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1969

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THE FAUNA OF THE TAPINOCEPHALUS ZONE

(BEAUFORT BEDS OF THE KAROO)

L. D. Boonstra

South African Museum, Cape Town

(With 22 figures)

[MS. received 1 May 1969]

CONTENTS

PAGE

A review of previous work : 5 : : : I

The fauna as a whole : 5) 3

The non-reptilian elements in the ernie and the fore 29

Petroleum in the rocks of the Tapinocephalus zone . oy 29)

Taxonomic revision . : ; : : : a 29

Pareiasauridae ‘ : : : ce 2G

Anomodontia : : : : : . 5 eS

Dinocephalia_ . : : : : . 5 Gi

Anteosauridae 2 . : : : 5), 8B

Titanosuchidae . § : : : dR

Tapinocephalidae . : : . 0 eo

Styracocephalidae . : : ¢ 3 - 44

Dicynodontia . : 5 5 : : ouEAA:

Endothiodontidae . : : : : snl

Dicynodontidae . : : : : ey ZS)

Theriodontia : : é : : : 5h ANS)

Gorgonopsia_.’ é 3 : : NAS

Hipposauridae 4 5 : : j |), 49

Galesuchidae. : 6 : : : a) 50.

Therocephalia . 2 : : : é 450

Pristerognathidae . : : : : eo kee 5O.

Lycosuchidae 3 : : : é Me in5S

Alopecodontidae . : : : ; bad GB

Scaloposauridae . 6 : : 5 ol 53

Systematic list of the tetrapods . 6 : Sal As

Tapinocephalus zone faunas outside South Niven : ST

A comparative table : . ety G%8)

Origin of the Tapinocephalus zone Sepa eynaes ; a 162

Descendants of the fauna of the Tapinocephalus zone . 64

Acknowledgement . 64.

References and Biblioemphy, of the Tarinereniaies zone. 64

A REVIEW OF PREVIOUS WORK ON THE Tapinocephalus ZONE AND ITS FAUNA

The oldest record in print of the discovery of fossil bones in the rocks of

the Karoo bears the date 1831. This is in the form of a letter printed in the

South African Quarterly Journal —a letter signed by C. H. Crisbrook and addressed

to Dr. Andrew Smith, Corresponding Secretary to the South African Institution.

Crisbrook writes that while visiting Beaufort in 1827 he stayed with Mr.

Baird, Ex-Landrost, who, amongst other mineral specimens, gave him a piece

of a fossil tooth, which had been found by the son of Fieldcornet de Klerk.

Ann. S, Afr. Mus. 56 (1), 1969: 1-73, 22 figs.

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

Crisbrook in the company of Baird, De Klerk and his son visited the site

and found other specimens of which some were collected. There is no record

of what became of these specimens nor can it with any degree of certainty be

determined from Crisbrook’s letter where the site was situated. The locality

indications given are inadequate and in important points contradictory.

Ruling out what would appear to be discrepancies I believe that these first

specimens are from the Tapinocephalus zone within the Koup.

In any case, if not from the Tapznocephalus zone, these specimens are

undoubtedly the first recorded Karoo fossils—antedating those found by

A. G. Bain near Fort Beaufort by 11 years. |

In the same publication mention is also made of ‘Specimens of Fossil

Bones of a large Mammiferous Animal from the Karoo; presented by the

Rev. Dr. Adamson’. These most probably also came from the Tapinocephalus

zone, but the data available are less convincing than those of the De Klerk

discovery.

The A. G. Bain specimens received by the British Museum in 1853,

although mostly from higher zones, do include some specimens from the Great

Karoo. These are rather poor specimens but they undoubtedly are all dino-

cephalians from the Tapinocephalus zone. Lydekker (1890) considers that the

British Museum specimen R1708 figured by Owen as Tapinocephalus atherstonet

was also presented by A. G. Bain, 1853. |

In a letter dated 18 January 1856 A. Rose-Innes of Beaufort West writes

that he is sending by ‘mail cart’ two fossils to the South African Museum.

There is no further record of these specimens and they may, of course, have

come from either the Endothiodon or Cistecephalus zones exposed near Beaufort

West.

In the following year, 1857, according to the museum register, Dr. Way

presented the proximal end of a tibia. This specimen from Uitkyk is still in

the collection of the South African Museum bearing the number 737 and was

identified in 1955, thus nearly a century later, by Boonstra as a tibia of the

titanosuchian reptile, Jonkeria ingens.

For the next 14 years no record could be traced of any work in the

Tapinocephalus zone. ‘Then in 1871, with the discovery of gold in the district

of Prince Albert, the area becomes geologically important and at least two of

the investigators of the mineral possibilities were also interested in fossils, viz.

Dr. Atherstone and T. Bain. In a parliamentary report of that year Atherstone

and Bain mention the occurrence of fossil bones on the farms Varsfontein,

Spreeufontein and Janwillemsfontein. In the Cape Monthly Magazine Atherstone

again refers to this area.

These specimens mentioned in 1871 were collected by Atherstone and

sent to London, where they were shortly afterwards described by Sir Richard

Owen in his Catalogue of the fossil remains of Reptilia of South Africa, and one

constitutes the type of Tapinocephalus atherstonei, and another is the type of

Bradysaurus bombidens. Another specimen was erroneously referred by Owen to

THE FAUNA OF THE TAPINOCEPHALUS ZONE 3

his ‘Pareiosaurus’ but subsequently made the type of Phocosaurus megischion by

Seeley in 1888. Other specimens also referred to his ‘Pareiosaurus’ by Owen

are really bones of a titanosuchid.

This confusion with respect to the bones collected by Atherstone appears

to be mainly due to the fact that Atherstone did not keep his various finds

separately but lumped bones derived from an area together to constitute a

single specimen and this initial mistake was apparently aggravated by faulty

labelling in the British Museum.

The collection which the British Museum purchased from T. Bain in

1878 includes specimens from the Tapinocephalus zone. A bone (49389) described

by Seeley in 1889 as an ulna is really a tibia of Titanosuchus. Another specimen

from Koedoeskop was made the type of Titanosuchus ferox by Owen in 1879.

Fragments (49404) from Palmietfontein constitute Broom’s type (1905) of

Cyniscodon lydekkeri. It also includes the Palmietfontein specimen of Bradysaurus

bombidens described by Seeley in 1888. No. 49425 was described by Seeley as

the type of Glaridodon, but referred to Titanosuchus by Lydekker in 1889. A

specimen from Letjiesbos (49407) has been referred to Lycosaurus (Boonstra

19344).

In 1880 the British Museum purchased a further collection from 'T. Bain.

A specimen from Vindraersfontein (R848) is the proximal end of a femur of

Brachypareia. Five other specimens from the Koup are poorly preserved elements

of Dinocephalia. R872 was made the type of Hyorhynchus platyceps by Seeley

in 1889.

In 1881 H. W. Oakley, assistant at the South African Museum, was sent

to the Koup on receipt by the museum of reliable information of the occurrence

of fossil bones in that area. The results of this mission were mentioned in the

Report of the South African Museum for 1881. Numerous fragmentary remains

were found and these were entered as ‘Fossil Bones of Dicynodon’, but in the

preserved, identified in 1955 by Boonstra as that of Titanosuchus. A further

result of Oakley’s mission was that on his reporting hearsay evidence of the

occurrence of further skeletons the Colonial Government in its budget for

1882-3 voted a sum of £200 for collecting Karoo fossils. With these funds

available T. Bain was employed to collect for the South African Museum in

the Tapinocephalus and higher zones in 1883. But before Bain’s collecting trip

the museum, according to the register, received from J. R. Joubert a collection

of 11 specimens, all from ‘near Beaufort West’. From this collection the types of

Ictidosaurus angusticeps, Scymnosaurus ferox, Dicynodon jouberti, Pristerodon brachyops

and Dicynodon pseudojouberti were later described by Broom (1903), 1g05a) and

Boonstra (1948b). The collecting trip undertaken by Bain in 1883 on behalf of

the museum was not a very successful undertaking and the museum today

possesses only two specimens which were probably obtained by Bain. Only

one, later described by Seeley as Delphinognathus conocephalus, is from the

Tapinocephalus zone.

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

In 1884 Dr. Exton presented to the British Museum a flattened tibia

(R519) which may be tapinocephalian

In 1888 Seeley published his account of Pareiasaurus (now Bradysaurus)

bombidens based on the Palmietfontein specimen, which T. Bain had sold to

the British Museum in 1878. A number of bones from Varsfontein, by Owen

thought to be pareiasaurian, were in this paper by Seeley made the type of

Phocosaurus megischion.

In 1889 Seeley erroneously described as a pubic bone a coracoid of

Titanosuchus from Koedoeskop and also correctly a humerus, femur and fibula;

from Varsfontein a titanosuchid ulna and from Janwillemsfontein he figured

the tibia thought by Owen to be pareiasaurian and now considered

titanosuchian.

In the same year Seeley visited South Africa, thus being the first trained

vertebrate palaeontologist to collect personally in the Karoo. He spent some

time collecting in the Tapinocephalus zone where he was accompanied by T.

Bain. At Tamboerfontein he was given a skeleton by J. S. Marais, now known

as the Tamboer specimen of Bradysaurus bombidens. Later Sarel Marais guided

Seeley to another skeleton now known as the De Bad specimen of Bradysaurus

bain. This specimen was excavated with the help of T. Bain, J. S. and Sarel

Marais and their respective sons on 12 August 1889. In the Transactions of the

South African Philosophical Society Seeley pays very warm tribute to the help he

received from all the local farmers—his specimens were taken to the station

by mule-wagon. A couple of sentences are here quoted. “The Dutch farmers,

ever on the alert for natural history phenomena, .. . at the first indication

that we were likely to visit a certain spot, every specimen that could be in any

way of interest to us was obtained, so that our labours were very much lightened’

and further on, “This carrying away was perhaps a little more difficult than

we in Cape Town had imagined. We were in the open veldt, where there is

no possibility of getting assistance . . . ; there are few facilities for bringing

away the specimens in the best possible condition, but owing to the aid which

was never wanting and never grudged in the least, we were able to gather up

the fragments, which filled several large cases, and a procession of mule wagons

bore away...’.

Every palaeontologist who has collected in the Koup will fully endorse

this warm tribute of Seeley’s.

In addition to the above, Seeley obtained from L. Pienaar of Weltevreden

a specimen described in 1892 as the type of Eunotosaurus africanus. A specimen

from De Cypher was made the type of Pnigalion by Watson in 1914 and another

from Tamboerfontein became the type of Anteosaurus.

