral vertebrae,

pelvis, and the hind limbs are seen in ventral aspect, while in

the smaller slab the caudal vertebrae are seen in lateral aspect

mostly as moulds. In the Bloemfontein specimen besides an imperfect

pelvis and hind limb there is preserved a very fine series of 54

caudal vertebrae.

In the type specimen 13 dorsal vertebrae are seen on the ventral

surface — all crushed and weathered. The length of the posterior

vertebrae is 44 mm. each, and the breadth across the articular

surface is about 24 mm. Three sacral vertebrae are displayed. The

length of each of the first two is 45 mm., and the maximum breadth

378 Annals of the South African Museum.

of the centra 30 mm. They are stouter than, and not compressed

medially so much as, the dorsal vertebrae.

There are 20 caudal vertebrae preserved, having a total length of

710 mm., but the anterior caudals are not present. The anterior

centra of those preserved are almost as high as long, having a length

of 33 mm. and a height of 30 mm.; posteriorly the centra become

lower so that the 13th has a height of 13 mm, to a length of 30 mm.

Similarly the dorsal spines and the haemapophyses become shorter

posteriorly. The 2nd haemapophysis preserved is 90 mm. long, the

15th 42 mm.

Both ilia are preserved, but incompletely. The anterior spine is

missing, but the posterior spine of the left bone is long. The height

of the upper border above the end of the postacetabular process is

about 80 mm. The posterior spine was probably squarely truncate.

Both pubes are present, lacking their distal ends. The shaft was

38 mm. broad and thin. The proximal end is broadened and the

pubic foramen is large and rounded. Both ischia are present in

contact with each other and with the pubes; but they lack the distal

ends. The proximal mesial parts of the bones are very thin. The

ischium passes very strongly backwards.

Both hind legs are preserved in a flexed condition, but the feet

are not in good condition.

The femora are only seen from the anterior side. The bone is

slightly S-shaped, bent more proximally than distally. The right bone

is 270 mm. long, the left 280 mm. The shaft of the larger bone

is 35 mm. wide at its narrowest. The distal end of the right femur

is 55 mm. broad. The 4th trochanter is not visible on either bone.

The right tibia is almost entire. It is 260 mm. long as preserved,

but lacks the distal end. The proximal end is 70 mm. broad, with

a somewhat sharp inner anterior end. The anterior face of the bone

is flattened just above the middle, but becomes rounded distally.

The shaft is 27 mm. broad at its narrowest and 25 mm. thick.

The left fibula is 250 mm. long, slightly bent, and slender. The

ends are only slightly expanded.

Of the metatarsals 4 are preserved on the left side, probably I-IV.

They are seen on their under surfaces. Their lengths, taken in

order, are 71, 102, 410 and 95 mms.

This form is thus seen to be slightly larger than Thecodontosaurus

browni.

Type. S. Af. Mus. Cat. No. 3712.

Locality. Ladybrand, O.F.S.

Horizon, Cave Sandstone,

Fauna and Stratigraphy of the Stormberg Series. 379

GYPOSAURUS CAPENSIS Broom.

1906. Broom. Hortalotarsus skirtopodus. Trans. S. Afric, Phil.

Soc: XVI, 3; p. 2017 Pit.

1914. Broom. Ann. S. Afric. Mus. VII, 4. p. 293.

The type consists of a partial skeleton in a block of sandstone,

in which the following portions are displayed: Eleven dorsal and six

caudal vertebrae, a few ribs and some abdominal ribs, part of the

right scapula, both ilia, the right pubis and ischium, the right femur

and fibula, tarsus and pes. The bones are only partly preserved,

and the ends are imperfectly ossified.

In his 1914 paper Broom placed the form in a new genus on the

following grounds. .““Among the chief characteristics of this new

genus the most noteworthy is the remarkable shape of the ilium,

The iliac crest has a greater anterior extension than in any other

known carnivorous Dinosaur, and the preacetabular process is rela-

tively small. The femur is relatively stout, and has the trochanter

quartus small and high up, and the trochanter major almost rudimentary”.

The animal is of the same size as Thecodontosaurus skirtopodus.

The ungual phalanges of the foot seem to be shorter than in that

species, but the general proportions of the remainder of the foot are

very similar,

Type. Partial skeleton in sandstone. S. A. Mus. Cat, No, 990.

Locality. Ladybrand, O. F. 8.

Horizon. Cave Sandstone.

ARISTOSAURUS ERECTUS van Hoepen.

4920. van Hoepen. Ann. Transvaal Mus. VII, 2. p. 77. Plates IX—X.

The form is a small one, the type consisting of the larger portion

of an animal exhibited on a slab and counter-slab of sandstone.

A portion of the left maxillary and left dentary is preserved.

The crowns of the teeth are broad, flat, and have the edges serrated.

The number of dorsal vertebrae is probably 14, and possibly 15.

There are certainly two sacral vertebrae, and possibly a third. Van

Hoepen says of the type “As appears from many loose sutural

connections, our animal is a young individual, and has not yet

reached the stage of life in which it possesses a true third sacral

vertebra.” The shoulder girdle is described as consisting of two

scapulae and two coracoids. The coracoid has a supracoracoid

foramen, which is fairly large and close to the middle of the scapular

380 Annals of the South African Museum.

border. The proximal end of the humerus is very broad, its upper

border convex. Below the processus lateralis the lateral border is

strongly concave. The shaft of the bone is narrow. The ulna is

just over two-thirds the length of the humerus.

The spina posterior of the illum is broad and truncated, the spina

anterior short. The acetabulum is well in the anterior half of the

bone. The pubis has a regularly concave lateral border and a

straight medial border. The pubic plate is narrowest near its distal

end, but it is slightly broader at the extreme end. The appearance

of the pelvis has suggested to van Hoepen that there was a complete

longitudinal symphysis between the pubes and ischia of the two

sides of the animal.

The femur is pronouncedly sigmoidal in lateral view; the fourth

trochanter is in the upper half of the bone. The tibia has a very

large proximal end, and the tuberositas tibiae did not project very

far. There is no thickening at the distal end of the bone. Thre

hind feet are entire in the type. The fourth metatarsal is slightly

longer than metatarsal Il. Metatarsal III is more slender than

the latter.

Discussing the affinities of the type van Hoepen says: ,,The nearest

relations of our form are amongst the Plateosauridae and Anchisauridae.

and it is clear that it belongs to either one or the other. The

Plateosauridae ave all medium sized to large Dinosaurs with fifteen

dorsal vertebrae, whereas our form is small and has most probably

fourteen dorsal vertebrae, agreeing in this respect with the Anchi-

sauridae. There is further agreement with the Anchisauridae in the

relation of the lower arm to the humerus; radius and ulna are

longer than half the humerus. The length of the shaft of the humerus.

stands to the length of the whole humerus as 58:93 or 0°62. This

relation brings our form in close proximity of Thecodontosaurus

antiquus. Taking all these facts into consideration it seems clear

that our form is an Anchisaurid.

Comparison with Anchisaurus shows that the dorsal vertebrae are

comparatively longer, and that the pubes of the two forms differ

greatly. Massospondylus is a much larger form. The distal ends of

its ischia are coalesced, and each is more or less triangular in section.

In our form the distal ends of the ischia are flattened through

pressure, but it is difficult to accept that their section was originally

triangular. Moreover, they are not coalesced. The relations of the

ileum of Massospondylus carinatus are different from those in our

form, for it is longer than the latter with regard to its shortest

height above the acetabulum, Relatively the dorsal vertebrae of our

Fauna and Stratigraphy of the Stormberg Series. 381

form are longer than those of Massospondylus carinatus. The relations

of the lengths of the metatarsals in Massospondylus harriesi is diflerent

from that in our form. In Massospondylus harriesi metatarsal [1 is

longer than metatarsal IV, whereas in our form metatarsal II is

shorter than metatarsal IV.

In comparing with Ammosaurus and Gyposaurus I need only refer

to the great difference in the ilea.

The only other genus of the family is Thecodontosaurus. Super-

ficially there is great resemblance between our form and the known

species of Thecodontosaurus. A closer study, however, reveals remar-

kable differences.

A comparison of the ileum of our form with, that of Thecodonto-

saurus antiquus shows that in the latter the spina posterior is much

more produced. The acetabulum cuts deeper into the ileum of our

form, which resembles the Plateosauridae in this respect. The

highest point of the acetabular concavity is situated much nearer

towards the middle of the bone than in our form, and this is another

point of resemblance with the Plateosauridae. The ilium of our form

is manifestly different from that of Thecodontosaurus cylindrodon, and

also in the direction of the Platessauridae.

The pubis of our form differs considerably from that of Thecodonto-

saurus antiquus, as far as the latter is known. In our form the

lateral edge of the pubis is regularly concave, whereas in Thecodonto-

saurus antiquus its upper end is sigmoidal. There is also great

difference in the shape of the pubic foramina. The shape of the

proximal end of the ischium of Thecodontosaurus antiquus, as far as

preserved, is quite different from that of our form, a. difference

which is best understood from a comparison of the figures.

Another difference becomes conspicuous when the leneth of the

humerus is expressed in lengths of dorsal vertebrae. Taking one of

the hinder vertebrae v. Huene came to the following results: In

Thecodontosaurus antiquus the humerus is about five times as long

as the vertebra, and in Thecodontosaurus skirtopodus about four and

a half times. In our form the length of the eleventh dorsal vertebra

is 29 mm. The length of the right humerus is 93 mm.. which

means that the humerus is only 3:2 times as long as the vertebra.

Therefore, the humerus of our form is relatively much shorter than

that of Thecodontosaurus antiquus and of T. skirtopodus.

There is great difference between the ischium of Thecodontosaurus

minor and that of our form.

The points of difference enumerated above show sufficiently that

our form does not belong to any of the known genera of the Anchi-

382 Annals of the South African Museum.

sauridae. It, therefore, represents a new genus, for which I propose

the name Aristosaurus n. g. With the species Aristosaurus erectus n. sp.

Aristosaurus erectus is much more highly specialised than Thecodonto-

saurus, Ammosaurus, Anchisaurus and even than Massospondylus.

The build of the pelvis, and especially the position of the ischium,

shows adaptation to a usually bipedal mode of locomotion. The same

may be concluded from the far forward position of the acetabular

concavity in the ileum. The position. of the trochanter quartus

seems to be very low down on the femur. Its upper end is 41 mm.

from the proximal end of the bone. The length of the trochanter

is at least 18 mm. Therefore the lower end of the trochanter is

situated at more than 59 mm. from the proximal end of the bone,

which means very near to the middle of the femur. However, con-

clusions may not be drawn from this fact, because exact measure-

ments cannot be obtained.

The humerus is much shorter in relation to the body than in the

other Anchisauridae. The anterior extremity is also relatively, much

shorter in relation to the posterior one than in all other Anchisauridae

excepting Anchisaurus solus. As in the Plateosauridae the tibia of

Aristosaurus is much longer than the humerus. This is also the

case in Anchisaurus solus. In the other Anchisauridae it is the reverse.

All this tends to show that <Aristosaurus is an Anchisaurid, highly

specialised in the direction of the Plateosauridae, and of the bipedal

mode of locomotion.”

Type. In the Transvaal Museum.

Locality. Rosendal, Senekal Dist., Orange Free State.

Horizon. Bottom of Cave Sandstone.

Although not specifically identifiable, the two bones in the col-

lection of the Rhodesian Museum, Bulawayo, which were sent me

for inspection by the Director of the S. Rhodesia Geological Survey,

should be mentioned here as they are of interest in being the only

known fossils from the Forest Sandstone of that region.

One bone, from Dingaan farm, Bubi District, is the distal end of

a left fibula, measuring 120 mm. in length as preserved. The

dimensions of the distal articular surface approximate to those of

Thecodontosaurus skirtopodus (Seeley), but the bone differs from any

of the described fibulae of this family or of the Massospondylidae

or Plateosauridae in having a shaft subcircular in section, whereas

those of the described species are more oval in section.

The other is a complete dorsal centrum from Waterfall farm,

Bubi district. It differs from that of Gyposaurus africanus (Broom)

Fauna and Stratigraphy of the Stormberg Series. 383

in that it is as high as broad, The length is 43 mm, and the lower

border is moderately concave. The dimensions make it possible that

it belongs to the same species as the other specimen. The articular

surfaces for the arch are beautifully displayed. There is a fragment

of an arch showing a transverse process with a very concave under

surface and incomplete neck and caudal vertebrae together with

fragments of ribs.

Fam. MASSOSPONDYLIDAE von HueEne.

1914. von Huene. Fossilium Catalogus I, 4, p. 15.

MASSOSPONDYLUS CARINATUS Owen.

1854. Owen. Cat. Foss. Rept. Mus. R. Coll. Surgeons, p. 97.

1890. Lydekker. Cat. Foss. Rept. Amphib, Brit. Mus. IV, p. 246.

1895. Seeley. Ann. Mag. Nat. Hist. (6) vol. 45, p. 102.

4906. von Huene. Geol. u. Pal. Abh. N.F. Bd. VIII, hft. 2, p. 36,

Pls. XITI-X VI.

4911. Broom. Ann. §. Afric. Mus. VII, 4, p. 294.

The following details of the type specimen are taken from von

Huene’s account, checked by examination of casts of the type which

are in the South African Museum.

The cervical vertebrae are elongate, with a keeled under surface

as in Plateosaurus. The zygapophyses are long anteriorly, and

posteriorly are elongate with oblique articular surfaces. The neural

spine is short, low, and thin.

The centrum of the (1st dorsal vertebra is characterised by the

extraordinarily high thin keel on the ventral surface. A few of the

other dorsal vertebrae are known. They are compressed in the

middle and the ventral side is rounded.

The anterior caudals are very short and high, and broadly rounded

below. The later ones are provided with a keel below and carry

posteriorly a ventral groove, which divides partially the face for the

articulation of the haemapophysis. The middle and hinder caudals

are longer, but do not reach the length of the cervicals.

The scapula is small and slender and is characterised by a high

alar process at the distal end on the upper side; the coracoidal half

of this process is thin and concave from without, as in Plateosaurus,

but more strongly so; the medial side of the process is flat. The

distal end is broadened.

In the humerus the proximal end is broad, the upper border

obliquely bent down to the processus lateralis which is sharply cut

384 Annals of the South African Museum.

off from the ridge, but is not bent so-strongly forwards as in many

other Triassic Theropoda. The caput humeri lies on the medial angle

and is posteriorly thickened; the highest part of the upper border

is similarly thickened. The distal end is broad and has its condyles

directed strongly forwards.

The first metacarpal is an extraordinarily compact broad short

bone. The proximal surface is triangular.

The ilium has a sharp but short anterior process; the spina

posterior is long and moderately broad. The upper surface is bent

inwards in the middle. The ridge above the acetabulum is not so

roof-like as in many other genera. The crista interior on the inner

side of the posterior process is not very prominent.

The middle and the distal end of the pubis is broadened and thin,

the lateral and distal edges being thickened.

The femur is slender and lightly bent from front to back. The

tibia has its distal end compressed from front to back, while the

proximal end is convex medially and has a large tuberosity laterally.

The anterior process is somewhat bluntly rounded. On each side

of the tuberosity the lateral edge is hollowed out.

Von Huene separates the genus from Plateosaurus on account of

its Thecodontosaurus-like tibia. The scapula, ilium, pubis, femur and

hand are more strongly built than in Thecodontosaurus.

Type. Isolated bones in British Museum.

Locality. Beaucherf, Harrismith, O.F.S.

Horizon. Red Beds.

MASSOSPONDYLUS HARRIESI Broom.

Text figs, 21-29.

4911. Broom. Ann. S. Afric. Mus. VII, 4. p. 299. Plates XV—XVII.

1920. van Hoepen. Massopondylus browni. Ann. Transyv. Mus. VII, 2.

p. 148. Plates XVII-XXIII.

In his original description Broom says ‘This Dinosaur resembles

Massospondylus carinatus sufficiently closely to suggest the advisability

of placing it at least provisionally in the same genus. The remains

consist of an imperfect humerus, a nearly perfect radius and ulna,

and a perfect manus, as well as portions of the femur and tibia,

and a number of toe bones all of one individual, and the perfect

pes of another individual.”

Re-examination shows that the remains supposed to constitute the

first individual really contain two individuals. The supposed femur

is in reality the distal half of a humerus, larger than that belonging

Fauna and Stratigraphy of the Stormberg Series. 385

to the complete fore-arm; and doubt is thus thrown upon the identity

of the tibia and bones of the hind foot with the forearm, although

all are marked with Mr. Walker’s collecting number “I”. Certainly the

supposed femur is no part of the type animal, and must be considered

separately.

The pes described by Broom is admittedly part of another individual ;

but comparison of it with a few portions of a pes marked “IP” and

found with the fore-limb renders it probable that it belongs to the

same species as the latter. I consider, however, the fore-limb as

the type of the species.

Comparison with M. carinatus is confined to examination of the

distal end of the humerus, the 1st. metacarpal, and the proximal end

of the tibia. Broom has figured the 1st. metacarpal, displaying its

general similarity with that of M. cavinatus, the inner distal condyle

of each being small compared with forms such as Gryponyxr. The

distal end of the humerus is somewhat distorted, but the condyles

seem to bear a similar relation to one another to those of M. carinatus.

The tibia approximates more closely in size to that of M. carinatus

than does the fore-limb, and probably belongs to a larger individual,

possibly the same as that from which the larger humerus (described

as femur) belongs. The proximal end of the tibia is slightly longer

and considerably thicker in M. carinatus; the greater thickness being

mainly due to the more prominent development of the tuberosity on

the lateral face. The anterior process is similar in each. The

posterior border of the bone is more concave proximally in M, carinatus

than in M. harriesi.

The larger humerus consists of the distal portion from the condyles

to the lower end of the deltoid crest. In shape and general pro-

portions it corresponds closely with the humerus of the type fore-

limb, the distal end being crushed in the same direction as the type.

The width of the distal end as preserved is 114 mm. There is a

broad shallow groove between the condyles on the posterior face,

and a deeper excavation on the anterior face of the bone. Its size

corresponds roughly to that of the humerus of M. carinatus,

The right pes has been fully described and figured by Broom. In

size it stands much closer to the fragmentary remains of M. carinatus

than does the fore-limb, and is thus of a larger individual which

shows few features distinguishing it from M. carinatus.

Whilst collecting in the Herschel Division of the Cape Province

I was fortunate enough to obtain from the hill overlooking the

Blikana Trading Store (and thus from an horizon in the upper third

of the Red Beds) an almost complete specimen of an animal which

386 Annals of the South African Museum.

can most easily be referred to this species. The caudal region had

weathered off and was not obtained; but the remainder of the

skeleton was found lying articulated with the exception of the head,

which was at a distance of about 3 feet from the front ofthe neck.

The specimen is one of the most complete known from the Stormberg

Beds and has thus been thought worthy of a fairly full description,

as it throws light upon the real position of these medium-sized forms.

The specimen is in the collection of the South African Museum —

Catalogue no, 5135.

Skull. The skull was lying detached from the neck at a distance

of some 3 feet from it and was, unfortunately, shattered by a blow

from a pick before its presence was realised. It has been possible,

Meee

Fig. 22. Massospondylus harriesi, Br.

Restoration of a side view of skull (5135).

(The teeth are omitted from this restoration). X 4.

however, to reconstruct several portions of it from the fragments

collected.

The basicranial region exhibits strong likenesses to that of T’heco-

dontosaurus antiquus and is also fairly close to Sphenosuchus acutus.

The occipital condyle is rounded and intermediate in size between

that of Sphenosuchus and of Thecodontosaurus. It is formed almost

wholly of the basioccipital, the suture between that bone and the

exoccipital running as in Sphenosuchus. Anterior to the condyle the

bone thins and then expands rapidly, its lower surface at the same

time curving strongly downwards to form a long transverse basi-

occipital ridge. This ridge is not furnished with a median notch

like that of Sphenosuchus. Its upper surface forms the floor of the

brain-case for some considerable distance, the suture with the

exoccipital being plaimly seen on each side. The anterior portion

of the bone is furnished with a median medullary ridge.

Fauna and Stratigraphy of the Stormberg Series. 387

The suture between the basioccipital and basisphenoid is not

traceable, but I am inclined to believe that it runs along the trans-

verse ridge. If this be so, the structure is diflerent from that of

Sphenosuchus, where the suture runs along either side of the inter-

tympanic foramina which lie wholly in the basioccipital, and agrees

with that in Plateosaurus and Thecodontosaurus. 'The median ex-

cavation is smaller than in Sphenosuchus. The distance between the

hinder end of the basisphenoid and the pterygoid apophyses of the

bone is comparatively greater than in Thecodontosaurus. These

apophyses are as in the European genus. Seen from above the

anterior portions of the two apophyses meet superiorly to form a

crest in advance of the circular hole for the hypophysis cerebri

(pituitary fossa). The basisphenoid is furnished laterally just behind

the root of the pterygoid apophysis with a deep groove corresponding

in position to the recessus basisphenoidei of T’hecodontosaurus figured

by von Huene. The bottom of this pit is presumably pierced by the

carotid foramen opening into the pituitary pit. Superiorly the relation

of the basisphenoid with the bones of the side-wall of the brain

case are obscure.

The foramen ovalis is only presented in section on the left side.

It occupies the same position as in Thecodontosaurus, perforating the

side-wall of the brain behind the sella turcica.

The exoccipital is of the form seen in Thecodontosaurus. Its lower

border is grooved proximally, the groove running inwards and forwards

to the foramen lacerum for the exit of nerves IX—-XI. This opens

into the bottom of the side-wall of the brain-case. Separated from

this foramen by a thin plate of bone is a larger opening, the foramen

jugulare for the passage of the VIIIth nerve. Mesial to the internal

opening of this foramen is a shallow pit in the brain case. The

opening for the XIIth nerve is only seen in vertical section passing

through the exoccipital above the level of the condyle. Behind and

above the foramen lacerum the side wall of the brain is furnished

with two small foramina, presumably venous.

The exoccipital articulates with the basioccipital below and the

supraoccipital above. It has a strong paroccipital process.

Save in details of relative sizes, this basicranium is similar to that

of Thecodontosaurus, and differs in certain features more from

Sphenosuchus.

A portion of the right side of the skull and lower jaw is preserved,

attached to a part of the top of the skull. The snout is missing.

The orbit was large, its length as preserved being 54 mm. Its

anterior border is formed by the pillar-like lachrymal which divides

388 Annals of the South African Museum.

the orbit from the antorbital vacuity. The limits of the prefrontal

are doubtful, but it seems to be a small bone wedged between the

lachrymal and frontal and extending forwards over the antorbital

vacuity.

The frontal is a large bone meeting its neighbour in the middle

line to form with it the whole of the interorbital space. It forms

the whole of the upper border of the orbit and extends back to

form part of the border of the anterior temporal vacuity. The

postfrontal, if present, as a separate bone, is small and forms no

part in the formation of the orbit although it may be part of the

anterior border of the upper temporal vacuity. There is some doubt

as to its separate identity and [I am inclined to consider it as fused

with the frontal as in Sphenosuchus.

The jugal forms the lower border of the orbit and meets the

lachrymal and maxilla anteriorly. The maxilla, as preserved, carries

4 or 5 flattened teeth. The teeth are seen better, however, in the

lower jaw.

A length of 110 mm. of the lower jaw is present, the front missing.

From the post-articular process the upper border rises in a regular

high curve, concave at first and then convex, to a point below the

middle of the orbit, whence it passes downwards and forwards.

The lower border is almost straight. The teeth carried by the dentary

are variable in size, large and small teeth apparently alternating.

The teeth are flattened with the upper halves of the anterior and

posterior borders serrated, as in Thecodontosaurus. The largest tooth

seen is 7 mm. long and 3°5 mm. broad. There are 7 serrations in

a distance of 4mm. The cross-section of the tooth is an elongate oval.

Vertebrae. The cervical vertebrae are somewhat crushed and

incomplete. The later cervicals agree closely with those of Masso-

spondylus carinatus. The middle cervicals are very elongate, compressed

in the middle, with a median ventral keel in the anterior half. The

zygapophyses are long with very oblique articular surfaces. The

neural spine is thin, low and long. The transverse process is in the

form of a long ridge at the level of the top of the centrum. The

whole body of the bone is slightly curved. The most complete bone

gives the following measurements :

Length of centrum . ; : : 105 mm.

Height of centrum . ‘ ; ; 40 mm.

Greatest length : : : ; 133 mm.

Max. height (probable) . ; : 85 mm.

What is probably the last cervical has a centrum 73 mm. long

Fauna and Stratigraphy of the Stormberg Series. 389

and 44 mm. high. The centrum is strongly compressed laterally

and has a very sharp median ventral keel. The side of the centrum

is furnished with a prominent parapophysis placed just behind the

middle of the bone for the articulation with the rib. The anterior

Fig. 23. Massospondylus harriesi, Br.

Left scapula and coracoid (5135).

A. External view. B. Ventral view.

cervicals have very shallow and elongate centra and comparatively

massive processes. They are, with the exception of what is probably

the first, very elongate; but the preservation is not sufficiently good

to admit of full description. The anterior cervicals are very bird-like

in general appearance.

390 Annals of the South African Museum.

Shoulder Girdle. Both scapulae are preserved, but are somewhat

incomplete at the distal end and each lacks the end of the supra-

coracoidal wing. The proximal end is very similar to that of the

smaller scapula assigned by von Huene to Massospondylus carinatus.

The glenoid cavity is the same size, the posterior half of the articular

surface for the coracoid slightly larger and the shaft of the bone a

little broader. The distal end expands fairly considerably, but not

so much as in Sphenosuchus acutus.

The chief measurements are:

Probable greatest length . 5 : 280 mm.

Greatest width . : : : , 104 mim.

Minimum width of shaft . : 5 40 mm.

Greatest thickness at articulation with

coracoid . : : 30 mm. (left) 43 mm. (right)

The left coracoid is almost entire, the right one represented by

the posterior half. The inner surface is concave, the outer slightly

convex except for a strong thickening in’the middle of the lower

portion of the bone. This thickening takes the form of a strong

short longitudinal swelling which rapidly subsides into the bone at

its upper and lower ends. This thickening is for the reception of

the coraco-brachialis muscle, according to von Huene. In front of

the coracoidal portion of the glenoid cavity the border is concave

to the anterior ventral angle. The anterior border is regularly curved.

The left coracoid is 135 mm. broad, 80 mm. high, and has a maximum

thickness at the scapular surface of 30 mm.

Attached to the upper portion of the anterior border of the left

coracoid is a small piece of bone lying generally in a plane at right

angles to that of the coracoid and bent in a convex manner when

viewed from in front. In collecting the shoulder girdle a number

of pieces of thin bone were found in situ in the region of the two

coracoids. These have been fitted together as much as possible and

seem to give indisputable evidence of the presence of two clavicles

and an interclavicle. These bones are too fragmentary to permit of

full description. The largest fragment is a piece which I take to

be the interclavicle. As preserved it is 86 mm. long. The bone is

a thin plate broadening somewhat posteriorly, and possibly anteriorly,

slightly convex from side to side ventrally save in the anterior portion

where there is a prominent longitudinal swelling for the attachment

of muscles, similar in form to that on the coracoids. The dorsal

surface is flat, so that the edges of the bone are thin. Lying at the

Fauna and Stratigraphy of the Stormberg Series. oot

anterior end of the dorsal surface is a fragment of a clavicle, which

here at its distal end is a thin plate of bone.

Humerus. The left humerus is slightly larger than the right and

has the deltoid portion bent into the body of the bone slightly more

than the other. Both humeri are complete and are of the general

type seen in the Vienna specimen of Thecodontosaurus skirtopodus,

although the bottom of the deltoid crest is just below the middle of

the bone. The deltoid crest is sharply cut off from the lateral edge

Fig. 24. Massospondylus harriesi, Br.

Right humerus (5145).

A. Anterior view. B. Medial view. <a:

of the proximal portion by an abrupt change in the outline of the

bone; and the process is but slightly bent from the general plane

of the upper half of the bone.

The outer edge of the proximal end is strongly thickened to form

a rounded knob about 385 mm, from the medial angle.

The medial edge of the bone forms a regular sweeping concave

curve from the caput humeri to the inner condyle. The distal end

is strongly broadened, and on the inner face there is a shallow

depression between the two condyles. Anteriorly there is a medial

392 Annals of the South African Museum.

ridge which bounds one side of a shallow hollow lying above the

inner condyle. The outer condyle is slightly larger compared with

the inner than in the type of the species.

The chief measurements are:

Right humerus Left humerus

Greatest length . ; : = 219) mom, 223 mm.

Breadth at distal end ’ : Dias 18 + 5,

Width at narrowest part of shaft DOM N: DON

Width at top of deltoid crest . 305 ,; a. Se

Greatest width of proximal end oe Si”,

From lower end of deltoid crest

to furthest part of distal end 108 ,, LOe es

Length of deltoid crest. : DDC, 3o) eae

Thickness of head. ; ‘ 163% a A ee

Max. thickness of proximal end Py ares PAS) =

Thickness at inner condyle ‘ DON cee 24 ,,

Thickness at outer condyle F 24 =,, 26);

Radius. Both radii are preserved, the left bone being crushed

proximally. The right is slightly shorter than the left, but each

has the same general form. The right radius is 132 mm. long, the

left 155 mm., each being considerably shorter than the ulna.

The proximal articular surface is saddle-shaped. Below it the

bone rapidly thins, so that the whole bone is a straight cylindrical

shaft with expanded ends. The anterior edge is provided with a

rounded ridge. At the distal end the posterior-medial face has two

knobs, presumably for muscle articulation, a larger medial one and

a smaller posterior one. At the proximal end the medial groove for

the insertion of the humero-radialis muscle is very prominent.

Breadth of proximal surface. : 06 nim.

Thickness of proximal surface . : 2m

Breadth of distal end : : ? By Oe

Thickness at distal end. : ; 23 ,,

Minimum width of shaft . : : 16

Uina. Both ulnae are preserved, but the left is somewhat im-

perfect. The right gives the following measurements:

Greatest length. : : : - 152 mm.

Width at proximal end . : : 02

Width at distal end. : . : 40

Fauna and Stratigraphy of the Slormberg Series. 393

Width at narrowest part of shaft. Alf isourn

Thickness of proximal end : : ol 5,

Thickness of distal end . : ‘ > Opaeee

The distal quarter of the bone is twisted to the left as in the

type. The bone is more elongate than that of Plateosaurus quenstedti

and the proximal end is more expanded. The proximal half is

somewhat similar to that of Plateosaurus erlenbergensis, but the

articular surface slopes more strongly and the lateral angle lies

nearer the anterior point. The medial border of the proximal end

is concave and below it the surface of the bone is hollowed out.

The anterior border of the bone is concave, the posterior border

straight except at the distal end, where it curves outwards and

slightly backwards.

Fig. 25. Massospondylus harriest, Br.

Right ulna (5135). X 4.

Hand. On the right side there are preserved the three carpale,

the dst. metacarpal and the whole of the (st. digit. Of the left

hand there are the 1st. and 2nd. carpale, all the metacarpals, and

the 1st. digit. In addition there is an isolated claw of the right hand.

The 1st. carpale articulates distally with the dst. metacarpal and

the 2nd. carpale. It is an irregularly oval-shaped thin bone, with

a maximum length measured from the mediai end to the lateral end

of 39 mm. in the left hand and 36 mm. in the right. The thick-

ness between the dorsal and ventral edges is 25 mm. (24 mm. on

right side). Between the proximal and distal surfaces the distance

is 143 mm. (142 mm. on right).

394 Annals of the South African Museum.

The 2nd carpale is a small bone, articulating proximally for most

of its length with the 1st carpale, distally with the 2nd metacarpal,

and medially ‘with the 1st metacarpal. Its dorsal surface is small,

its palmar face rectangular with an area of 23 mm. X 12 mm.

(18 x 13 on right). The distance between the dorsal and palmar

surfaces is 19 mm. (46 mm. right).

Fig. 26. Massospondylus harriesi, Br.

Left pubis (5135).

Ventral view. X 3.