Lydekker in his Catalogue of 1890 listed and sorted out the specimens then

in the British Museum, correcting a number of errors with regard to specimens

from the Tapinocephalus zone.

In 1892 Seeley described the type of Delphinognathus conocephalus, which he

had on loan from the South African Museum. This dinocephalian is thus the

THE FAUNA OF THE TAPINOCEPHALUS ZONE 5

oldest described type of Karoo vertebrate housed in a South African institution.

In 1895 Seeley described Pristerognathus polyodon collected by himself at

Tamboerfontein in 1889, and in the same year published a photograph of the

mounted De Bad specimen of Bradysaurus baint. In a book entitled Creatures of

other days by Hutchinson, published in 1896, there is a drawing by J. Smit of

Bradysaurus baini reconstructed in the ‘flesh’. As the pose follows that of the

Seeley’s mounted skeleton the result is most unsatisfactory, but the drawing

in Swinton’s book (1948), half a century later, with the same pose, is

inexcusable.

In the first annual report of the Geological Commission, 1896, Schwarz

writes: ‘In the lower beds of the Gouph a large quantity of Paretasaurus bones

were discovered, all of them referred to P. Bainiz, but owing to the absence of

facilities for excavating them they were left till more favourable circumstances

should occur, one bit only, consisting of 16 vertebrae with the pelvis in situ,

was taken for the sake of identification. . . . I have found them all over the

country from Spreeufontein, Prince Albert, to directly under the Nieuweveld

Mountains, as at Hottentots River and Knoflock’s Fontein, and the same species

occurs throughout. Just south-east of Fraserburg Road Station I have, however,

obtained P. bombidens, Owen, and at the Prince Albert Goldfields a variety of

large forms, some of which are new. P. Bainii is the commonest form.’ In 1898

Webster presented some Bradysaurus vertebrae, found by him near Prince Albert

Road, to the South African Museum.

From August 1902 Rogers and Schwarz spent over three months in the

Koup. Rogers (1903) writes: “During our journey in the Gouph, we heard

of a large skeleton in the rocks near van der Byl’s Kraal, and when we reached

that place we found that the skeleton was on Hoogeveld, Lot A, ground which

now belongs to Mr. J. P. Snyman, of Knoflock’s Fontein. Mr. Snyman very

kindly took us to the spot where the bones lay, and allowed us to remove them.

In the matter of the transport of these bones, which weighed some 700 lbs., to

our camping place, and thence to the railway, we were greatly helped by

Mr. M. van den Bergh, of van der Byl’s Kraal. . . . The bones were . . . cleared

of their matrix by Miss Wilman, under the superintendence of Dr. Broom, who

found that they are an almost complete skeleton of Paretasaurus serridens ... Qwen.’

(Now the type of Embrithosaurus schwarzi.) “The Commission decided to have

the fossils, collected since the commencement of the survey in 1896, named and

described . . . and to have the descriptions published in a special volume of the

Annals of the South African Museum’.

Rogers & Schwarz (1903) mention that many fragmentary bones were

encountered. The better examples collected were examined by Dr. Broom.

These are: from Grootfontein a small species of Paretasaurus, from Buffel’s Vlei

an interclavicle probably of Titanosuchus, from Drooge Fontein a new fossil

reptile — possibly allied to Titanosuchus, from possibly Knoflock’s Fontein (per

J. P. Snyman), the type of Glanosuchus macrops and from Paarde Bosch a partial

humerus possibly of Lycosaurus.

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

In 1903 Broom described a skull, presented by the Rev. van der Merwe to

the Victoria College (now the University of Stellenbosch), with the designation

Lycosuchus vanderrieti. In the same year Broom described what is now the type of

Embrithosaurus schwarzi, collected by Rogers and Schwarz (S.A.M. 8034), also

Titanosuchus cloetei from the ‘Gamka River’ presented to the South African

Museum by Mr. Justice Cloete; and the 1881 J. R. Joubert’s types of Ictidosaurus

angusticeps and Scymnosaurus ferox.

In their report for 1903 Rogers & Du Toit announce the discovery at

Knechts Banken near Calvinia of ‘Lamellibranch shells and fish scales, together

with Glossopteris’; the first two were identified as Palaeomutela and Palaeoniscus.

This locality is probably in Ecca beds. In his contribution to this report

Schwarz mentions (p. 92) some localities where bones have been found, viz.

Prince Albert Road Station, Spreeufontein, Kleinwaterval and Seekoeigat.

Also in 1903 the South African Museum received a collection from J. M.

Bain which had been collected by his father, T. Bain; this includes, besides the

type of Pristerognathus baini, four specimens of Dicynodon jouberti, one of Robertia

broomiana two of ? Titanosuchus and a pareiasaurian bone. The latter is from

Springfontein and all the others have the locality unrecorded. From C. de

Villiers came a dinocephalian femoral fragment from Spes Bona, 23 miles south

of Beaufort West.

In 1904 Broom collected the type of Dicynodon megalorhinus and P. H.

du Plessis discovered at Seekoeigat the bones which Broom designated as the

type of Scapanodon duplessist.

In 1905 the Rev. J. H. Whaits, a railway clergyman and then rector at

Prince Albert, discovered his first specimen, a Lycosuchus, in the Tapinocephalus

zone at Fraserburg Road. In the same year P. H. du Plessis discovered at

Seekoeigat specimens which became the types of Alopecodon priscus and A.

rugosus and also two Pristerognathus, one Glanosuchus, one Dicynodon megalorhinus,

seven poor dinocephalian specimens and fish scales. On Bokfontein he found

what became the type of Pelosuchus priscus and on Waterval a poor skull of

Alopecodon. Mr. J. L. Cairncross, operator of a drilling machine, discovered on

Sandvlakte the specimens Broom made the types of Eccasaurus priscus and

Archaeosuchus cairncrossi, incorrectly maintaining that they came from the Ecca

beds, whereas in fact they are forms from the Tapinocephalus zone. At Prince

Albert Road Station Cairncross also found fish scales and a skull of Dicynodon

pseudojoubertt.

In 1905 Rogers in his classification of the Karoo rocks retains Seeley’s

(1892c) triple division but calls them the Lower, Middle and Upper Beaufort

beds. Broom (1905)) retains the name Pareiasaurus zone for the lowest part of

the Lower Beaufort and proposes that the middle part be called Endothiodon

zone and the upper zone A‘stecephalus zone.

In 1906 Cairncross found a Palaeomutela and some poor pareiasaurian and

dinocephalian bones near Prince Albert Road, and Whaits discovered the

type specimens of Trochosaurus acutus and Hyaenosuchus whaitsi at Rietfontein,

THE FAUNA OF THE TAPINOCEPHALUS ZONE 7

Prince Albert and of Pardosuchus whaitst and Rhinesuchus whaitsi at Fraserburg

Road, together with a specimen of Pristerognathus at Rietfontein, Prince Albert,

and at Fraserburg Road a specimen of Cynartognathus paucioridens and a tapino-

cephalian at Fraserburg Road. The above types were designated as such by

Broom in 1908.

In 1907 Broom listed 16 types occurring in the Paretasaurus (now

Tapinocephalus) zone.

In 1908 Whaits contributed an Eunotosaurus from Rietfontein, Prince

Albert, a Jonkeria from Letjiesbos and some pareiasaurian and dinocephalian

fragments from Prince Albert Road. In 1909 A. R. E. Walker, of the South

African Museum, collected at Roggekloof, Sutherland, the specimen made

the type of Pristerognathoides roggeveldensis by Boonstra and also a Bradysaurus

and some dinocephalian bones. He also excavated a pareiasaurian found by

Gordon on his farm Hottentotrivier. In this year Broom (1g0g9a and c) listed

20 species in the Paretasaurus (now Tapinocephalus) zone and described as

Tapinocephalus the skull in the Seeley collection, which Watson in 1914 made

the type of Mormosaurus seeleyt.

Rogers in 1910 again worked for months in the Tapinocephalus zone in the

districts of Beaufort West, Fraserburg, Sutherland and Laingsburg and the

sheet map 13, commenced by Schwarz in 1896, was published. In the report

of the Geological Commission for this year only one specimen from the

Tapinocephalus zone is mentioned, viz. an Alopecodon from Skaapskooi, south-

west of Fraserburg, together with fragmentary pareiasurian remains from the

Moordenaar’s Karoo and Komsberg.

Broom, in 1910, published his important comparison of the Permian

reptiles of North America with those of South Africa; and also described a

skull in the British Museum collected by Seeley at Tamboerfontein as a

Titanosuchus (now the type of Anteosaurus magnificus).

In 1911 Broom described the type skull of the Moschops material found

by Whaits on Spitzkop, Moordenaars Karoo and subsequently sold by Broom

to the American Museum of Natural History and also Eriphostoma microdon

found by Whaits near Fraserburg Road.

During 1912 the good skull and limb-bones of Tapinocephalus (S.A.M.

2343) were collected at Uitkyk by P. le Roux, M.P. for Beaufort West. Broom

(19126) designated as types of Taurops macrodon a snout found at Bosmanshoek

in the Komsberg by Whaits and Pristerognathus (now Cynariognathus) platyrhinus

from Grootfontein, Beaufort West. During 1912 Watson, with the help of a

grant from the Percy Sladen Fund, collected in the Karoo using for con-

veyance, what Haughton describes as ‘a decrepit horse and trap—the former

capable at most of 10 miles per day’. Two of the important specimens which

Watson collected in the Tapinocephalus zone—an Embrithosaurus skull and the

type of Broomia perplexa—both came from Hottentotsrivier, Beaufort West.

In 1913 Haughton undertook his first collecting in the Tapinocephalus zone.

On Abrahamskraal he collected the types of Galesuchus gracilis and Trochosaurus

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

intermedius and also a specimen of Dicynodon jouberti. On Uitkyk he obtained a

Brachypareia. Working with Whaits two specimens of Anteosaurus and a tapino-

cephalian were collected at Vivier. Whaits found the type of Struthiocephalus

whaitst at Vivier.

In this year Broom described three specimens obtained for the Albany

Museum by Whaits on Hottentotsrivier as the types of Pareiasaurus (later

Koalemasaurus) acutirostris, Scylacognathus parvus and Scymnorhinus (now Broomia-

saurus) planiceps. In the Annals of the South African Museum Broom also described

a pareiasaurian manus and pes found by G. Gordon on Hottentotsrivier and

excavated by A. R. E. Walker in 1909. In the same year Haughton published

photographs of the skull of Tapinocephalus found by Le Roux on his farm Uitkyk

in 1912. Watson (19145) also published a paper on the Beaufort beds.

Broom, in 1914, wrote his important comparison of the South African

dinocephalians with the American pelycosaurs and designated as the type of

Moschognathus whaitsi a specimen that had been collected by Whaits in the

Beaufort West district. In the American Museum Journal Broom published

photographs of Bradysaurus whaitst and of a shoulder girdle of Tapinocephalus

atherstonet now known to be that of a titanosuchian. In his Croonian lecture

Broom figured postcranial bones of Moschops and Moschognathus and listed 19

genera of tetrapods derived from the ‘Pareiasaurus’ zone and erroneously two

dinocephalian genera were said to be from the Ecca beds.

Watson published a number of papers in 1914 on his work in the

Tapinocephalus zone. The first was on the Embrithosaurus skull he had obtained

at Hottentotsrivier; the second on the zones of the Beaufort beds; the third on

the nomenclature of the pareiasaurians in which he designates the specimen

collected by Schwarz in 1902 as the type of Embrithosaurus schwarzi; the fourth

paper was on the Dinocephalia, in which specimens in the British Museum

were made the types of Mormosaurus seeleyi, Pnigalion owent, Lamiasaurus newtoni

and he also described a skull as Titanosuchus which he later made the type of

Anteosaurus magnificus; in a fifth paper Broomia perplexa was described; sixthly,

Eunotosaurus africanus was fully redescribed and lastly in a paper on the

therapsids a specimen collected by T. Bain was described under the name

Lycosuchus? (now Scymnosaurus watson.)

In 1914 Watson proposed that the lowest zone of the Beaufort beds be

called the Tapinocephalus zone instead of Pareiasaurus zone.

During 1914 Haughton and Whaits collected together on Janwillems-

fontein and obtained two specimens of Bradysaurus seeleyi, a humerus of Fonkeria

ingens, a skull each of Scymnosaurus ferox and Pristerognathus and an unidentified

therocephalian. Haughton excavated a skull and much of the skeleton of

Struthiocephalus whaitst on Abrahamskraal and from La-de-da he obtained the

skull which became the type of Moschosaurus longiceps.

In 1915 Rogers collected a skuil which has since been identified (Boonstra

1954) as Pristerognathoides minor on Jakhalsfontein, Prince Albert. Whaits

collected the type of Riebeeckosaurus longirostris at Vivier Siding and at Prince

THE FAUNA OF THE TAPINOCEPHALUS ZONE 9

Albert Road he found a specimen of Dicynodon jouberti and an Emydops longiceps,

at Klipbank the type of Pristerognathoides minor, at Letjiesbos a lycosuchid skull

and on Brand Doorns (Leegte?) a titanosuchid jaw and a gorgonopsian and

at Hottentotsrivier a lower jaw of Brachypareia watsoni. Haughton, in 1915, on

Abrahamskraal obtained a skull of Robertia broomiana and fragments of an

Anteosaurus, figured by Boonstra in 1955, and from Uitkyk brought the limb-

bones associated with the Tapinocephalus skull that had been found by P. le

Roux and also a beautiful moschopid fibula. Also in 1915 Broom published

his catalogue of the specimens he had sold to the American Museum; this

contained two pareiasaurs, eleven dinocephalian specimens, and six thero-

cephalians, all of which had been found by Whaits.

Haughton in 1915 described the types of Struthtocephalus whaitsi (found by

Whaits in 1913), and Trochosaurus intermedius and Titanosuchus dubius found by

himself in 1913. Broom in 1915 named a number of poor specimens in the

British Museum that had been purchased from T. Bain in 1878; the new names

are Simonhinella baini, the paratype of Ictecephalus polycynodon, Cerdodon tenuidens,

Cyniscodon lydekkeri and Scymnosaurus watson. In September of this year Haughton

described a skull found by himself at La-de-da as Moschosaurus longiceps and

another collected in 1913 at Abrahamskraal as Galesuchus gracilis.

In 1916 Van Hoepen described a skull and partial skeleton from Abrahams-

kraal under the name Jonkeria truculenta. During this year Haughton collected

in the Koup and found at Abrahamskraal seven pareiasaurs, four dinocephalians

and one therocephalian; on Leeurivier he obtained two pareiasaurs and four

dinocephalians; on Bloukrans two dicynodonts, on Welgemoed the type of

Jonkeria haughton and a good Bradysaurus seeleyt; on Stinkfontein a Scymnosaurus

ferox and the type of Alopecideops gracilis; on Lammerkraal the type of Struthio-

cephaloides cavifrons and a poor titanosuchid; from Rietfontein another titano-

suchid and from Rietkuil a Koalemasaurus. W. van der By] also presented a skull

which later became the type of Bradysaurus vanderbyli.

During 1917 Haughton spent two weeks in the Koup and obtained 16

pareiasaurs, nine dinocephalians, four therocephalians, two dicynodonts and

on Voélfontein and Bloukrans fish scales. Cairncross collected a Dicynodon

jouberti at Prince Albert Road.

In 1918 Haughton described the specimen found by Whaits at Klipbank

as Alopecognathus (now Pristerognathoides) minor and figured the braincase of a

tapinocephalid which had been found by P. le Roux in 1905. In this year

A. L. du Toit discussed the zones of the Karoo system. During 1919 Van der

Byl collected a Dicynodon joubertt and an unidentified gorgonopsian and a

dinocephalian and Haughton in his review lists 40 genera in the Tapinocephalus

zone.

In 1920 Gregory published a notice of the mounted skeleton of Moschops

capensis Broom and Joleaud proposed the name Broomisaurus for Broom’s

Scymnorhinus.

In 1921 Watson re-examined the type of Scymnosaurus watsoni and proposed

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

the name Anteosaurus magnificus for the specimen previously described as a

Titanosuchus, and Broom reconsidered the tarsus of Broomia perplexa.

Romer, in 1922, in his paper on the locomotor apparatus, discussed the

conditions in Moschops and Propappus. Abel in his Lebensbilder in 1922 discussed

some palaeobiological aspects of life in the Karoo.

Broom, in 1923, described a skull referred to Scapanodon duplessist collected

by Van der Byl on Abrahamskraal, also two skulls from Kookfontein found by

M. J. van Wyk were designated as the types of Dinophoneus (now Jonkeria) ingens,

another skull found by Van der Byl on Abrahamskraal became the type of

Dinartamus vanderbyli and a lower jaw was described as Titanosuchus cloetei (later

this became the type of Phoneosuchus (now Jonkeria) angusticeps and a further

jaw became the type of Enobius strubenr). Also during 1923 C. le Roux collected

on Boesmanskop the types of Dolichopareia angusta and Nochelesaurus alexandert

and on Abrahamskraal the type specimen of Brachypareia watsoni and at Zwarts

Siding an Embrithosaurus schwarzi and on Lammerkraal an unidentified

therocephalian and on Bloukrans the type of Pristerognathoides vanwykt.

Haughton in 1923 accompanied Case on a journey through the Karoo on

which the latter based some of the views expressed in his paper of 1926.

In 1924 Haughton accompanied Von Huene through the Karoo and the

latter obtained specimens from the Tapinocephalus zone which are now housed

at Tubingen, and described in 1931. Von Huene’s observations during this

trip were incorporated in his compilation of 1925. Haughton (1924a) also

described as Galesuchus gracilis the specimen collected by himself on Abrahams-

kraal in 1913, and in his bibliographic list (19245) he listed 56 species as

definitely from the Tapinocephalus zone. In 1924 Broom described a young

pareiasaurian skull found by Van der Byl on Abrahamskraal and a new

species Pareiasaurus (later Nochelesaurus) strubent from the same locality. C. le

Roux collected in 1924 a Dolichopareia angusta, a Pristerognathus and a Propappus

on Abrahamskraal.

In 1925 Broom described a skull found by Van der Byl at Abrahamskraal

as the type of Pristerognathus vanderbyli and another became the type of

Alopecodon minor; and from Lammerkraal Ictipareia brevirostris and the specimen

collected for the South African Museum by C. le Roux on Bloukrans in 1923

became the type of Pristerognathus (now Pristerognathoides) vanwyki. In 1925

Haughton mentions a specimen of Rhinesuchus whaitsi from Bloukrans.

Von Huene’s ‘Lebensbild’ (1925) was a very useful compilation of the

then known facts on the fauna of the Karoo. It was palaeobiologically orien-

tated and included a zone map—the first published —but being based on the

inadequate survey of the area was inaccurate in many respects, in particular

as far as the limits of the Tapinocephalus zone is concerned. The fauna of the

Tapinocephalus zone was listed as consisting of one invertebrate, one fish, one

stegocephalian, seven cotylosaurs, one archichelonian, 21 dinocephalians,

three dromasaurians, three dicynodonts, three gorgonopsians, 21 therocepha-

lians, one cynodont? and three incertae sedis. This makes a total of 66 species.

THE FAUNA OF THE TAPINOCEPHALUS ZONE sit

Rogers in 1925 published a sheet map with an explanation of the country

near Laingsburg and mentions only three localities of fossil finds.

In 1926 Haughton in an address, ‘Palaeontology in South Africa’, listed

68 species from the Tapinocephalus zone. Gregory also published a detailed

account of the skeleton of Moschops and also of some features of Moschognathus

and reviewed the work on other described dinocephalians. In his study on

the environment of tetrapod life, Case incorporated his observations made in

the Tapinocephalus zone during 1923.

In 1927 Broom described the important skull of Anningia megalops found

by Van der Byl on the farm Bloukrans, Prince Albert.

In 1928 Broom described the type skull of Tauwrocephalus lerouxt found by

J. J. le Roux and his brother on Abrahamskraal and of Criocephalus vanderbyli

found by Van der By! on the same farm and he also figured a partial skull of

Tapinocephalus found by M. J. van Wyk on Ganskraal, Prince Albert. Bones

found by Haughton in 1917 on Sewefontein, Prince Albert, were in error

figured as those of Tapinocephalus, whereas they are in fact titanosuchian

(Scapanodon septemfontis).