The 14st metacarpal has a maximum length of 42 mm., a proximal

width of 38 mm., and a distal width of 35 mm. The proximal end

is almost triangular in shape, the ulnar edge having a length of

25 mm. In shape it compares closely with that of Massospondylus

carinaius, but the distal articular surface is thicker than in the

larger form. Also the ulnar proximal edge of the dorsal surface is

pointed and not flattened as in M. carinatus.

The 2nd metacarpal has only the proximal half remaiming. Its

Fauna and Stratigraphy of the Stormberg Series. 395

end is 20 mm. broad and 24 mm. thick. On the radial side it has

dorsal and palmar knobs with a hollow between, and on the ulnar

side there is also a groove for muscle insertion. The dorsal side is

slightly hollowed, but the palmar side is flat. The shaft of the bone

is slender and circular.

The 3rd metacarpal is 53 mm. long, 29 mm. broad proximally,

and 21 mm. broad distally, and the thinnest part of the shaft has

a breadth of 143 mm. For the 4th and 5th metacarpals the corres-

ponding measurements are 42 mm., 20 mm., 16 mm. and 9 mm.,

and 28, 18, 13, and 41 mm. respectively.

In the dst digit the 1st phalanx is 37 mm. long. Its proximal

width is 31 mm., its distal width 23 mm. It is so twisted that the

claw curves strongly inwards and but slightly downwards. The claw

has a length as preserved of 65 mm., and when complete was

probably 75 mm. long. It is strongly curved and compressed dorso-

ventrally. The greatest breadth at the proximal end is 36 mm. and

the thickness, which is greatest on the radial concave side of the

claw, 24 mm. There is a large bony process on the radial side

at the proximal end for the insertion of the flexor tendon.

Pubis. Both pubes are preserved, entire save for the ischial

articular surfaces. The head of the right pubis is somewhat distorted

and the bone has a maximum length of 270 mm. The maximum

length of the other bone is 285 mm. The breadth at the distal end

is about 50 mm. and the thickness 23 mm.; but the breadth decreases

to 43 mm. and the thickness to 12 mm. in the middle of the bone.

The medial portion is very thin; the lateral border fairly sharply

rounded. At a distance of 240 mm. from the distal end the public

plate is turned abruptly downwards at right angles in its medial

portion while the lateral part, now forming the pubic neck, is curved

more gently upwards and outwards. The subacetabular process ts

50 mm. long and 30 mm. broad, The pubic foramen is large.

The pubis is seen to be longer than was supposed by von Huene

for Massospondylus carinatus. The distal end is somewhat thicker

than in the Plateosauridae.

Femur. The femur fairly slender and slightly bent backwards at

its distal end. The proximal end is bent strongly inwards, more so

than in M. carinatus. The trochanter minor is not defined, but the

trochanter major stands out as strong, somewhat curved, crest on

the anterior face of the bone. The chief measurements are:

Greatest length. : ; : . doo mm.

Width at head . ; ‘ 5 : 79

396 Annals of the South African Museum.

Thickness at head. : : ; 32. mm.

Distance from head to top of trochanter

major : : : : DD,

Width at top of trochanter major, Tile Fy,

Fig. 27. Massospondylus harriesi, Br.

Right femur (5135). x 4.

A. Anterior view. B. Medial view.

The 4th trochanter is a fairly short high thin crest on the posterior

face, lying well within the upper half of the bone.

Fauna and Stratigraphy of the Stormberg Series. 397

Distance from head to top of troch-

anter IV ; : . ; ; 88 mm.

Distance from head to bottom troch-

anter IV ; : : 5 Be OAS

The distal end of the bone is bent backwards and swells considerably

to form the condyles. The lateral condyle is narrow and turns

. 28. Massospondylus harriesi, Br. (5135).

Right tibia lateral view.

- , outline of proximal end

Left tibia outline of proximal end X 4

ab pe

outwards, and from it a sharply rounded ridge runs up the bone a

distance of about 80 mm. The inner condyle is broader and less deep,

and stands higher than the lateral condyle. Between the two is a

narrow, deep groove, narrower than that of M. carinatus. The

articular surface is inclined at an angle of about 70° to the anterior

398 Annals of the South African Museum.

slope of the bone. On the anterior face there is a short median

depression above the condyles.

Maximum thickness at distal end. 79 mm.

Thickness of lateral condyle. ‘ 2055

Thickness of medial condyle — . : SOD.

Depth of lateral condyle . ; : Nd? 56

Depth of medial condyle . : : O° %,;

Tibia. ‘The right and left tibiae are complete. ‘The proximal portion

of the right has been flattened laterally, so that the head is con-

siderably narrower than that of the left. The outline of the head

of the bone differs in the two. The right tibia has a concavity both

before and behind the lateral tuberosity as in Massospondylus carinatus ;

the left tibia has the anterior concavity, but behind the tuberosity

the edge is regularly convex to the posterior point. It can thus be

seen that considerable care must be taken in distinguishing forms

by such features as the shape of the tibial head, which is variable

even in an individual.

The general shape of the proximal end approximates closely to

that of M. carinatus and needs no detailed description. As a whole

the bone is slender with the distal end but slightly expanded. The

anterior condyle lies above the posterior and looks both outwards

and downwards. The chief measurements are:

Right Left

Greatest length . 3 . 900 mm. 298 mm.

Length of proximal end . 100 ,, oi,

Width of proximal end. SOM Gla

Minimum width of shaft . 24, 23 eae

Minimum thickness of shaft 348) op Down

Length anterior border,

distal end E ; 5 DO Opa

Length posterior border,

distal end- . : : 4D ,, AV,

Max. thickness, distal end 4D, oT

Fibula. The right fibula and most of the left are preserved. The

bone is slender, flattened laterally, slightly expanded at the end and

almost straight. The inner face at the proximal end is hollowed out

with a well-marked sharp crest bounding it anteriorly. In the mid-

part of the bone the medial face has a slight longitudinal ridge

anteriorly. At the distal end the anterior and posterior faces are

Fauna and Stratigraphy of the Stormberg Series. 399

flattened and the lateral edge is broadly rounded. The lateral side

has a strong longitudinal crest running for a short distance up from

the condyle. In rear of it is a shallow groove, bounded posteriorly

by another short prominent ridge. The distal articular surface is

convex.

Fig. 29. Massospondylus harriest, Br. (5135).

A. Right fibula, lateral view.

B. Right astragalus. Front view.

C} re sf . Top view.

All X#.

Right Left

Total length of bone. - 290 mm, =

Length of proximal surface ASt 46 mm.

Width of proximal surface QE 30 t

Minimum width of shaft . 41h AS

Length of distal surface. 42 ,, Lo

Width of distal surface. 29 21

400 Annals of the South African Museum.

Hind Foot. The right hind foot was found in situ, while most of

the bones of the left foot were recovered from the talus below the site.

Astragalus. — The astragalus is preserved on both sides of the body.

The. maximum width from side to side is 72 mm. in the right bone

and 76 mm. in the left. The maximum thickness in the medial half

is 43 mim. (left, 40 mm.) and the maximum height in that half 34 mm.

(left, 30 mm.) The lateral portion shows a high flattened upper

portion for articulation with the anterior distal condyle of the tibia

and posteriorly a low elongate narrow surface. Below the anterior

articular face the bone is undercut posteriorly. The lower anterior

angle is a short rounded prolongation, the posterior corner being

broadly rounded. The lateral face shows a small shallow concavity

for articulation with the calcaneum.

The thickness at the lateral end is 31 mm. (left, 27 mm.), the

height 37 mm. (left, 57 mm.), and the breadth of the anterior upper

surface 17 mm.

The under surface is broadly convex with a faint median ridge

running from left to right.

Caleaneum. — A bone which | take to be the calcaneum is preserved

in connection with the right foot. It is roughly triangular in outline,

with a small lateral protuberance. Its upper surface is concave, its

lower convex. Its greatest width is 36 mm. and its maximum thick-

ness 24 mm.

Tarsalia, — The only other bone of the tarsus preserved is

tarsale III, which is found in both feet in articulation with the third

metatarsal. In section it is triangular, and its upper surface slopes

from the back inwards and forwards so that it is thickest in its

posterior and lateral parts. Its under surface is hollowed with a

short medial anterior peg to fit closely to the proximal surface of the

metatarsal. Its anterior edge is 28 mm. long, its medial face 31 mm.,

and its other face 41 mm. Its greatest thickness is 20 mm.

The following table gives the chief measurements of the metatarsals

and phalanges of the right foot. The 3rd. digit is missing ; it was

not preserved with the remainder of the animal, and was possibly

lost during life.

Length. Prox. breadth. Distal breadth.

Metatarsal I 86 mm. 33 mm. 30 mm.

Digit 4. ist phalanx 51 _s,, 29%, 23 55

claw IPE es DOP is. --

Metatarsal IT AD See _- —

Digit 2, dst phalanx 48 ,, 36s, BDA

Fauna and Stratigraphy of the Stormberg Series. 401

Length. Prox. breadth. Distal breadth.

2nd phalanx 36 mm. 29 mm. 25 mim.

claw G15. 24. —

Metatarsal IIT. VAD 305" ,5 Some

Metatarsal IV. AD ates 9 ea hey te

Digit 4. 1st phalanx 41 ,, SON as 260.5,

2nd phalanx 36 _,, 29 25° 5

3rd phalanx 27 _,, Alain Oey?

4th phalanx 23. ,, DAO as AO ls:

claw Lh ae AS? —

Metatarsal V. Cie BN] 33 —

Digit 5. 1st phalanx 17 ,, ee _—

Comparison of these with the type pes of Massospondylus harriesi

shows that the phalanges of the digits have almost indentical measure-

ments in the two animals. The metatarsals in this animal have

different lengths, however, those of M. harriesi being somewhat longer,

except in metatarsal [.

The metatarsals show few distinguishing features. The proximal

end of metatarsal Il has the plantar edge considerably longer than

the dorsal, while the medial and lateral faces are both hollowed out

for articulation with the neighbouring bones. Metatarsal I looks almost

entirely inwards. Metatarsal [V is a much flattened bone, and its

distal end is slightly twisted inwards. Metatarsal V thins rapidly at

its proximal end and from the lateral face, and hes almost entirely

behind metatarsal IV. Its distal half shows the usual pads for the

reception of tendons or muscles, especially on the plantar surface.

Among the collection of the Durban Museum submitted to me is

an incomplete animal from Foutanie, Fouriesburg, O.F.S. (the type

locality) consisting of a right scapula, a right humerus, a left coracoid,

a left fibula and digits 1, 2, 3, of a left pes. These closely correspond

with the type and must be placed within the species. The right

scapula has the ventral edge a little straighter than in the Blikana

specimen, being thus closer to the specimen described by van Hoepen

as Massospondylus browni. The lett fibula has a maximum length of

about 285 mm. and a proximal surface 43 mm. long, and 20 mm.

wide. The bone is flattened laterally, especially at the distal end.

Its inner face at the proximal end is concave, its outer face convex.

The proximal end is slightly more slender than in the Blikana specimen.

The digits of the left pes correspond very closely to the type in size.

It seems to me that van Hoepen is probably in error in assigning

402 Annals of the South African Museum.

the remains described by him from St. Fort, Bethlehem, O.F.S. to

Massospondylus browni. The only possible comparison that can be

made is between the femora. and it is apparent that, not only is the

proximal end of the type of M. browni (.e. Thecodontosaurus browni)

stouter than in van Hoepen’s specimen but the trochanter quartus is

nearer the proximal end. Moreover, the type is a considerably smaller

animal, and has been assigned by von Huene, whose judgment cannot

lightly be set aside, to the genus YVhecodontosaurus. There can be

no doubt that Dr. van Hoepen’s species belongs to the genus Masso-

spondylus, and the question arises whether it is a specimen of M.

harriesi to which it closely approximates in size.

Comparison can first be made with the almost complete specimen

of M. harriesi from Blikana. This shows that the humerus is almost

exactly the same shape, and very slightly smaller; the ulna differs

in that the Cape specimen has a slightly more slender shaft and has

a posterior prolongation at the proximal end, which may have been

worn off in van Hoepen’s specimen; the radius of the former has

a more slender shaft, swelling somewhat more abruptly at the ends;

the scapula and coracoid are very similar, save that the distal end

of the St. Fort specimen is apparently slightly narrower, thus

coming closer to the type; the femur of the Blikana specimen is

more strongly bent in anterior view and straighter in lateral view

— but the St. Fort animal is considerably crushed; the tibia and

fibula both show slight differences, and the bones of the feet are

slightly more slender and longer in the St. Fort animal.

When compared with the type of M. harriesi, however, the differ-

ences in the foot are not so great; and in view of the fact that

van Hoepen’s specimen in closer to the type than is the South

African Museum animal which I have referred to M. harriesi, I

cannot refrain from including it in the same species. Four speci-

mens of M. harriesi are thus known — the type, the South African

Museum animal, the bones described by Dr. van Hoepen, and a

partial skeleton in the Durban Museum. Of these, the first and last

are from the one locality, the third from a closely-neighbouring area,

and the other from some distance away and a slightly lower hori-

zon. From them we can see the amount of individual variation

that may occur within one species; and such variation should be

taken into account when the identification of forms is under con-

sideration.

Type. Bones of fore limbs — 8. A. Mus. Cat. No. 3394,

Locality, Foutanie, Fouriesburg, O. F.S.

Horizon. Top of Red Beds.

Fauna and Stratigraphy of the Stormberg Series. 403

MASSOSPONDYLUS SCHWARZI, Sp. nov.

The bones upon which this species is founded were unearthed

by Professor Schwarz at Makomoreng, Mount Fletcher, C. P. They

are in the collection of the South African Museum (Cat. No. 5134).

The species differs from M. harriest and Aetonyx palustris in the

comparative size of the metatarsals. Taking the first metatarsal as

a standard, the second and third are longer in this form than in

the other species, in this respect the foot approximating most to

M. harriesi. In addition, the phalanges of the digits are relatively

longer than in the other forms, those of the third digit being actually

as large as in M. harriesi which has a third metatarsal 20 mm. longer

than in this new form.

These differences seem sufficient to justify the provisional erection

of a new species for these remains; they certainly cannot be placed

in any described species until we know more concerning the limits

of individual variation among these Dinosaurs. The distal end of the

tibia is somewhat more swollen than in MM. carinatus, but as the foot

bonse seem to approximate more closely to Massospondylus than to

Aetonyx, 1 shall provisionally name the form Massospondylus schwarzi.

The following table gives the chief measurements of the bones of

the foot:

Greatest length. Proximal width. Distal width.

Metatarsal I 71 mm. 33 mm. 30 mm.

dst phalanx 41 ,, SA eee 23,

claw imperfect 19; —

Metatarsal IT 418 mm. prob 25_,, 33,

dst phalanx 47 ,, Ss; Die

2nd phalanx 31 ,, 2A, 1S 5e NER

claw imperfect AS —

Metatarsal IIT 135 mm. 22) -:, aH os

dst phalanx 47 ,, Hs oe Dt ass

2nd phalanx 34 ,, 23.5 AO) ee

3rd phalanx 30 ,, oe aS eer

claw imperfect AM) —

Associated with the foot are some portions of the leg and sacrum

which are too imperfect for accurate determination. :

Type. Incomplete pes and distal end of tibia. S.A. Mus. Cat.

No. 5134,

Locality. Makomoreng, Mount Fletcher, C.P.

Horizon. Red Beds.

4O4 Annals of the South African Museum.

AETONYX PALUSTRIS Broom.

Text fig. 30.

4911. Broom. Ann. 8. Afric. Mus. VII, 4. p. 304. Pls. XV and XVII.

The type consists of ‘ta few imperfect dorsal vertebrae, a good

scapula and coracoid, a good humerus, a good radius and imperfect

ulna, the greater part of each manus, the upper end of one tibia,

and the almost complete right pes”.

A little extra description can be added to that given by Broom.

There are portions of three cervical vertebrae attached to one another,

of which the middle one is almost complete. The centrum has a

leneth of 70 mm. and a height at the articular surface of about

Fig. 30. <Actonyx palustris, Br.

Right Metatarsal. I. (Durban Mus.).

Distal end, front view.

35 mm. The maximum height of the bone is 68 mm. As in Mas-

sospondylus the centrum is elongate and compressed with a concave

ventral surface; but the median ventral keel is not nearly so prominent

as in the larger form. The neural spine is slightly higher than in

M. carinatus, but is of the same type. The zygapophyses, as far as

they can be seen, agree with those of Massospondylus. The transverse

processes are small.

Several vertebrae from the posterior part of the tail are preserved,

seven of them in association, and others now isolated but forming,

according to Mr. Walker’s field numbers, a series of 8. The former

are more complete than the latter.

The length of the centrum in the first series diminishes from 25 mm.

to 23 mm. with the last having a length of 20 mm. The maximum

ya)

height of the largest bone is 22 mm., and of the centrum 11 mm.

Fauna and Stratigraphy of the Stormberg Series. 405

All are slightly expanded at the ends, with the ventral surfaces

rounded. They call for little comment, being of the type seen in

Sellosaurus fraasi, figured by von Huene, with the exception that the

zygapophyses stand somewhat higher from the centrum in Aetonyx.

The dorsal spine is very small.

In the other series the centra only are preserved. They vary in

length from 33 mm. to 26 mm., and are thus anterior to those

already described. The ventral surface of the centrum is rounded

but in its posterior half it is somewhat flattened with two short

incipient keels flanking the flattened portion. These keels arise from

the small oblique ventral face of the hinder end, which carried a

small chevron.

Type. Partial skeleton, S.A.Mus. Cat. No. 2768, 2769, 2770.

Locality. Foutanie, Fouriesbure, O. F.S.

Horizon. Top of the Red Beds, Stormberg Series.

There are in the Durban Museum collection several bones which

I ascribe to a large specimen of this species. They are from Fou-

tanie, Fouriesburg, but it is uncertain whether all belong to the

one animal although the probability les im that direction. They

consist of metatarsals I-III of the right side, metatarsal I of the

left side, the distal end of a fibula and the distal end of a tibia

the latter somewhat doubtfully assigned to this species.

The metatarsals of the right foot are approximately of equal

length with those of the type Massospondylus harriesi; but they are

more slender. The 1st metatarsal differs from that of Massospondylus

in having a narrower proximal end, the medial upper edge beimg

more convex, and in the slope of the distal end beimg much more

oblique with the lateral portion of the articular surface very much

swollen. The total length of the left metatarsal I is 82 mm.

The distal end of the fibula differs from that of M. harriesi in

having a more flattened shaft and less prominent ridges at the

distal end on the posterior face.

DROMICOSAURUS GRACILIS v. Hoepen.

1920. van Hoepen. Ann. Transv. Mus. VII, 2. p.103. Pls. XITI—XVI.

This is a form slightly larger than Massospondylus carinatus, but

closely allied to Massospondylus and Aetonyx. The type consists of

fragments of humerus and radius, a fairly complete cervical vertebra,

406 Annals of the South African Museum.

some caudal vertebrae, the pubes, the ischia, a femur, a tibia, a fibula

and some foodt-bones.

Van Hoepen has pointed out certain differences between this and

previously described forms. The outstanding features seem to be

judging from the description and figures given, the straightness of

the femur and the great height of the anterior end of the proximal

surface of the tibia. Further, the lateral edge of the pubis is much

straighter than in Massospondylus harriesi or in Gryponyx africanus,

the only known South African forms of similar size in which it is

satisfactorily preserved.

In connection with the original description and comparison of this

form with Massospondylus harriesi it should be noted that the tibia

and ‘femur? described by Broom, upon which van Hoepen based

his comparisons, are probably not bones of the type at all, as pointed

out in my re-description of M. harriesi in the present paper. Never-

theless, the tibiae of the two forms do vary in shape, the head of

that of D. gracilis being much more inclined to the axis of the shaft

than that of M. harriesi.

Type. Partial skeleton in the Transvaal Museum.

Locality. Naauwpoort Nek, Bethlehem, Orange Free State.

Horizon. Red Beds.

Fam. PLATEOSAURIDAE von HuEne.

PLATEOSAURUS STORMBERGENSIS Broom.

4915. Broom. Bull. Amer. Mus. Nat. Hist. XXV p. 162, figs. 48, 49.

A species of a large size, founded on a right femur, a right first

metacarpal, portions of vertebrae and portions of the pubes.

The first metacarpal is much longer than broad, 99 mm. long and

56 mm. wide distally. The bone is more elongate than in Masso-

spondylus or Gryponyx.

The femur differs from that of P. cullingworthi in its thinner

proximal end, its narrower distal end, and the slightly lower position

of the trochanter quartus. The medial distal condyle also appears

to be much stouter in P. cullingworthi. It differs from Gryponyx in

that the trochanter quartus does not lie wholly in the upper half of

the shaft; but it is not so low down as in Euskelesaurus or Melanorosaurus.

The pubis has a broad anterior plate.

Two phalangeal bones collected by Dr. D. R. Kannemeyer at Witkop,

Jamestown, C.P. — the type locality and now in the South African

Museum (Cat. No. 1875) probably belong to this species. One has

a length of 55 mm., a proximal width of 32 mm., a proximal height

Fauna and Stratigraphy of the Stormberg Series. 407

of 28 mm. and a distal width of 30° mm. At the distal end the

medial portion of the articular surface is bigger than the lateral.

The other bone, which is probably the first phalanx of the first digit

of the left foot, has an axis which curves outwards at the distal end.

The greatest length is 53 mm., the proximal width 33 mm., and the

maximum distal width 32 mm. At the distal end the palmar surface

is much broader than the dorsal surface.

Type. In collection of American Museum, New York. (Cat. No. 5605.)

Locality. Witkop, near Jamestown, Aliwal North, C.P.

Horizon. Base of Red Beds.

PLATEOSAURUS CULLINGWORTHI, sp. nov.

Text figs. 31—35,

At Kromme Spruit, Herschel, C. P. a number of fragments of

large Dinosaurian bones belonging to 3 or 4 individuals were found

weathered out down one of the slopes of a steep kopje formed of

the basal rocks of the Red Beds. At one poimt near the top of

the same kopje bones were found in situ. Excavation revealed a

‘“nocket” of isolated bones belonging to two individuals — a larger

and a smaller — of apparently the same species. Some of the

weathered bones, including two femora, have also been associated

with these remains; the remainder belong obviously to an animal

of heavier build, having large and heavy dorsal vertebrae and have

been described as a species of Huskelesaurus. The bones from the

pocket, together with the two femora and one or two other bones

are considered to belong to a new species of Plateosaurus which |

have named in honour of Mr. C. W. Cullingworth to whose energy

some of the finds are due.

Although this form shows some differences from the more typical

members of the genus from Europe, especially in the greater length

of the humerus compared with that of the other bones, it has not

been considered advisable to separate it generically from Plateosaurns,

Vertebrae. Cervical. Judging from the curvature of the ventral

border of the centrum, the only cervical vertebra preserved (S. A.M.

Cat. No, 3345) is probably the 3rd. or 4th. or possibly the 5th.

Taking a line at right angles to the end faces, the posterior ventral

point les 34 mm. below the anterior ventral point. The total length

of the centrum is 157 mm. Its ends are roughly circular, the

anterior 57 mm. high, the posterior 61 mm. The posterior end is

more deeply concave than the anterior, and both have their ventral

408 Annals of the South African Museum.

borders rounded off. From the anterior end there runs a strong

ventral median keel which gradually dies away until it disappears

in the posterior third of the centrum. The whole vertebra is com-

pressed in the middle. The small diapophysis lies on the upper

half of the centrum, 40 mm, from the anterior border. Below and

anterior to it is a small horizontal parapophysis. Between the two

is a fairly deep groove. The prezygapophyses extend well in front

of the centrum and have slightly convex facets. Seen from in front

they form between them a wide V-shaped groove for the reception

of the preceding postzygapophyses. Their lower borders are hori-

zontal, so that each process thickens rapidly laterally. The post-

zygapophyses are slightly shorter than the anterior processes. The

facets are slightly concave. The whole process is strongly built.

The neural spine is missing; but its base is long and narrow, and

the spine must have been low.

Dorsal. An anterior dorsal centrum (Cat. No, 3345a) has a length

of 102 mm.; its ends are 80 mm. high and 70 mm. wide. In the

middle the body is strongly constricted, having a width of only 24 mm.

The ventral border is concave and has a sharp longitudinal keel.

The ends are concave, equally so, and their ventral borders are

rounded off. The parapophysis for the capitulum of the rib is midway

along the body in its upper half, directly below the transverse process.

The transverse process is 50 mm. long, placed above the middle of

the body. Its upper surface is flat and horizontal, and its distal end

is considerably thickened so that its articular surface is roughly

triangular with a width of 356 mm., looking downwards and outwards.

The anterior border of the process passes directly into the outer

border of the prezygapophysis; its hinder border is proximally emar-

ginate. The width between the facets of the transverse processes is

115 mm. The prezygapophysis extends in front of the centrum, and

is double the length of the postzygapophysis. The general shape of

the processes is of the normal Plateosaurian type.

Two other dorsal centra are preserved (Cat. No. 3356) having the

following measurements:

Length . ; : . 105 mm. 110 mm.

Height of anterior end. Sonne 90,

Height of posterior end 1oeues o2 5;

Width of anterior end. (Ghoje Fee ieee

Width of posterior end G3a5;, We Bes

Median width : : 310) ee Bil bee

Fauna and Stratigraphy of the Stormberg Series. 409

Caudal. There are two imperfect caudal vertebrae, both from the

mid-caudal region. One is 67 mm. long, has the anterior end of the

centrum 52 mm. high and 48 mm. wide, the posterior end 50 mm,

high and 44 mm. wide, and is considerably constricted in the middle

width a minimum width of 24 mm. The ends are somewhat concave

and the ventral borders of the ends are bevelled off, the anterior

more than the posterior. The anterior zygapophysis are small and

upwardly directed. The transverse processes arise at the top of the

centrum and stand out horizontally.

In the other, the centrum is 75 mm. long with its end 48 mm.

high and broad. It is but little compressed and the ventral surface

is flattened. The prezygapophyses are directed strongly upwards, while

the transverse processes are small and arise from behind the middle

of the centrum. In neither vertebra are the neural spines or the

postzygapohyses preserved.

Scapula, (Cat. No. 3348.) The proximal part of a left scapula is

present, lacking the glenoid cavity and the coracoidal articular surface.

The greatest breadth at the proximal end was probably 150 mm.

The inner face of the scapula is flat, curving strongly inwards

proximally; the outer face is lightly convex. Above the glenoid

cavity the bone thins rapidly; the anterior border is uniformly thin

save for a slight thickening near the coracoid. The supracoraco-

scapular concavity is well-marked and large and the deltoid crest

strongly developed. There is also preserved a fragment of the upper end

which shows that distally the bone was expanded as in Plateosaurus,

The narrowest part of the bone has a breadth of 71 mm. and a

thickness of 24 mm.

Humerus. Two left humeri are preserved, one (Cat. No. 3542) of

a larger, and one (Cat. No. 3350) of a smaller individual. The two

agree closely with one another in general characteristics, although

the smaller bone has been flattened so that the processus lateralis

is not so strongly bent as in the larger bone.

From within the bone is seen to be very slightly S-shaped. The

proximal part is very broad, the shaft thin, and the distal end

broad. The proximal and distal articular surfaces are strongly

inclined to one another. The head of the humerus lies somewhat

within the middle line. The anterior edge of the proximal articular

surface is a regular curve; the posterior edge has two saddle-shaped

prominences due to thickenings at the inner angle and at the caput

humeri. This latter is at the extreme end of the bone. At the

outer side of the broad, concave bicipital fossa is the processus

lateralis, a strong crest lying parallel to the inner border of the

410 Annals of the South African Museum.

upper half of the bone, and whose anterior surface is inclined to

the axis of the bone, the upper half pointing medially, the lower

end outwards. The lower end of this crest at its Junction with the

shaft lies half-way down the humerus.

The distal end is broad, but the condyles are not greatly thickened,

the whole articular surface being much narrower in comparison

Fig. 31. Plateosaurus cullingworthi, Htn.

Humerus No. 3342. X }.

with its length than in the European species of Plateosaurus or in

Pachysaurus. Both condyles are somewhat thickened in front,

but practically not at all on the posterior face. On the anterior

surface there is a marked concavity above the condyles in the

median line.

The following table gives the chief measurements of the two bones

and compares them with those of the humerus of Plateosaurus

reinigert :

Fauna and Stratigraphy of the Stermberg Series. 4\1

No. 3342 No. 3350 P. reinigeri

Total length . ; . 455 mm. 405 mm. 400 mm,

Length from lower end of

lateral process to distal

end of bone. 5 . D2 3 20 aes: 200)-.,

Length of lateral process J0Re 90s + 1005-,,

Thickness of lateral process oot > eee 25 «,,

Distance between upper

end of lateral process and

inner angle of bone . 155 ,, 1867. (180) *;,

Thickness of caput humeri OSes Des, DO! 53

Thickness of shaft . ‘ Do Lid a5, Do: 45

Breadth of shaft. ; O3tt.. 53 Cs, _

Breadth of distal end . 166. ,, LS iis es ee Lae

Thicknessat median condyle 51

Thickness at lateral condyle 5v4 5O 50) =;

Thickness between condyles 32 ,, 23°"; ye

From the humerus of Gresslyosaurus the bone differs in that its

proximal edge does not form such a high bow, its inner edge is

straighter in the proximal half, and the lateral process is much more

sharply bent over. The proximal edge is more curved than in the

other species of Plateosaurus or in Pachysaurus; and the distal end

is not provided with sharp longitudinal ridges as in Teratusaurus,

Radius. A left radius (Cat. No. 3347) has its lower end turned

towards the ulna. The upper part is flattened and broadened, having

slight median longitudinal grooves on both its medial and lateral

surfaces. The proximal articular surface is saddle-shaped, concave

in front, convex behind. From the higher posterior angle of the

bone a strong crest passes downwards and curves outwards, ending

abruptly on the posterior part of the lateral surface 60 mm. down

the shaft. The proximal end is 83 mm. long and 43 mm. broad;

the distal end 60 mm. long and 53 mm. broad. The narrowest part

of the shaft has a diameter of 35 mm. The distal end is trapezoid

in shape, and its posterior half is obliquely inclined upwards and

backwards.

Ulna. A left ulna (Cat. No. 3351) is compressed from side to side,

slightly bent with the medial face convex. The proximal end is bent

forwards and has an anteriorly directed prominent point. The hinder

portion of the proximal end is shghtly higher than the anterior portion,

forming an incipient olecranon process. The lateral edge of the ar-

ticular surface is 108 mm. long and is slightly concave. From the

4A2 Annals of the South African Museum.

median angle to the anterior point the length is 88 mm., and to the

posterior corner 68 mm. At the broadened end of the shaft the

lateral surface is concave, while below it is flat; the medial surface

is convex. The distal end is 70 mm. broad and 40 mm. thick. The

whole length of the bone is nearly 300 mm. In general aspect the bone

approximates closely to that of Plateosaurus except that the anterior

portion of the upper articular surface is longer and narrower.

There is also preserved the proximal end of another left ulna

somewhat smaller than the other. The median angle is slightly more

forward, the distances from it to the anterior and posterior points

being equal.

Ischium. A right ischium lacking the posterior end and the pubie

articulation, and the iliac articular portion and the distal end of a

Fig. 32. Plateosaurus cullingworthi, Htn.

Right ilium No. 2780. X }.

left ischium probably belong to this species. The whole length of

the bone was probably between 370 and 400 mm. The iliac articular

surface is 135 mm. long and 85 mm. broad. Its inner edge is straight,

its outer edge regularly convex. The groove on the hinder portion

of the bone is not very pronounced. It begins 100 mm. below the

proximal end. The broken end of the style shows a triangular cross-

section, the medial face being 70 mm. long and the width of the

bone 40 mm. The anterior angle is more acute than the lateral.

The distal end of the bone — as seen in the fragment of the left

side — is but slightly swollen. The posterior border curves slightly

forwards distally. At the extreme end the inner edge is 95 mm.

long and the bone is 63 mm. thick.