In 1928 Boonstra, without any own means of conveyance and relying

wholly on the friendly farm-owners for transport from farm to farm, started

on a collecting trip from Beaufort West. On Boeteka he collected a partial

femur of Phocosaurus; on Mierfontein an ilium and humerus of Jonkeria and a

pareiasaurian humerus and ulnae; on Leeufontein two fair specimens of

Bradysaurus seeleyi, on Boesmansrivier the type specimen of Styracocephalus

platyrhynchus and three poor pareiasaurs; on Mynhardtskraal six pareiasaurs

including the one which yielded the first completely preserved manus, and

postcranial bones of a struthiocephalid and of Keratocephalus; and on Klein-

koedoeskop the type skull and skeleton of Hipposaurus boonstrai, a good

Nochelesaurus skull and a pareiasaurian pelvis and a fibula of Parascaponodon.

J. CG. Avenant of Die Walle, part of Kleinkoedoeskop, was hired to take the

specimens to Letjiesbos station and Boonstra got him interested in locating

fossils. The result was that early in the following year Boonstra went up to

excavate a number of fossils located by Avenant on Koedoeskop and on this

trip some of the neighbouring farms were also worked with the help of Avenant’s

donkeycart. From Kleinkoedoeskop the specimens which later became the

types of Jonkeria koupensis, Scymnosaurus major and Robertia broomiana were

collected in addition to four pareiasaurs, eight dinocephalians, 40 dicynodonts,

four therocephalians and a gorgonopsian; from Rietkuil were obtained what

later became the type of Pristerosaurus microdon, a Scymnosaurus and a Pristerog-

nathus, a gorgonopsian and three dinocephalians and from Boesmansrivier an

Anteosaurus snout. Arrangements were then made to hire Avenant’s donkey-

wagon for a three-month trip commencing in March. This trip yielded the

following: from Die Vlei a Pristerognathus and two other therocephalians; from

Stinkfontein an Alopecognathus and another therocephalian; from Sewefontein

a lower jaw of Brachypareia watsoni; from Voélfontein the specimen which later

12 ANNALS OF THE SOUTH AFRICAN MUSEUM

became the type of Parascapanodon avifontis, three Bradysaurus seeleyi with feet,

a Bradysaurus baint and two Embrithosaurus, a good Anteosaurus abelt skull and

six other dinocephalians, a Scymnosaurus and two fragmentary stegocephalians;

from Seekoeivlei four Bradysaurus and two Fonkeria haughtont; from Vindraers-

fontein a dinocephalian femur; from Boorfontein and Klipbanksfontein each

a gorgonopsian; from Knoffelfontein a complete skeleton with feet of a

Bradysaurus seeleyt; a Bradysaurus vanderbylt, Brachypareia rogersi, a pareiasaurian

femur and a fair skull of Scymnosaurus major found by the owner, Snyman; from

Kruisvlei the specimens which years later became the types of Avenantia

kruisvleiensis and Anteosaurus acutirostris and in addition a Bradysaurus seeleyt

and the mass entombment of cranial and postcranial elements of at least a

dozen Moschops individuals. On the Merweville Commonage a femur of

? Tapinocephalus was found; on Melkbosfontein four Bradysaurus baini—one a

giant with all four feet—were collected; on Welgemoed an Embrithosaurus

with feet; on Wilgerfontein a poor pareiasaur and dinocephalian; on Saai-

rivier the type humerus of Jonkeria parva; on Jacobskraal a tapinocephalid, a

therocephalian and an amphibian; from Wolwefontein (Wakkerstroom) a

paratype of Parascapanodon avifontis, two Fonkeria, a tapinocephalian and a

moschopid, two pareiasaurians and a scylacosaurid were collected.

Rietfontein, Prince Albert, yielded a beautiful moschopid fibula; Droé-

fontein a dinocephalian as also Elandskop; on the last farm worked, Veld-

mansrivier, a scapulo-coracoid of Parascapanodon and a skull of Brachypareia?

rogerst were found.

In 1929 Broom in his review of the Titanosuchidae designated the

following new types: Jonkeria vanderbyli for a specimen collected by Broom. on

Abrahamskraal, Jonkeria crassus for a poor specimen found by Van der Byl

near Kruidfontein Station, Dinosphageus (now Fonkeria) haughtoni for a skull

and some postcranial elements collected in 1916 by Haughton on Welgemoed,

Phoneosuchus (now Jonkeria) angusticeps for a lower jaw found by Van der Byl

on Abrahamskraal, Jonkeria pugnax (now ingens) for a skull found by M. J.

van Wyk on his farm Kookfontein and Anteosaurus (now Pseudanteosaurus) minor

found by Broom near Merweville.

Haughton (1929c) described the following therapsids: an imperfect skull

found by Boonstra on Boesmansrivier was made the type of Styracocephalus

platyrhynchus, a skull found by himself on Abrahamskraal in 1920 as Eoarctops

vanderbyli and a second gorgonopsian Hipposaurus boonstrai has as type specimen

a skull of the nearly complete skeleton collected by Boonstra on Kleinkoedoeskop.

In the same part of the Annals of the South African Museum Haughton &

Boonstra attempted a classification of the Pareiasauria and the following new

names were proposed for forms from the Tapinocephalus zone: Bradysaurus

seeleyi and Bradysaurus vanderbyli, Bradysuchus (now Bradysaurus) whaitsi,

Nochelesaurus alexandert and Pareiasaurus strubent became Nochelesaurus strubent,

Dolichopareia angusta and P. acutirostris became Koalemasaurus acuticostris,

Brachypareia watsoni and in this new genus was also placed the former Propappus

THE FAUNA OF THE TAPINOCEPHALUS ZONE 13

rogerst and lastly Platyorpha broomt.

Haughton (19296) also described the braincase in a number of pareiasaurs.

Boonstra in two further papers in the series of ‘Pareiasaurian studies’

described the first complete manus in the specimen of Bradysaurus vanderbyli

collected by himself at Mynhardtskraal and some incomplete pedes.

In December 1929 Broom figured the shoulder girdle of Pristerognathoides

minor.

During 1929 two major scientific conferences were held in South Africa,

viz. the joint meeting of the British and the South African Associations for the

Advancement of Science and the XV International Geological Congress.

Before, during and after these meetings some of the visiting palaeontologists

collected in the Tapznocephalus zone. They were W. Janensch of Berlin, Othenio

Abel of Vienna and A. S. Romer with the veteran American collector Paul

Miller as his assistant, whereas D. M. S. Watson was more interested in the

younger zones.

Abel, spending a few days under the guidance of L. D. Boonstra, obtained

a few bones of pareiasaurs and dinocephalians and a paper published in 1930

contained nothing new. Romer and Miller worked from two camps -— Stink-

fontein and Hottentotsrivier. I quote two paragraphs of a letter from Romer

giving his views on the nature of Tapinocephalus zone matrix and collecting in it:

‘We found collecting in the Tapinocephalus zone pleasant and

comparable in many ways to conditions in our western states. We camped

out, with a cook and general camp helper. The terrain in the zone is

relatively flat, and fences few, so that, much as in many western areas,

we found little difficulty in reaching any locality by car — particularly

useful in picking up ponderous pareiasaur or dinoceph materials. The roads

were few and poor—parts of the Main Cape to Johannesburg road were

no more than a pair of ruts in the veldt, if that—but that, again, was a

familiar condition in our own West at that time. The local farmers were

friendly and cooperative. And the total lack of rainfall during our visit

to the Gouph was, of course, a boon to the collector (although not to the

sheep farmer).

The one great disadvantage to collecting in this zone is, of course,

one with which you are all too familiar, but for which we were not fully

prepared—the exceedingly hard nature of the mudstone matrix. I

remember all too well our first day in camp at Stinkfontein. Miller and I

set out to prospect in the morning and met again for lunch. Both of us

had seen an excellent fossil prospect, and went out in the afternoon to

develop our finds. When we met again for supper, both of us were exceed-

ingly discouraged. We ruined our chisels, dulled our picks, and made

almost no impression on the rock!’

A list supplied by D. Techter gives the ‘booty’ taken to Chicago as: about

a score each of Dinocephalia and Pareiasauria, three Amphibia, at least six

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

Therocephalia and a number of anomodonts. Except for a few described by

Broom and Olson (Struthiocephalus millert and Rhinesuchoides tenuiceps) and.

Moschoides romeri (Byrne) and Brachyprosopus broomi Olson and some studied, but

specifically unidentified by Olson, ‘the vast bulk of the material has never

been studied, catalogued, or even adequately prepared’ (letter dated 24 May

1956 from D. Techter to the author).

Janensch collected for the Palaontologisches Museum of the Humboldt

University in Berlin the following specimens: seven pareiasaurs, three dino-

cephalians and two therocephalians.

In 1930 Broom published a figure of Scylacognathus parvus.

Boonstra’s study of the pareiasaurian mandible was published as a joint

paper with Haughton and the material studied included 27 specimens from

the Tapinocephalus zone. Later in the same year Boonstra’s work on the pareia-

saurian hind-limb was again published as a joint work with Haughton and

was based on 26 specimens from the Tapinocephalus zone and some from younger

zones.

Haughton’s paper before the International Geological Congress included

a discussion of the fauna of the Tapinocephalus zone.

In 1931 Von Huene’s paper appeared in which he gave an account of

the most important specimens he had collected in 1924. The Tapinocephalus

zone forms described were: the type specimen of Keratocephalus moloch from

Abrahamskraal; the type skull of Dicynodon haughtonianus from Bloukrans and

further a skull of Bradysaurus baint and a lower jaw of Nochelesaurus strubent,

both from Abrahamskraal, as also postcranial bones of a Jonkeria and tapino-

cephalids and lastly ten specimens of Dicynodon joubertt and four of Dicynodon

megalorhinus, all from Bloukrans.

In 1931 Broom proposed the new generic name, Cynariognathus, for the

specimen in the American Museum collected by Whaits on Grootfontein,

Beaufort West, previously described as Pristerognathus platyrhinus. |

In 1931 Boonstra, travelling over the Tapinocephalus zone, managed to

collect the type of Alepognathus angustioriceps on Kroonplaas, a pareiasaurian

humerus on Boesmansrivier, some dicynodonts on Kroonplaas and a struthio-

cephalid scapula on Grootfontein in the Fraserburg district. In the same year

Van der Byl presented a Criocephalus skull from Abrahamskraal to the South

African Museum. In 1932 a partial skull of Keratocephalus moloch was obtained

from Fraserburg Road.

In February, 1932, Boonstra’s account on the fore-limb of the pareiasaurs

was published which included reference to 12 specimens of Bradysaurus bainq,

seven of Bradysaurus seeleyi, three of Bradysaurus vanderbyli, six of Bradysaurus

sp., one of Platyorpha, one of Nochelesaurus strubent and two of WNochelesaurus

alexanderi, three of Dolichopareia angusta, five of Embrithosaurus schwarzi, and

four of Brachypareia. Hans Reck collected in the Koup during 1932.

In October, 1932, Boonstra published on the palaeobiology of and the

divergence in the Pareiasauridae and on the hyoid apparatus in the Pareia-

THE FAUNA OF THE TAPINOCEPHALUS ZONE 15

sauria. In the same year Broom’s Mammal-like reptiles of South Africa was

published by Witherby and in this book the following Tapinocephalus zone

forms were included: one anningiamorph, 32 dinocephalians, 33 thero-

cephalians, two dromasaurians, six gorgonopsians, one burnetiamorph and

three anomodonts.

During 1933 Boonstra wrote on the distribution of the -pareiasaurians in

the Koup and on a method used in the excavation of these reptiles.

In the same year Grossarth collected for the Alte-Akademie in Munich.

In April, 1934, Boonstra described the cranial osteology in the series of

pareiasaurian studies in which 49 skulls from the Tapinocephalus zone were

studied. This was followed by an account of the dermal armour and the

vertebral column and ribs. In the whole series of pareiasaurian studies the

various specimens were assigned to a stratigraphic position within the zones,

viz. low, mid and high, and this was done on the then prevalent idea that the

rocks in the Koup were lying in the main horizontally and that the present

day topography indicated the stratigraphic position. Rossouw & De Villiers’s

work (1952) have since shown that this was false.

In June, 1934, Van Hoepen, in his paper on the classification of the

Dicynodontia, proposed new generic names for some of the forms from the

Tapinocephalus zone—but these have not been accepted by later workers and

fall away.

In July, 1934, Boonstra published three papers in the Annals of the South

African Museum. The first (1934a) included an account of the postcranial

skeleton of Hipposaurus boonstrai found by himself in 1928; the second (19345),

on the Gorgonopsia in the British Museum, included a re-study of the type of

Cyniscodon lydekkert; and in the third paper (1934c) the following Therocephalia

from the Tapinocephalus zone in the British Museum were re-examined:

Alopecodon, Cynariognathus, Pristerognathus, Scylacosaurus, Scymnosaurus, Trocho-

saurus, Cerdodon, Ictecephalus and the doubtful Hyorhynchus and Theriodesmus.

Towards the end of 1934 Broili & Schréder described a gorgonopsian

specimen found by Grossarth in the previous year on La-de-da and made this

skull the type of Pachyrhinos kaiseri.

During January, 1935, Boonstra published three papers in the American

Museum Novitates on specimens in the American Museum. In the first it was

proposed that Broom’s Pareiasaurus whaitsi, later made the genotype of Brady-

suchus by Haughton and Boonstra, be considered a species of the genus Brady-

saurus ; in the second paper the following Therocephalia from the Tapinocephalus

zone were re-studied: Alopecodon, Alopecognathus, Cynariognathus, Scylacorhinus

falkenbacht synonym of Scylacosaurus sclateri and Trochosaurus and thirdly a new

account was given of the type specimen of the gorgonopsian, Eriphostomna

microden.

During 1935 Broili & Schréder published two papers in which specimens

from the Tapinocephalus zone are described: from Brakwater, Schréder brought

a dinocephalian, which was named Titanognathus (now Anteosaurus) lotzi; in

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

the second paper a new form of endothiodont, Brachyuraniscus reuningi, from

La-de-da, collected by Grossarth in 1933, is described.

In October, 1935, Broom described Lycedops scholtzi, collected by C. S.

Scholtz on Abrahamskraal and in ‘Notes on some species of pareiasaurian

reptiles’ he wrote about the forms from the Tapinocephalus zone. Finally, in

1935 Boonstra collected a pareiasaurian skull on Voélfontein.

Early in 1936 Broili & Schréder described Cynariognathus seeleyi on a skull

found by Grossarth in 1933 on the farm La-de-da.

In May, 1936, Broom described a skull found on Stinkfontein under the

name Dinosuchus (now Anteosaurus) vorsteri, a snout found by Van Hoepen on

Abrahamskraal many years before became the type of Dinopolus airox, and an

imperfect skull was made the type of Trochorhinus vanhoepent. A skull found by

Broom at Letjiesbos was figured as that of Pristerognathus baini, and a specimen

discovered by J. A. Galloway at Luttig became the type of Cynariognathus

gallowayt.

In August, 1936, Boonstra redescribed both the tapinocephalian and the

titanosuchian cranial material housed in the American Museum. In September

Broom (1936a) gave a fuller description of Dinosuchus (now Anteosaurus) vorstert.

In November Broili & Schroder described a skull found by Grossarth on

Klein-Waterval under the name Pristerognathus peyeri.

In the Philosophical Transactions Broom figured sections through a number

of therocephalian skulls including that of a new species — Trochosaurus dirus.

During 1936 Camp collected in the Koup but I have not been able to get

a list from him of the specimens collected.

In June, 1937, Broili & Schréder described some anomodont skulls that

had been collected by Grossarth on La-de-da under the name Dicynodon (now

Oudenodon) huenet and synonyms of this species under the names Dicynodon

broomi and D. grossartht. In September Broom (19370) described the skull of

Elliotsmithia longiceps which had been obtained in about 1917 by Van Hoepen

from Abrahamskraal, and a skull from Grantham as Struthtocephalus rheedert

and thirdly a skull from Klipbank under the name Dicynodon gamkaensis.

In October, 1937, Olson & Broom described some specimens in the Walker

Museum, Chicago, that had been collected by Romer and Miller in 1929,

viz. a skull of Struthiocephalus millert and one of Rhinesuchoides tenuiceps, both from

Stinkfontein.

In November Broom (1937a) gave the name Alopecognathus megalops to a

skull found by Van der Byl in 1929 on Abrahamskraal.

In December Olson fully described a skull collected by Romer and Miller

on Hottentotsrivier under the name Brachyprosopus (now Brachyuraniscus) broomt.

Also in 1937 Byrne named a skeleton collected by Romer and Miller on

Hottentotsrivier in 1929 Moschops romert.

Finally in 1937 Boonstra collected the types of Anteosaurus major on

Boesmansrivier and of Anteosaurus abeli on Kruisrivier in the Sutherland district.

In July, 1938, Olson sectioned and described an unidentified thero-

THE FAUNA OF THE TAPINOCEPHALUS ZONE 17

cephalian skull collected in 1929 by Romer and Miller on Sewefontein.

During 1938 Boonstra, collecting in the Koup, found on Deesweesfontein

the type skull of Aoupia koupensis and limb bones of Phocosaurus; on Buffelsvlei

a paratype of Anteosaurus crassifrons and three tapinocephalids, a pareiasaurian

skull on Skoppelmaaikraal, and on Boesmansrivier a pareiasaurian skull, a

femur of Phocosaurus and some limb-bones of Parascapanodon.

During May, 1939, Boonstra collected on Buffelsvlei five therocephalians

belonging to the genera Pristerognathus and Scymnosaurus and a Glanosuchus

macrops, a Dicynodon pseudojouberti and a weathered titanosuchian skull; from

Koedoeskop a jonkeria haughtoni; from Bloukrans per P. J. Rossouw a Dicynodon

joubertt and a Bradysaurus skull from Mynhardtskraal.

In December, 1939, Boonstra published his review of a century of work

on the Karoo.

In 1940 Von Huene included in a compilation the then known forms

from the Tapinocephalus zone, viz. one labyrinthodont, 13 Pareiasauria, one

archichelonian, three anningiamorpha, 21 titanosuchids, 15 tapinocephalids,

two dromasauria, 30 pristerognathids, three alopecopsids, two ictidosuchids,

two scaloposaurids, nine Gorgonopsia, one burnetiamorph, nine anomodonts

and Tamboeria maraisit. In 1940 Boonstra described Rhinosuchus avenanti from a

skull found by Avenant on Mynhardtskraal. Romer & Price in their ‘Review

of the Pelycosauria’ included the following forms from the Tapinocephalus zone

in the Varanopsidae?—Elliotsmithia and Anningia, and Byrne made some

comparisons of his Moschoides with other therapsids.

Finally during 1940 Boonstra collected on Mynhardtskraal the type of

Paranteosaurus primus, Anteosaurus levops, a paratype of Parascapanodon avifontis, a

good Bradysaurus seeleyi skull, a juvenile Struthiocephalus whattst, a femur of

Titanosuchus ferox, two Jonkerias and a Galesuchus; on Cypher some therocephalian

limb-bones, parts of an endothiodont and a Dicynodon joubertt; on Voélfontein

a Bradysaurus; on Kroonplaas the type of Cynariognathus paucioridens and nine

dicynodonts and on Kleinkoedoeskop four Jonkerias, one Anteosaurus abeli, a

Struthiocephalus whaitst, a Pristerognathus and four dicynodonts; on Bulwater a

good skull of Anteosaurus vorstert; on Buffelsvlei a Struthtocephalus, a Struthio-

cephaloides and a moschopid femur and from A. C. Bothma of Die Krans he

obtained a new species of Moschops and from Aasvoélbos a Scapanodon

septemfontis.

In 1941 Broom reconsidered the position of Broomia and Eunotosaurus and

H. Zinn collected a specimen of Dicynodon pseudojoubertt from Dikbome in the

Laingsburg district.

In March, 1942, Watson in a review of the Permian and Triassic tetrapods

discussed the dinocephalians of the Tapinocephalus zone. In November Camp;

Taylor & Welles proposed for Dinosuchus the name Broomiasuchus but this skull

really is a species of Anteosaurus. Von Huene suggested that the vertebra

described by Seeley under the name Tamboeria maraisi may be that of a pristerog-

nathid therocephalian. During a short collecting trip in 1942 Boonstra collected

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

on Dikbome the type skull of Anteosaurus laticeps and a skull of Struthiocephalus

whaiist.

In a comparative paper on the cranial morphology of the therapsid

suborders, Olson in 1944 studied the following skulls from the Tapinocephalus

zone: two anomodonts from Hottentotsrivier and another two from Stink-

fontein, of the former one is the geno-holotype of Brachyprosopus (now

Brachyuraniscus) broomi, a therocephalian from Sewefontein and a pristerognathid

from Hottentotsrivier. These specimens were all collected by Romer and

Miller during 1929.