Femur. Two right femora (Cat. Nos. 3602, 3603) have been built

up from fragments found weathered down the slopes. One is larger

Fauna and Stratigraphy of the Stormberg Series. 413

than the other, but the two are of closely similar shape and proportions.

The larger (3602) has a length of 600 mm. The breadth at the

proximal end, from the border of the caput femoris to the lateral

border is 175 mm. The proximal end is strongly rounded, and its

shape approximates closely to that of Plateosaurus erlenbergensis. The

upper end of the bone is 85 mm. thick, but it is compressed between

the upper surface and the trochanter major. At the trochanter major

Fig. 33. Plateosaurus cullingworthi, Htn.

Right ischium. X 3.

it swells again, but not to such an extent as in Kuskelesaurus. The

shaft is shghtly S-shaped.

The lower end of the 4th trochanter lies 505 mm. below the top

of the bone. The trochanter is slightly curved, concave medially,

and lies nearer the medial than the lateral side of the bone. The

distal end is broadened and thickened. The medial condyle is consi-

derably larger than the lateral. Above the condyles on the hinder

face the intercondylar fossa is shallow and broad ; between the condyles

it is narrower and deeper. The medial condyle is a large rounded

4A4 Annals of the South African Museum.

boss. The lateral condyle is equally high, but it is narrower, and

from it a well-marked fairly sharply rounded ridge passes on to the

hinder face of the bone. The anterior face of the bone is flat in its

lower half. The lower articular surface has a maximum width of

175 mm., and a height of 105 mm.

The femur differs from that of P. stormbergensis in the position of

the 4th trochanter and in the greater thickness of the caput femoris.

Tibia. The right tibia (Cat. No. 3341) is a complete bone, whose

total length is 440 mm. The proximal surface has a length of 175 mm.

Fig. 34. Plateosaurus cullingwortht, Htn.

Right femur (3602). X 7.

from the median condyle to the anterior point, and a maximum

breadth of 99 mm. The surface is higher on the inner border than

on the outer. At the distal end the anterior process stands at least

90 mm. higher than. the narrower, prominent posterior malleolus.

The anterior border of the distal end is 141 mm. long, the inner

border 83 mm., and the posterior border (parallel to the anterior)

76 mm. long.

This bone is shorter than the tibia of Gryponyx africanus, but is

much more robust. The head is the same length but wider. The

distal end is very much stouter.

Fauna and Stratigraphy of the Stormberg Series. 4A5

Metatarsals, The second and third metatarsals of the right side

(Cat. No. 3343 and 3344 respectively) are preserved. The proximal

end of the second is quadrangular, having both its longer edges

concave, The larger axis of the parallelogram has a length of 92 mm.,

the shorter 53 mm, From the angles sharp ridges run down the

bone. The length of the bone is 216 mm., while the distal end has

a greatest breadth of 69 mm.

Lyn?

VW) Y

Fig. 35. Plateosaurus cullingworthi, Htn.

Right tibia (3341). Outer view.

The length of the third metatarsal is 226 mm. The proximal

end is narrowly triangular, 89 mm. long and having a base of

o2 mm. The distal end has a breadth of 66 mm.

A number of bones from the Red Beds just above the village of

Lady Grey, C. P., seem to belong to this species. They can be

correlated with the type by means of a left tibia, somewhat smaller.

than that described, but of similar shape and relative proportions;

a right humerus, of the same size as No. 3350; the distal ends of

the two ischia; the distal end of the right and the proximal end of

416 Annals of the South African Museum.

the left femur, comparable in size with No. 3603; and part of the

left scapula. In addition there is an ilium and part of a pubis,

together with vertebrae, ribs, and other fragments.

Ilium. The greatest length of the right ilium is 325 mm. Most

of the upper border is missing, but it was probably only slightly

curved. The posterior process is truncated obliquely, its hinder

border measuring 65 mm. The anterior process is fairly long, its

lower border rounded, its upper border sharp and thin, The ace-

tabulum is 165 mm. wide and 90 mm. high. The supra-acetabular

crest is prominent around the anterior upper portion of the border,

Fig. 36. Gryponyx africanus, Br.

Left pelvis of type.

A. Latero — ventral view.

B. Lateral view — ventral surface to right.

rounded on its outer and upper surfaces; but it dies away poste-

riorly. The preacetabular process is 1430 mm. long and 65 mm.

broad on the medial face.

Pubis. The larger portion of a right pubis is preserved. It lacks

the anterior edge, the distal end and the subacetabular portion,

The greatest length of the remainder is 360 mm. The posterior

border is strongly concave, so that the plate broadens considerably

distally.

Co-types. Portions of two animals in South African Museum.

Catalogue numbers as in description.

Loéality, Kromme Spruit, Herschel, C. P.

Horizon. Base of Red Beds.

Fauna and Stratigraphy of the Stormberg Series. 4AT

GRYPONYX AFRICANUS, Broom.

Text figs. 56-38.

1911. Broom. Ann. S. Afric. Mus. VII, 4, p. 294. Pls. XIV-XYV.

In addition to the original description the following points may

be noted.

Vertebrae. ‘Two posterior dorsal vertebrae have been developed.

The centra are each 75 mm. long; the anterior articular surface of

the first is 55 mm. broad, the posterior surface of the second

Fig. 37. Gryponyx africanus, Br.

Left femur of type.

A. Anterior view. B. Medial view.

63 mm. broad and 85 mm. high. The middle portion of the cen-

trum has a minimum width of 28 mm. The maximum height of

the vertebra was 180 mm. The dorsal spine is thin, long and

fairly low. <A figure is given herewith.

Pelvis, The pelvis of the type is crushed, and has been restored

and figured by Dr. Broom.

The two pubes form a complete symphysis. They are seen in

the type from the ventral side. The greatest length of the left

pubis is 440 mm. The pubic neck has a minimum width of

58 mm., and is situated at a distance of 95 mm. from the articular

surface for the ium. Its maximum thickness is not more than

20 mm. At the articulation with the ilium the pubis is 65 mm.

4A8 Annals of the South African Museum.

broad. The pubic foramen is large, oval in shape, with a long

diameter of 49 mm. and a short diameter of 35 mm.

The pubic plate is long, comparatively narrow and thin — the

lateral border sharply rounded and concave, the medial border thin

and straight. At the distal end the plate widens and thickens, so

that the distal surface is 70 mm. long and 40 mm. wide. At its

narrowest part the plate is 55 mm. wide.

Femur. The right femur is markedly S-shaped and_ strongly

bowed, the anterior face bemg convex. The length is between 535

mm. and 540 mm. The head of the bone is missing. Above the

level of the trochanter major the outer edge curves regularly to the

proximal surface and is parallel to the inner edge. The top of the

trochanter major les 100 mm. from the proximal surface. The

upper end of the fourth trochanter is 175 mm. from the proximal

end and the lower end about 260 mm. so that the trochanter lies

wholly in the proximal half of the bone.

The distal end is 97 mm. broad. The inner edge is 140 mm,

long, the outer edge 115 mm. The inner condyle is thicker than

the outer, and the sulcus between them is deep and narrow. The

ridge running from the outer condyle to the posterior face of the

bone is longer and more prominent than that from the inner

condyle.

Tibia. Dr. van Hoepen has pointed out that the figure given by

Dr. Broom and designated ‘touter view of left tibia” is in reality an

inner view of the right tibia.

The proximal end of the tibia seems to be somewhat flattened

from side to side and its width was possibly somewhat greater than

the 71 mm. given by Broom. The highest point of the bone lies

a little in advance of the anterior corner on the antero-medial

border. Instead of the medial border of the proximal end having

a regularly convex outline its middle portion is concave. as shown

in the figure. The lateral border has a pronounced concavity be-

tween the tuberositas tibiae and the lateral condyle. The anterior

half of the proximal surface is concave; the posterior portion is

also concave, and the two concavities are separated by a saddle.

The medial border is higher than the lateral. The shaft narrows

very rapidly below the articular surface and expands but slightly

distally, increasing only in width and not in thickness. Below the

proximal articular surface the lateral face is broadly grooved long-

itudinally, and separated from the posterior face by a rounded ridge

which carries a small boss of bone. The anterior proximal edge of

the bone is rugose.

Fauna and Stratigraphy of the Stormberg Series. 419

The shaft has a minimum width of 46 mm. and a minimum thick-

ness of 44 mm.

The distal articular surface has a maximum width of 90 mm. and

Fig. 38. Gryponyx africanus, Br.

Right tibia of type.

Latero — posterior view.

Posterior view.

Outline of proximal end.

Outline of distal end.

a maximum thickness of 53 mm.

The posterior edge is 70 mm.

long. The anterior condyle stands about 45 mm. above the posterior

condyle, and its surface faces downwards and outwards.

The sulcus

between the condyles is narrow and fairly deep.

420 Annals of the South African Museum.

Type. Pelvis and hind limbs, right and left manus, and vertebrae.

(S.A. Mus. Cat. No. 3357, 3358, 3359.)

Locality. Foutanie, Fouriesburg, Orange Free State.

Horizon. ‘Top of Red Beds.

GRYPONYX TRANSVAALENSIS, Broom.

4912. Broom. Trans. Geol. Soc. S. Afr. XIV, p. 82, Pl. XIII, figs. 3, 4.

4920. van Hoepen. Ann. Transv. Mus. VII, 2. p. 102.

This is a very imperfectly known species, the type consisting of

a claw phalanx and the distal end of a metatarsal.

The species is apparently about four-fifths the size of Gryponyx

africanus; according to Dr. Broom “‘it differs in having a much less

developed tubercle for the flexor tendon which makes the flexor surface

less curved and gives the whole claw a less hooked appearance”. Van

Hoepen, however, considers that ‘“‘the greater part of the tuberositas

for the flexor tendon is broken away”. He further thinks that the

claw belongs, not to the right hand, but to the left; and that it differs

from his Massospondylus browni (i. e. M. harriesi) im that the proximal

end of the lateral side ridge lies relatively much higher with regard

to that of the medial side in G. transvaalensis.

Type. Claw of manus and a metatarsal, in Transvaal Museum.

Locality. Wiepe 1258, N. Transvaal.

Horizon. Bushveld Sandstone.

GRYPONYX TAYLORI nov. sp.

Text fig. 39.

The remains forming the type of this new form were discovered

in 1915 in the neighbourhood of Fouriesburg, Orange Free State, in

an exposure near the top of the Red Beds. They consist of the

pelvic girdle and sacral vertebrae found in conjunction.

Vertebrae. Three sacral vertebrae were found in situ between the

two sides of the pelvis, somewhat flattened from side to side. The

first centrum is not fused to the second.

The second and third vertebrae are of equal length, and each slightly

longer than the first. The centra are higher than wide with concave

ends. The ventral border of the first is more excavate than those

of the other two; the centrum of the third is more excavate post-

erlorly on its lower border than anteriorly. The lower border of

each is sharply rounded, but not keeled nor pointed. The first centrum

is 79 mm. long, the others each 84 mm. The posterior surface of

the third centrum is 82 mm. high.

Fauna and Stratigraphy of the Stormberg Series. 424

The second sacral rib is missing; but it is apparent that the base

of the third sacral rib is larger than the second, and the second than

the first. Distally the sacral ribs are fused to form one long surface

for the support of the ilium. The base of the first lies on the anterior

part of the centrum just below the neural arch and only covers a

small part of the body; that of the third occupies at least half of

the anterior half of the body and extends up to the strong transverse

process. Inferiorly the proximal part of this rib is strong and rounded ;

superiorly it thins considerably and then widens out to meet the

under side of the fairly wide, horizontal, and backwardly directed

transverse process. The anterior face of the rib is thus fairly strongly

concave, the posterior face also being concave, but less so. The first

rib is essentially of the same character, but is much smaller.

Of the zygapophyses only the prezygapophysis of the first vertebra

is well seen. It is strong with a flat upper surface facing somewhat

inwards. The first postzygapophysis is closely fixed to the prezyga-

pophysis of the second vertebra. .

The neural canal is high and narrow. The neural spines are higher

than the centra, thin, broader above than below, and slightly back-

wardly directed.

Ilium. The left ilium lacks only the posterior process, which is

present in the type of the genus.

The anterior spine is short and sharp, its lower border rounded and

fairly thick, its upper border sharp and thin. Its lower border is

straight. The preacetabular process is long and strong, widest at the

end. In cross-section it is triangular, the apex of the triangle —

formed of an obtuse angle — being on the outer side of the bone.

The upper half of the process has a strong sharp ridge on its outer

edge, which ridge gradually becomes less pronounced as it continues

round the acetabular border until it disappears altogether just behind

the mid-point of the upper border of the acetabulum.

The postacetabular process is short and broad, its inner surface

flat, its outer surface broadly rounded. The hinder border is concave.

The body of the bone thins away rapidly above the acetabulum,

and the upper border is bent inwards between the anterior and

posterior spines.

The acetabulum is high and narrow. More than half of it is formed

by the ilium.

Ischium. The expanded proximal portion of each ischium is

preserved. The surface for articulation with the pubis is long and

narrows below; both it and the iliac surface are thickened. The

hinder surface of the bone is thickened and carries a longitudinal

429 Annals of the South African Museum.

median groove which dies out about 50 mm. below the posterior

upper border of the bone. The anterior border is thin.

Pubis. The left pubis is a long bone with an expanded proximal

portion and a thickened distal end. The middle portion is slender,

with a rounded and thickened anterior border having no longitudinal

Fig. 39. Gryponyx taylori, Htn.

Left side of pelvis. X }.

groove, and thinning posteriorly. The posterior border is missing,

but the bone seems to be narrower than in Seeley figure of Masso-

spondylus. The pubic foramen is large and oval in shape.

The chief measurements of the pelvis are as follows:

Width of acetabular opening . : = 101mm:

Length of preacetabular process. ae Oe ere

Fauna and Stratigraphy of the Stormberg Series. 423

Greatest width of preacetabular process . 71 mm.

Width of postacetabular process — . : ODP.

Length of anterior spine . ‘ : : 637) 5;

Length of iliac surface of ischium . : The) ee

Length of ischio-pubic suture . : ee LOG -.;

Length of pubis ; : : . a) A

Length of iliac surface of pubis —. : S05;

Thickness of distal end of pubis. , Tes

In size this pelvis is almost identical with that of the type of

Gryponyx africanus, but it differs in the possession of a much larger

anterior iliac spine and in the different slope of the upper iliac border,

agreeing therein both with Massospondylus and Plateosaurus.

In his discussion of Massospondylus in “Die Dinosaurier der Euro-

peaischen Triasformation” von Huene lays emphasis on the fact that

the distal portion of the pubis is short and thick, thus differing from

all Plateosaurs. In this specimen, however, the pubis is long and

slender distally as in Gryponyx and, therefore, the form cannot be

placed in the Massospondylidae. In spite of the difference in the

shape of the anterior spine of the ilium I am inclined to place it in

the genus Gryponyx, naming it after Mr. H. M. Taylor, while

collecting with whom [ discovered the remains.

Type. Pelvic girdle and sacral vertebrae. S. Afr. Mus. Cat. No. 3453.

Locality. Fouriesburg, O. F.S.

Horizon. Top of Red Beds.

EUSKELESAURUS BROWNI, Huxley.

1866. Huxley. Quart. Journ. Geol. Soc. XXIII, p.1.

1894. Seeley. Ann. Mag. Nat. Hist. Ser. 6, Vol. XIV, p. 317.

1906. von Huene. Geol. und Palaeont. Abh. N.F. Bd. VII, Hft 2,

p. 123.

4911. Broom. Ann. 8S. A. Mus. VII, 4. p. 292.

The type specimens of this species are in the British Museum and

in the Museum (histoire naturelle in Paris. They consist of frag-

mentary vertebrae, femur, tibia and fibula, and pubis. The fragmentary

nature of these remains renders comparison with other specimens

unsatisfactory for specific identity. The type bones have been fully

described and discussed by von Huene, who arrives at the following

conclusions.

Euskelesaurus browni is not only larger than most other Plateosaurs

but there are certain characters which, on the one hand, distinctly

424 Annals of the South African Museum.

separates it from the other genera, and on the other hand brings it

very close to them. The shortness of caudal vertebrae, especially

the posterior ones, does not occur to such a degree in other Plateosaurs,

likewise the shortness and compression of the phalanges of the foot

and consequently also of the metatarsals. The greatest similarity 1s

with Gresslyosaurus. The fourth trochanter les in the lower half

of the femur as in Gresslyosaurus, certainly rather lower than in that

genus; but the whole femur is, in proportion to the enormous vertebral

column in its central part, conspicuously shorter than in Gresslyosaurus

and the other Plateosaurs. The tibia is extremely strong at the

proximal end. The dorsal vertebral centrum is higher than in other

Plateosaurs. The third sacral centrum is pointed below. The troch-

anter major of the femur is larger than in other Plateosaurs.

Type. In British Museum.

Locality. ‘“Stormberg”, C.P. Almost certainly from the Kraai River.

Horizon. Base of Red Beds.

EUSKELESAURUS CAPENSIS (Lydekker).

1889. Lydekker. Orinosaurus capensis Geol. Mag. Ser. 3, vol. VI,

p. Joo,

1906. von Huene. Geol. und Palaeont. Abh. N.F. Bd. VII, Hft. 2,

p. 129.

This is known only from the proximal end of a tibia and a small

portion of a femur. It is larger than the type species. The form

of the proximal end of the tibia is characteristic, and it is possible

that the species is generically distinct from E. browni. Too little is

known of it, however, to warrant such a separation being made, and

none of the bones in the South African Museum collection can be

assigned to the species.

EUSKELESAURUS AFRICANUS sp. Nov.

Text figs. 40-41.

From Kromme Spruit come a number of large bones found weathered

down the slope of the kopje that yielded Plateosawrus cullingworth.

Of these, a number of vertebrae, two attached ischia, and some ilia

are the most completely preserved. There are also portions of tibiae

and of a femur which are, however, not sufficiently complete to

admit of satisfactory comparison with those of Huskelesaurus. The

animals represented by these bones must have been somewhat smaller

than E. brownt and I have therefore decided to keep them in the

already-established genus and to designate them as Huskelesaurus

Fauna and Stratigraphy of the Stormberg Series. 425

africanus n. sp. The type is in the South African Museum (Cat.

No. 2608),

Sacral vertebrae. Two fused sacral vertebrae are preserved, lacking

the sacral ribs and the neural spines, They are probably the 1st

and 2nd sacrals,

iy /

lig

Nh:

Nii

Ischia. Posterior view. X 4.

The anterior centrum has a leneth of 1440 mm.; its front end is

flat and has a height of 120 mm. and a breadth of 130 mm. The

sides of the body are swollen and the ventral surface is broadly

rounded -— not so much so as in the succeeding centrum, The sacral

rib is represented only by a section across its junction with the body.

426 Annals of the South African Museum,

This is elongate with a convex anterior edge and a concave posterior

edge, while the upper edge is nearly straight and lies just. below the

level of the zygapophysis. The prezygapophysis is shortand its upper

surface is concave, looking inwards and upwards.

Fig. 41. Huskelesaurus africanus, Htn.

Left ischium, lateral view. X }.

The succeeding centrum had a length of 145 mm. Its posterior

face was 130 mm. high and 140 mm. broad. Its ventral surface

was very broadly rounded and without any trace of median keel.

The sacral rib itself is lacking, but it must have been very large

and strong, larger than that of the first sacral. Its base is on the

centrum extending over the greater part of the side of the body from

Fauna and Stratigraphy of the Stormberg Series. 4917

the front to the back and from the transverse process down to within

35 mm. of the ventral surface. The upper surface of the transverse

process is flat and horizontal, the process passing outwards and back-

wards. The posterior zygapophysis is short; its under surface is

concave facing outwards and downwards. The base of the neural

spine is 100 mm. long and not more than 25 mm. wide. The width

between the tips of the transverse processes was probably 290 mm.

Ischium. Both ischia are present, lacking only the subacetabular

expansions.

The greatest length of the bone is 570 mm. Ventrally the two

bones were probably in contact throughout their length; dorsally

the distal ends have a straight contact for 250 mm. and then diverge

gradually. Just above the point of contact on the posterior side the

shaft has a breadth of 50 mm.; at the distal end it broadens out to

100 mm. The shaft is flat posteriorly, while its outer and anterior

borders are concave. The most characteristic feature of the bone is

the very pronounced thickening at the distal end — a thickening

greater than that seen in any other known Plateosaur. Whereas at

its narrowest the bone has a thickness of only 40 mm., just above

the distal end it swells to a thickness of 140 mm. and a breadth of

100 mm.

The distance between the inner sides of the proximal ends of the two

bones is 1440 mm. The medial face of the proximal portion is concave,

the lateral face convex. The greatest breadth across the iliac articular

surface is 410 mm. The groove on the hinder face of the bone is

about 1450 mm. long and is well-marked.

Type. Sacral vertebrae, ischia. (S. Af. Mus. Cat. No. 3608.)

Locality. Kromme Spruit, Herschel, C. P.

Horizon. Base of Red Beds.

GIGANTOSCELUS MOLENGRAAFFI, van Hoepen.

4916. van Hoepen. Verhand. Geol.-Mijn. Genoot. Ned. en Kolonien.

Geol. Serie III, p. 107, text fig. 3.

Founded on the distal end of a right femur. The bone is some-

what larger than that of Huskelesaurus, but is relatively thicker and

has a narrower sulcus between the two distal condyles. The chief

interest of the specimen lies in the fact that it is of a type that occurs

apparently only near the base of the Red Beds in the Cape Province.

Type. Distal end of a femur in the Transvaal Museum.

Locality. Waakdoornbult, 544. Waterberg Dist., Transvaal.

Horizon, Bushveld Sandstone.

428 Annals of the South African Museum.

EUCNEMESAURUS FORTIS, v. Hoepen.

4920. van “Hoepen. Ann. Transv. Mus. VII, 2. p. 93. Pls. XI,

ST in fel:

The type of this form consists of the proximal half of a femur,

a complete tibia, a proximal portion of a pubis, portions of dorsal

and caudal vertebrae and some fragments, of a large form.

In addition to the comparisons made in the original description,

the following may be added. The tibia approximates in size to that

of Melanorosaurus readi, but the proximal end appears to be thicker

(although in M. readi it is somewhat incomplete). Further, the

proximal end slopes down to the shaft more gradually in £. fortis

than in M. readi, and the shaft is more slender in the latter form.

The differences, however, are not very acute and may well be merely

specific. Until further comparisons are possible, however, there can

be no great harm done in retaining separate generic names for the

two forms. The femora, as far as they can be compared, seem

different. The proximal end is broader in M. readi, owing to the

fact that the lateral border does not pass so abruptly into the proximal

border in £. fortis. In the latter, too, the fourth trochanter seems

to occupy a slightly more medial position. In the former character,

Eucnemesaurus occupies an intermediate position between Plateosaurus

and Melanorosaurus.

From Plateosaurus cullingworthi the tibia of Eucnemesaurus differs

markedly in general shape, the shaft of the former being much more

slender in its middle portion; the articular surfaces are also of

different shape.

Type. In the Transvaal Museum.

Locality. Zonderhout, near Slabberts, O. F.S.

Horizon. Red Beds (probably about half way up.)

MELANOROSAURUS, gen. nov.

This genus is characterised by the following features: Vertebrae

lighter and smaller than those of Gresslyosaurus and Euskelesaurus

when compared with the length and size of the femur; humerus

with lateral process sharply bent over, proximal edge forming a

moderately high bow, less marked than in Gresslyosaurus; femur

with straight shaft, whose lateral border forms approximately a right

angle with the proximal surface at the upper, outer corner, and with

lower end of fourth trochanter below the middle of the femur; distal

end of tibia broader in front than behind.

Fauna and Stratigraphy of the Stormberg Series. 429

MELANOROSAURUS READI sp. nov.

Text figs. 42-47.

The bones forming the type of this species were found on the

northern slope of the mountain Thaba ’Nyama (“Black Mountain”),

lying between Josana’s Hoek and Josana’s Nek in the district of

Herschel in the Cape Province. They were lying isolated and em-

bedded in a soft red mudstone below a sandstone band not far

above the base of the Red Beds. The bones consist of a tibia, a

fibula, part of the pelvis, some vertebrae and metatarsals, together

with a femur lying partly embedded in the overlying sandstone and

the proximal half of a humerus found weathered down the slope.

They are in the collection of the South African Museum (Cat. Nos.

3449, 3450). I can collate these remains with no hitherto-known

species, and have much pleasure in naming them after Mr. B. Read,

former Principal of the Bensonvale Training School, of whose kind-

ness, display of interest, and hospitality I have a lively recollection.

Another individual (S. A. M. Cat. No. 3532) belonging, apparently,

to the same species — but somewhat smaller than the type — is

represented by some bones excavated below the Rooi Nek, between

Kromme Spruit and Majuba Nek, Herschel, from an horizon about

one-third the way up the Red Beds. These remains include a

scapula and a complete humerus, and thus add to the knowledge of

the form.

Vertebrae. With the type several isolated vertebral centra were

obtained. The centra are all considerably longer than high. Their

end surfaces are oval in shape with the larger axis vertical. One —

probably a posterior cervical — has a sharp ventral surface, keeled

at either end. The dorsals are rounded below; all the vertebrae

are lightly built compared with those of Gresslyosaurus. The length

of a dorsal centrum is 110 mm., the height of its anterior surface

95 mm.; while the width at the middle of the centrum is 45 mm,

The ventral surface of the bone is not acutely concave.

Scapula. Among the bones from Rooi Nek an almost complete

scapula is preserved. The bone has been flattened, so that its origi-

nal curvature is lost. The upper end is somewhat expanded and is

thinner than the proximal end. The anterior border is thin. Proxi-

mally the posterior border is rounded and comparatively thick, but

distally it is thin. The glenoid cavity is not deep, but it is broad

and fairly high. The articular surface for the coracoid is very

broad. The greatest length of the bone is 450 mm. The width at

430 Annals of the South African Museum.

the distal end is 175 mm., and that at the narrowest part of the

shaft is 75 mm.

Humerus. The Rooi Nek specimen contains an almost complete

right humerus. Its proximal edge forms a somewhat higher bow

than is seen in the incomplete bone of the type from Thaba ’Nyama;

but this latter shows signs of having been somewhat abraded before

its final entombment. Nevertheless, the proximal end is not so

strongly arched as in Gresslyosaurus, and the distal end differs con-

Fig. 42. Melanorosaurus readi, Htn.

Right humerus inner view (3532). X 4.

siderably from that genus. The lower end of the lateral process

lies nearly half-way down the bone.

The distal and proximal articular surfaces are inclined to one

another at an angle of about 45°.

The distal end has been somewhat flattened, but it is broad and

thin. The lateral condyle is considerably smaller than the medial,

Fauna and Stratigraphy of the Stormberg Series. ASA

and there is a well-marked broad depression between them on the

anterior face. The posterior surface of the distal end is almost flat,

the lateral condyle being somewhat thickened.

The bone is 500 mm. long. The distance from the lower end of

the lateral process to the distal end of the bone is 260 mm. The

shaft is 55 mm. thick and 67 mm. broad. The distal end is 1455 mm.

broad, the thickness at the lateral condyle is 65 mm., and at the

medial condyle 65 mm.

Radius. The left radius presents no features of unusual interest.

It is slightly shorter than the ulna, having a length of 280 mm.

The proximal and distal ends are each 89 mm. wide, while the shaft

at its narrowest is 43 mm. wide.

There is also preserved a right radius, somewhat smaller than the

Fig. 43. Melanorosaurus readi, Htn.

Right ulna of type. X }.

corresponding bone of the other side, It is 250 mm. long. The

proximal end is flattened, while the distal end is thicker and stronger.

In both bones the proximal articular surface is concave in front and

convex behind as in Plateosaurus and other forms.

Ulna. The right ulna is complete, being 300 mm. long. The

proximal end is expanded and has a triangular articular surface, of

which the anterior and posterior angles are sharper than the lateral

angle. The anterior point is directed forwards, the posterior one

backwards and outwards. The posterior half of the surface is strongly

convex in its middle portion, being raised considerably above the

432 Annals of the South African Museum.

anterior half of the surface. There is thus a strong boss forming

an incipient olecranon process. The maximum length of the proximal

surface is 120 mm., its maximum width 60 mm.

The posterior border of the bone is bent in an S-shaped curve,

the anterior border beimg regularly convex. In the former feature

the bone differs considerably from that of Plateosaurus erlenbergensis

which has a straight posterior border. The thinnest part of the shaft

occurs about 10 em. from the distal end, where it has an oval cross-

section whose diameters are 35 mm. and 45 mm. The distal end

is 95 mm. broad.

Ilium. (In the Thaba ’Nyama specimen one ilium is complete. In

the Rooi Nek specimen one ilium is complete and the other almost

so. The shape of the ilium is most like that of Massospondylus carinatus.

Fig. 44. Melanorosaurus readi, Htn.

Right ilium of type. X }.

The upper edge is fairly straight and the anterior spine is not so

long as in Gresslyosaurus. The acetabulum is bounded in its upper

anterior quadrant by a sharp ridge which dies away towards the

end of the preacetabular process. The postacetabular process is short ;

its posterior border is straight, except at the lower corner, where it

is provided with a hook-like projection. The posterior iliac spine is

short and high. The body of the bone is slightly bowed inwards.

The height of the bone from the bottom of the postacetabular process

to the upper edge is 275 mm.; the greatest length is 4145 mm. The

width of the acetabulum is 240 mm., its height 420 mm.

Pubis. An incomplete pubis is preserved, which shows that the

anterior portion is a broad, flattened plate.

Fauna and Stratigraphy of the Stormberg Series. 435

Femur. This bone was in doubtful association with the other remains,

and may possibly belong to another form. It is short compared with

the length of the humerus, shorter than in Gresslyosaurus. It agrees

with Gresslyosaurus and Huskelesaurus in that the lower end of the

Ath trochanter is below the middle of the bone. In outer view the

bone is straight and is considerably expanded distally. The width of

the distal end at the lateral condyle is greater than that at the medial

condyle; the groove between the two condyles is broad and fairly

shallow.

The length of the femur is 620 mm., the breadth at the proximal

end 170 mm. The lower end of the 4th trochanter is 350 mm.

Fig. 45. Melanorosaurus readi, Htn.

Right femur. XX 35.

below the top of the bone. The minimum width of the shaft is

95 mm., its minimum thickness 73 mm. The width of the distal

end is 175 mm.; the thickness at the lateral condyle 145 mm., and

at the medial condyle 80 mm.

This bone, when compared with the tibia, is shorter than in other

members of the Plateosauridae. In Euskelesaurus, however, where

the femur is short and stout, the tibia is not known; but it is possible

that the two bones in that genus bear somewhat the same relation

to one another as in this form. It is in the size of the vertebrae

that the two forms differ so markedly.

Tibia, The tibia is a massive bone 45 cm. long with a large

434 Annals of the South African Museum.

proximal end, a straight shaft, and a broadened distal end. The

tibial tuberosity is weathered away to a certain extent, but must

have been prominent. The proximal articular surface has a greatest

length of 195 mm., and a greatest width probably of about 100 mm.

The inner border does not stand much higher than the outer. The

inner border is convex with a shallow concavity at the middle.

Between the tibial tuberosity and the anterior and posterior ends of

Fig. 46. Melanorosaurus readi, Htn.

Left tibia, outer view. X 2.

the surface there is, in each case, a shallow concavity. The anterior

portion of the surface is higher than the posterior, and is not excess-

ively prolonged, but is bluntly rounded.

Just below the middle of the bone, where it is most slender, the

shaft has an antero-posterior diameter of 80 mm., and a lateral diameter

of 60 mm. The anterior surface is provided with a sharp keel in

its lower half which passes down to the anterior condyle. The post-

erior surface is rounded, the outer and inner surfaces flattened.

At the distal end the anterior border has a length of 130 mm.

Fauna and Stratigraphy of the Stormberg Series. ABS

measured along the lower border and stands about 40 mm, above the

posterior border, which is 90 mm. long. The inner border is strongly

rounded with a length approximately equal to that of the hinder border.