In 1945 Hesse collected the following specimens at Prince Albert Road:

a Pristerognathoides vanwyki, a Robertia broomiana, a Dicynodon pseudojoubertt and a

D. joubertt.

During 1946 Boonstra collecting with the help of J. du Plessis, on Dikbome,

the type skull of Struthiocephaloides cavifrons and, on Koringplaas in the Laings-

burg district, the type skull of Anteosaurus cruentus. In the Sutherland district

Boonstra obtained from Esperance a Robertia broomiana and a Dicynodon pseudo-

jouberti, and on Elandsberg a Dicynodon pseudojoubertt, two tapinocephalids and

an unidentified tapinocephalian; in the Fraserburg district on the farm Selery-

fontein an unidentified therapsid and a Dicynodon were found and on both

Rietfontein and Kookfontein a Dicynodon joubertt.

Boonstra in 1947 collected the following: on Dubbelfontein (Brits Eigen-

dom) 58 Dicynodon skulls and two weathered therapsid skulls; on Lammerkraal

through Mr. J. Pienaar, the owner, the type skull of Agnosaurus pienaari and

in addition eight Dicynodons, a Scymnosaurus and four other therocephalians;

from Ve'dmansrivier three gorgonopsians and a Pristerognathus; on Perdewater

six skulls of Pristerognathus and 21 Dicynodons.

In 1948 Watson discussed certain cranial features of the tapinocephaloids

and titanosuchids. In the same year Boonstra reviewed the anomodonts of the

Tapinocephalus zone and erected the new genera Robertia and Koupia and the

new species Dicynodon pseudojouberti and as the name was preoccupied, proposed

that Broili & Schréder’s Dicynodon huenei be named D. broiliz (now Oudenodon

huener). Localities of all the specimens from the zone in the South African

Museum are given. Boonstra in a semi-popular book gave an account of the

Tapinocephalus zone and included reconstructed life scenes depicting Bradysaurus

seeleyt, Keratocephalus and a titanosuchid and a tapinocephalian (executed by

Hesse) and a!so a Moschops in the flesh.

Finally, in 1948, Boonstra collected the following specimens: on Bloukrans

a paratype of Parascapanodon avifontis, on Steenboksfontein two Dicynodons; on

Skroefpaal a good Jonkeria skull that had been located by Rossouw; on Michau’s

Request a mass of Dicynodon skeletons that had been located by Jooste; on

Vindraersfontein the type of Theroides cyniscus, a Fonkeria and an Anteosaurus;

on Bosluiskraal and Skoorsteen a number of Dicynodons and on Lammerkraal

three skulls of Pristerognathus, one of which had been found by the owner, J.

Pienaar. Boonstra also accompanied P. J. Rossouw, who was mapping the

THE FAUNA OF THE TAPINOCEPHALUS ZONE 19

area, to indicate the spots where previous finds had been made. Zinn also

collected two Dicynodons on Dikbome. In 1948 the Geological Survey presented

a tapinocephalian femur found at Arthurskraal—one of the most easterly

localities of the Tapinocephalus zone that have yielded fossils.

In 1951 F. P. Grobbelaar presented a Dicynodon jouberti found at Fraserburg

Road to the South African Museum. Boonstra described a skull of Keratocephalus

moloch from Fraserburg Road which had been obtained in 1932, and to this is

appended a note by Von Huene on the Tiibingen specimen. Brink & Kitching

described the snout of Walteria skinneri which came from the farm Elandsfontein.

Finally for this year Boonstra collected: on Dikbome a hind-limb with

pes of Bradysaurus and five pristerognathids, on Steenboksfontein the type of

Ptomalestes avidus a tibia of Parascapanodon, two therocephalians, two Dicynodons

and a Mormosaurus; on Droéfontein a Dicynodon and an interclavicle of

Struthiocephalus, on Bosluiskraal a Pristerognathus; on Buffelsvlei (assisted by J.

Marais) the type of Maraisaurus parvus, a paratype of Anteosaurus crassifrons, a

Keratocephalus, a Dicynodon, a moschopid and a Pristerognathus; on Vereniging a

skull of Brachypareia; on Skoppelmaaikraal a Pristerognathus.

In 1952 Boonstra published a number of short papers in which were

described: the type skull of Struthtocephalus akraalensis, Hipposaurus major, with

a life-sized reconstructed model of Hipposaurus boonstrai, the type skull of Anteo-

saurus abeli, a redescription of the type skull of Struthiocephalus whaitsi, the type

skull of Struthiocephalus (now Struthiocephaloides) duplessisi, the type skull of

Struthiocephaloides cavifrons, the type skull of Agnosaurus pienaari, the type skull

of Riebeeckosaurus longirostris, that of Avenantia kruisvleiensis and the type skull of

Struthionops intermedius. Also in 1952 Rossouw & De Villiers published their

sheet map of Merweville and the explanation thereto in which the known

fossils occurring within the area mapped are listed with the localities of each

indicated and the Tapinocephalus zone sub-divided into three horizons, with

tables indicating the distribution of the fossils.

Boonstra in 1952 collected the following specimens: on Nuwefontein

(Roxana) in the Nieuweveld a complete skeleton of Bradysaurus seeleyi, and a

fair skull of Anteosaurus abeli, both located by Jooste; on Kruis van Bloemfontein

a Eunotosaurus picked up by F. D. Visser; and on Gatsplaas, Prince Albert, a

skull of Struthiocephalus and some Dicynodon skulls. Mr. J. J. Hugo of Vanaswegens-

fontein, Loxton, also sent in some teeth of probably Struthiocephalus, from what

was up to that date the furthest north-easterly site recorded in the Tapinocephalus

zone. .

In 1953 Toerien published an important paper in which a basis for the

classification of the anomodonts is presented. All the known forms from the

Tapinocephalus zone are considered and the following new forms described:

Brachyuraniscus merwevillensis from Buffelsvlei collected by the C. J. van der

Horst Expedition in 1945, Broilius antjiesfonteinensis collected by Kitching in

1946 on Antyjiesfontein, Dicynodon vanderhorsti from Antjiesfontein, also found

by Kitching, as also Dicynodon antjiesfonteinensis and Dicynodon schroederi, collected

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

by Kitching on Klein-Waterval.

In the same year Boonstra attempted to bring the known titanosuchians

into taxonomic order and published a photograph of a life-sized reconstruction

of the head of Anteosaurus. He also published two photographs of the life-sized

reconstruction of Hipposaurus and an account of the cranial morphology of

Struthiocephalus with a life-sized reconstruction of the head, and finally in this

year discussed the articulatory region in some pristerognathids and proposed

the following new forms: Theriodes cyniscus, Pristerognathus roggeveldensis,

Maraisaurus parvus and Alopecognathus angusticeps.

In a lecture on Gondwanaland during 1953 Haughton discussed the

Tapinocephalus zone and its fauna. In this year the owner of the farm Sandrivier,

R. L. Jack, found part of the skull of a Tapinocephalus atherstonet.

In 1954 Boonstra studied the pristerognathids in the South African

Museum and described the following new forms: Scymnosaurus major, Ptomalestes

avidus, Pristerosaurus microdon, Cynariognathus paucioridens, the new generic name

Pristerognathoides for Alopecognathus minor, and Pristerognathus roggeveldensis and

Pristerognathus vanwyki, and erected a new species Pristerognathoides parvus.

In a study of the genus Anteosaurus Boonstra (1954a) named the following

new forms: A. acutirostris, A. crassifrons, A. major, A. laticeps, A. cruentus, A. levops,

A. minusculus and later also Micranteosaurus parvus and Paranteosaurus primus, and

for Anteosaurus minor the new generic name Pseudonteosaurus.

In February, 1954, Boonstra made a skull, collected by Cairncross in 1918

near Prince Albert Road, the type of a new species of scaloposaurid, under

the name Blattoidealestes gracilis.

In January, 1955, Boonstra described a specimen, collected by Haughton

in 1916 on Abrahamskraal, under the name Struthiocephalellus parvus. In a

detailed comparative account of the girdles and limbs of the South African

Dinocephalia Boonstra also proposed the following new names for new forms:

Scapanodon septemfontis, Parascapanodon avifontis, Fonkeria koupensis, Fonkerta parva

and Jonkeria rossouwi and divided the South African Dinocephalia into four

infra-orders, viz. Tapinocephalia, Titanosuchia, Anteosauria and Styracoce-

phalia.

In 1955, in his book Evolution of the vertebrates, Colbert suggested the splitting

up of the Dinocephalia, thus anticipating Watson & Romer (1956). Colbert

published only one figure of the dinocephalian skull, and this of the inade-

quately known Delphinognathus, and quite erroneously shows a fossa between

jugal and quadratojugal.

In 1956 Watson & Romer, in their classification of the therapsids, split

the sub-order Dinocephalia — placing the infra-order Titanosuchia composed of

the Brithopodidae, Anteosauridae and Jonkeriidae in the sub-order Therio-

dontia and the infra-order Dinocephalia (Tapinocephalia) composed of the

Deuterosauridae, Moschopidae and ‘Tapinocephalidae in the sub-order

Anomodontia.

This arrangement is followed by Romer in his Osteology of the reptiles, but

THE FAUNA OF THE TAPINOCEPHALUS ZONE 21

Von Huene in his textbook retains the Unterordnung Dinocephalia with a

Famielienkreis Tapinocephaloidae (Moschopidae, Struthiocephalidae, Tapino-

cephalidae) and a Familienkreis: Titanosuchoidea (Deuterosauridae, Brithopo-

didae, Titanosuchidae, Jonkeriidae) and a Familienkreis Burnetiamorpha

consisting of Styracocephalus and Burnetza.

Kermack in 1956 studied the tooth replacement in some therocephalians

from this zone housed in the British Museum.

Boonstra and Zinn collected eight Therocephalia, eight Pareiasauria, 20

Dinocephalia, four Anomodontia, one amphibian and two fishes on a trip in

the Tapinocephalus zone.

Also in 1956 Boonstra published figures of skulls of Tapznocephalus,

Phocosaurus, Mormosaurus and Keratocephalus and a photograph of a life-sized

reconstruction of Bradysaurus seeleyt exhibited in the South African Museum.

During 1957 Boonstra and Zinn undertook four short collecting trips in

the TYapinocephalus zone obtaining 24 pareiasaurian, 61 dinocephalian, 14

therocephalian and three anomodont specimens. On one of the trips S. Fourie,

post-graduate student of Stellenbosch (later palaeontologist at the National

Museum in Bloemfontein), was taken along.

In the same year Boonstra established the new species Moschops koupensis

and figured other moschopids in the museum collection and showed that in

Delphinognathus there is no lateral foramen but a notch between jugal and

quadratojugal.