Fibula. The left fibula is preserved entire, although the bone

surface is somewhat shattered. Its greatest length is 477 mm. The

proximal end is expanded, its inner surface slightly concave, its outer

surface convex. The greatest breadth of the proximal end is 140 mm.,

while it is but 44 mm. thick. The anterior border of this end is

thin and sharply rounded, while posteriorly the bone is thicker. The

upper edge is higher behind than in front.

The shaft is flat on the inner side and strongly rounded laterally.

It has a diameter of 50 mm.

Fig. 47. Melanorosaurus readi, Htn.

Metatarsal III. Front view. X }.

The distal end is slightly swollen in an antero-posterior direction.

The lower surface is 85 mm. long and 53 mm. broad, convex on the

inner side and obliquely flattened laterally. The anterior face of the

distal end is broadly rounded, while the posterior face has a sharp

ridge on its inner side, sloping away to the outer face as a flat surface.

Metatarsai. One of the metatarsals is preserved, probably the 3rd.

It is astraight bone having a length of 200 mm. The proximal articular

surface is triangular, the anterior angle bemeg slightly obtuse. The

surface is 80 mm. broad, and has a maximum thickness of 40 mm.

At its thinnest part the shaft is 44 mm. broad and 26 mm. thick.

The distal articular surface is 62 mm, broad. Distally, the anterior

surface of the bone is very flat, while posteriorly it is slightly concave

with a shallow wide median groove just above the articular surface.

Type. Incomplete skeleton. S.A. Mus. Cat. Nos. 3449, 3450.

Locality. Thaba ’Nyama, near Bensonvale, Herschel, C. P.

Horizon. Near base of Red Beds.

A436 Annals of the South African Museum.

OrpER ORNITHISCHIA Seeley.

GERANOSAURUS ATAVUS Broom.

1914. Broom. Ann. 8. Af. Mus. VII, 4, p. 306. Pl. XVII, fig. 24.

“The collection of bones consists of badly crushed fragments of a

skull with the anterior part of lower jaw fairly well preserved, some

slender birdlike hind-limb bones, and a number of very imperfect

vertebrae. The vertebrae seem too large to have belonged to the

skull, and there being thus some doubt about the bones being those

of one animal | think it better to describe the jaw-bones alone and

to make them the type.

As preserved, the lower jaw has the left dentary fairly complete

with a considerable portion of the right and the predentary nearly

perfect.

The predentary has its upper surface displayed, which is concave.

It is 42 mm. long and the same in width. The outer and anterior

edges are sharp and doubtless formed a horny beak.

The dentary as preserved measures 73 mm. in length, and there

is probably but little missing from the posterior end. The anterior

half bears 9 teeth which have rounded roots in sockets. The teeth

in the fragment of maxilla have flat chisel-shaped crowns with the

outer face feebly ridged. Probably those of the mandible were similar

in this respect. The most remarkable thing about the dentition is

that the most anterior of the teeth is larger than the others, and

may be looked upon as a canine. The total length of the series is

35 mm. Most of the teeth have a diameter of between 3 and 4mm.,

but the anterior tooth has a diameter of 5 mm.”

Associated with the lower jaw, on another small slab of stone is

the imperfect mould of a tibia, fibula and some of the bones of the

foot which bear nearly the same relation to the size of the Jaw as

does the tibia to jaw in Nanosaurus. It seems probable, therefore,

that they are of the same species as the type jaw.

The tibia is long and slender, apparently agreeing with that of

Nanosaurus in that it is compressed proximally with a somewhat

triangular cross-section, while its distal end is more rounded in section.

The proximal end is nearly all preserved. The anterior portion of

the articular surface is higher than the posterior part and is cons-

iderably narrower, the tuberositas tibiae being well defined. The

lateral process is rounded and strong. The shaft is slender; the

distal end is only preserved in the form of a mould of the lateral

surface. The total length is 146 mm.; the length of the proximal

Fauna and Stratigraphy of the Stormberg Series. 437

articular surface was about 25 mm.; the shaft at its narrowest probably

had a width of not much more than 10 mm.

The fibula lies on the outer side of the tibia and closely appressed

to it, crossing it at a low angle. It was apparently shorter than the

tibia and very slender, being widened at its proximal end.

Part of the pes is preserved, consisting of a smal] tarsal bone and

the most of one digit, probably the second. The metatarsal is

19 mm. long posteriorly and 14 mm, long anteriorly, the distal sur-

face being inclined to the axis of the tibial shaft. The whole axis

of the digit is inclined to that of the tibia, so that the animal

appears to have been digitigrade. The first phalanx is short, its

proximal end broader than the anterior end; the second phalanx is

18 min. long. The claw is incomplete, but was long and compara-

tively slender.

Type. Incomplete lower jaw — S. Afr. Mus. Cat. No. 1871.

Locality. Top of Barkly Pass, Elhot, C. P.

Horizon. Cave Sandstone (base of).

( 438 )

PATE

S LRADIGH Ar EY

Moreno BeEbs.

The general features of the Molteno Beds are described by Rogers

and du Toit in ‘The Geology of Cape Colony” as follows:

“The Molteno Beds are first met with at a point a little to the

east of Steynsburg and form the higher-lying ground in the Division

of Molteno; they extend along the fort of the Stormbergen into

Herschel, the Orange River Colony, and Basutoland, and along the

base of the Drakensbergen through East Griqualand into Natal.

The formation consists of sandstones. shales and mudstones, the

softer beds being much like those of the Ecca and Beaufort, grey,

greenish or bluish in colour, but without the calcareous concretions

so abundant in the lower groups.

Fossil plants are in places abundant, but seem if anything to be

more plentiful in the lower half of the Molteno Beds; silicified wood

is common in some of the sandstones.

The sandstones of the Molteno beds are quite unlike any that

occur in the lower groups of the Karroo system. In general appea-

rance and in the character of the surface to which they give rise,

they resemble the Table Mountain Sandstone more closely than any

other in the Colony, but they are coarser in grain and much looser

in texture. In most localities the quartz grains are coated with a

later deposit of quartz with more or less perfect crystalline faces

which reflect light well, so that the rock sparkles in the sunlight.

To such varieties the term “glittering sandstone” has been appro-

priately given.

Grains of felspar are abundant in these sandstones, sometimes in

such quantity that the rock can almost be termed an arkose. The

loose texture of the Molteno sandstone has allowed the felspar to

weather considerably, and the dull white grains of weathered fel-

spar are always conspicuous constituents of the sandstones, more

especially in the finer grained varieties. Rounded or spherical nodu-

Fauna and Stratigraphy of the Stormberg Series, 439

les, hollowed out in the centre when the hard outer shell has been

broken through, are quite a characteristic feature of the Molteno

sandstones. The nodules are formed by the oxidation of pyrites and

the deposition of some of the resulting iron compounds in a spheri-

eal zone about the lumps of decomposed sulphide. The hard shell

is thus due to the addition of the hydrated iron oxides to the

cementing material usually present.

The lowest of the coarse glittering sandstones has been termed

the ‘‘Indwe Sandstone’, and forms a reliable bench-mark from

which the horizons of the different coal outcrops can be defined.

The finer grained varieties of sandstone .... are of a yellowish

erey or cream colour and furnish a good building stone.

The coarse gritty sandstones occasionally become conglomeratic,

the pebbles consisting principally of vein-quartz and of quartzite.

A peculiar feature is the occurrence in the Molteno sandstones of

smooth rounded or oval pebbles usually a few inches across but

occasionally ranging up to boulders a couple of feet in diameter.

They are, as a rule, scattered irregularly through the sandstones,

but in the Molteno Division they are particularly abundant along a

certain horizon and form a bed of conglomerate a few feet in

thickness. The pebbles are sometimes found resting upon a_ coal

seam and partly imbedded in the base of the sandstone overlying

the coal. The pebbles are almost entirely of white or brownish,

sometimes glassy, quartzites like those of the Witteberg or Table

Mountain series. They are most abundant to the south-west of the

Stormbergen. Some of these pebbles show pitting externally due

to the formation of cubes of pyrites a layer of which occurs just

below the surface.

In the Molteno Beds there are numerous outcrops of coal, but

the workable seams are restricted to three well-defined horizons.

The lowest one is that of the Indwe seam; to this belong the coals

at Indwe, Cala and that near Engcobo. The second is about eighty

feet higher and is known as the Guba seam. The uppermost is the

horizon of the Moltena seam, 300 ft. above the Indwe seam.

The layers of coal seldom exceed twelve inches in thickness (in

the Guba seam there is one about 25 inches thick), but as several

usually occur alternating with thin bands of black shale it is pos-

sible to extract from three to four feet of coal in mining operations.

On all three horizons these composite seams appear to occupy a

number of detached areas, in between which the coal is either

replaced by shale or else is entirely absent. In most cases this is

due to non-deposition of carbonaceous material, but sometimes to

4AO Annals of the South African Museum.

erosion of the matter deposited, contemporaneous erosion, a pheno-

menon which is seen in thousands of cases throughout the Karroo

beds. At Indwe the upper layers of coal and shale are in places

missing, and the surface thus denuded is overlain by massive sand-

stone with pebbles at its base.

The coals of the Molteno beds are usually laminated and contain

very thin streaks of shale; they are coals which where formed very

probably at a considerable distance from the spot where the plants

erew, and the alternation of thin layers of coal and silt evidently

points to the vegetable matter having been deposited over the floor

of the basin in the same manner as the silt.”

In addition, it is well to give a few details with regard to the

stratigraphy im the various Divisions in which the beds have been

studied.

Glen Grey, Queenstown d Wodehouse Divisions. In this area the

Molteno Beds are essentially arenaceous in character more so than in

the country to the north. The Indwe Sandstone forms the most

important subdivision, and sometimes the second thick sandstone from

the base, the Gubenxa Sandstone, can be identified. There is a

general thickening of the beds towards the south and south-east, as

is evidenced by the fact that the thickness of the strata below the

Indwe Sandstone is 150-250 feet in Aliwal North, 450-500 feet at

Sterkstroom and Indwe, 700 feet at Cala, and 1000 feet at Lady

Frere.

The Indwe coal-seam is composed of a number of bands of coal

and shale, which are constant in character throughout the Indwe

mining area. The upper layers were in certain spots removed by

contemporaneous erosion. The ‘‘wash-outs” must have been formed,

according to Du Toit, by streams or currents of water flowing over

the seam of coal and shale while they were in a soft incoherent state.

The sandstone underlying the coal undulates, and thin layers of coal

and shale were spread over the surface and perpetuated the irregularities

beneath them. Erosion was most vigorous in the original gentle

troughs of the seam, and thus the top coals in these troughs are

no longer present, their place being taken by sandstone and grits.

In the Northern portion of Wodehouse near the top of the Beds

is a hard fine-grained white sandstone with a few Thinnfeldia fronds.

A similar bed is found at the same horizon in Elliot.

Aliwal North d& Herschel. The thickness’ of the Molteno Beds in

these Divisions is probably about 1000 feet. The Indwe Sandstone

Fauna and Stratigraphy of the Stormberg Series. AMA

lies about 200 feet above the upper limit of the Burghersdorp Beds

and forms a very prominent horizon, showing the same characters

as in the south. The Indwe Coal is not present, except that it is

represented in places by a hard black carbonaceous sandstone with

thin streaks of coal. The Cala and Gubenxa coals are represented

by 8 feet of mixed coal and shale. These lie 300 feet above the

Indwe Coal.

The upper beds are very similar throughout and consists of coarse-

grained pebbly sandstone, usually finer in texture than is the case

further south at Indwe in the Transkei. Higher up are fine-grained

sandstones with shales and mudstones, sometimes reddish in colour,

followed by soft pinkish felspathic grits.

~~ BASALT

(32; CAVE SANDSTONE ~ SEEN

ABouT 150-250’ CAVE SANDSTONE :

LG SL ON THABA'NCHU MOUNTAIN

= RED & BLUE

ABouT 450’ RED BEDS SANDSTONES

& SHALES ON SLOPES oF

= = THABA'NCHUS HILLS

5 Tet 2's: COARSE GRITTY SANDSTONE BE CUTE EES RU

aie & GREY SHALES & MUDSTONES WITH OCC COALY SEAMS

: YELLOW FINE GRAINEQ SANDSTONE

=S=— — & BLUE SHALES & MUDSTONES

MALTENO BEDS

ABOUT 300’

———ABouT MIDDLE SEEN IN CUTTING ON Rwy. JUST

500’ BEAUFORT BEDS E. OF THABA'NCHU STN.

THIN SANDSTONES & 4 BLUE & GREEN SHALES & MUOSTONES

SEVERAL THOUSAND FEET THICK

BETWEEN THABA'NCHU

& BLOEMFONTEIN

Fig. 48. Section in Thaba’ Nehu District, O.F.S.

Orange Free State. Little detailed work has been done on the

Stormberg Beds which le in the Eastern Free State between the

Orange River and Harrismith. It is certain that the Molteno Beds

are absent at the latter place where the Red Beds rest unconform-

ably on the Upper Beaufort Beds; and there is probably a gradual

thinning out of the formation northwards from the Orange River,

similar to that which has been traced by Du Toit in Natal. At

Thaba’ Nehu the beds are about 300 feet thick; the lower two-thirds,

according to a section supplied by the Irrigation Department, consists

of blue and grey shales and mudstones with occasional coaly seams,

the upper third of coarse gritty sandstone. At Verkikersberg, S.W.

of Memel, in the extreme north-east of the Orange Free State, the

Molteno Beds are also absent.

Transkei. In the Transkei the maximum thickness varies from

1800 feet in the south of the area to 1400 feet in the north. The

Beds are essentially arenaceous. They consist of thick layers of coarse

442 Annals of the South African Museum.

pebbly felspathic sandstone separated by bluish and grey softer

weathering, fine-grained sandstones, mudstone, and shale. The sand-

stone contains a good deal of felspar, fragments of which are frequently

'/, in. in diameter, while the grains of quartz frequently exhibit partial

restoration of the crystal faces, giving rise to ‘‘glittering sandstones”.

Small pebbles of vein-quartz are abundant in certain layers. Sporadic

pebbles of granite, graphic granite, and white fine-grained quartzite

occur, more commonly near the base of the series and frequently

just above, or even resting on, a coal seam.

The Indwe Sandstone and Gubenxa sandstone are well represented.

The top of the series here is a more than usually coarse pebbly sandstone.

Griqualand Kast. Du Toit has described the Molteno Beds in this

area from the Divisions of Maclear, Mount Fletcher, Qumbu, and

Mount Frere. The beds bear a close relation to those in the Transket.

In no place are they more than 1800 feet thick and are essentially

arenaceous. [ron pyrites is found in irregular layers in the sand-

stones in a few localities.

Du Toit describes the beds as having the following characteristics,

“The sandstones contain a good deal of felspar, usually rather de-

composed. The pieces of felspar may, in cases, attain a diameter of

half an inch, while in a railway cutting, about a mile south of Ugie

Station, small pebbles of granite and graphic granite were observed

in addition, The grains of quartz frequently show partial restoration

of the crystal faces, and the light reflected from these facets gives

the rock a sparkling appearance in the sunlight; hence the appropriate

name of “glittering sandstone” given to them. Small pebbles of white

or blue-black quartz up to an inch or thereabout in length are most

characteristic of these sandstones, and some portions are so pebbly

in character that they can almost be termed conglomerates.’

Further to the north-east, in Mount Currie on the Natal Border,

the thickness has diminished to about 1100 feet, even to 950 feet

just over the border. The layers of coarse pebbly sandstone are thinner

and spaced closer together than in Maclear. “Otherwise the formation

is the same, e.g coarse-grained pebbly sandstones and grits — usually

false-bedded and sometimes conglomeratic and crowded with smooth

water-worn quartz pebbles — rather felspathic and passing into grey

finer-grained types: most of the sandstone sparkles or ‘‘glitters” in

sunlight owmeg to the reflection of light from the faces of quartz

crystals”. The pebbles in the Indwe Sandstone range from a few

inches up to a foot in length. At one place there is a ferruginous

conglomerate full of quartzite pebbles. Below the Indwe Sandstone

Fauna and Stratigraphy of the Stormberg Series. AAS

bluish green and grey mudstones and thin fine-grained felspathic

sandstones occur. The higher horizons of the Series consist of grits

and pebbly sandstones, alternating with blue, grey and buff mudstones

and shales.

Natal. Along the eastern slopes of the Drakensberg ranges du Toit

has traced the Stormberg Series from the Cape Border to Van Reenen’s

Pass and Harrismith. At Hlatikulu Hill at the head of the Bushmans

River in the Division of Estcourt he has measured a thickness of

{40 feet of Molteno Beds consisting almost entirely of grits. At the

base the beds are false-bedded with ferruginous nodules; at the top

they consist of white coarse grits; and there are few thin shaly beds.

At Bezuidenhout’s Pass on the Orange Free State — Natal border

the Molteno Beds are absent; and they are similarly absent on the

Platberg near Harrismith.

Rep Beps.

Elliott. The Red Beds reach their maximum thickness in the

neighbourhood of the Barkly Pass where they are about 1600 feet

thick. The sandstones are fine-grained and irregularly distributed.

Bands up to 30 feet in thickness are common and are usually red

or purple in colour when unweathered. Conglomerate beds as a rule

are absent; but in one or two localities beds of sandstone contain

abundant white quartzite and quartz pebbles. The shales and mud-

stones which form the bulk of the Red Beds are red, purple, green

or white, and occasionally violet in colour. It is noticed that the

thicker sandstone bands very often lie upon eroded surfaces of the

softer beds.

Wodehouse. The base of the Red Beds contains coarse ‘‘glittering

sandstones” like those of the underlying Molteno Beds. The thick-

ness in the district is very variable. Just north of Indwe there are

1500 feet of sediments; but 12 miles to the west the thickness

diminishes to 650 ft. Further to the south it is again increased.

Again, between Dordrecht and Jamestown the thickness is generally

650 feet, while to the north-east in the lower Waschbank it is

900 ft. to 1000 feet.

The lower portion consists of banks of rather coarse-grained gritty

sandstone (pale yellow or bluish in colour) with interbedded yellow

and buff mudstones and thin purplish red sandstones, shales, and clays.

The sandstone beds are thicker, coarser, and closer together than

in Elhot,

Annals of the South African Museum.

—

The upper 400 ft. or so are brillantly coloured shales and sand-

stones, Calcareous rocks are frequent, occurring usually as irregular

nodules and concretions in mudstones.

Aliwal North. The thickness measured at Kraai River Poort is

900 feet. while to the north on the Herschel-Aliwal Boundary and

at Lady Grey it has fallen to 600 feet. The general features are

similar to those found further south. At the base of the beds a thick

yellow sandstone separates the Molteno Beds from the overlying

purple and red shales and mudstones. The succession Is well displayed

at Lady Grey, where Kynaston noted that the sandstones appear to

become finer-grained as one ascended in the series. Occasional layers

of caleareous concretions occur, which sometimes contain bone—

fragments.

Barkly East. Only the top portion of the Red Beds is seen in

this mountainous district. The beds have a general dip to the South,

whilst to the north in Herschel, they dip in the opposite direction.

Du Toit noted the presence of much silicified wood on the farm

Glencoe, the tree stems being of considerable size.

Herschel. The Red Beds are well displayed along the north side

of the Wittebergen. The thickness at Palmiet Fonte (near the

Basutoland Border) and between Kromme Spruit and Majuba Nek is

at least 900 feet.

At the base are red and purple mudstones, weathering to light

blue, which have yielded a number of large Dinosaur bones. At the

summit. at Dulcie’s Nek, is a bed of limestone. Between, the strata

are made up of very brillant red shales and clays with prominent

red and occasionally buff sandstones. The sandstone band which occurs

at about one-third the way up the succession as seen at Josana’s

Hoek appears to be persistent and will probably prove a useful

‘“bench-mark” for palaeontological purposes.

Pebbles are very uncommon in the sandstones. During two fairly

extensive collecting expeditions which examined most of the outcrops

in the district only two reddish small semi-rounded pebbles of quartzite

were seen.

Fossil wood is found in certain localities, occurrimg in the thin

sandstones. Some of the trunks are large One section 9 feet long

now in the South African Museum has a diameter of over 2 feet.

It was found near the base of the formation at Kromme Spruit lying

in a thin sandstone at a low angle to the bedding planes. Other

Fauna and Stratigraphy of the Stormberg Series. AND

also

specimens of similar and even greater diameter were obtained

from near the base of the beds — at Blikana.

‘

Orange Free State. The Red Beds are well exposed on the eastern

border of this province, being especially well seen between Thaba

*Nehu and Modderpoort, around Ficksburg, and between Fouriesburg

and Harrismith. Fossils have been collected at Fouriesburg, but no

details of the Beds are available. Red colour predominates, but at

Ficksburg the base of the formation consists of very thick grits —

which, however, may prove to be a local excessive development of

the Molteno Beds,

The Red Beds are exposed on the sides of the Thaba ’Nchu hills

Cave Sand slona

While medium grained Sandstone

oe S aeited and greensk shales, muds/ones and soft while grits

—__ -Read fo pu rplish Soft felspalhie grits Red. Beds

a — ~-Fed and purple Shales

~~+-Hard gritty sands/one

Fig. 49. Verkijkersberg. S.W. of Memel, O.FS.

about Tweespruit. They are about 450 ft. thick and are predominantly

argillaceous, consisting of red and blue shales with at least one prom-

inent sandstone band.

Du Toit has studied the section shown by the Platberg, near

Harrismith and he has kindly sent me details of this unpublished

study. At the base of the mountain are red and purple shales with

medium-grained sandstones, representatives of the Beaufort Beds —

a slightly lower horizon of which has yielded a Middle Beaufort fauna

on the Harrismith Commonage. Lying above these are 290 feet of

strata which he assigns to the Red Beds, there being thus an unconformity

between the Beaufort Beds and the Red Beds, the Molteno Beds being

absent. At the base of the Red Beds is a thin but gritty sandstone,

This is followed in succession by over 100 feet of soft purple shales,

AAG Annals of the South African Museum.

a massive fine-grained sandstone with thin soft bands, and at the

top are red and purple shales. It must be pointed out that the un-

conformity is not evidenced by any apparent discontinuity in succession,

nor is there any difference in dip between the two formations.

Further north, at Verkijkersbere, S.W. of Memel, the thickness of

the Red Beds has fallen to 160 feet. The red clays of the Upper

Beaufort Beds are succeeded by a hard gritty sandstone, often quartzose

at the base and this in turn by a fine-grained sandstone. Then

follows a series of red to purplish soft felspathic grits, red and greenish

shales and mudstones and soft white and red grits. The top of the

formation is formed of white medium-grained sandstones. The basal

sandstone is variable in thickness and rests sharply on red mudstone:

Mx - - Columnar Dolerile

fs "3 -- -Cave Sands/ane

|--/assive fine-grained sands/one with Shin Soft bands Rel Ber.

-Soft purple shales

ed — — Sandslona . TAin but grilty

A Red and purple shales

= ” Medium -grained Sand s5/ane

Fig. 50. E.S.E. corner of the Platberg, Harrismith, O.F.S.

portions of it, especially at or near the base, are quartzose grits and

show false-bedding dipping in a South-Westerly direction.

Midway between Harrismith and Memel, on Tandjes Berg, the Red

Beds are intermediate in thickness.

Transkei. In the Transkei the thickness is uniformly 1200 feet.

The sandstones are gritty at the base of the series, occasionally carrying

isolated boulders of quartzite; higher up they become finer-grained

and yellow. The softer mudstones and sandstones and shales are

of brilliant red, purple, and blue tints, weathering pale. Many of

the so-called “buff” sandstones owe their colour to weathering, being

red on a freshly-fractured face.

Griqualand East. In Maclear and the Divisions bordering it to the

North and East the thickness of the Red Beds diminishes from South

to North. having a maximum of 1200 feet. Du Toit has described

a section showing the full succession shown in the ascent from

Beaufort /

Fauna and Stratigraphy of the Stormberg Series AAT

Pot River to Tent Kop in Maclear. This shows predominating purple

and red shales, mudstones and soft sandstones often of remarkably

brillant colourmge — which bleach on exposure. Coarse grits and

occasionally pebbly sandstone occur at the base, and not uncommonly

there are found boulders of quartzite like those of the Molteno Beds.

The most prominent sandstones are usually white in colour, and in

places contain nodules of iron pyrites or marcasite. Many of the

sandstones when fresh are red; they are commonly full of porous

patches which represent spots originally rich in calcareous material.

Clay-pellet conglomerates are not uncommon at the base of certain

sandstones. Fossil wood is occasionally seen. Vertebrate fossils are scarce,

In Mount Currie the thickness is only 400 feet. The beds, as else-

Sr asale

~ Grelty Zone

"5 ek ag ae Cave Sandsléne

---Massive medium gramed $.p wilh gritat base

=-~While and dark red. $.§, with red muds/ones

“\- —- While coarse gril

sabes false-bedded, with fern nodules

} Upper Beaufort Beds

Fig. 51. Hiatikulu Hill, near Table Mountain, Natal

where, consist of several bands of fairly fine-grained sandstone alter-

nating with blue, red, and purple mudstones and soft sandstones.

Natal. At Hlatikulu Hill the Red Beds have a thickness of 430 feet

resting on the Molteno Beds. Shales overlie the top white coarse

erit of the Molteno Beds, and contain one or two gritty bands. They

are succeeded by white and dark red. sandstone with red, purple and

blue mudstones and soft sandstones; these by dark purple mudstones.

Then comes a massive medium-grained sandstone with grit at the

base, and between it and the Cave Sandstone are reddish shales.

Further north, at Bezuidenhout’s Pass on the O.F.S. border the

thickness has diminished to 400 ft. The formation rests directly on

purple mudstones of the Upper Beaufort Beds and has at its base a

bed of gritty and pebbly sandstone which, in its lower portion jhas

quartz-pebbles up to '), an inch in diameter as well as pellets of shale

and some sandstone fragments. The mass of the Red Beds is made

-~Dark hurple muds/one Red Beds

Molteno eds

4AS Annals of the South African Museum.

up of an alternating series of sandstones, which get finer-grained

towards the top, and purple mudstones. The sandstones are white

and the two uppermost bands contain flattened pellets of pale blue-

green shale.

. . a > ®

The junction between the Upper Beaufort Beds and Red Beds appears

to be an unconformable one, but is without angular discordance.

CAVE SANDSTONE.

Elliott. The Cave Sandstone here is a massive yellowish fine-grained

felspathic sandstone of remarkable uniformity of texture. It is slightly

stratified at its base, and sometimes more so at its summit; but

lamination is usually absent. Some sections show false-bedding on

200' ae Feseea. ct eh Cave Sandsfone

| _- Whihsh fine-grained sandslone B

a >Purple mudslone and red gritty Sandslone R eds.

Viery \- 77+ -Whélish medium-grained Sands/ane

_Whilith gritly f-s.

2\- ~__ Redand purple muds/snes

“= \-- - Grify and pebbly S.S.

=~ - Purple mucds/ones Upper Beaufort Bec

Fig. 52. Bezuidenhout’s Pass, O.F.S. border.

an extensive scale. When fresh, the rock is pinkish or reddish in tinge.

Its maximum thickness, as displayed at the Barkly Pass, is 800 feet.

Occasionally the sandstone is replaced over short distances by thin

yellow sandstones and purple mudstones; similarly in Matatiele it is

replaced locally by red clay, often mottled with green or alternating

with light blue clay. At the top, the rock is sometimes interbedded

with lavas and ash.

Barkly East, Wodehouse, Aliwal and Herschel. In these districts the

rock is a fine-grained sandstone, usually pale yellow in colour, but

varied by white, buff, pink, and blue. It is generally unbedded, but

in places its upper portion is laminated. False-bedding is not un-

common, and is sometimes extensive.

In places the sandstone is split up by layers of ash and_ basaltic

lava. The thickness is very variable, ranging from 150 feet to 800 ft.

Du Toit, in his Report on these Divisions, compares the Cave Sand-

Fauna and Stratigraphy of the Stormberg Series. 4AQ

stone with the massive white Jurassic sandstone from the Grand

Canon region of Arizona.

Under the microscope the rock is very uniform in texture, chiefly

made up of grains of clear quartz only slightly rounded. Angular

fragments of orthoclase and microcline occur, as well as plagioclase,

small grains of zircon, epidote, and sometimes tourmaline.

At Siberia, in Wodehouse, a shale-band near the base of the

formation has yielded a fragmentary fish, many specimens of Lepidurus,

numerous Cyzicus and Ostracods, and several forms of insect. In

other places silicified wood is not uncommon.

Structurally, the sandstone is seen to be eroded and faulted before

the main outpourings of the lava.

At Lady Grey, Aliwal North Division, where the Cave Sandstone

is very well seen, the lower bed of sandstone is approximately 80 feet

thick and is underlain by a few feet of soft reddish sandy shales.

This bed is without sign of bedding planes, but exhibits well-marked

vertical jomting. Above it is 400 ft. of sandstone clearly exhibiting

a rude stratification in its upper portion and noticeable false-bedding

at more than one horizon.

Orange Free State. Along the western border of the Drakensberg

mass the Cave Sandstone constantly appears above the Red Beds.

On Thaba’Nchu mountain it varies in thickness from 150 feet to

250 feet. Its thickness at Harrismith is 250 feet, where it is overlain

by columnar dolerite, and at Verkikersberg 500 feet. Its features

are constant, a creamy or white massive sandstone forming the main

body of the rock. At Fouriesburg the base of the formation is red

and purple passing into white, so that it is impossible to draw a

lithological distinction between the Cave Sandstone and the top of

the Red Beds. From this level have come the types of Gryponyx

africanus and Massospondylus harriesi as well as other lightly-built

forms. A shale-band in the Cave Sandstone of Harrismith yielded

Cyzicus draperi in various stages of growth.

Transkei. At the extreme North-east of the district the formation

has a thickness of 800 feet; but a few miles away it thins to 50 feet.

There is here an interruption of deposition caused by volcanic out-

bursts. At Tent Kop the Cave Sandstone is altogether absent, the

lavas resting directly upon the Red Beds. These ashes dip beneath

the Cave Sandstone of the neighbouring areas.

Lithologically the Cave Sandstone is uniform throughout the area

white to cream in colour, sometimes deep pink or red towards the

ADO Annals of the South African Museum.

base. It is almost invariably fine-gramed and is composed of grains

of quartz-sub-rounded to angular in outline — with grains of felspar

and mica and small crystals of zircon, garnet, and rutile.

Griqualand East. The thickness in this area is very variable. In

the North-East, near the Natal border, it has a maximum thickness

of 800 feet, while westwards it thins in one place to 50 feet. In

the Tsitsana Reserve its thickness is 300 feet yet at places near by

falls to 100 feet. This variability is due in part to the outpouring

of lavas before the close of the formation of the sandstone — in

places sandstone is found intercalated with lava-flows.

The sandstone is generally uniformly white to creamy in colour

occasionally being pink or red especially towards the base. The basal

portion is well-bedded and at a few places rests unconformably on

the Red Beds. The base frequently exhibits false-bedding.

Natal. At Hlatikulu Hill the Cave Sandstone is 190 feet thick,

and at the top has a gritty zone overlain by basalt.

According to Churchill (1898) the thickness varies in the stretch

between the head of the Bushman’s River and Mont aux Sources

from 200 feet to 600 feet, while at the south end of Thaba’Mhlope

it is 800 feet thick. The rock there is compact, hard and gritty,

usually cream or white in colour, but sometimes light red, and

occasionally carries a “few round, hard sandstone nodules, often cont-

aining a little pyrites’. At the base of the formation is a 6-10 ft.

thick bed of a rather friable, light-coloured marly sandstone resting

on a 5—15 ft. thick bed of nodular sandstone. This may be taken

as the base of the Cave Sandstone as it rests on a “deep, pink,

earthy layer”.

Transvaal,

The Stormberg Series in the Transvaal is preserved as a number

of outliers, forming the Bushveld Series, of which the most important

occur on the Springbok Flats, the Komati Poort Coalfield, in the

area North of the Zoutpansberg, and in the Limpopo Valley, The

features of the series in each of the areas will be briefly outlined.

Springbok Flats. Mellor in 1905 gave an account of the sandstones

of Buiskop and the Springbok Flats, which are overlain by the

Bushveld Amyedaloid. He stated that the sandstones of the Springbok

Flats are universally rather fine in grain and uniform in texture,

rarely, if ever, gritty, and contain no conglomerates or pebblewashes

except, possibly, at the extreme base. They are peculiarly massive

Fauna and Stratigraphy of the Stormberg Series. 451

and homogeneous and only rarely show traces of bedding planes, and

then only at wide intervals. The colour varies from deep red to

almost pure white or cream. The red sandstone is usually hard and

quartzitic; where weathered and soft, it loses its red colour and

Fig. 53. Sketch-map of Transvaal to show distribution of Bushveld Series.

becomes yellowish or white. The sandstone is distinguished by well-

marked vertical joimting.

West of Naboomspruit Station the sandstone rests directly on granite,

but elsewhere upon the Coal-Measure Grits. It has a thickness there

of about 30 feet. Its upper portion is massive, but towards the base

it shows frequent traces of stratification, and the lowest beds consist

of distinct bands of sandstones alternating with coarse gritty and

A452 Annals of the South African Museum.

conglomeratic beds which are composed of debris and fragments of

eranite. In places the fine-grained sandstones lie directly on the

eranite without intervening conglomerates.

At Slypsteen Drift the sandstones are even in texture, fine-grained

and often massive; some beds show a tendency to lamination and

are at times false-bedded. Wm. Anderson, in a paper published in

1912, described the occurrence of fossiliferous beds ‘exposed in the

water-channel of the Compies River, in the vicinity of the store at

Stypstee Drift, Springbok Flats, Waterberg District”. ‘“Stypstee Drift”

is presumably a misprint, and should read “Slypsteen Drift”. Regarding

this occurrence Anderson writes “As this is the only postion in which

I observed outcrops of these fossiliferous sedimentary beds, I have

no evidence as to their probable lateral extent or distribution, because

the country to the north-east, west and south-west chiefly consists

of extensive areas of alluvial deposits, through which occasional out-

crops of Recent calcareous rocks are not uncommon .... To the

westward of Stypstee Drift, under the alluvials of the Springbok Flats,

these fossiliferous sedimentary beds probably become associated with

the amygdaloidal basalts, which form a portion of the Upper Karroo

Series, and are well developed in the western and south-western

portion of the Flats. It is, however, probable that this occurrence

of fossiliferous Upper Karroo beds is not continuous with those of

the west, but has been formed in an isolated basin. Similar, probably

contemporaneous, sandstones and shales occur at the foothills of the

western limit of the Springbok Flats, the sandstones occasionally

attaining a considerable individual thickness, as at Buiskop, to the

north of Warmbaths, where the rock has been extensively quarried

for building purposes.” According to Anderson, the beds at Slypsteen

Drift rest unconformably on the Red Granite of the Bushveld. In

the main section the lower exposed strata consist of practically

horizontal exceedingly fine-grained, light grey, argillaceous shales.

The sandstones above are markedly false-bedded and much jointed

vertically. ‘In all cases the fossil bones occur as the nuclei of

ferruginous nodules and not in a free state in the sediments. They

occur more frequently in the nodules from the sandstones, but are

more fragmentary than are those from the shale nodules. On all

the exposed outcrops these beds do not show the slightest evidences

of deposition under violent climatic conditions, although the presence

of frequent bone fragments in the ferruginous nodules, which have

evidently been much worn by attrition before they were deposited

in the sediments, would rather incline one to the idea that there

must have been around this lake-basin areas in which considerable

Fauna and Stratigraphy of the Stormberg Series. 453

erosion took place, in producing the material for these sedimentary

beds. It is, however, difficult to explain why the eroded bones should

find a place in these quietly deposited sediments, while the rock

products of the same erosion are not present either as conglomerates

or as isolated pebbles or boulders”. Anderson saw no sign of plant

remains; he pointed out that the ferruginous material segregated

around the bones which varied in size from an inch to three feet

in length. He considered that the bone-bearing nodules were con-

fined to more or less distinct horizons on which the individuals are

fairly plentiful. Large bones and small bones are mixed together

indiscriminately — the two chief horizons being one amongst the

lower shales and the other some distance higher.

Dr. Broom, who examined these remains, could not be certain of

their specific identity. He considered it not improbable that they

were all representative of one species, pointing out their resemblance

to Gressly saurus and their possible identity with Huskelesaurus brownt

or Huskelesaurus capensis.

Neither Mellor nor Kynaston recorded determinable fossils, but

in a bore hole at Ludlow 2555 specimens of Cyzicus (= Estheria) (?)

were found in sandstone. Van Hoepen has described also a new genus

of Theropod — Giygantoscelus — from bones in the Transvaal Museum

which came from Haakdoornbult 544 in the district of Waterberg to

the west of Pienaars River Station.

Komati Poort Area. The Bushveld Sandstone of the Komati Poort

coalfield was described by Kynaston in 1906, In that area, fine-grained

sandstones with distinctive features lie between the Coal Measures

and the amygdaloidal basalts.

The sandstones are usually without signs of stratification. They

are very fine-grained, even-textured throughout, soft, pale greyish or

yellowish, sometimes pinkish in colour, sometimes mottled with darker

spots. At one outcrop numerous spherical concretions up to 9 in.

in diameter were seen, consisting of a hard shell around a softer

interior.

Towards the base, the series becomes calcareous, and is often crowded

with irregular lumps or nodules of finely crystalline limestone. Below

this occur thin-bedded, soft, dark-red and greenish sandy shales and

marls, the red colour predominating. The total thickness of the

series is JOO feet. No organic remains have yet been discovered in

the beds.

Northern Transvaal and Valley of the Limpopo. Mellor, in 1908,

454 Annals of the South African Museum.

described the geology of the N.W. Zoutpansberg District in a Memoir

of the Transvaal Geological Survey. He found that the Bushveld

Sandstone series is especially conspicuous along the Limpopo Valley

in the neighbourhood of Rhodes’ Drift, from which point it extends

east and west for many miles, forming prominent ridges rising 200

to 300 feet above the river. The sandstones vary in colour from red

to yellowish white, are fine-grained, even in texture and sharp to

the touch. They are usually extremely massive and frequently re-

presented by a single bed from 30 ft. to 50 ft. in thickness without

divisional planes. Occasionally, however, false-bedding on a very

large scale traversing the full thickness of the rock and dipping at

angles as high as 20 degrees is brought out by weathering.

Below the upper and harder portion of the massive sandstone and

erading upwards into them there is almost invariably found about

15—20 ft. of sandy or marly mudstones, usually light green to purplish

in colour, which contain numerous concretionary masses of limestone,

varying from an inch to 2 or 3 feet in diameter. This lower marly

portion of the sandstones usually weathers out into caves. Downwards

the marly rock passes into red or purple sandy shales and soft sand-

stones, calcareous in places, and frequently mottled with light green

patches. The average thickness of this lower series is about 200 feet.

‘Where the sandstones form very prominent ridges and kopjes they

are frequently found to have been much hardened by secondary

silica, usually deposited along the numerous joint planes. In some

cases the jomt planes and fractures are so numerous that the whole

rock becomes a breccia.”

Dinosaur remains have been discovered in these beds. On the

farm Wiepe 1258, Mr. Bowker found a number of bones which

were described by Broom as Gryponyx transvaalensis.

In the west of the Waterberg Division of the Transvaal Bushveld

Sandstones occur in the area between the Limpopo and its tributary

the Pongola and apparently extend a little way across the river into

the Bechuanaland Protectorate. According to information supplied

by Dr. du Toit the sandstone rests on Coal Measures which them-

selves lie unconformably on Waterberg Beds or the Old Granite;

and Nelson’s Kop is capped with a volcanic flow. The Nelson’s

Kop sandstone — more fully described later — is an approximation

to the Forest Sandstone type.

Dr. du Toit has recently presented to the South African Museum

a few bones which he obtamed from the beds at Slypsteen Drift

on the Limpopo. They are, unfortunately, unrecognisable specifically,

consisting of isolated phalanges and a portion of a caudal vertebra.

aie

Fauna and Stratigraphy of the Stormberg Series. ADD

Their size, however, is comparable with that of similar bones of

Plateosaurus or one of the larger genera of South African Theropoda.

Southern Rhodesia.

Molyneux (Quart. Journ. Geol. Soc. 1903, LIX, p. 279) gave the

first provisional classification of the Upper Karroo rocks of Southern

Rhodesia. He suggested the following: —

“Thaba ’Sinduna Series 200 ft. Sandstones and volcanic rocks of

Thaba ’Sinduna and Shiloh.

Forest Sandstones . . 1000 ft. Fine sandstones of the forest-

country, with sandy clay. Tra-

vertine on the surface. Bubi,

Gwampa and Sikonyanla basalts,

Conglomerate basement near the

Djombi River.

Escarpment Grits. . . 400 ft. Coarse red sandstones, with sub-

angular pebbles, as seen in the

ereat escarpment which _ stret-

ches from the Mafungabusi Moun-

tains to near Wankie.”

Maceregor (in litt.) says that the beds classed here as Forest

Sandstone are largely Kalahari, and therefore of more recent age

than the Stormbere.

In an account of the geology of the region round Wankie, Light-

foot (1914) found 100 feet of Forest Sandstones — whose top was

not seen — lying on 300 ft. of the Escarpment Grits. The ‘Forest

Sandstone” of this area is said by Macgregor to be equal to his

Nyamandhlovu Sandstone. Lightfoot conjectured a break at the base

of his so-called Forest Sandstone and, if Macgregor be correct, this

break would probably correspond in time to the formation of the

Forest Sandstone of the Bulawayo area.

_ Macgregor (1916) described the Forest Sandstone of the typical

area North of Bulawayo. He divides the sediments into four groups;

4. Nyamandhlovu sandstones intercalated in basalts.

3. White sandstone.

2. Red marls.

1. Basal beds, resting on old schists ete.

Maufe (1919) described briefly the Upper Karroo Rocks of the

Amanxele Hills in the Bembesi basin north of Bulawayo. There

456 Annals of the South African Museum.

buff-coloured sandstone lies unconformably on a floor of granite and

greenstone schists.

The latest paper by Molyneux (1919) described the succession in

the Pasipas area N. of Bulawayo. He divides the sediments as

follows: —

SanGdstonen seen eee oeante

eee Sees eae) SOU elite

Arid climate. .

Nyamandhlovu

\ pasalt 2 eee cee OOHiG:

oreuP. | Lava flows and

Aeolian sands. Transition sandstone 40 ft.

Upper division. Fine, — pulverulent

sandstone, interst-

MEV GEM eG Gul ees AUR Ahi

Middle division . Marlsand sandstones 30 ft.

Fine calcareous sand-

Lower division . stone with calecrete.

Limestone . . . 1410 ft.

Basal beds. . . Partially sorted ar-

kosewy,. soe ear LOE

Forest

Sandstone

In petrological features, the Escarpment Grits consist throughout of

a “coarse grey grit, containing banks of pebbles, which are often as

large as eggs. It is not bedded very regularly, false bedding being

common, and the rock usually breaks along a pebbly bed. The pebbles

are all well rounded, and are mostly of quartz, but granite and gneiss

pebbles occur occasionally. The rock is well jointed, and the joint

faces are usually coated with a dark brown ferruginous skin.” (Lightfoot)

The Forest Sandstone of the typical area shows the following

features. The Basal Beds are conglomerates containing a large per-

centage of calcium carbonate, and are typically white in colour.

Besides containing derived pebbles, siliceous concretions are very

plentiful. The under surface of the beds is very irregular and in

places the hollows are filled with a fine red marly sandstone lying

beneath the conglomerates. In a specimen kindly sent me by

Mr. Macgregor a small piece of bone was detected. This is the only

fossil so far recorded from this conglomerate. The Red Marls have

a maximum thickness of 4 feet. They are pinkish red to chocolate

brown in colour, laminated, and very friable. Hitherto they have not

been found to contain fossils. The White Sandstone follows the Red

Marls. It is a massive white rock composed of angular grains of sand

cemented in an opaline matrix. No bedding planes occur except just

Fauna and Stratigraphy of the Stormberg Series. 457

above the marls, where there is evidence of a minor unconformity

showing contemporaneous erosion, In one place strong current bedding

indicating currents from the south is to be seen. In another place an

exposed surface is ripple-marked and apparently sun-cracked. Cal-

careous concretions and nodules are common. The upper part of the

bed is usually dead-white, but it is sometimes stained pale pink.

Fossils have only been found at two places. These were vertebrae

and portions of a fibula of a small Dinosaur, belonging probably to

either Thecodontosaurus or Gyposaurus. The Nyamandhlovu Sandstone

is intercalated with sheets of basalt which overlie the preceding bed

unconformably. The sandstone is false-bedded, of a deep-red to pale

chocolate colour, and generally coarse-grained. It is well laminated

and cleaves into broad flags. The grains of the sandstone are well

rounded and of fairly uniform size. (Macgregor.)

The Forest Sandstones of the Wankie area, as described by Light-

foot, are much finer-grained than the Escarpment Grits, and contain

pebbles only rarely. The sandstone is of a deep red colour and is

very much false-bedded, forming flaggy bands. It rests on the Es-

carpment Grits in irregular patches, some of which are large, and

these when viewed from a distance appear to be outliers. In thin

section the rock closely resembles the slides made from the stone at

Pasipas, near Bulawayo.

The rocks of the Pasipas area were described by Molyneux. He

describes the sandstone as made up almost entirely of quartz grains

of a common size of 0°2 mm. in diameter, a few reaching 0:7 to 1 mm.

The rock is generally soft and easily excavated and, unless indurated

by the process of silicification or protected by the sheets of overlying

basalt, disintegrates rapidly. The normal rock is composed of fine

erains of quartz joined by interstitial felspathic clay in the upper

division and by carbonate of lime in certain lower beds. The basal

beds generally represent the weathered state of the rocks of the Ar-

chaean complex; at other places there is a variable amount of sorting

by the action of moving water (storm water). The lower division of

Molyneux consists of unbedded deposits either with calcareous cement

or lenticular beds of limestone, the deposits being mainly sandstones

occasionally with calcareous balls and nodules, with a thin layer of

pale brown or pink marl. The middle division consists of coloured

marl interbedded with sandstones. The upper division is characterised

by the non-calcareous nature of its interstitial clay.

Mennell (1904) says that the sandstone of Thaba ’Sinduna seen

under the microscope shows sand grains that are often beautifully

rounded and include fragments of perfectly fresh felspar (microline

458 Annals of the South African Museum.

etc.) which indicates that it is largely of windworn material.

A section made from a rock kindly supplied by Mr. Macgregor

shows distinctly two sizes of grains a large rounded grain of dia-

meter about 05 mm., and a somewhat polygonal, smaller grain which

is also rounded, of about 0-1 mm. diameter. The latter make up the

bulk of the rock, which is from Esipongweni, about 20 miles N. of

Bulawayo and about 30 feet below the first basalt. The larger grains

are nearly all quartz. The smaller grains consist partly of fresh

felspar, mainly microcline. Some of the quartz grains show strain

shadows under crossed Nicols. There are occasional small grains of

apatite and what is possibly tourmaline. The appearance of the grains

is closely similar to those of the present-day Kalahari sand, save that

there is no ferruginous coating.

Molyneux has pointed out that towards the top of the Forest

Sandstone the beds contain increasing quantities of coarser rounded

erains mixed with the others, As the Transition Sandstone is reached

the grains are still coarser and dark grey in colour and are all well

rounded, cohering together by oxide of iron.

Apart from the fragmentary Dinosaurs mentioned, the only Storm-

berg fossils found in 8. Rhodesia are plant remains which have re-

cently been discovered by Mr. Macgregor and have been sent to

Professor Seward for determination.: I am indebted to Mr. Macgregor

for a preliminary note on this occurrence (in litt.). He writes “I had

the fortune to obtain some fossils from the lower deposits (at Soma-

bula). There are forms resembling Thinnfeldia and Phyllotheca so |

take them to be Stormberg provisionally. I believe the whole deposit

except a surface implementiferous rubble to be Karroo, but there

are two types very distinct at Willoughby though interbedded at

other places. The older deposits are red and pink mudstones and

micaceous sandstones with white fossiliferous sandstones occurring as

lenticles, and gravel with very well rounded pebbles at the base.

The newer deposits cut channels through the older sometimes into the

granite beneath. They are essentially coarse ill-consolidated arkoses

with inbedded gravels. The chief poimt is that the base of the deposits

resembles the base of the Forest Sandstone and the upper beds re-

semble the Escarpment Grits. There is nothing at all like the white

Forest Sandstone. The fossils were at the bottom”. Macgregor cons-

iders the Escarpment Grits to be probably of the same age as the

Somabula gravels as they have the same highly rounded quartz and

quartzite pebbles.

Finally, Molyneux has seen white sandstone with basalt on top

overlying Escarpment Grits in the Mafungabusi district, 60 mile N.N.W.

Fauna and Stratigraphy of the Stormberg Series. 459

of Somabula. If this white sandstone is Forest Sandstone, then the

succession of the beds seems clear. The Escarpment Grits (with a

white sandstone carrying Thinnfeldia below them in the Somabula

area) are followed by the Forest Sandstone (seen at Mafungabusi),

which is white in its lower part and red above, and this passes up

by progressively coarser sandstones of more and more rounded grain

through the Transition Sandstone into the Nyamandhlovu Sandstone.

Where the Escarpment Grits are absent, as in the district north of

Bulawayo, the base of the Forest Sandstone is a conglomerate

followed by a Red Marl. If, on the other hand, the sandstone of

Mafungabust represents the top of the Forest Sandstone or the Trans-

ition Sandstone, then the Escarpment Grits of the Wankie area would

seem to equal, in part at least, the Forest Sandstone of the type

area. In any case, the correlation with the Stormbergs of the Cape

seems clear. Thinnfeldia is characteristically a Molteno plant, although

it occurs sporadically in the Red Beds; and the Dinosaurian remains

are closely comparable with those from the top of the Red Beds or

the Cave Sandstone. The lithological changes are also comparable

with those seen in the Cape. Coarse pebbly sandstones typically give

place to finer-grained sandstones with a local intermediate develop-

ment of red marls; and the climatic changes are in the same direction

as those in the south of the continent, although at any given time

conditions seem to have been more arid in the north than in the south.

Writing of the Forest Sandstone Molyneux says “The rocks of the

group show none of the usual planes of bedding of aqueous deposits,

but are thick deposits of sand grains of uniform size without any

sorting of coarser material... The basal beds are made up of

debris resulting from the weathering of granite in situ or which may

have been partially sorted into different sizes by storm waters, and

in this manner the deeper hollows of the Archaean landscape were

levelled up. From this stage onward there is evidence of climate of

increasing aridity. The angular shape of the smaller grains in the

rock shows that there has, in the lower division especially, been little

abrasion or rounding in transport. The deposits are no doubt due to

winds that carried dust and particles from a very dry area. To do

this atmospheric movements need not have been different from those

of to-day ... The calcrete that occurs in the lowest beds of the

Forest Sandstone points to a climate of semi-aridity, but as time went

on desert conditions approached and there seems to have been pro-

gressive dessication of the whole country. The marls probably re-

sulted from the deposition of fine material through the drying up of

standing water — not a lacustrine condition in the manner of lakes

460 Annals of the South African Museum.

connected with silt-bearimg streams, but after the type still known

in Africa as that of promiscuous pans or “‘vloers” described by du

Toit. During the stage of the upper division aridity increased; there

was not sufficient moisture to segregate the lime, but the vegetation

that existed was still of sufficient growth to prevent the accumulation

of dunes. But complete desert conditions were not long delayed, for

during the transition period the wind-borne debris was sorted by

wind rolling into sand dunes. The Nyamandhlovu epoch must have

been one of complete desolation. Volcanism manifested itself in the

flow of the sheets of basalt of great extent. Dust continued to fall

and would be absorbed in the moving lava, but directly that flow

ceased in any locality and the surface hardened the wind resumed

its work of sorting and rolling, with the consequent formation of the

interbedded sandstones.” It is worthy of note that the ‘current-

bedding” of the Nyamandhlovu sandstones is paralleled by similar

phenomena in the upper portion of the Cave Sandstone, especially

when it is interbedded with lava flows.

Maufe has shown that in places where the Forest Sandstone les

on granite and greenstone schist floors, the rock apparently fills up

valleys in the old pre-Karroo floors, beimg in part banked up against

the sides of those valleys.

Belgian Congo.

The basin of the Congo forms a gigantic depression which is formed

of tilted and folded sedimentary rocks and schists of Palaeozoic and

Pre-Cambrian age resting upon Archaean granites and gneiss. At

the bottom of this hollow is a series of continental formations which

rest unconformably upon the older rocks. These le, in the main,

horizontally or dip at feeble angles. They have been grouped by

Prof. Jules Cornet into

3. Lubilache Beds (‘Lubilash” of some authors)

Lualaba Beds

1. Kundulungu Beds.

The Kundulungu Beds are considered by Cornet to be of ‘*‘Permo-

Carboniferous” age; but the other two groups are thought by the

same author to be the equivalents of the Upper Karroo Beds of

South Africa, and it is these which must be considered here.

Ball and Shaler (1910) describe the whole series above the Kun-

dulungu as the Lubilache Series with a vertical thickness greater

than 1500 feet, while in the Kasai region their Lubilache rests on

Fauna and Stratigraphy of the Slormberg Series. 461

old rocks without the intervention of the Kundulungu series, and

has a thickness of fully 700 ft. near Luebo and of 800 ft. on Lomani

with a dip towards the centre of the basin. Mathieu hints that

Ball and Shaler may be correct in surmising that the Lualaba and

SSS ert |!

BSF;

me |

Lubilacs Beds.

Lualala Beds.

Kunde lunga Dads.

Ancient folded rocks

Fig. 54. Sketch-map of Belgian Congo to show distribution of Karroo System

(from Revue Zool. Afric.).

Lubilache Beds may possibly be only phases of the one series.

Passau, Horneman and others agree, however, with Cornet in sepa-

rating the two series, and Passau has shown that Cornet separated

the Lualaba from the Lubilache by means of a distinct carbonaceous

shaley zone below the sandstone of the latter. These carbonaceous

shales occur over a wide area, e.g. at Stanley Falls, the Lomani

462 Annals of the South African Museum.

Valley, at Lusuna and Lusambo along 5° Lat., and also in the east

in Kasai.

Lualaba Beds. The Lualaba Beds have been described by Cornet

as consisting of clays, grits, sandstones, limestones with little coher-

ence and often oolitic, etc., and have been shown to cover a large

area in the Congo Basin.

Studt in 1913 gave the following as an ideal complete section of

the Lualaba Beds:

12. Mottled clays & mudstones — black, white and red . 100 ft.

11. Coarse red micaceous & felspathic gritty, pebbly sand-

StOMESTES piss ic tt to) a ee pees Ree Oem iy dere Letts

10. Greyish calcareous shales & thin aragonite bands . . 10 ft.

9. Black shales with pyritic coals, & coarse felspathic grits 140 ft.

8. Light grey calcareous false-bedded & banded sandy shales — 80 ft.

7. Pyritic black shales and gritty sandstones. . . . . 10 ft.

)

6. Light grey calcareous banded & fluted sandy shales . 60° ft.

5. Red pebbly sandstones, grits & conglomerates with

intercalated beds of brown shale ..... . . 160 ft.

4. Light pink to grey sandstones & shales. . . . O0mt

3. Pyritic black shales & coals with intervening lenathne

OTIS ee coat ee Mion oe ; ee crud ceeer ad AU ang

2. Light red to yellow atten eae saniletanes &shales 20 ft.

Ll; ‘Grey*calcareous ibreccia. 1. anes) <5 s-e-tiee) ere en ee OO it

Studt’s bed no. 12 is not the carbonaceous shale mentioned by

Passau, and it is possible that his beds 10, 11 and 12 may belong

to the Lubilache series.

Passau in a paper on the beds in the area north of the Kasai

along the Ulindi and Elila subdivides the succession as follows: —

o. Banded arenaceous shales with indeterminate

Lubilache ) plants.

4. Clayey banded shales wlth nodules and pebbles.

3. Black clay shales — graphitic, carbonaceous and

micaceous with plant fragments.

2. Clayey shales with pebbles,

1. Soft greenish clayey sandstones passing into

conglomerate with very large inclusions gener-

ally of subjacent rocks.

Lualaba

He states that the Lualaba has a more restricted occurrence than

the Lubilache, on account of the overlap of the latter on the former.

Hitherto, the Lualaba Beds have yielded only a few fossils — fish

Fauna and Stratigraphy of the Stormberg Series. 403

remains and Entomostraca — which have been briefly described by

Leriche.

At Kilindi, at the junction of the Lualaba and Lindi rivers, fish

remains were found ina soft, whitish argillaceous-calcareous sandstone

which lies horizontally — the ‘middle calcareous shale zone” of Passau.

This zone may correspond with bed 8 of Studt’s classification. Leriche

described the remains as Peltoplewrus maeseni.

From white limestones at Kindu on the Lualaba two forms de-

scribed as Pholidophorus corneti and Lepidotus (?) sp. by Leriche were

obtained. In the latest paper on the Palaeontology of the Congo (1920)

Dr. Leriche describes a large number of fragmentary fish remains

under the name of Lepidotus congolensis, a species already described

by Hussakof on material from the Congo basin. These remains are

from calcareous clay from near the base of the Lualaba Beds in the

vicinity of Stanleyville. He also describes a single scale attributed

to Colobodus from just north of Ponthierville. Cornet also mentions

the discovery of fish debris in bituminous shale in the neighbourhood

of Ponthierville.

Entomostraca are common in some localities, e.g. at pots on the

railway between Stanleyville and Ponthierville, almost covering the

surface of the layers on which they are found. Leriche examined

them, and described a Phyllopod Estheriella lualabensis Leriche, and

an ostracod Darwinula globosa var. stricta R. Jones. This latter variety

occurs in the Rhaetic of Scotland and Leriche considers the Congo

specimens to be indistinguishable from the Scotch form. The same

form also occurs at Songa, 43 km. below Ponthierville, in bituminous

shales; on the banks of the Oviatoku (a tributary of the Lualaba)

in beds full of the ostracod Metacypris passaui Leriche; at Bamanga,

14 km. below Ponthierville; and at Kindu in clear limestones. From

a study of these forms Leriche concluded that the Lualaba Beds were

of Upper Triassic age. Cornet, on non-palaeontological grounds,

correlated them with the Beaufort Beds of the Karroo System, con-

sidering, however, that the Lualaba Beds were a complex whole, in

which one should be able to distinguish several zones.

Messrs. Ball and Shaler obtained fossils from several localities in

the series (which they described under the name of the ‘“Lubilache’’)

viz., indeterminable plants from shale at Niangwe, 200 ft. above the

base of the series: ostracods of the genera Cypris, Candona and pos-

sible other Cypridae, a valve of Hstheria, and fragmentary fish-remains

from limey shales 10 miles below Stanleyville and 150 (?) ft. above

the base of the series; and a possible Estheria valve from soft sand-

stones at Sangula at the confluence of the Buschinmai and Sankuru

464 Annals of the South African Museum.

rivers. (Of the organic nature of the last, Cornet is doubtful; and

Leriche places the locality in the Lubilash formation). Ulrich, from

an examination of these fossils says ‘*The bed from which the fossils

were procured is Mesozoic and Jura-Triassic, rather than later.”

Ulrich further states that if the fossils indicate anything concerning

the climate it would be that it was relatively moist and cool, and

that the water in which they were deposited was either fresh or

brackish.

Lubilache Beds. Studt describes these beds as being “reddish to

white friable sandstones and conglomerates containing numerous bands

of concretions so penetrated by secondary silica as to have the ap-

pearance of pebbles or boulders of granular quartzite, flint, or jasper,

while agate, onyx, and chalcedony are also common. The conglome-

rates ‘often contain black pyritic shale pebbles.” Ball and Shaler

also emphasise that ‘tas a rule massive bedding predominates in the

sandstones, as does a reddish colour; grayish and white beds are,

however, not uncommon”. Passau considers the thickness of the

sandstone to be from 3500-400 metres. In the Lubilash Beds at

Kitari, a red shale has yielded numerous carapaces of a Phyllopod

assigned by Leriche to Estheria sp. The same bed has also been

said to yield ostracods “recalling Darwinula globosa”, a variety of

which is described from the Lualaba Beds. The Estheria is said to

be sharply defined from Estheriella of the Lualaba Beds by the ab-

sence of radial ribs. The red shale in which the specimens were

found occurs as a thin band in the thick sandstones of the Lubilash.

Correlation of the Lualaba and Lubilash Beds with South African

zones is difficult on account of the paucity of the fossil remains from

the Congo. Of the fish from the Lualaba the only genus found in

South Africa is Pholidophorus which has been described from the

Burghersdorp Beds (Upper Beaufort). Lepidotus is a Semionotid;

Semionotus is a Cave Sandstone form. Colobodus is a 'Trias-Rhaetic

genus. The phyllopod Hstheriella lualabensis is apparently close to

E'stheria greyi from the Middle Beaufort Beds of Cradock. It seems

possible, therefore, that part at least of the Lualaba Beds may be

the equivalent of the Middle and Upper Beaufort Beds of the Union.

The lithological nature of the Lubilache Beds suggests instantly

correlation with the Stormberg Beds. Cornet first suggested this cor-

respondence, and Maufe and Molyneux both see resemblances between

the Lualaba Beds and the Forest Sandstone group of Southern Rhodesia.

Maufe writes, in criticism of Studt’s later view that the Lubilache

was of Waterberg age, ‘The author’s description of the Lubilash Beds

of Katanga might well be a description of the forest sandstones of

Fauna and Stratigraphy of the Stormberg Series. 405

Southern Rhodesia”; while Molyneux says “I agree that the Lubilash

series much resemble (in petrological features) the escarpment grit

and forest sandstones. Thus it must be that the Lubilash are my

escarpment-forest series, and of Upper Karroo age. The chalcedonic

segregations, agates, and other silifications, and the friable nature of

his fine sandstones are remarkably akin to features of the forest

sandstones of Mafungabusi Mountains, where there is no doubt that

they overlie the coal measures.” As has been shown, the Forest

Sandstone is almost certainly the equivalent of the Cave Sandstone.

More palaeontological evidence is, however, greatly desirable, as simil-

arity of petrological features merely means similarity of conditions

of deposition and need not of necessity imply similarity of age. The

Kstheria sp. described by Leriche seems to have a fairly close resem-

blance to the Cave Sandstone form from the beds at Siberia C.P.;

and their occurrence in a thin shaley band in massive sandstones is

certainly significant when compared with the mode of occurrence of

the latter.

Most writers on the geology of the Belgian Congo follow Cornet’s

original idea of the lacustrine origin of the Lualaba and Lubilache

series. For example, Ball and Shaler picture their ‘“‘Lubilache series”

as having been formed in a lake which was probably subsiding in

the middle, with low land to the west, north and south but hilly

ground to the east — the hills possibly rising to a height of 2000 feet.

Between these latter ran valleys, some deep enough and narrow enough

to be worthy of the name of ‘‘fiords”. No where was the lake deep.

Cross-bedding was common, as well as abrupt changes from sandstone

to shale. There was deep disintegration of the rocks of the shore-line

due to weathering. Cornet, however, writing in 1910 remarked with

regard to the “Lac Lubilachien” that his views had undergone modi-

fication since 1893. The conception of Lake Lubilache was a simple

one and provisionally admissible at that time. But the Lubilache

series, which is far from being limited to the stretch of the actual

basin of the Congo, is much more complex than was at first supposed.

It is possibly in part of lacustrine origin; but dunes and aeolian

sediments generally play an important role in it. The ‘“gres poly-

morphes”, which are so characteristic of the system are certainly

desert formations. Passarge follows Cornet in considering the Lubi-

lache Beds as the products of a desert climate.

er)

ior)

Annals of the South African Museum.

GENERAL CONSIDERATIONS.