In 1958 Brink described a good skull of Struthtocephalus kitchingi.

During 1959 Boonstra, with Zinn, C. Gow and R. Rau as assistants,

during two trips in the Koup collected two pareiasaurs, 71 therocephalians,

381 anomodonts, 41 dinocephalians (including seven rather poor specimens of

Styracocephalus of which hitherto only the unique type specimen was known),

three stegocephalians, one Eunotosaurus and four gorgonopsians.

In 1960 three collecting trips were undertaken by Boonstra and Zinn,

with Gow and H. Boonstra accompanying them on one trip each. The haul

consisted of 31 dinocephalians, 21 therocephalians, 92 anomodonts, 11 pareia-

saurs, one stegocephalian and a fairly good skull of the rare Hipposaurus.

During 1961 Boonstra had the opportunity of visiting Moscow where he

studied all the Russian material related to the Tapinocephalus zone forms of

South Africa.

In 1962 Boonstra published a paper on the dentition of the titanosuchian

dinocephalians and showed that these forms, although retaining a canine,

were really herbivores.

In the same year Boonstra and Zinn explored both the southern and

northern outcrops of the zone east of longitude 22° 35’ and found the exposures

poor in the southern part but good in the north, but both areas yielded only a

couple of scraps. It would thus appear that the fauna of those times were

concentrated in the area to the west of this line.

In 1963 Boonstra in his paper ‘Early dichotomies in the therapsids’,

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

including data obtained during his Russian visit, emphasized the fact that the

therapsid fauna of the Tapinocephalus zone was already greatly diversified and

that the main branches must have arisen in Ecca times. In a further paper,

‘Diversity within the South African Dinocephalia’, the relationships of the

four families (Anteosauridae, Titanosuchidae, Tapinocephalidae and Styraco-

cephalidae) were stressed as also the herbivorous habit of the Titanosuchidae.

It was also suggested that the various species of Struthiocephalus represented

growth stages of a single species.

Since 1964 Boonstra has published a series of studies on the girdles and

limbs of the early Therapsida and also an account of a very well preserved

skull of Struthiocephalus, where the relations of the stapes could be determined.

The studies on the locomotor apparatus of the early therapsids were con-

cluded in 1967 with a paper entitled ‘An early stage in the evolution of the

mammalian quadrupedal walking gait’.

In 1967 a paper appeared by Cys based on a therocephalian specimen

collected by Camp.

In 1968 Boonstra published a paper on the basicranial axis of the Dino-

cephalia based on serial cross-sections cut by circular diamond-studded saws.

In the same year Boonstra also published a paper, “The terrestrial reptile

fauna of Tapinocephalus zone age and Gondwanaland’.

THE FAUNA AS A WHOLE

Since Owen in 1876 described the first specimen from the Tapinocephalus

zone under the name Tapinocephalus atherstonet, 152 species of reptiles have

been named from this zone in South Africa by 15 authors.

The number of specimens collected from this zone is only very approxi-

mately known. I have not been able to obtain accurate lists from the various

institutions, in South Africa as well as overseas, which house such collections.

The South African Museum houses the largest collection, viz. 1,553 specimens.

The other South African institutions together house approximately 77 of

which the Bernard Price Institute has 58. In overseas institutions there must

be over 400 specimens; in London there are at least 104, in Chicago 93, in

New York 41 and an unknown number in Washington, California, Munich,

Berlin and Vienna. It is thus very probable that well over 2,000 specimens have

been collected from this zone.

To indicate the relative occurrence of the various groups of reptiles I

give here a table of the number of specimens in each group in the collection of

the South African Museum:

Dicynodontia 705

Therocephalia 4.22

Dinocephalia 439

Pareiasauria 160

Gorgonopsia 22

Eunotosauria 5

THE FAUNA OF THE TAPINOCEPHALUS ZONE 23

All the known specimens from this zone in South Africa come from the

western part of the main Karoo Basin, lying approximately between longitudes

20° to 23° and latitudes 31° 20’ to 33°. The area concerned cannot be more

than about 20,000 sq. miles, which gives a yield of about 1 specimen per

10 sq. miles. The Koup, lying between the Nuweveldberge and the Swartberg,

has yielded well over 90% of the specimens. The rest come from the Moorde-

naar’s Karoo and a strip of the Nuweveld stretching from the Klein Roggeveld

eastwards through the towns of Sutherland, Fraserburg and Loxton.

It is not known with certainty whether rocks of this zone extend north-

eastwards into that part of the main Karoo Basin which extends into the

Transvaal and Natal to form a narrow outcropping belt between the under-

lying Ecca beds and the overlying higher zones of the Karoo beds. If so, the

basin in which the sediments of Tapinocephalus zone times were deposited would

be very large indeed, but on the inadequate facts I believe that the zone

wedges out. The present extensive drilling could supply the much needed

stratigraphic data as a compensatory by-product if the search for oil in the

Karoo beds proves unsuccessful, as I believe it will. At present we do know

that the zone crops out east of longitude 23° as two belts lying respectively

north and south of a westward tongue of younger Beaufort beds.

I have undertaken collecting trips eastwards along both these belts of

outcrops; in the south as far as King William’s Town and in the north as far

as Deelfontein and found some scraps of bone. In the northern belt good

exposures are present, but the southern belt is affected by the Cape Foldings

and the beds lie at steep angles. This, together with the nature of the vegetation,

reduces the extent of the exposures for collecting purposes, but even so some

identifiable specimens would have been found if present in the numbers known

from the western part of the basin.

We thus have to conclude that during Tapinocephalus zone times only the

western part of the Karoo Basin was populated by a reptilian fauna living

under conditions in which entombment was possible.

It is thus evident that without the help of the geological cartographer the

palaeontologist has to work at a serious disadvantage. Only parts of the southern

limits of the Tapinocephalus zone have been accurately mapped by our Geological

Survey, but this institution, whose function it is, could very easily map the

rest of the Ecca—Tapinocephalus zone boundary, although some other lithological

criterium would have to be used for the northern boundary where the zone

has a facies different to that in the south. Nowhere has the boundary between

the Tapinocephalus and Endothiodon zones been accurately mapped. The Geo-

logical Survey has, moreover, done little to determine the conditions under

which the sediments of this zone have been formed. Differences in the nature

of the deposition from west to north-eastwards may very well account for the

lack of entombment if in fact the fauna extended thus far. Only for the Merwe-

ville sheet — 198 —have adequate details of the nature of the zone been given

and in the Skoorsteenberg — 166, Gamkapoort — 3321B, Prince Albert — 3322

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

and Grahamstown — 136 sheets less full details have been given. Here, Rossouw

& De Villiers have found it possible to establish a threefold subdivision on

lithological grounds and this has been a great help to the palaeontologists

studying this fauna. If only we could get the contiguous areas of the zone mapped

in this manner!

In the type area the Tapinocephalus zone is from six to seven thousand

feet in thickness and consists of a succession of sandstones and mudstones with

bands of chert and bands and lenses and nodules of calcareous and marly

rocks. These are not continuous but wedge out laterally in irregular series of

overlapping lenses. For a more detailed account of the lithology I refer you to

the work of Rossouw, De Villiers and their colleagues. Here it will suffice to

state that the beds were laid down in a fairly warm, rainy climate over a fairly

level flood plain area, with periodic flooding and the formation of lakes,

marshes and shallow pans or vleis which silted up with the inclusion of the bones

of the reptiles living in or around these water patches.

Little is preserved of the flora in the form of leaves but fossil (silicified)

wood is fairly abundant. There must, however, have been a well-developed

Glossopteris flora considering the large number of herbivores in the fauna, but

here again it is evident that plants and bones seldom tend to be preserved in

the same beds. It has been suggested that the Ph values of the sediments have

a bearing on this phenomenon.

An account of the nature of the preservation of the remains of the different

groups constituting the fauna will be of interest in showing the nature of some

of the ecological conditions prevailing at this time.

The pareiasaurs are mostly found in blue mudstone as coniplete skeletons,

whose position indicates that the animals were entombed standing on their

legs with the head held high as if they were trapped in soft mud. The skeletons

occur singly and there is no suggestion of general catastrophes, but rather of

individual non-violent deaths. With the number of carnivores present in the

fauna the presence of so few disarticulated pareiasaur skeletons is remarkable.

Only once have I encountered the remains of a really juvenile animal and this

was disarticulated. A number of half-grown skeletons are known, but most

are mature.

In the case of the Dinocephalia, I know of only one skeleton (that of

Struthocephalus whaitst) entombed in the usual pareiasaur condition. In both

the tapinocephalids and the titanosuchids (both herbivorous groups) the

skeletons after death apparently lay exposed on higher ground with the bones

becoming widely scattered and then finally embedded in near-by depressions,

filling up with argillaceous or arenaceous sediments, mostly as individual

disarticulated bones, usually of the same animal. But at a number of places a

thin bed has produced over a fairly small area a mass of bones of a number of

separate individuals, e.g. bones of Moschops at Spitskop and at Kruisvlei,

Struthiocephalus at Skoenmaker, Criocephalus at Moddergat and a number of

struthiocephalines at Die Krans. At Kruisvlei the large number of bones of

THE FAUNA OF THE TAPINOCEPHALUS ZONE 25

Moschops was associated with a skull and some limb-bones of an anteosaurid.

Of the anteosaurids, which were slinking carnivores, it is remarkable how few

of the postcranial bones have been preserved, in contrast to the many large

skulls.

It would appear that one has to postulate the presence of carrion eaters

in addition to flowing water to account for the scattering and subsequent loss

through over-exposure and decomposition of the postcranial parts. There was,

of course, no hyaena-like destruction of the bones.

Of the Therocephalia from the Tapinocephalus zone I know of only one

skeleton which lay practically complete on its ‘belly’. This is an Alopecognathus

from Lammerkraal. Otherwise the finds have been of isolated skulls or partly

or wholly disarticulated postcranial parts of separate individuals. But at a

number of places fairly thin lenticular beds, that had been formed in shallow

depressions, have produced disarticulated remains of a number of individuals

of the same or of different species. These have usually been associated with

parts of small dicynodont forms.

The gorgonopsians are rare in this zone and the finds have been mostly

of isolated skulls, but the type specimen of Hipposaurus boonstrai consisted of a

neatly curled up complete skeleton overlain by a second skull.