CAPE-ORANGE FREE STATE-NATAL AREA,

Molteno Beds. The chief features upon which stress must be laid

in considering the Molteno Beds of this area are as follows:

The formation dies out to the North, thinning from 2000 feet at

its most southerly outcrop to 140 feet along the northern Natal-

Basutoland border and disappearing altogether at Harrismith; so that

its greatest North-South extent is just over 200 miles, an average

diminution of 10 feet per mile. How much further south it once

extended we cannot tell; the supposed occurrence of Molteno beds

at the top of the Great Winterberg, on the Fort Beaufort-Tarka

Divisional boundary has been disproved by a recent investigation

undertaken by the author. But that they extended well south of

their present outcrop is shewn by the occurrence on the coast near

Port St. John’s of a down-faulted patch at least 1600 feet in thickness.

The beds consist of sandstones, shales and mudstones which are gray,

greenish or bluish in colour, and lack any prominent calcareous con-

cretions or bands. In the south there is a great preponderance of

arenaceous beds, but towards the north the argillaceous deposits play

a more important, though still rather subsidiary part; it must be

noted that in Natal also the shales are inconspicuous. The sandstones

are coarse in grain, loose textured, and contain abundant felspar.

In the south they are coarser than in the north. The sandstones

occasionally contain nodules formed by the oxidation, and subsequent

hydration in the outer layers, of iron pyrites. Occasional conglomerates

occur, containing irregular boulders and pebbles which sometimes

rest on coal-seams partly imbedded in overlying sandstones. Such

pebbles are most abundant to the south-west.

The workable coal seams all occur in the lower portion. The coals

are in thin layers alternating with thin black shales. They occupy

detached areas, their absence from some areas being partly explicable

by non-deposition and partly by contemporaneous erosion.

The only fossils are plants, which seem more abundant in the

lower half of the formation. Animal remains have not yet been

definitely identified, although Dunn has recorded the presence of bones

at Molteno.

These features all point to deposition under deltaic conditions,

either those of a true delta or of an aerial delta.

Barrell has pointed out that the determination of ancient true

Fauna and Stratigraphy of the Stormberg Series. 467

delta deposits from the study of the strata requires the demonstra-

tion of evidence that both subaerial and subaqueous sediments were

deposited — the subaerial on the landward side of the strand-line

and the subaqueous on the seaward (either marine or epicontinental)

side. There can be no doubt that during the deposition of the Mol-

teno Beds the land which furnished the rock-waste lay to the south

and possibly in part to the east of the present outcrop; and, if a true

delta had existed, subaqueous deposits would be expected towards the

north. It would seem, however, as if the greater part of the features

of the beds can be explained better by premising merely terrestrial

conditions. At any rate, it can be said that terrestrial conditions

will explain the features without calling in the aid of any large body

of water.

Consider the features of aerial deltas in which purely terrestrial

deposits are being laid down. Such features vary according to the

nature of the climate, whether it be constantly rainy, intermittently

rainy, semi-arid, or arid. The late Joseph Barrell worked out the

criteria for each type, and to him we owe the following general survey.

In constantly rainy climates the deposits on piedmont slopes brought

down by rivers and floods carry a large amount of humus and forest

erowth. The soil and subsoil are continually wet and drainage carries

away the soluble elements. Thus deposits will contain little or no

iron, magnesia, lime, potassium or soda and, on the other hand, will

be rich in carbon. They will, on account of the diminished evapor-

ation and constant saturation, be white, black, or gray in colour.

Coal-seams will be abundant, moisture being necessary for their

formation. Barrell says “It may be concluded that the broad as-

sociation of carbon with sediments which are thoroughly decomposed

and leached throughout is the mark of continuously rainy climates

which are tropic or at least warm temperate; with sediments which

are imperfectly decomposed and incompletely leached the mark of

more or less continuously rainy climates which are in addition cool

or cold.”

When the climate is intermittently rainy much of the humus in the

soil is yellow or red, and the clays are slightly calcareous. When

yellow or brown flood-plain deposits are buried and lithified the up-

stream portions will be found somewhat more arenaceous, varying

from red to brown sandstones and usually including red, green and

occasionally black shales. Over the terminal land portions of the

deposit, the sandstones should be finer-grained and the quantity of

shales increase, becoming more gray, green and black. Thick wide-

spreading coal beds are impossible: but in occasional swampy areas

468 Annals of the South African Museum,

coaly shales will enclose and preserve the vegetation that grew in

the swamps. Elsewhere vegetable growth, being alternately wet and

dry, soon decays and is destroyed; casts of leaves and trunks in the

lighter-coloured shales and sandstones being the only evidence that

remains.

Under conditions of semi-aridity, the thorough seasonal oxidation

of nearly all deposits except those made in permanent pools, lakes

etc. results in the marked dominance of deep-red and brown shales

and sandstones, a moderate amount of variegated shales and a few

containing carbon, Lime will exist disseminated in noticeable amount

through both shales and sandstones, and may occasionally give rise

to markedly nodular or solid calcareous strata. The microscope should

show a noticeable amount of felspar in the finer portions of the rock,

as well as mica. “The most marked chemical distinction of sub-arid

flood-plain deposits from those of truly arid regions is found in the

small quantity of evaporation deposits of calcium carbonate, gypsum,

and salt, but especially of the two latter. Lime may be quite abundant,

as shown by the kankar of the Indo-Gangetic plain, its importance

depending largely upon the quantity in solution in the river water”.

In river deposits of semiarid climates casts of logs are most likely

to be preserved in the sands deposited in the neighbourhood of stream

channels. Away from these channels wetting and oxidation would

tend to destroy the logs.

In the flood-plain deposits of arid climates fluvial, pruvial and aeolian

formations are all of wide occurrence. The most distinctive structures

are: (1) the presence of mud-cracks filled with aeolian sands, the

mud-flakes being usually polygonal plates upturned at the edges (cf.

mud-flats of Orange River, at Kheis, as described by Dr. A. W. Rogers) ;

(2) interbedding of fluvial and aeolian sands; (3) the presence of

scattered and facetted pebbles.

It can be readily seen that the features shown by the Molteno

Beds are intermediate in character between those postulated for flood-

plain deposits laid down in constantly rainy climates on the one hand

and intermittently rainy climates on the other. The southern more

arenaceous facies represents the upstream portions; the northern the

downstream terminal beds. Du Toit has shewn that the thinning

northwards to the Natal border is due to actual thinning of the

various members, while north of that some of the members (the

upper) are missing, This is what would be expected from the sup-

position of a land to the south contributing rock-waste which was

laid down in the form of a fan at the foot of the mountains, the

amount of material becoming less as time progressed. The absence

Fauna and Stratigraphy of the Stormberg Series. 469

of red sediments and calcareous sediments precludes the possibility

of any arid or very dry intervals.

The abundance of felspar in the sandstones proves that decomposition

was not very complete, and argues a somewhat cool climate with

discontinuous rainfall. The presence of boulders and pebbles can be

explained by occasional torrential downpours following on periods

of comparative quiescence in which vegetation was able to flourish.

It is possible that part of the lower Molteno Beds at least are

formed of material that was accumulated on the land surfaces during

Upper Beaufort times. During the comparatively cold periods of the

Beaufort fairly deep disintegration of the rock-surfaces to the south

took place, resulting in the formation of a mass of undecomposed

felspar, quartz and other grains which, on the incoming of Molteno

conditions accompanied by earth-movement, formed the material

which built up the foundations of the Molteno Beds by transport

northwards.

The nature of the coal-seams and of the fossil plants associated

therewith is of considerable interest. The conclusion expressed by

Rogers and du Toit as to the formation of the seams requires some

modification. In the first place the often perfect preservation of the

plants, which are mainly delicate and fragile fronds of ferns and

fernallies, precludes the possibility of their having been transported

far from their place of growth by river-torrents. There is little or

no maceration shown by the fossils, except occasionally in the sand-

stones, and it is more reasonable to suppose that the plants were

buried near to the swamps in which they grew and flourished.

Such swamps occupied discontinuous areas in the region; for, al-

though contemporaneous erosion is the explanation of the lack of

coal seams in certain parts, their absence in others is most likely due

to an original absence of swampy ground. Such discontinuity in

swamp conditions is a feature of flood-plains in an intermittently

rainy climate rather than in continuously rainy regions. It is just

possible that the conditions were nearly the same as those described

by Davis from Turkestan where there is an alternation of inundations

and drought. During the former, and following it, vegetation temp-

orarily springs up, withers, and gives place to desert conditions until

the next flood. Such alternation would possibly result in thin coal

seams intercalated in thin sandstones or shales; but conditions of too

great dryness would not be favourable to the formation and preser-

vation of plant fossils.

Red beds. The most striking feature of the Red Beds is the colou-

470 Annals of the South African Museum.

ration of the sandstones, shales and mudstones which is predominantly

red or reddish-purple. Even those sandstones which appear pale in

the field prove, when freshly fractured, to have a decided red colour ;

and frequently, as along the northern face of the Wittebergen near

the village of Herschel, and at Lady Grey, the mudstones are_bril-

liantly crimson. In the Herschel division and in Aliwal North, the

clays near the base are frequently blue or even grey; but only the

few feet or so lying directly above the Molteno Beds show this

characteristic; they soon give place to typically red deposits.

The climatic significance of red is a subject which has engaged

the attention of several workers, especially in America, where the

development of the Permo-Carboniferous Red Beds and the Triassic

Red Beds has stimulated research. Barrell, discussing the subject

in 1908, considered that red colours of sediments are due to oxidation

at the time of origin of the sediments, ferric oxide being a component

part of the accumulating deposits. He pointed out that in moist

climates, heat and exposure all tended to the production of red _ soils,

but that red was also a feature of some deserts. He concluded that

the chief condition for the formation of red shales and sandstones is

merely the alternation of seasons of warmth and dryness with seasons

of flood, by means of which hydration is accomplished, This supple-

ments decomposition at the source of the rock-waste and that which

takes place in transportation in rivers. Wetting, drying and oxidation

decompose the original iron minerals and remove all traces of carbon.

Red shales and sandstones may thus originate in rainy, sub-arid or

arid climates without any close relation to temperature, and typically

as fluvial and pluvial deposits upon land; but the origin of such rocks

is most favoured by climates which are hot and alternately wet and dry.

The same author in 1913 pointed out that redness in rocks is no

criterion for the the separation of humid from arid climates, although

red beds are frequently the accompaniments of aridity.

C. W. Tomlinson, in 1906, dealt with the conditions of origin of

the Red Beds of Western U.S.A. He states “Where alternations of

light- and dark-red strata occur, the more’ deeply coloured beds are

in most cases of finer grain than the others. The occurrence of

coarse-grained massive buff sandstones in a series of maroon or cho-

colate shales has been noted by many writers. This association holds

true in many other Red Beds besides the group here under con-

sideration, Thwaites reports it as an almost unfailing relation in the

Lake Superior sandstone series of Northern Wisconsin, and Geikie

mentions its existence in the Triassic New Red Sandstone of Gt. Britain”.

Further, ‘the colour of prevailing red strata in the Red Beds series

Fauna and Stratigraphy of the Stormberg Series. 471

is due to the presence of ferric oxide. A grey or green colour sig-

nifies a low proportion of ferric oxide, and usually a preponderance

of ferrous over ferric compounds”. Tomlinson considers the theories

which propose respectively that the iron has subsequently been in-

troduced into the sediments from igneous magmas, and that it is due

to subsequent deposition from meteoric water; neither of these theories

has much evidence in its favour. Wherever intrusions have affected

Red Beds, their effect tends to be, not to heighten, but to destroy

the red colour; while the more impervious strata are usually redder

than the more pervious. The American beds, according to the same

author, owe thei colouration to the presence of ferric oxide which

was transported as such and deposited almost wholly as a mechanical

sediment, chiefly as a coating to sand-grains; but beyond saying that

the mauguration and cessation of red bed sedimentation was probably

connected closely with climatic and topographic changes involved in

the orogenic history of the continent no definite statement as to

climate is made.

Case has also considered in detail the formation of the Permo-

Carboniferous Red Beds of Texas, and shows there is nothing in them

to oppose the most generally accepted hypothesis that the red colour

owes its presence to the mature weathering of iron-bearing rocks in

a fairly humid region, with alternations of relative drought and humi-

dity. The clays of Texas have a solidity and density which would

not be present if the colouration were due to oxidation and dehy-

dration of the iron subsequent to its deposition.

Again, in his latest work (1919) he says that flood-plain deposits

of arid regions are marked by the presence of highly oxidised or

carbonated minerals with a lack of hydrous oxides or sulphides. This

is largely due to the normally low water-table, which permits the

penetration of air deeply into the soil, and the exposure of the animal

constituents to oxidation or carbonation. Also, the lack of vegetation

on an arid flat means a lack of carbon. The common result is the

prevalence of a red colour, the presence of gypsum associated with

the remains of terrestrial animals, and a lack of plant remains.

There is thus a consensus of opinion with regard to the theory

that the iron oxide, which gives the red colour, is due to the de-

composition and oxidation of iron-bearmg minerals and that it was

deposited in the red strata mechanically in its oxidised condition; but

its presence alone is not sufficient to make any pronouncement as to

whether the climate was humid or arid. If the red is not associated

with blue or grey or black, then the probability inclines towards the

arid climate or, more nearly, the semi-arid.

472 Annals of the South African Museum.

Attention might be drawn here to the red sandy soil of Bechuana-

land as described by Rogers (1906). The sand is often of a deep

red colour, almost like brickdust, but on approaching an area in which

calcareous tufa predominates the colour is seldom so intense. An

examination of the material under the microscope shows that the

grains are more or less rounded, especially those of quartz, and vary

in diameter from a quarter to 1 mm, in diameter. The red colour

is due to a coating of oxide of iron, which is removed by boiling

with HCl. The great bulk of the material is composed of quartz

grains, which show trains of inclusions and cavities. Plagioclase 1s

not uncommon, and is present in the form of cleavage flakes, with

the angles somewhat worn. Chalcedony and agate are rather rare;

zircon and magnetite are abundant, while epidote is a common con-

stituent. The composition of the sand shows that it has not been

derived from the disintegration of the rocks of the district alone.

The felspar and magnetite have been contributed by the disintegration

of dolerites. It is probable that the quartz grains have been brought

down from the Transvaal and the Orange Free State, Cape and

Basutoland by the Vaal, Harts and Orange Rivers. The sand depo-

sited along river-banks will be blown over the country by the pre-

railing N.W. wind. The decomposition of the diabase and dolerite

and of pyrites in shales yield compounds of iron which can be taken

into solution and deposited as oxide in the cracks and cleavages of

the sand grains and around the grains themselves.

In the Red Beds of the Stormberg Series we find that save for

occasional blue clays and whitish sandstones at the base, the various

members are all red in colour, Carbonates occur, sometimes freely,

as nodules, and in places form beds of limestone.

In the Maclear division, for example, the sandstone is commonly

full of porous patches or small hollows representing spots originally

rich in caleareous material, with here and there limestone nodules.

Gypsum and salt are absent a feature which distinguishes the beds

from the Wichita Series of Texas. Conglomerates are rare, such as

do occur being generally at the base of the formation.

At the base of the sandstone layers, however, bands of clay-pellet-

conglomerate are not uncommon. Such bands indicate a certain

amount of unconformity, the clay-pellets being formed by the rolling

and rounding of the possibly dessicated upper layers of partly con-

solidated mudstone which lie beneath the sandstones. That the mud

was dried by exposure before successive sediments were laid upon

it is indicated by the occurrence of layers showing sun-cracks (cf

examples in the South African Museum from Fouriesburg, O.F.S.);

Fauna and Stratigraphy of the Stormberg Series. 473

while du Toit has collected and recorded mudstones from the Red

Beds carrying worm-tracks, and the South African Museum possesses

examples of large Dinosaur tracks from Morija, Basutoland,

There is a pronounced tendency for the sandstones to become finer

in grain towards the top of the series. The author found in Herschel,

near the head of Bamboes Spruit, a local development near the top

of the Red Beds of a rock which has the appearance of being an

ancient silcrete or surface quartzite; and from a slightly lower hori-

zon a slightly reddish quartzite pebble was obtained, about an inch

long, wich looked like a ‘dreikanter” whose edges had been some-

what rounded. Save for occasional silicified logs fossil plants are all

but absent (Thinnfeldia and Schizoneura occur sparingly each having

been found hitherto at one locality only); and the fauna is entirely

a land one. Here it might be remarked that Tomlinson states “No

actual remnants of organic matter are reported to have been found

in red strata” — the tendency of organic matter is to turn red into

green. It is not known whether this statement still holds good for

America; but in the Red Beds of the Stormberg Series all the fossil

reptiles, with the exception of some large bones from the base, have

come from red clays and fine-grained soft felspathic red sandstones.

In most cases the bones are found disarticulated; but at Blikana in

Herschel, to mention one instance, a complete articulated skeleton

of the smallish form Massospondylus harriesi was found lying on its

side in red strata. The red muds especially are occasionally spotted,

somewhat sparsely, with green; and bones from Fouriesburg, O. F.S.

are surrounded by a thin layer of greenish rock; but there can be

no doubt the green colour is due to subsequent reduction of the

iron oxide by the agency of animal matter after deposition.

Moody (Quart. Journ. Geol. Soc. 1905), discussing the variegation

in colour of the Keuper Marls in England, considers — mainly on

chemical grounds — that “the variegation of marls is not to be

explained by the assumption that bleaching of the red rock has

occurred through reduction of ferric oxide and the loss of iron”; and

he considers that the even distribution of ferric oxide in the’ English

Triassic rocks is probably due to the action of chalybeate waters

permeating the whole of the sandstone and part of the overlying

marls. However, the intense colouring of the mudstones as well as

the sandstones of the Red Beds of the Stormberg Series and the

occasional association of green with fossil bones renders this theory

a somewhat improbable one to account for the colouration in

South Africa.

Microscopic study of the finer-grained sandstomes shows that the

474 Annals of the South African Museum.

sand-grains are not of the “millet-seed” type, but are sub-angular

in outline with their edges and corners slightly rounded. They are

uniform in size; and the ferric oxide acts as a cementing material

coating the grains. Mica is commonly present.

All these features agree closely with those postulated by Barrell

for flood-plain deposits in a semi-arid climate; the basal portion of

the Red Beds thus forming a link with the Molteno Beds — the

climate gradually changing from an intermittently rainy to a semi-

arid one, the aridity increasing as time went on.

A consideration of the horizontal and vertical distribution of the

animal remains is of some interest. The base of the Red Beds has

yielded fossils chiefly in the district of Herschel. They are all large,

heavy-limbed forms such as Huskelesawrus browni and Plateosaurus

cullingworthi while a little higher in the same area are still large

forms such as Melanorosaurus readi. As far as we are aware, no

small, lightly-built forms have been found at the base of the forma-

tion. As one ascends the forms become smaller and apparently

much more agile. Massospondylus occurs from about half-way up

to near the top, both in Herschel and in the north at Fouriesburg.

At the latter place it is associated with the somewhat larger but

still light-limbed Gryponyx africanus and a small Thecodontosaurus.

The southern end of the mass has yielded little from the base of

the Red Beds but large bones have been noted by du Toit in Elhot

and Maclear; the small Thecodontosaurus minor is from the upper

half, as are the extraordinary Cynodonts T’vritheledon and Lycorhinus,

while the supposed Predentate Geranosaurus is from the summit of the

formation. Erythrochampsa is from the very top also, and is another

very lightly-built small form, characterised by an armoured back.

Two features display themselves in this survey. The first is that

the animal type became progressively more agile; the other that the

majority of the fossils come from the centre and northern half of

the present exposures. This latter feature may be disproved by sub-

sequent finds; the southern portion of the area has not been searched

to any appreciable extent: and the outcrops are frequently grass-

covered and in bad condition for fossil-hunting. There can be no

doubt, however, that the first is of some importance; and it gives

assistance to the argument for a climate which gradually became

more arid. As conditions became more rigorous, there would have

been given an impetus towards agility and the ability to travel

longer distances, in search of sustenance; and it is significant that

deserticolous animals of to-day are long-limbed in structure.

It seems probable that the various animals did not live far from

Fauna and Stratigraphy of the Stormberg Series. ATS

their place of entombment. Although complete skeletons are not

common — one articulated skeleton only from the Red Beds is known

to the writer — the majority of the described forms are known

from a number of associated bones, some of which are frequently

found still articulated. Isolated bones are rare, and few show signs

of rolling or long transportation. At the base of the Beds, remains

of several large animals were found together in bluish clay near

Kromme Spruit, Herschel — apparently swamp-lovers whose remains

were washed by moving waters into some quiet swampy spot: a

supposition whose probability is increased by the discovery near by

of a large silicified log. In general, the bones of an incomplete

skeleton of a single animal occur together in one spot — a fact

which would be difficult of explanation if transportion over a long

distance be postulated,

Cave Sandstone. We have seen that the Cave Sandstone is a

massive fine-grained rock of varying thickness with bedding planes

but feebly developed. and that in the basal portion only. The rock

is generally white or cream-coloured, but often it is pink or red

and at its base is sometimes coloured as deeply as the underlying

Red Beds. Athough the massive portion of the formation is unbedded

it is often traversed by vertical joints.

The following description by du Toit of a specimen of the Cave

Sandstone from Rocky Dell, Maclear, C. P., may be taken as typical

of the bulk of the formation. (see Geol. Comm. Rept. 1910, p. 88).

“The rock is composed of grains from ‘05 to ‘O8 mm. across of

quartz and felspar, the former predominating. They vary in outline

from sub-rounded to angular and are sometimes elongated splinters

with sharp edges. Some of the quartz grains are quite clear, others

contain needles of rutile and dusty inclusions. The felspar consists

of orthoclase and plagioclase, either fresh or clouded — and_ kaoli-

nised; microcline is absent. There are flakes of somewhat altered

biotite mica, muscovite mica, and a good deal of secondary

white mica (sericite) in the felspar, around quartz grains, and some-

times within the quartz itself. A characteristic feature of the Cave

Sandstone is the presence of grains of zircon and in the slide there

are a number of worn crystals of this mineral, together with some

colourless garnet, and some grains of rutile. The groundmass of the

rock is fairly abundant, cloudy and dusty and probably for the most

part kaolin; in places it has a pinkish colour corresponding to the

pink mottling of the sandstone in the hand specimen”. “Another

section from the summit of the Hlankomo Mountain, Mount Fletcher,

476 Annals of the South African Museum.

shows nearly similar features, but microcline felspar is present in

addition and the rock contains somewhat more mica’.

Two main views have been put forward as to the mode of origin

of the Cave Sandstone. We will consider first that of Professor Schwarz

as developed in a paper in the Trans. S. Afr. Phil. Soc. Vol. XVI

(1905) p. 30.

Schwarz considers that the Cave Sandstone is a tuff, blown out of

the volcanic vents which opened the period of Drakensberg volcanic

activity, a portion of which still, in some cases, remains in the throats

of the vents. “If” he says “the Cave Sandstone was formed before

the production of the material in the pipe, then the latter ought to

be made up of portions of all the rocks which the vent traverses,

but we find that this is not the case, and that the material in the

pipe is identical with that of the Cave Sandstone. The Cave Sand-

stone, however, contains 83°5°%) of Si0,, with grains of quartz,

microcline, plagioclase, zircon, rutile, tourmaline, chlorite, garnet and

epidote, while the lavas of the Drakensberg — some of which flowed

out of the pipes — are basic in composition”. Therefore, says Schwarz

in effect, the pipes must have tapped deep-seated rocks such as the

eranites and crystalline schists — outcrops of which occur in Natal —

and the triturated material from them forms the Cave Sandstone.

The author of this theory does not believe that the Cave Sandstone

was formed by ordinary denudation from land-surfaces, as no land

surface. of a requisite nature was near enough at hand. He agrees

that the Molteno Beds are the detritus of a granitic region, presum-

ably of the southern prolongation of the Madagascar ridge; but argues

that the break in the deposition of the Molteno Beds marked the

disappearance of the source of the supply. In conclusion he states

“In the Cave Sandstone we have many peculiar features that could

be explained by the supposition that it flowed from the crater mouths

as a mud. It is hard otherwise to account for the immense thickness

of the embedded mass; it is hard to explain the sudden change of

great thicknesses of the white rock to a red clayey material; and

still more mysterious is the pseudo-bedding that one can see at

N’quatsha’s Nek, where the stratification is just such as would be

produced had the whole been stirred round in a gigantic pot like a

pudding”.

The same author, in his ‘‘Causal Geology” (1910) speaks of the

Cave Sandstone as a ‘“‘non-voleanic tuff’; and in ‘South African

Geology” (1912) says the Cave Sandstone” consists of rounded grains

of quartz and felspar which have been corroded on the surface and

enveloped with minute scales of talc; thus the ordinary aspect of a

Fauna and Stratigraphy of the Stormberg Series. 477

sandstone is entirely masked and the rock has the appearance of chalk.

The sandstone is not an ordinary sediment for the fine coating of talc

scales would have been soon rubbed off the sand grains if they had

been dragged along the sea floor by currents, and the coating could

not have formed after the sandstone was consolidated .... The non-

volcanic material torn from the granite walls of the chimneys would

issue as fine sand, and the corrosive action of the hot gases in the

vent would account for the alteration of the grains on their surfaces”.

The features shown by the Cave Sandstone, when considered with

the other members of the Stormberg Series, do not seem to call for

any such mode of origin as is outlined here. It seems that the

presence of Cave Sandstone infilling some of the pipes led mainly to

the formation of the theory. Du Toit has studied a peculiar pipe

of this nature and has published the following description,

THULE TRIG. STATION

8323

et So eee eS

Volcanic Ne

& o oO

Molteno

Fig. 55. Diagrammatic section through Voleanic Neck of Thule — after du Toit.

Length about 4 miles.

“A very interesting occurrence in view of the information which

it yields upon the conditions of deposition of the Cave Sandstone,

is the large voleanic neck on the farm Thule (Griqualand East),

almost on the crest of the Drakensbergen. It is nearly a mile across

with somewhat irregular outline, and occupies a hollow hemmed in

by Cave Sandstone, while several narrow ravines have trenched the

area and laid bare good sections. The peculiar feature in this neck

is that the Molteno Beds and Red Beds are lying quite undisturbed

right to the very edge of the pipe, but the Cave Sandstone, on the

other hand, commences to show an inward dip at a distance from

the margin of the latter, varying from about a quarter of a mile

on its eastern side to well over a mile on its west side. The sheet

of sandstone curves downward over the denuded edges of the Red

Beds, and in places possesses dips of as much as 40°, passing below

dark greenish agglomerate at the extreme south-west end, but

appearing to abut against the volcanic infilling at other points.

The material within the pipe consists of medium-grained agglo-

merate with fragments of basalt and amygdaloid of various kinds,

478 Annals of the South African Museum.

but the bulk of the stuff is of sedimentary origin, and in places it

passes into a pale yellowish-green sandy rock with few foreign

fragments, some of this passing by degrees into Cave Sandstone.

Fragments of Cave Sandstone and red and purple shales and sand-

stone are present, blocks of basalt up to 6 feet across and Jumps of

Molteno grit as much as 7 ft. in length.

The history of the pipe must have been somewhat as follows: —

At the close of the period of deposition of the Red Beds a small

volcano came into action on this spot and gradually enlarged its

boundaries, portions of the margin (composed of Red Beds) slipping

down at intervals into the vent. Then the Cave Sandstone com-

menced to be formed and was deposited at an angle over the denuded

hollow encircling the agglomerate neck. Sediment would become

mingled with the fragmental material within the pipe, while occasional

volcanic explosions would account for the ashy streaks and patches

in the Cave Sandstone at several points close to and outside the

pipe. On the south-west margin the fragmental matter most probably

represents an indipping patch of stratified ash overlying the sandstone”,

This occurrence is by no means an isolated one; and the matter

will be furthered discussed later in the section.

The other difficulties which stood in the way of Professor Schwarz’s

acceptance of the Cave Sandstone as a product of ordinary denu-

dation seem to disappear if we consider that the climate underwent

a further approach to aridity, and that from the onset of Molteno

times there was a gradual secular climatic change in one direction

throughout the formation of the deposits of the Stormberg Series.

We have seen that the base of the Cave Sandstone is generally

red and often bedded — this indicating a continuance of the flood-

plain deposits of the semi-arid Red Beds time. The massive portion

of the Cave Sandstone has, however, as du Toit (14918) has pointed

out, practically all the features of an aeolian deposit such as the

Pleistocene loess of the Northern Hemisphere.

In its typical form the loess of Asia and Europe is a fine-grained

deposit, consisting of minute particles of hydrated silicate of alumina,

quartz, felspar, mica, and other minerals, more or less cemented by

calcium carbonate, the segregation of which gives rise to concretions.

It is also often impregnated with alkaline salts, and nearly always

stained yellow with ferruginous matter. Its homogeneity of compo-

sition and structure contrasts it with all water-deposits. There is an

almost complete absence of any stratification, and the particles of

mica in it are uniformly distributed without orientation. A charac-

teristic feature is its well-marked vertical jointing. Its fossils consist

Fauna and Stratigraphy of the Stormberg Series. 479

almost entirely of land animals, distributed evenly throughout its mass,

Great numbers of bones of mammals occur locally — mostly belonging

to forms abounding on steppes and grassy plains.

The consolidation of such a deposit would very conceivably result

in such a rock as the Cave Sandstone. The grains in loess, like

those of the Cave Sandstone, are not rounded as those of desert sands,

but are described as “angular or sub-angular”. One striking diffe-

rence, however, is in the much greater abundance of hydrated silicate

of alumina in the loess as compared with the Cave Sandstone: another

in the number of tubules ‘‘marking the site of the roots of countless

generations of plants”; and a third in the absence of land-shells in

the Cave Sandstone.

Taking into account its various peculiarities, von Richthofen con-

cluded that the loess of China is the product of aeolian influences

acting during periods of long duration and under different conditions

of climate, of which three are distinguished in North China: (1) an

erosion period, in which the surface of the country was sculptured

by erosion and denudation into the figure it still retains beneath the

covering of loess; (2) a Steppe period, in which the conditions of the

saline steppes of Central Asia were extended over the whole of Northern

China; (3) a loess period, now existing, in which the former Steppe-

districts become converted into Loess-districts by the gradual accu-

mulation of dust at the surface, the dust being held in place by the

growth of vegetation. This theory of the formation of the loess can

be accepted for Northern China; but it does not fit exactly all the

requirements of Cave Sandstone formation.

In the first place there is the difference in composition between

the two rocks. The Cave Sandstone is felspathic; but the felspar is

but little decomposed and there is no such alteration product as

hydrated silicate of alumina in any large quantity. Sand grains play

the predominant part in the formation of the deposit. The chemical

nature of the sandstone and the comparative angularity of its frag-

ments indicate fairly arid weathering of the original land from which

the material was derived. In this, there are not true ‘‘loess-conditions”’.

Secondly, there is no indication in the Cave Sandstone of the little

tubes representing former grass-roots, nor any sign of extensive vege-

tation.

Silicified trunks of trees have been recorded from various localities

in the Cave Sandstone, as for instance at Morya and Masitisi in Basuto-

land; but most of the occurrences, together with the doubtful occur-

rence of coaly material near Belmore, Barkly East, seem to be in the

upper, sometimes laminated portion of the deposit.

480 Annals of the South African Museum.

This may also be looked upon as as mark of aridity. Richthofen

considered vegetation necessary for holding in place the dust of which

the loess is formed; but this is only so when the area of depositon

is itself subject to constant winds. If the dust is deposited as a

result of the slackening of the force of the wind which carried it,

then vegetation is not a necessary adjunct; and the lack of vegetable

remains argues conditions adverse to their growth.

The third difference is the absence of land-shells in the Cave

Sandstone. Most terrestrial mollusca require moisture for their de-

velopment and maintenance; and, although their absence from a

land deposit is a piece of purely negative evidence, it lends additio-

nal weight to the theory of aridity when so many other facts point

in the same direction.

In spite of these minor discrepancies, however, the ‘loess’ seems

to be the nearest among recent deposits to which we can liken the

Cave Sandstone. It is of interest to note that near the base are

highly irregular calcareous concretions similar to the ‘“‘loess-dolls”

figured by Wright.

The occurrence of mammalian bones in the loess is paralleled by

that of reptilian remains in the Cave Sandstone. All the forms

hitherto discovered are light-limbed probably cursorial forms —

Gyposaurus, Thecodontosaurus and the like, together with Notochampsa.

As has been shown, this latter was erroneously supposed to be a

Crocodile, but is in reality a light-limbed long-snouted armoured

Pseudosuchian. There is nothing of the water-loving, marsh-living

type in this assemblage; they are all very conceivably the inhabi-

tants of an arid or semiarid clime, adapted for comparatively rapid

transit from place to place. In no case has a complete skeleton

been found; but, on the other hand, isolated bones are rare and

the usual occurrence is in the form of articulated portions of a ske-

leton. It should be noted that, in the case of Notochampsa, the

skeleton is nearly complete, even some of the fragile bones of the

fingers and toes being preserved. Had the conditions been moist, it

is improbable that such would have been preserved in a sandstone-

water percolating through the porous sediment would have rapidly

dissolved the bones which have been preserved, as we believe, by

the rapid accumulation of a dry wind-borne coarse siliceous dust.

That the climate was not absolutely arid is shown by 4) the fact

that the deposit is in no sense a ‘“‘desert-sandstone” in the Cape —

O.F.S. area, and 2) the very rare local developments of greenish

and bluish shales containing a water-fauna. Of these, the best-

known is the shale-band in the Cave Sandstone at Siberia in the

Fauna and Stratigraphy of the Stormberg Series. 481

Wodehouse Division. Here is a 20 feet zone of finely-bedded blue

and green shale containing abundant fossils among which have been

recognised one imperfect fish, several examples of insects and nu-

merous examples of a small Ostracod, a Cyzicus in various stages of

growth, and of an <Apus-like Phyllopod, Lepidurus. The crustacean

part of this fauna is all of a semi-arid facies, capable of living in

damp or even dry mud for many months of the year even as they

or their near relations do in the Karroo to-day. Fish are also found

in what is supposed to be Cave Sandstone near Ficksburg, O. F.S.,

from which place slabs have been quarried with numerous specimens

of Semionotus capensis on each, the occurrence seeming to indicate,

from the fish point of view, a sudden catastrophic dessication of

their living quarters.

Further, the Rev. S. S. Dornan has recorded that many fossil

fish occur in the Cave Sandstone at Masitisi, Basutoland, as well as

silicified trunks of trees,

It is of interest here to consider a little more closely the physical

conditions of deposition, other than climate, of the Cave Sandstone

in the Cape-O. F.S. area.

Although in places there is a gradual transition from the Red

Beds to the Cave Sandstone indicating absolute continuity of depo-

sition from one to the other especially in the North, in other places

there is undoubted unconformity so that, while in some parts of the

area deposition was going on, in other parts erosion was being

effected at a greater rate than deposition.

As an extreme instance of this we may cite the fact that at

Glenelg in the Maclear District the Cave Sandstone rests directly on

pebbly grits of the Molteno Beds — the Red Beds having entirely

disappeared. Here it does not seem certain whether the Red Beds

were eroded subsequent to the deposition of the main mass of the

formation and before the deposition of the Cave Sandstone, by means

of a strong scouring agency, or whether the absence of Red Beds is

to be explained by a continuous ‘contemporaneous erosion”.

We have already referred to the unconformity which occurs in

the neighbourhood of the ancient volcano at Thule. At Tent Kop,

in the Maclear district, a bed of volcanic ash rests on the Red Beds

and underlies the Cave Sandstone; while other examples might be

cited showing that volcanic activity began, at any rate in the South,

before the deposition of the Cave Sandstone.

At Tent Kop the sandstone overlying the Ash bed is full of small

angular inclusions of indurated sandstone and shale.

That volcanic activity was rife during the formation of the Cave

482 Annals of the South African Museum.

Sandstone is evidenced by many examples, especially in the south

of the region and towards the top of the formation. In Maclear

two beds of sandstone are intercalated in the lavas. It is interest-

ing, too, to note that the ash here contains fragments of Molteno

Sandstone up to 5 feet across, and a few blocks of pre-Karroo quart-

zite. These have evidently been torn off from below and thrown

out by volcanoes in the neighbourhood,

Further north, in Barkly East and the neighbouring districts, in

a few places the Cave Sandstone is entirely absent, the volcanic

flows resting directly on the Red Beds. In the Barkly East Division

the formation is split up by numerous beds of lava and ash, both of

which are sharply defined from the sandstone. At Siberia in Wode-

house, three bands of sandstone are intercalated in the lavas. At

the head of Bamboes Spruit in Herschel there are four such bands.

Near the large Belmore volcano in Barkly East the Cave Sandstone

contains numerous masses of vesicular lava up to four feet across,

the lower surfaces of which consist of pipe amygdaloid. Du Toit

has described a section near Barkly East Township showing pinkish

sandstone resting on a bed of ash and abutting at one end against

a sheet of doleritic lava; in the sandstone are embedded rounded

masses of altered doleritic lava. At Waterfall in the Barkly Division

there is a section showing alternation of volcanic and sedimentary

material, both the sandstones and the lavas having a_ lenticular

character; there are pipe-amygdaloids at the base of some of the

lava-flows and rounded portions of lava in the beds of sandstone.

It is important to notice that in all the occurrences cited in the

last paragraph the intercalation of sandstone and volcanics occurs

above the main mass of the Cave Sandstone; also that the sandstone

in this upper portion of the formation is usually well laminated.

This fact, coupled with the occurrence of pipe amygdales in the lavas

where these rest on the sandstone, seems to show that the conditions

of deposition of the sandstone were somewhat different from those

of the main mass and were somewhat moister and less arid. How-

ever that may be, it is apparent that the Cave Sandstone was not

deposited uninterruptedly; in places the deposition was hindered by

outbursts of ashes and of lavas from the many volcanoes of the area.

Du Toit remarks ‘‘While the Cave Sandstone accumulated freely in

the north and was followed by lava flows, the ejection of ash in the

south was so considerable that full development of the sandstone was

prevented in that region, and it was only at a slightly later period

that lavas commenced to be erupted there’.

The large Modderfontein volcano in Aliwal North came into existence

Fauna and Stratigraphy of the Stormberg Series. 483

while the Cave Sandstone was being laid down, for in an outlier of

Cave Sandstone near by the rock is full of small fragments of shale

and sandstone that were blown out from the volcano.

Very little, or no, work has yet been done upon the area north of

the Orange River, but it is probable that the Cave Sandstone there

will be but little affected by volcanic flows — the volcanoes seemingly

having come into being progressively from south to north in point

of time.

The work of du Toit in the south and centre of the area has shown

that, presumably in connection with the outbreak of volcanic activity,

there was a certain amount of faulting and folding which has affected

the sediments but not the overlying lava flows.

In Maclear, the dips of the Cave Sandstone show that folding took

place during, and probably also somewhat previous to its deposition

and that they had ceased by the time that the area became flooded

with lavas. There are local disturbances of both the Red Beds and

the Cave Sandstone, but the overlying lavas are undisturbed, A certain

amount of subsidence caused the formation, too, of local hollows which

were filled with ash.

At Siberia, in Wodehouse, faulting on a small scale affects the Red

Beds and Cave Sandstone, but not the volcanics. In the Barkly East

Division the main bed of Cave Sandstone and the ash-bed are aflected

by small folds, producing domes and basins in the strata, and

ac-

cording to du Toit — there is clear evidence of a great zone of sub-

sidence formed during the eruption of the earlier lavas. A generalised

section along the Kraai River shows the Red Beds and the main-

body of Cave Sandstone with the ash-bed bent into an asymmetrical

undulating trough, while the lavas with the thin intercalated sand-

stone beds lie horizontally and undisturbed except in the east.

It is of interest briefly to consider the probable changes which took

place during Stormberg times in the land area from which these

sediments were derived.

That a land mass lay to the south and probably partly to the east

of the present area occupied by the Stormberg beds cannot be doubted.

How far to the south it lay we cannot say; but the occurrence of a

fair thickness of Molteno Beds near Port St. John’s indicates the

presence of a land-mass to the south of the present continent. Of

its petrological characters little can be said. The constituent minerals

of the Stormberg sandstones indicate derivation from granitic or

granitoid rocks in part. The lithological character of the original

rock from which sediments are derived is always, however, a matter

484 Annals of the South African Museum.

mainly of speculation unless the sediment is near the region of erosion

and other conditions are favourable for preserving the mineral con-

stituents of the original rock or rocks in an unaltered condition.

Chief interest centres around the probable physiographic changes of

the parent land-mass.

A general survey of the Stormberg sediments reveals a sudden in-

terruption of deposit of shales and fine-grained sandstones which took

place throughout Upper Beaufort times and the displacement of that

type of sediment by a series of grits, coarse-grained sandstones, con-

glomerate bands, and somewhat irregular large lenses of shales (some-

times coaly). This type was followed by a series of sandstones,

reddish in colour and becoming progressively finer-grained, inter-

bedded with red clays; and that by the fine-grained Cave Sandstone.

This succession can be profitably contrasted with that of the Siwaliks

of the Himalayas as described by Medlicott and Blanford (1879),

They say ‘Sandstone immensely preponderates in the sub-Himalayan

deposits, and is of a very persistent type from end to end of the

region and from top to bottom of the series. Its commonest form

is... of a clear pepper and salt grey, sharp and fine in grain, ge-

nerally soft, and in very massive beds. The whole Middle and Lower

Siwaliks are formed of this rock, with occasional thick beds of red

clay and very rare thin, discontinuous bands and nodules of earthy

limestone, the sandstone itself being sometimes calcareous and thus

cemented into hard nodular masses. In the Upper Siwaliks conglo-

merates prevail largely; they are often made up of the coarsest shingle,

precisely like that in the beds of the great Himalayan torrents... The

mountain torrents are now in many cases engaged in laying down

great banks of shingle at the margin of the plains, just like the

Siwalik conglomerates; and the thick sandstones and sandy clays of

the Tertiary series are of just the same type of form and composition

They conclude that the

”?

as the actual deposits of the great rivers.

Siwaliks were laid down as a fluviatile outwash from the rising

Himalayas.

We see here that the succession in the Siwaliks is the exact op-

posite of that shewn by the Stormberg Beds —- at least to the top

of the Red Beds. In the Siwaliks fine-grained sandstones and red

clays give place to coarse-grained sandstones, conglomerates and coarse

shingle-beds. If Medlicott and Blanford are correct in assigning the

Siwaliks to fluviatile deposition from a rising mountain mass, then

it may be presumed that the Stormberg Beds are possibly sediments

derived from a land-mass slowly lessening in altitude.

This presumption is further buttressed by certain theoretical con-

Fauna and Stratigraphy of the Stormberg Series. 485

siderations which have been fully set out by Barrell (1908) and which

will be briefly noticed here.

In youthful topographic stages rock-breaking dominates over rock-

decay; in topographic old age the reverse is normally true. In arid

climates, however, and to a lesser extent in cold climates, where chemical

action plays a less important part than in humid and wet climates,

there is dominance of disintegration over decomposition even in to-

pographic maturity or old age. Again, in the youthful stages of to-

pography the waste is carried away from the higher ground as soon

as disintegration frees it from the rocks, in arid and subarid climates

by the streams arising from torrential rains, and is deposited in inland

basins or on piedmont slopes. Under arid and subarid conditions this

waste is coarser, less decomposed and has less true clay than sim-

ilarly situated waste of more pluvial climes. Even in maturity, the

mountains of arid regions retain their nakedness, roughness and

sterility and give rise to disintegrated waste rather than decomposed

material. As, however, the difference in altitude between the parent

mass and the sediments formed from it diminishes, so will the trans-

porting power of the torrents lessen, and the sediments will tend to

become finer in grain — the larger fragments of disintegrated matter

being unmoved by the slower and less powerful streams.

In the old age of arid regions wind erosion becomes increasingly

more important than water erosion, so that the products of erosion

in old-age, when the desert surface is approximately flat, are chiefly

wind-borne loess and dune-sand.

A number of instances might be cited of present-day regions in

which one or more of these stages is seen; it is enough to quote

the Eastern Persian area described by Huntington, the aerial deltas

arising from the decaying high-lands of Arizona, and the sand-filled

valleys and ‘island mountains” of Namaqualand in South Africa.

It would seem that the evidence afforded by the nature of the sedi-

ments, coupled with these theoretical considerations and present-day

examples, tends to point to the conclusion that the land-mass from

which the Stormberg sediments were derived, suddenly rejuvenated

at the end of Beaufort times, was in process of planation during

the deposition of the beds until the intense volcanic activity put an

end to sedimention in the area.

On the other hand, there is little or no direct evidence of local

or regional earth-movement such as would have caused sudden reju-

venation at the end of Beaufort times; and the sudden onset of the

conglomeratic and coarse type of sedimentation which characterises

the Molteno Beds may have been due to more intense precipitation

486 Annals of the South African Museum.

of moisture resulting in torrential downpours and a consequent rush

of swollen streams at certain seasons. Even if that be a partial

explanation: of the phenomenon, it is certain that the land to the

south must have had a marked elevation above that upon which the

sediments were deposited. No intensity of rain-fall would cause

pebbles and coarse sand to be washed from one area on to another

at approximately the same level ; im other words, subaerial conglo-

merates are not features of wellmatured or senescent stream-deposits.

Transvaal Area.

The correlation of the various outliers of the Bushveld Sandstone

with one another and with the Stormberg Series of the Cape has

been based mainly upon petrological grounds and upon the fact that

they underlie the amygdaloidal lavas. It has been supposed that

they are but remnants of a once extensive mass; but, however that may

be, it is certain that they were laid down upon an uneven surface.

In the Springbok Flats area, north of Pretoria, the series is bent

into a double basin, and rests either directly on the Red Granite or

upon highly inclined and sometimes much broken and faulted sedi-

ments of the Waterberg System. The basal beds of the Bushveld

Series here are frequently composed of debris and fragments of

granite, felspar, and quartz with interstitial partly decomposed

felspathic material — derived from the Granite — and_ occasionally

of large rounded quartzite pebbles, probably from the Waterberg

Conglomerate. These coarse beds are confined to the edge of the

Series. Above them are in general, layers of red marl or shale;

and they are followed by the true Bushveld sandstone. In the west

and south of the area. however, the red marl overlies the so-called

“Coal-Measure Grits’.

In the Komati Poort area the beds dip to the east, sometimes at

an angle of 10 degrees; and this dip has been held to account for

the occurrence of the series at a level so much below that of the

Springbok Flats. Here the ‘‘Bushveld Sandstone” les, apparently

conformably, upon the ‘‘Coal Measure Series” — which le themselves

upon the older Granites, schists, and altered sediments of the Swazi-

land System in the following order: — Pale, hard, often coarse

felspathic grits, often massive with pebbly bands, at the base, fol-

lowed by sandy micaceous shales, carbonaceous shales, grits and

sandstones. Between these latter and the fine-grained sandstone

yellow shales occur at one or two points. The upper portion of the

Coal-Measure Series carries Glossopteris.

Garrard has traced this series in a north-south belt through eastern

Fauna and Stratigraphy of the Stormberg Series, 487

Swaziland to within 25 miles or so of the Zululand coal-field. In the

latter (St. Lucia field) the coal-bearing beds are separated from the

Basalts by horizontal beds of calcareous sandstone. At Somkele the

coal is considered by du Toit to be of Beaufort age, overlain by red

and purple shales which he thinks are: Middle or Upper Beaufort.

In the N. W. Zoutpansberg the Bushveld Sandstone rests either

on the old gneisses and schists or upon a variable thickness of Coal

Measures underlain in places by Glacial Conglomerate.

The palaeontological evidence seems to bear out the correlation

of the Zoutpansberg and Springbok Flats members with the Red

Beds and Cave Sandstone of the Cape Province area. Two questions

remain to be settled definitaly — 14. Is the fine-gramed sandstone

of the Komati Poort area really Bushveld Sandstone? 2. Is there

any Transvaal representative of the Molteno Beds? Kynaston has

already covered most of the ground, but the questions can be briefly

discussed.

4. The only recorded fossil from the Komati Poort area is Glos-

sopteris sp. from below the fine-grained sandstone. Here, in the

southern extension of the belt in Swaziland, and in Zululand upper

coal-seams occur near the top of the ‘‘Coal-Measures”. In the

St. Lucia coal-field the following plants have been found: — Glossop-

teris browniana var. indica, G. browniana var. angus'fulis, G. damudica,

G. retifera, G. acuta. G. spatulo-cordata, Phyllotheca sp. and Tueniopteris

spathulatum, This flora was held to be an Ecca one by Etheridge

jun.; but there seems little doubt that the assemblage is, as a whole,

typically Lower Beaufort. These beds are, however, separated by a

wide belt from the amygdaloidal lavas at Somkele, a belt which is

not yet at all known. If the coal-bearing beds at Komati Poort are

the equivalent of those at St. Lucia then two possibilities arise: —

(a) that the fine-grained sandstone at Komati Poort is true Bushveld

Sandstone and lies unconformably on the Beaufort Beds — just as

the Red Beds lie unconformably on the Beaufort Beds in the N. E.

Free State; or (b) that the fine-grained sandstone is itself of Upper

Beaufort age. The general nature of the sandstone, its mode of

weathering, its non-variability over a large district, and the known

unconformity at Harrismith and in the neighbourhood lend strong

support to the former alternative. The matter awaits further treat-

ment based upon more detailed field work; but it is important to

note that the similarity of the sandstones in the different areas points

to formation under similar climatic conditions; and these conditions

do not seem to have occurred in the southern area until the onset

of Red Beds times,

488 Annals of the South African Museum.

2. This question has been partly answered above. In the Cape

the Molteno Beds are a local phase corresponding to a certain type

of climate and certain geographical conditions which enabled coarse

pebble-beds, finer sandstones, and carbonaceous shales to accumulate

in a huge subaerial delta over a plain situated at the foot of an

upraised land mass to the south, In the High Veld of the Transvaal

there is no corresponding mass of sediment. During the deposition of

the Beaufort Beds in the south of the Union, the Transvaal area was

subjected to erosion and denudation (deposition only taking place

within certain basins), the sagging of the geosyncline in the south

possibly being compensated by a rising of the northern area. This

period of erosion, however, came to an end at some point during

Stormberg times; and in some of the basins formed on the unequal

land surface debris accumulated, to be soon covered by thin shales

and the Bushveld sandstone. There is evidence to show that warping

and sagging took place in the region north of Pretoria during Karroo

and post-Karroo times, which would account for the overlaps round

the margins of such basins as have been preserved. The basins, in

fact, owe their preservation to the sagging which took place. In

spite of minor differences visible in specimens from different places

the Bushveld Sandstone is of so uniform a character that it is

reasonable to suppose that the patches that remain are but relics of

a once more widely-spread layer that stretched up into Southern

Rhodesia. Like the Cave Sandstone it was subaerial and probably

mainly aeolian; and once the basins were filled up by the driven

and blown sand, the material spread out to cover a larger and

larger surface. We cannot estimate its maximum extent, nor con-

jecture what peaks of older rock stood up above the sandy levels.

We shall note later that there seems to be a lateral change from

a fine-grained “‘loess-like” deposit to a somewhat coarser, rounded

desert-sandstone. In this area the basal conglomerates may be the

equivalent of the Molteno Beds — they are equally the result of the

action of torrential rains able to produce temporarily streams strong

enough to carry fair-sized pebbles; while in the basal part of the

reddish clays north-west of Pienaars River Station the occurrence of

a very thick compound seam of coal lends colour to this suggestion.

The climatic conditions which gave rise to the Red Beds of the

Cape were shortlived in the Transvaal; and the climate rapidly

changed to one comparable with that reigning in Cave Sandstone

times further south; the onset of such conditions was possibly ear-

lier in the north than in the south. As a general conclusion, the

further north one goes from the Cape area the greater is the evi-

Fauna and Stratigraphy of the Stormberg Series. 489

dence of aridity furnished by the Bushveld and Forest Sandstones;

and it is of interest here to examine the .evidence afforded by a

microscopic examination of the rocks from the various localities.

The closest approximation to the true Cave Sandstone is possibly

afforded by the fine-grained sandstone from Komatipoort. This was

described by Kynaston as follows: — The rock ‘shows numerous

small quartz grains, of a generally uniform size, and on the whole

angular and sub-angular rather than rounded, embedded in an

exceedingly fine-grained matrix, in which individual grains are bar-

ely distinguishable. Grains of plagioclase felspar may occasionally

be noted..... The microscopic characters agree very closely with

those of the fine sandstones from the Springbok Flats, the propor-

tion of matrix to individual quartz grains being somewhat higher in

the latter.”

The Buiskop sandstone is of fairly fine grain, and there are very

few grains which are coarser than the general texture of the rock.

A considerable amount of ferruginous cementing material is present

in the red varieties; and the grains of quartz, although not splintery,

are polygonal in outline with rounded corners. The red variety is

very similar to the sandstones of the Red Beds.

A specimen of sandstone from Nelson’s Kop, in the west of the

Waterberg District near the Limpopo and Pongola Rivers, contains

a larger and a smaller. The larger

two types of quartz grains

grains are less prevalent than the other; they are rounded without

any trace of angularity and they vary in size from a grain having

diameters of O°3 mm. and 022 mm. to one with diameters of O°6

mm. and 0-4 mm. The smaller grains, of a diameter of 0-15 mm.

and under, are not so obviously rounded, but none of them are

splintery, although thin sections are somewat polygonal. Most of

the grains are coated with a thin reddish-brown layer of iron oxide,

while there are scattered grains of the same material. Felspar is

much less common than in the Forest Sandstone. The whole sec-

tion is reminiscent of that figured by Molyneux as a typical section

of the Nyamandhlovu Sandstone. For the sample I am indebted

to Dr. du Toit, who also supplied me with a piece of Bushveld

Sandstone from Castle Kopjes, on the main road from Louis Tri-

chardt to Messina.

The Castle Kopjes rock is light in colour and consists of equidi-

mensional grains of quartz with a very occasional grain of plagio-

clase. The grains are fairly well rounded, smaller than those of the

Nelson’s Kop stone and similar in size to those of Buiskop.

Reviewing the features of these Transvaal specimens, it would

490 Annals of the South African Museum.

appear that, in general, they are intermediate between the true

Cave Sandstone of the South and the Forest Sandstone and Nya-

mandhlovu Sandstone of Rhodesia — the more southerly outcrops

partaking more of the nature of the former, the westerly and nor-

therly ones more of the character of the Rhodesian rocks. Thus we

ean picture the loess-like formation of the south giving place gra-

dually to the true desert sands of the north, the fragments of the

Transvaal deposits which now remain to us representing the tran-

sition stage between the two types and shewing an intermingling of

the fine-grained angular fragments from the south with the rounded

desert sands of the north.

Age of the Stormberg Series.

Until fairly recently the age of the Stormberg Series was accepted

as ranging from Rhaetic to Lower Jurassic; within the last few

years, however, there has been a tendency to throw back the age

along the time-scale and to consider the Cave Sandstone as no later

than Rhaetic.

The evidence in favour of the older view was mainly two-fold,

based on the nature of the plants of the Molteno Beds and that of

Notochampsa from the Cave Sandstone. The latter was thought by

Broom to be a true crocodile allied to Pelagosaurus; and as croco-

diles did not appear until the Jurassic in other parts of the world

the Cave Sandstone was deemed Jurassic. The Molteno plants were

considered by Seward to be of Rhaetic age; and the Series was thus

spaced between those limits.

Detailed examination of the evidence leads, however, rather to the

acceptance of the later view. In the first place the Rhaetic age

assigned to the Molteno plant forms is based upon what is by no

means clear evidence and in view of recent discoveries the conclu-

sion may need considerable revision. Dr. du Toit is studying a large

amount of material mainly collected by himself, and although his

results are not ready for publication he has permitted me to say

that the undoubted association of Glossopteris, Chiropteris, Pterophyl-

lum, Callipteridium and apparently Rhexoxylon with the typical Mol-

teno genera Baiera, Thinnfeldia, and Taeniopteris give a very pro-

nounced Keuper appearance to the flora.

The vertebrates which it should be remembered come from. still

higher horizons, when studied in detail, also bear a Triassic aspect.

The three Cynodont forms are specialised relics from the Upper

Beaufort Beds. These latter, which underlie the Molteno Beds, have

Fauna and Stratigraphy of the Stormberg Series. 491

yielded the Stereospondyls Capitosawrus and Trematosaurus, forms

which are found in the Bunter (L. Triassic) of Germany.

Among the Archosauria the crucial Notochampsa istedana has been

shown to be not crocodilian but a Thecodont lymg apparently be-

tween Aetosawrus and the Crocodilia. The former occurs in the

Keuper of Germany, the latter first appear in the Lower Jurassic;

this would denote a Rhaetic age for the Cave Sandstone which has

yielded the intermediate form.

Erythrochampsa from the Red Beds has a crocodilian pelvis; but

too little is known of the form to be certain of its affinities, and it

has thus no bearing on the discussion. Sphenesuchus from the Red

Beds is a Pseudosuchian of somewhat specialised form, It is pro-

bably somewhat later than Aetesawrus (Keuper), although it should

be noted that the highly specialised Schleromochlus 1s found in the

Lettenkohle (Upper Muschelkalk) of Scotland.

In Europe, Thecodontosturus ranges from the Lower Muschelkalk

to the Middle Keuper: the larger Gresslyosaurus and Plateosaurus

occur in the Upper Keuper and Rhaetic. In South Africa the order

of appearance is reversed. The larger Euskelesaurus and Plateosaurus

are lower Red Bed forms while the lighter-limbed Thecodont-saurus,

Massospondylus, Gyposaurus and the like occur in the Upper Red

Beds and Cave Sandstone. Massospondylus has been shown to have

retained a primitive form of shoulder-girdle; and it seems probable

that these light-limbed animals migrated to the Stormberg region

only when conditions became sufficiently arid for them in the south.

In spite of the slight differences between the detailed succession of

the Saurischia in the European and South African regions there is

no doubt as to the general similarity of the fauna; and since none

of the European genera survived beyond the Rhaetic additional

weight is lent to the theory that the Stormberg Series is not later

than Rhaetic.

The occurrence of Geranosaurus is paralleled by that of the Pre-

dentate Nanosaurus in the Rhaetic of North America; while the

Cave Sandstone fish Semionotus occurs in the Keuper of Europe.

Finally, the general aridity which, as we have seen, prevailed

increasingly through Stormberg times is a characteristic of the

Triassic epoch; and in the absence of any evidence to support the

theory of the Jurassic age of part of the Series we must conclude

that Stormberg sedimentation began in Middle Triassic time and

ended in the Rhaetic.

Professor Schwarz considers that the Stormberg Beds should be

divorced entirely from the Karroo System and that, if they are not

492 Annals of the South African Museum.

sufficiently important to warrant their promotion to a system of

their own, they should be grouped with the Uitenhage System.

His reasons for this view, as stated in Proc. Geol. Soc. 8. Africa

1919, are open to question. ‘The most important of all faunal

changes, the passage of the vertebrate remains from reptilian to

mammalian” is not yet proved to be a feature of the Stormberg

Beds; the fauna is a reptilian one, with members of the Beaufort

Theriodontia still living; the Dinosaurs are, as we have seen, Triassic

and not Jurassic nor Cretaceous forms; and Glossopteris is still a

feature of the Molteno Beds.

Important evidence bearing on the age of the sediments has

recently been forthcoming from South America. The beds of Karroo

age in the State of Parana, Brazil, show the following succession : —

6. Eruptivas da Serra Geral. . . « . - . . . 600 metres.

5s SATENILO des BOUUCAtIi | we mee omer: Eieide kit Uaieeiesey Onan me

A Serie sili vRASLO vet eae. Auer s meee release Oem

Calcareo Rocinha ... . = ee

3. Serie Passa Dois Grupo Estrada Nova. .. 150 _,,

Grupos lraty, Sees poms Olena

DP eSenie Luloaraon. (oe hnsy os as. Sone ar ee OU) ns

{io Serremltanane a) ceaw hs eeies Laie ere eee OU) a

The Itarare Series is glacial, and its upper portion contains a marine

fauna of Lamellibranchs and Gasteropods.

The Tubarao Series contains a Glossopteris flora.

The Iraty Group contains the reptiles Mesosaurus and Stereosternum.

The Rio Rasto group has yielded the reptiles Scaphonyx fischeri and

Erythrosuchus sp. and is presumably approximately equivalent to our

Upper Beaufort Beds. Recently Holdhaus (1918) has described from

the same Series the lamellibranchs Solenomorpha similis, S. intermedia,

S. altissima, S. deflera, and Sunguinolites elongatus — an assemblage

which has led him to class the beds definitely as Permian.

The Botucati sandstone, sandwiched as it is between the beds con-

taining Erythresuchus and the thick volcanic outpourigs which, like

those of the Stormberg, made an end of sedimentation, can but cor-

respond to the Stormberg sediments of South Africa, Following as

it does conformably upon strata now classed as Permian there can

be little doubt that it is not later than Triassic in age — a result

in accordance with that now obtained from our consideration of the

South African deposits.

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2902. 2:9 11:

do. The Volcanoes of Griqualand East. Trans. S. Afr. Phil. Soc. XIV, p. 98. 1903.

Stupt F. EK. The Geology of Katanga and Northern Rhodesia. Trans. Geol. Soc.

S. Afr. XVI, p. 44. 1913.

Tomiinson C. W. The Origin of Red Beds. Journ. Geol. XXIV, pp. 153, 238.

Trevor T. G. & Metior E. T. Report on a Reconnaissance of the North-Western

Zoutpansberg District. Transvaal Mines Dept. 1908.

acutus (Sphenosuchus)

Aetonyx.

africanum (Striatotegmen)

africanus (Kuskelesaurus)

africanus (Gryponyx) .

africanus (Phthartus) .

angustidens (Lycorhinus)

ANOMODONTIA . ae:

antiquus Ee padiier)

APODIDAE .

Apudites

Apus. .

Archaeogryllodes

ARCHOBAUBIA .

Aristogaurus .

atavus (Geranosaurus).

Austroblatta .

browni (Euskelesaurus)

browni (Massospondylus).

browni (Thecodontosaurus) .

Caloblattina

Candona

capensis (Euskelesaurus) .

capensis (Gyposaurus) .

capensis (Semionotus) .

carinatus (Massospondylus)

CoLEOPTERA

Colobodus .

congolensis (Lepidotns)

corneti (Pholidophoras)

CRUSTACEA. .

cullingworthi (Plateosaurus)

Cypris . ;

CYZICIDAF .

Cyzicus .

Darwinula.

INDEX.

463

Dictyopyge. ‘

draperi (Cyzicus)

Dromicosaurus

dubius (Thecodontosaurus) :

ENTOMOSTRACA

EPHEMERIDAE ,

erectus (Aristosaurus) .

Erpetosuchus .

Erythrochampsa .

Estheria :

Estheriella .

Euestheria .

Euparkeria.

Euskelesaurus

formosa (Helichthys) .

fortis (EKucnemesaurus)

Geranosaurus .

Gigantoscelus .

globosa var. stricta (Darwinula) .

gracilis (Dromicosaurus) .

GRYLLIDAE

Gryponyx .

Gyposaurus

harriesi essospgndy us)

Helichthys. . . . .

Hongaya

INSECTA. .

istedana (Notochampsa) .

Lepidotus .

323,

PAGE

340

326

405

aie

326

331

aig)

Soll

367

327

463

326

344

423

340

428

436

427

463

405

336

417

379

384

340

333

330

358

338

Lepidurus .

leviseuri (Pedeticosaurus)

Liadoblattina

longipes ( Erythrocham psa)

Inalabensis (Estheriella) .

Lycorhinus.

maeseni (Peltopleurus).

mangaliensis (Cyzicus)

MASSOSPONDYLIDAE .

Massospondylus .

Melanorosaurus .

Mesoblattina .

MESOBLATTINIDAE .

Mesoblattopsis

Mesoblattula .

Metacypris . :

minor (Thecodontosaurus)

minuta (Estheria) .

molengraafh (Gigantoscelus).

monus (Pachygenelus) .

murchisonae (Cyzicus).

Nanosaurus

Notochampsa .

NorocHAMPSIDAE

ORNITHISCHIA.

Ornithosuchus

ORTHOPTERA . .

ovata (Estheria).

Pachygenelus .

palustris (Aetonyx)

passaul (Metacypris)

Index.

PAGE

370,

328

366

333

367

463

343

463

327

383

428

331

333

463

374

327

427

340

327

355

358

436

345

331

327

340

404

463

497

PAGE

Pedeticosaurus 366

Peltopleurus . 463

Pholidophorus 463

Phthartus . 331

PIsScEs ‘ 338

PLATEOSAURIDAE. 406

Plateosaurus . 404

PLECTOPTERA . 331

Protogryllus 336

readi (Melanorosaurus) 430

REPTILIA : ae 340

Rhaetoblattina 333

Rhipidoblattina . 333

riconoi (Tritheledon) 341

Samaroblatta . 333

Schleromochlus ; 365

sch warzi (Massospondylus) . 403

Semionotus. 338

skirtopodus (Thecodontosaurus) 370

SPHENOSUCHIDAE. : 344

Sphenosuchus. . . 344

stormbergensis (Archaeogryllodes). 336

stormbergensis (Lepidurus) . 328

stormbergensis (Plateosaurus) : 406

stowiana (Estheria). 326

Striatotegmen . 331

taylori (Gryponyx) . 420

TELEOSTOMI : d 338

THECODONTOSAURIDAE . 370

Thecodontosaurus 370

THERIODONTIA O40,

transvaalensis (Gryponyx) 420, 455

Triassoblattina 332

Tritheledon. 341

i Sue AS |

if a :

Or 7

( 499. )

12. — On some Gorgonopsian skulls in the collection of the South

African Museum. — By S. H. Haucurton B. A., D. Sce.,

KF. G. 8., Hon. Curator of the Palaeontological Collections.

(With 8 Text-figures).

In a recently published paper (“The Bases of Classification of the

Theriodontia’, Proc. Zool. Soc. 1921, p. 35) Watson has described

in some detail the Gorgonopsian and Therocephalian types in the

possession of the British Museum which are sufficiently well pre-

served to show those features of the skull which have an important

bearing on the interrelationships of the Therapsid Reptiles: and has,

at the same time, drawn attention to the fact that a number of the

types belonging to other Museums are somewhat imperfectly known.

The following notes are based upon a study of Gorgonopsid skulls

which are im the collection of the South African Museum, and are

mainly observations additional to those which occur in the original

descriptions of the various specimens.

The forms discussed are:

1. Galesuchus gracilis Htn. — Tapinocephalus zone.

2. Scylacops capensis Br. — Lower half of Cistecephalus zone.

3. Aelurognathus tigriceps (Br. & Htn.) — Lower half of Cistece-

phalus zone.

4. Aelurognathus serratidens (Wtn.) — Lower half of Cistece-

phalus zone.

5. Gorgonognathus longifrons Utn. — Lower half of Cistecepha-

lus zone.

6. Arctognathus whaitsi Htn. — Middle of Cistecephalus zone.

7. Sycosaurus laticeps Htn. — Middle of Cistecephalus zone.

8. An unnamed form. — Middle probably of Cistecephalus zone.

GALESUCHUS GRACILIS, Htn.

4915. Haughton. Ann. S. Afric. Mus. XH, 3, p. 82. Fig. 10.

This form is of great interest as being the earliest known Gorgo-

nopsid. The type and only specimen came from the lower part of

the Tapinocephalus zone; and it is unfortunate that the specimen

is not more perfect. The snout is missing; the zygomatic arches

are almost weathered away; and the natrix is so intractable that

it has not yet been possible fully to develop the palate and_basi-

500 Annals of the South African Museum.

cranial region. Nevertheless, certain observations can be made beyond

those published in the original description.

As a whole, the skull is narrow and elongate, the orbits looking

outwards and the temporal fossae more outwards than upwards.

It is probable that the squamosal region was not very much _ broa-

dened. The snout is high, almost as high as broad in the canine

Fig. 1. Galesuchus gracilis, Htn.

a. Dorsal view of type skull.

b. Occipital view of left side of type skull.

region, and is square in cross-section, The occiput is very slopmg —

the slope is probably not very much exaggerated by post-mortem

deformation.

The dorsal and lateral surfaces of the skull have already been

described, and can be readily understood from the restored figures

viven. Interesting features are the great depth of the infraorbital

bar, which is almost wholly formed of the jugal; the greater anterior

extension of the prefrontal over the lachrymal, which is as deep as

On some Gorgonopsian Skulls, d01

long; and the small part which the frontal plays in the orbital border,

considerably less than in Gorgonops. The postfrontal is of the shape

seen in Scymnognathus whaitsi. There is no depression on the pre-

orbital surface.

The parietals are excluded from the border of the post-temporal|

fossa, but each sends a narrow process backwards to lie between the

squamosal and tabulare.

The squamosal is partly lost, but it can be seen that its posterior

surface, at the articulation with the tabulare and paroccipital process,

is produced backwards to form a ridge — the inner wall of the

auditory groove — as in Aretops: and that on the outer side of this

ridge there is a shallow vertical groove.

The basicranium is very deep. The condyle is weathered away;

but the basisphenoid tubera, which are close together, lie directly

below the occipital condyle. The median basisphenoidal keel is thin

and deep, lessening in depth anteriorly as it nears the level of the

pterygoid flanges. These lie below the postorbital bar.

Very few features of the braim-case can be determined. The brain-

cavity is small and shallow, the foramen magnum being high up

in the skull. The fenestra ovalis is large and low down, The epi-

pterygoid is a slender rod expanded dorsally ; its lower end is not seen.

As far as its features are discernible, therefore, the form shows

a mixture of primitive and more advanced characters — some of the

latter being sufficiently pronounced to prevent us from considering

Galesuchus as ancestral to the forms of the succeeding zone such as

Gorgonops or Arctops. The primitive features are (1) the high, square

section of the snout, (2) the laterally directed orbits, (3) the posterior

position of the pterygoid flanges, (4) the sloping occiput, (5) the deep

basioccipital and (6) the deep basisphenoidal tubera. More advanced

features are the lack of antorbital depression, the reduction in size

of the lachrymal, and the small part played by the frontals in the

formation of the supraorbital border.

SCYLACOPS CAPENSIS, Br.

19io.° (broom. Ann, -S.2Atric. Mus! xl; A. p:°8:

In the type specimen described by Broom the basicranial region

of the skull is somewhat crushed; but in a skull obtained at Well-

wood, Graaff Reinet by the Rev. J. H. Whaits (S. A. Mus. Cat.

No. 3444) most of the details of this region are perfectly displayed.

The skull in question differs from the type in being somewhat

smaller, in having wa slightly shorter snout and somewhat narrower

502 Annals of the South African Museum.

parietal region: but there can, I think, be no doubt that it is

specifically identical.

The foramen magnum is small, and placed about onethird of the

height of the skull above the base as seen in occipital view.

The basioccipital condyle is fairly large and rounded, although it

is smaller than that figured by Watson in Seymnognathus whaitsi.

The basioccipital is fairly thin posteriorly, but it swells out laterally

in front to form part of the wall of the fenestra ovalis. Medially

on the lower surface it is deeply and narrowly grooved — the groove

being a continuation of that in the basisphenoid. The bone supports

the exoccipitals above and the paroccipitals laterally. The exocci-

pitals are small, forming the lateral margins of the foramen magnum.

Fig. 2. Scylacops capensis, Br

a. Occiput of type skull.

b. Left side of snout of type skull.

The occipital plate is very concave, so that the outer ends of the

stout paroccipitals le some distance behind the foramen magnum,

The inner anterior end of the bone lies below the level of the basi-

occipital condyle, and forms part of the border of the fenestra ovalis,

The type skull shows the presence of large tabular bones on the

occipital plate.

The foramen jugulare has its lower end slightly below the top of

the basioccipital condyle, and looks partly backwards and_ partly

downwards.

The basisphenoidal tubera form the anterior border of the fenestra

ovales and are placed well forward. Between them the basisphenoid

is deeply hollowed; the ridges from them rapidly converge and

combine to form the median basisphenoid keel which passes forward

to meet the pterygoid.

In the type skull the septomaxilla rests on the premaxilla but has

a short articulation with the maxilla posterior to the septomaxillary

On some Gorgonopsian Skulls. 503

foramen. The nasals end anteriorly in a nearly straight) margin,

and externally overhang the nostril shghtly. The nostrils look mainly

forward,

Careful development of the palate shows that Broom’s figure is

substantially correct. The pterygoid has no anterior prolongation

and is widely separated from the posterior nares and prevomer by

the palatine. The deep median groove in front of the pterygoids is

partly covered in by processes of the palatines as in the Aeluros-

aurid figured by Watson in his 1921 paper; and there seems to be

a median suture throughout this groove, the ‘‘vomer” thus being

prevented from forming a part of the palatal surface.

AELUROGNATHUS TIGRICEPS. Br. and Htn.

1915. Seymnognathus tigriceps Broom and Haughton, Ann. 8. Afric.

Mus? XL) 4p. 26, Pl. VI, figs. 1—4.

1915. Seymnognathus tigriceps Broom. Proc. Zool. Soc. p. 225,

1914. Seymnognathus tigriceps Broom Phil. Trans. B. Vol. 206,

p. 20. PE WIp fis. 66.

The type skull is considerably crushed laterally, as is evidenced

by the bending backward of the right pterygoid flange; so that

probably the height of the snout is exaggerated when compared with

its width. Comparison of the photographs of the type in the Annals

S. A. Museum with Broom’s figure of 1913 shows that the verticality

of the snout in the latter is somewhat exaggerated. The snout. is,

however, not so rounded as that of Scymnognathus whaitsi. There is

no “step” between the lower edges of the premaxilla and maxilla. The

septomaxillary foramen is fairly large. The septomaxilla is large and

hasa large facial portion. The nostril is well overhung by the nasals,

Which have a straight front edge transverse to the axis of the snout.

The palate is so crushed that it is rather mdetinite. The palatines

are very large and the pterygoids comparatively small, the latter

having no anterior extension to the internal nares. The presence or

absence of a median bone on the palate is uncertain. The median

portion of the palate is deeply grooved between two prominent

ridges formed by the pterygoids and palatines, Between the pterygoid

flanges and the posterior nares the palate is shorter than im Seym-

nognathus whaitsi approximating more in this particnlar to that of

the Aelurosaurids. The ectopterygoid is large, and forms part of

the front face of the powerful pterygoid flange. The anterior part

of the mouth is considerably vaulted.

There is a weak preorbital depression on the fice.

504 Annals of the South African Museum.

AELUROGNATHUS SERRATIDENS, Htn.

L915. Seymnognathus serratidens Haughton. Ann, S. Afric. Mus.

XII, 3, p. &. Pl. XMI, figs. 2—4 Text fig, 14.

Fig. 3. Aelurognathus serratidens, Htn.

a. Left side of type skull.

b. Cross-section of snout just in front of canine,

showing form of prevomers.

c. Dorsal view of type skull.

d. Palatal view of type skull.

On some Gorgonopsian Skulls. 505

Since the publication of the original description of this species the

palate of the type has been almost fully cleaned; and its features

are such that it has been thought advisable to separate this and the

preceding form from the very different Seymnognathus whaitsi and to

erect the new genus Aelurognathus for their reception. It is possible

that this genus may also include other forms from the Cistecephalus

zone, such as minor and angusticeps, which have hitherto been as-

signed to the Kndothiodon zone genus Scymnognathus.

The median portion of the palate is excavated into a very narrow

and deep groove bordered by strong tooth-bearimg ridges formed by

the palatines and pterygoids. The median groove is seemingly pier-

ced by an interpterygoid foramen. The pterygoid has a short ante-

rior prolongation on the roof of this vaulted area, but there is no

median vomer seen on the palate. The posterior nares extend far

back — to the level of the last molar. The interchoanal bar is

ridged on its ventral surface. Superiorly it forms a thin median

plate with a grooved dorsal edge; this plate is pierced posteriorly

by a large transverse foramen. The relations of the prevomer,

pterygoid and palatine in their vertical portions seem to be as in

the skull which is later to be described as Aretognathus whaitsi.

The sphenethmoid meets the frontals and parietals and ventrally

touches the median plate formed by the pterygoids and basisphenoid.,

The ventral surface of the palate is essentially similar to that of

the Aelurosaurid figured by Watson. The pterygoid flange is very

massive; and as in Aelurosaurus the ectopterygoid forms the lateral

portion of its front face.

The snout is deep and of rounded section, higher than broad.

There is no “step” in the maxillary border. The septomaxilla has

a large facial portion with strong turbinal processes; the septo-

maxillary foramen is fairly large. The anterior end of the nasal

slightly overhangs the nostril. There is a pronounced antorbital

depression, more pronounced than in A. ftigriceps.

GORGONOGNATHUS LONGIFRONS, Htn.

1915, Haughton. Ann. S. Afric. Mus. XII, 3. p. 84 Pl. XIII,

figs. 1—3.

1918. Haughton. Ann. S. Afric. Mus. XII, 6. p. 209. figs. 56, 57.

1921. Watson. Gorgonognathus. Proc. Zool. Soc. p. 78.

The type specimen has been split longitudinally in the posterior

half, and shows the general shape of the brain-case, although it is

not possible to remove the matrix and expose the foramina for the

506 Annals of the South African Museum.

exit of the nerves. The basioccipital is somewhat thicker than in

Scymnognathus whaitsi and the floor of the cerebellum rises more

abruptly. The pro-otic has a long upper process which appears to

form a transverse plate forming the anterior wall of the cerebellum.

Superiorly on the side-wall of the brain-ease, the pro-otic articulates

with what seems to be a distinct epiotic. Just below its upper sur-

Fig. 4. Gorgonognathus longifrons, Hgn.

a. Lateral view of type skull.

b. Palatal view of type skull.

Dorsal view of type skull.

face the pro-otic is pierced by a large oval foramen whose longer

axis lies vertically.

The median part of the palate is deeply vaulted. The ridges form-

ing the boundaries of this vaulted area converge posteriorly, and

are the inner edges of swollen portions of the pterygoids and pala-

tines, which carry a few teeth, Lateral to these swellings the palate

On some Gorgonopsian Skulls. 507

is again vaulted, more broadly than and as deeply as the median

excavation. The pterygoids do not reach forward to the internarial

bar, although they have a slight anterior prolongation on the roof

of the median vault. [ am again unable to distinguish a median

vomer on the palate. It is true that no median suture is to be

seen; but the structure of the median plate, as revealed in an obli-

que section, seems to preclude the possibility of the median vomer

appearing on the palate. The pterygoids seem to be fused. The

pterygoid flanges are far forward. The palatines are large bones.

As Watson has pointed out, the dorsal surface of the skull is very

like that of Rhopalodon. The snout is shorter, but the orbits occupy

the same caudal position. The frontal forms part of the orbital

border. There is a strong rounded ridge running forward from the

supraorbital border, giving the prefrontal portion of the snout a

squarish form. The front of the snout is rounded in section.

The “step” in the dentigerous border is weak; but the palatal

portion of the premaxilla is considerably higher than the palatal

portion of the maxilla, which has a deep vertical portion forming

the outer wall of the internal nares.

ARCTOGNATHUS WHAITSI, n. Sp.

The South African Museum possesses a skull collected by the Rey.

J. H. Whaits at Houd Constant, Graaff Reinet, C. P. (S. A. Mus.

Cat. No. 4337) which is considered to be an'‘Arctognathid. The skull

is of medium size and is typically a Gorgonopsian in the features of

the lower jaw and intertemporal region. The upper surface is weath-

ered away, so that the limits of the dorsal bones are not discernable.

The dental formula of the upper jaw is probably 14c¢1m 4. The front

of the snout is weathered away, and only 3 incisors are preserved ;

but there is sufficient space between the front incisors to permit of

the presence in the whole skull of another tooth on each side. Inv

dental formula, therefore, the skull agrees with that of the genus to

which it is here assigned. The upper canine on the right side is a

replacing tooth, the previous tooth being in a process of absorption.

The molars are very small. All the teeth are finely serrated on the

posterior border,

The chief measurements of the type are as follows: —

Greatest length) 4. . 7.4% =. -emrca 470) mim:

Length from snout to front of orbit . ae 97

Annals of the South African Museum.

ore)

42. mm.

circa

Interorbital width .

Intertemporal width

Basal length .

SS aK

SS) )

WY4

\M eZ

Fig. 5.

Arctognathus whaitsi, Htn.

Palatal view of type skull.

On some Gorgonopsian Skulls. 509

The snout is as high as broad, and rounded in section. The

nostrils are almost terminal and as far as can be seen, were not

overhung to any extent by the somewhat straight front of the nasals.

The chief features of the palate can best be understood by rete-

rence to the figure reproduced herewith. The central vault is some-

what narrow and cordate, the ridges which bound it being formed

partly of pterygoids and partly of palatines. The lateral vaultung

is very pronounced, the highest portion of the vaults being at about

the middle of the ectopterygoids. The interchoanal bar is mainly

formed of the prevomers, the suture between which is plainly seen;

but the posterior narrow part of the bar has its under surface

formed by the palatines. These are large wing-shaped bones. The

pterygoids form most of the roof of the median vaulted area, but

are widely separated from the prevomers. In the figure given there

seems at first sight to be a small longitudinal median bone lying

on the roof of the palate between the pterygoids — 1. e., occupying

the position assigned by Watson to the vomer in Therapsid skulls.

But | am of the opinion that the line shown in the drawing as

bounding the left of this area is not a suture but a crack. It is

not continuous, whereas the other Jongitudinal line continues back

to the very small interpterygoid vacuity. This latter line I inter-

pret as the median suture between the two pterygoids, thus eli-

minating the presence of the vomer on the palate.

The ridges bounding the vaulted area carry no teeth.

The brain-case and occiput are not preserved, nor can the epi-

pterygoid be seen.

SYCOSAURUS LATICEPS, gen. et sp. nov.

The beautiful skull which forms the type of this new genus and

species was collected for the South African Museum by the Rey.

J. H. Whaits on the farm Zuurplaats in the Division of Graaff

Reinet, C. P. A number of interesting details are presented by it,

both on the dorsal and ventral surfaces.

The front of the snout is weathered, but the five incisors are

shown. They are large teeth, oval in cross-section, with their

pointed crowns serrated on the anterior edge. There is a diastema

of 19 mm. between the 5th incisor and the large canine, which is

closely followed by 5 small molars. Both canine and molars are

oval in cross-section and the latter certainly are serrated on their

anterior edges.

The anterior end of the nasal is missing and the details of the

nostril are not decipherable. The septo-maxillary foramen is com-

510 Annals of the South African Museum.

paratively small. The dentigerous border of the upper jaw is incom-

plete; but it is possible that there was a slight “step” at the canine,

The nasals are long and narrow with no posterior widening. The

lachrymal region is slightly depressed and somewhat overhung by a

preorbital ridge. The imtertemporal region is broader than the

interorbital. The pineal foramen is small and surrounded by a

g. 6. Sycosaurus laticeps, Htn.

a. Lateral view of type skull.

b. Dorsal of view type skull.

c. Palatal view of type skull.

small raised boss. It lies at the level of the postorbital bar and

just in front of the top of the occipital plate, which is shaped like

a half-bowl. The frontal is comparatively smal] and wholly excluded

from the orbital border by the postfrontal and prefrontal. The post-

frontal is as in Seylacops capensis. There is no preparietal, and in

On some Gorgonopsian Skulls. dd

this feature the skull differs from all the Gorgonopsia in which = the

structure of the top of the head is known. The median suture

extends the whole length of the top of the skull. About 13 mm.

in front of the pineal foramen it is crossed by the suture separating

the frontals from the parietals, the limits of which bone can be

wholly seen. The parietal is wholly excluded from the border of

the temporal opening by the Jarge postorbital which meets the squa-

mosal posteriorly. The temporal opening is bounded by the post-

orbital and squamosal except for its lower border, which is formed

mainly by the jugal. The jugal is a large bone, supported by the

maxilla in its anterior third, and overlapped by the postorbital half-

way and the squamosal posteriorly. The jugal extends right to the

end of the skull, almost touching the quadrate mass.

The palate is partially displayed. There is possibly a small, narrow,

elongate interpterygoid vacuity lying just behind the descending flanges

of the pterygoids. Immediately anterior to these flanges there is a

fairly shallow median groove which passes forward to the bar sepa-

rating the imternal narial openings. In its posterior half this groove

is narrow and its sides are formed of dentigerous keels of the ptery-

goids. At the bottom of the groove there is a median suture exten-

ding, apparently, from the interchoanal bar to the interpterygoid

vacuity. The suture is very evident in the anterior broader portion

of the groove where it abuts abruptly against the somewhat spongy

bone of the interchoanal bar. In the posterior portion it is not so

evident; but there are no signs of any lateral sutures separating a

vomer from the pterygoids, and the depth and narrowness of the

groove together with its acute dorsal angle would seem to preclude

any possibility of a vomer lying in that position. On the other hand,

the imterchoanal bar has every appearance of being single. It is a

spongy bone, sharply separated off posteriorly from the hard bones

of the palate. Anteriorly it broadens somewhat and shows the two

canals separated by a median keel on the ventral surface; but

nowhere is a median suture visible. Posteriorly its meets the

pterygoids.

The occipital plate shows a small triangular foramen magnum;

the basioccipital condyle is not so massive as in Arctops willistoni.

There is a large interparietal with a median keel which is continued

almost down to the foramen magnum. The tabulare is large and

lies partly against the parietal and partly against the squamosal.

The foramen jugulare looks almost wholly downwards. The whole

occipital plate slopes strongly backwards. The paroccipital process

is fairly broad but thin.

512 Annals of the South African Museum.

The basioccipital is thin. The basisphenoidal tubera are at least

as strong as those of Scymnognathus whaitsi. The median pterygoid

bar is shortened.

UN-NAMED FORM.

A somewhat weathered pre-orbital portion of a skull, collected

high up on the slopes above Oudeberg Hotel, Graaff Reinet, C. P.

(S. A. Mus. Cat. No. 3320) has been developed to show the features

Fig. 7. Un-named form. (S. A. M. Cat No. 3330).

Palatal view of skull.

of the palate. It has also been split longitudinally and in several

places transversely (at points indicated by the lines A, B, C, in the

figure), and most of the structure of the mid-region has thus been

elucidated,

Most of the interchoanal bar is formed of the prevomers whose

ventral surface is rounded anteriorly and becomes flatter and wide

posteriorly. In sections A and B the prevomers are fused; but im

section C they are separated by a distinct vertical suture. Between

them they clasp dorsally a very thin vertical median plate, occupying

the same position as the bone called sphenethmoid by Broom in his

On some Gorgonopsis Skulls. 515

figure of Seylacognathus parrus. Here however, this median plate

seems to be an anterior prolongation of the vertical plates of the

pterygoids fused together. In section C the median bone seems

continuous with the two side walls of the pterygoids which are here

supported by the palatines.

In the same section the pterygoids are seen to continue forward

between the palatines and the prevomers; although on the palatal

Fig. 8. Un-named form (S. A. M. Cat. No. 3330).

a. Median longitudinal section of skull.

b. Cross-section at AA.

c. Cross-section at BB.

d. Cross-section at CC.

surface they disappear some distance behind the posterior border of

the internal nares. There is no median plate of bone on the palate.

Moreover the parasphenoid prolongation of the basisphenoid seems

to end just above the plane of the pterygoid flanges, where it is

ol’ Annals of the South African Museum.

seen between the two thin ascending plates of the pterygoids. These

latter, as stated above, seem to face anteriorly and form the median

bone which is clasped by the prevomers. Anterior to it, the longi-

tudinal section shows another well-ossified plate in the nasal septum ;

and the pterygoid plate is pierced by a large opening just above the

internarial bar.

There is a true sphenethmoid, very similar to that of Déicynodon

seen on the posterior surface of the specimen. It forms the inner

wall of the orbit, and is seen to articulate with the frontal, prepa-

rietal and parietal. The body of the bone separates the orbit from

the anterior prolongation of the brain. A similar ethmoidal element

has been described by Watson in Scymnognathus whaitsi. The rib

along the ventral surface probably rested in the groove in the upper

surface of the parasphenoid which, in our specimen, is displaced to

one side.

Broom described a similar structure in the type of Aelwrognathus

tigriceps. He says the ascending plates of the pterygoid ‘‘pass up-

wards and clasp the front of the median sphenoid. In front of the

sphenoid they become ankylosed, and form a median plate which

extends forwards to meet the vomer” (i, e. the prevomers). ‘‘Whether

this large, thin, median plate is entirely made up of the fused ptery-

voids, or whether there is a median basi-cranial element as well,

cannot be made out in the specimen”.

One of the most interesting features which has arisen from a

re-study of the fine series of types in the Museum collection 1s con-

nected with the question of the presence or absence of a median

vomer on the palate. In the earlier descriptions of the Gorgonopsian

palate, such as that of Seylacops capensis by Broom, there was no

mention of a median bone on the palate between the pterygoids

such at occurs in some at least of the Cynodonts, e. g. Diademodon.

Broom considered then that the interchoanal bar was the true vomer

and that there were no paired prevomers. This position he dubiously

maintained in his Croonian Lecture of 1913, pointing out that “till

further specimens are studied it will be impossible to settle the

question conclusively”.

In his 1921 paper on the Classification of the Theriodontia Watson

figures and describes a median vomer forming part or most of the

roof of the median vaulted area of the palate in Gorgonops torvus,

Scymnognathus whaitsi and Arctognathus curvimola among the Gorgo-

nopsia; whilst, on the other hand, from my examination of material

for the present paper | have been unable to discover a single Gorgo-

On some Gorgonopsian Skulls. o15

nopsian form with an undoubted vomer on the palate. The evidence

presented is thus contradictory, and it becomes necessary to examine

it in detail.

In Gorgonops torvus the anterior part of the median interpterygoid

suture is “open, with a visible strip of matrix in it”. But it suddenly

ends and is with certainty not continued back in the middle line,

being apparently replaced by a pair of much less obvious sutures.

All the sutures on the palate other than those surrounding the median

vomer are obvious,

The median bone in Scymnognathus whaitsi is shown in Watson’s

figure in its posterior half only. It is said to correspond exactly in

position and relations with the posterior median bone in Gorgonops

and the back of the vomer in Diademodon. Watson, however, does

not definitely dissociate this bone from the one forming the inter-

choanal bar although he considers them to be probably distinet.

Arctognathus curvimola is also said to show a median vomer. In

this form “there is no trace of a suture down the midline of the

groove, and its roof seems to be formed by a median bone, which

terminates at the sudden end of the groove and must be bounded

by sutures with the ptervgoids along its edges: of these presumed

sutures nothing can be seen”? in the type.

It must be admitted that this evidence is somewhat unsatisfactory.

No stress can be laid upon the absence of a median suture, as in

some of the South African Museum specimens this suture is not

seen even though, as in the case of Gorgonognathus, the other

sutures of the palate are quite clearly marked. Both in ventral

aspect and in cross-section there seems to be evidence that the

pterygoids have become thoroughly fused together. On the other

hand the median suture is quite definitely present throughout the

entire distance between the imterchoanal bar and the median ptery-

goid in Sycosaurus: and, in consequence, until more definite proof

is given of its presence, | am unable to accept the vomer as for-

ming an integral part of the Gorgonopsian palate.

In this connection it is of interest to note that in a Cynodont

skull from near Burghersdorp, recently described as Cynidiognathus

(Trans. R. Soc. S. Afric. 1922) it is only the front part of the

grooved portion of the palate which is formed by a median bone,

the hinder portion being made up of the pterygoids only. The

median bone here seems to be the same bone which further for-

ward forms the median septum of the skull and which is probably

the fused prevomers of the Therocephalia and Gorgonopsia. Notice

was made in that paper, with his permission, of a recent discovery

O16 Annals of the South African Museum,

by Dr. Broom. He sectioned a small Gorgonopsian skull and found

that the bone which forms the median bar behind the palate and

which has consistently been called the basisphenoid is, in reality,

the vomer; and that the basisphenoid proper is a small ossification

lying between the basioccipital and vomer and not appearing on the

ventral surface of the skull.

[t is difficult to understand how, if it is the homologue of the

parasphenoid of the Amphibia, the vomer should appear in the

Therapsids as a median bone on the palatal surface. The evolu-

tionary tendency in the Amphibia and lower Reptiles is to crowd

the parasphenoid away from the palate by an enlargement of the

pterygoids in the mid-line and to make of it a thin vertical wall of

bone: and once it had attained this form, as in the Dinocephalia, it

is unlikely that a reversal would take place in order to force it once

again between the pterygoids on to the palate.

In a paper prepared for the Williston Memorial Volume and not

yet published, I have considered the same subject briefly. A small

skull named there Whaitsiella was considered to have a median

vomer and to be ancestral to the aberrant form Whaitsia which

definitely has no vomer. Further examination of Whartsiella leads

to doubt as to the presence of a median bone on the palate. The

little skull is much crushed and cracked, and the supposed sutures

may be cracks. The supposed bone les ventral to the pterygoids

and certainly has no connection with what has hitherto been known

as the basisphenoid: and it may very possibly be a flake of the

pterygoids. Certainly Whaitsia has no median bone on the palate;

and whether it be a Gorgonopsid or a Therocephalian it yields

quite definite evidence on that point.

Cynosuchns whaitsi almost certainly has no median vomer in the

posterior part of the palate, although there is no actual median

suture seen separating the pterygoids which have lost the anterior

prolongation; while Ictidopsis likewise lacks the bone. The accumu-

lation of evidence furnished by the specimens studied here seems

to lead to the view that the median “vomer” of the Cynodonts is

the homologue of the paired prevomers and is due to the backward

shift on the palatal surface of these bones which in the earlier

Therapsids make up the interchoanal bar, Their function as sup-

ports for Jacobson’s cartilage are usurped in the Cynodonts by the

palatal processes of the premaxillae: but they still play, in Cynidio-

gnathus, a part in the formation of the internasal septum.

If this view be correct then, in the line of evolution of the Cyno-

donts, we should see a progressive backward growth of the prevo-

On some Gorgonopsian Skulls. 517

mers on to the central part of the palate and, at the same time,

the retrogression of the pterygoids from the posterior border of the

internal nares. There does not seem to be clear evidence of this

change in the Gorgonopsia: but the occurrence of the prevomers as

an essential part of the Therocephalian palate behind the plane of

the internal nares is a significant fact bearing on the question of

the origin of the Cynodonts. It is desirable that more should be

known of the details of the Therocephalian palate; the later forms

like Akidnognathus and Moschorhinus ave at one with earlier genera

such as Iectidosuchus, Alopecognathus and Scylacosaurus in having the

prevomers passing back between the palatines to articulate with the

pterygoids,

Aelurognathus

Arctognathus .

capensis (Scylacops).

Galesuchus

Gorgonognathus .

gracilis (Galesuchus)

INDEX.

50]

499

505

499

laticeps (Sycosaurus) . .

longifrons (Gorgonognathus)

Scylacops

serratidens (Aelurognathus)

Sycosaurus .

tigriceps (Aelurognathus).

whaitsi (Arctognathus)

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ANNALS

SOUTH AFRICAN MUSEUM

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PARY VY, contaming :—

-13.—Some New Species of Anomodontia (Reptilia)—By R.

Broom, D:Se., C.M.Z.S., and 8S. H. Haventon, -B.A;

F.G.S., Assistant Director.. (With six Text-figures.)

14.—Investigations in South African Fossil Reptiles and

. Amphibia (Part 10), By 8. H. Haveuron, B.A., F.G.S.,

Assistant Director. :

10 —Descriptive Catalogue of the Anomodontia, with

especial reference to the examples in the Sonth

African Musewm (Part 1). (With three Plates

and twenty Text-figures.)

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15 —Investigations in South African Fossil Reptiles and Amphibia

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17. — The fauna and Stratigraphy of the Stormberg Series. —

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12. — On some Gorgonopsian skulls in the collection of the South

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F. G.S., Hon. Curator of the Palaeontological Collections.

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