Of the Dicynodontia a find at Mechau’s Request consisted of a number

of complete and partial skeletons in an arenaceous layer of about one foot in

thickness. The indications here are that we have to postulate the occurrence of

a local catastrophe. Isolated skulls occur throughout the zone, but by far the

greater number of dicynodont specimens have been found in widely separate

patches where the remains, also mainly skulls, have been eroded out of thin

lenses, these lenses having been formed in shallow pans into which the dis-

articulated skeletal parts from higher ground had been transported by water.

Often therocephalian remains are mixed up with the dicynodont material.

Although patches of initially soft mud surfaces, now indicated by the

presence of ripple marks in the consolidated rock, are not uncommon, not a

single track of any of the various groups of reptiles composing the fauna has as

yet been encountered in the whole of this zone.

The geographical distribution of the collected specimens in the type Koup

area shows some peculiarities, which are however readily accounted for. The

collector soon thinks of the fossil-bearing part of the Koup as divided into two

distinct areas, viz. a south-eastern and a north-western, with the railway line

roughly forming the dividing line.

Because of the proximity of Cape Foldings, the beds of the overlying

Tapinocephalus zone along the southern part of the area lie tilted, sometimes at

considerable angles. The beds are thus seen in section, with the result that the

likelihood of finding specimens is much smaller than it is in the northern

part where the beds lie nearly horizontally.

The presence of monoclinal folds tends to bring the lower levels of the

zone to the surface as one proceeds northwards. This applies specially to the

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

western part but less to the eastern part because the monoclines peter out

eastwards. The north-western part of the Koup thus has more of the lower

subdivisions exposed and these at low angles.

One would thus expect more specimens to have been collected from the

lower division of the zone, but the collection in the South African Museum

gives a contrary picture, viz. 555 specimens from the lower division and 769

from the upper divisions. (These numbers refer only to specimens whose

stratigraphical position is known with certainty.)

The expected condition prevails as far as the Dinocephalia and Pareiasau-

ria, large animals whose remains are mostly found singly, are concerned, viz.

362 low and 89 high.

It is the Therocephalia and Dicynodontia that cause the contradiction.

The relation of the numbers of these two groups in the two horizons is, for the

Tapinocephalus zone

Pareiasauridae

Eunotosauridae

-Anteosauridae

Titanosuchidae

Tapinocephalidae

Styracocephalidae

Galeopsidae

Endothiodontidae

Dicynodontidae

Hipposauridae

Galesuchidae

Pristerognathidae

Lycosuchidae

Scaloposauridae

Millerettidae

Varanopsidae

Fig. 1. Diagram to illustrate the relative num-

bers of the described species of the various

reptilian families recorded from the Tapino-

cephalus zone.

THE FAUNA OF THE TAPINOCEPHALUS ZONE P57

former 99 : 119 and for the latter group 102 : 567. The reason for this is that

in these two groups the mostly small skulls and snouts more often occur in

patches with a great concentration of specimens in the higher divisions, where

they are eroded out of lenses 4—5 inches in thickness, apparently formed in

shallow pans into which the skeletal remains have been carried by flood waters.

Such patches of greater concentration are rare in the lower division, and

only a few are known, e.g. one on the farm Kleinkoedoeskop. If we plot the

number of described species in the various families that constitute the fauna

as they occur in the three subdivisions of the zone we get a graphical picture

of the taxonomic composition of the fauna (fig. 1).

In the Dicynodontidae there are more species present in the middle

division than in the lower division.

In the Eunotosauridae, Hipposauridae, Lycosuchidae and Scaloposauridae

the lower and middle divisions have about the same number of species.

The Millerettidae and Varanopsidae occur only in the lower division,

whereas the Styracocephalidae and Galeopsidae are not known from the lower

division.

All the other families (Pareiasauridae, Anteosauridae, ‘Titanosuchidae,

Tapinocephalidae, Endothiodontidae, Galesuchidae and Pristerognathidae)

are more abundantly represented in the lower division.

The upper division of the zone is very poorly stocked.

It is manifest that the reptile fauna is preponderantly therapsid. The

pelycosaurs are represented by one advanced family—the Varanopsidae.

There is one primitive Chelonian? family and the cotylosaurs are well repre-

sented by an advanced family—Pareiasauridae. The sauropsids are poorly

represented by a single specimen of Broomia perplexa.

If we plot (fig. 2) the number of specimens in the South African Museum

collection in the various infra-orders as derived from the three subdivisions

we can readily see that the herbivores are represented in the lower division

by nearly equal numbers of pareiasaurs, titanosuchians, tapinocephalians and

Dicynodontia. In the middle division the former three groups are again about

equally represented but in greatly reduced numbers, whereas the Dicynodontia

are now very abundant.

The carnivores are represented by three groups—the Anteosauria,

Gorgonopsia and Therocephalia, with the latter predominant, especially in

the middle division.

In figure 3 I have plotted, not the number of specimens, but the number

of described species in the three categories—herbivores, carnivores and

insectivores. This tends to show that using the number of specimens rather

than the number of species gives a truer picture especially as far as the middle

division of the zone is concerned. For the zone as a whole, the fauna consists

of 53°% herbivores if one takes the number of recorded species as basis, but

according to the number of specimens this figure is 80%, which is ecologically

a better proportion.

ANNALS OF THE SOUTH AFRICAN MUSEUM

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THE FAUNA OF THE TAPINOCEPHALUS ZONE 29

The present day mammalian fauna of South Africa, as recorded by

Ellerman, gives the percentage of herbivores as 46°% according to the number

of species and 61% if the calculation is based on the number of genera.

This mammalian fauna consists of 125 genera and 221 species and the

reptilian fauna of the Tapinocephalus zone has recorded 96 genera and 152

species.

THE NON-REPTILIAN ELEMENTS IN THE FAUNA AND THE FLORA

It is clear that what we know of the fauna of this zone gives a picture very

far from reality.

One cannot believe that the four genera of Mollusca (Carbonicola, Kidodia,

Palaeomutela and Palaeanodonta) were the sole representatives of the Invertebrata

living at this time, but one can only speculate as to what teeming numbers of

these soft-bodied animals were the contemporaries of the dominant reptiles.

Of the fish inhabiting the ponds, lakes and streams, only very few and poor

specimens of palaeoniscids are known.

Even the Amphibia are poorly represented by a few specimens of the

family Rhinesuchidae.

Of the plants, which must have constituted a fairly rich flora in order

to have been able to nourish the numerous herbivorous reptiles, some more

than a ton in weight, there is preserved some fossil wood of the genus Dadoxylon,

and here and there some leaves of Glossopteris, Schizoneura and. Phyllotheca.

PETROLEUM IN THE Rocks OF THE Tapinocephalus ZONE

There is no evidence that the organic remains of the lowly animals we

presume to have lived during these times on their demise provided the source

of the hydrocarbons necessary for the formation of petroleum. The pseudocoal

filling veins at a number of places in this zone is a residual product of petro-

leum, but this has been shown to have infiltered from beds underlying the

rocks of the Tapinocephalus zone. The nature of these veins of pseudocoal clearly

shows that they were unsuited as reservoirs for petroleum. It has, moreover,

been shown that the texture and the structure of the rocks of this zone contra-

indicate the possibility of the existence of other reservoirs for petroleum which

may have arisen in adjoining beds (Rossouw, 1957).

TAXONOMIC REVISION

Pareiasauridae

In 1929 Haughton & Boonstra attempted a classification of the then

known pareiasaurs. As basis the structure of the teeth was considered. To

avoid the pitfalls presented by postmortem deformations the proportions of the

skull table, which would be least affected, were used, together with other

features as a criterion for further diagnosis. On these bases eight genera were

recognized in the fauna of the Tapinocephalus zone, viz. Bradysaurus, Bradysuchus,

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. Rhinesuchidae represented by Rhine-

suchus. Skull in dorsal and ventral view.

Nochelesaurus, Dolichopareia, Koalemasaurus, Brachypareia, Platyoropha and. Embritho-

saurus. Subsequently I realized that Platyoropha was indistinguishable from

Bradysaurus, and that Bradysuchus, founded on a feature recorded by Broom,

which is pathological, is also a synonym of Bradysaurus.

Now with more material available and a reassessment of the effects of

deformation on both the cranial and the postcranial skeleton, as well as such

factors as sexual dimorphism, and age, and the recent stratigraphical subdivision

of the zone a further reappraisement is suggested.

The genus Embrithosaurus stands on the basis of teeth with nine cusps

arranged in three groups of three. So also does Bradysaurus, with less than nine

cusps consisting of a terminal group of three cusps, an anterior group varying

from 1-3 and a posterior group of 2~—3 cusps. All the other genera have a cusp

arrangement, with only slight variations, similar to that of Bradysaurus and

can be included in this genus. Moreover, the genus Brachypareia appears to

have been based on specimens which may very well be considered to be imma-

ture. Discounting the features liable to deformation the other noted differen-

ces can at most be specific. The genus Bradysaurus could then include the four

species baini, seeleyi, angusta and strubent.

At the time when our ‘Pareiasaurian studies’ were written the prevailing

view was that the strata of the Tapinocephalus zone within the Koup, north of

the Folded Belt, lay more or less horizontally. Consequently it was thought

that the higher the altitude of the sites the higher was the level within the

zone. On this basis it was found that a number of species lived unchanged

from near the base to near the top of the zone. This was challenged by Broom,

who maintained that it was unlikely that species could remain unchanged for

this length of time, viz. according to Broom, 3-5 million years for the 1,900 ft

thickness of the zone.

In 1952 Rossouw & De Villiers gave the thickness of the zone as 6—7,000

ft and were able to establish a threefold subdivision of the Tapinocephalus zone

and, moreover, found that monoclinal folding brought the strata upwards as

one proceeded from south to north, so that in the north where the surface

THE FAUNA OF THE TAPINOCEPHALUS ZONE 31

Fig. 5. Pareiasauridae represented by

Bradysaurus. Skull in lateral, dorsal and

ventral view.

altitude is the highest the lower subdivision becomes exposed on the surface.

The stratigraphical level of all the fossil sites had thus to be determined anew.

Of the 70 specimens of pareiasaurs listed by Rossouw the distribution is

as follows:

Upper subdivision 1s

Middle subdivision 8

Lower subdivision 61

32 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. Varanopsidae represented by

Anningia. Skull in lateral and dorsal view.

The specimens entered into the South African Museum registers are

distributed as follows:

Upper subdivision )

Middle subdivision ng]

Lower subdivision 128

There is some doubt as to whether the specimens listed above as from the

middle part of the zone are correctly so placed. The localities are mostly

defined as to the farm and some of these large farms have outcrops of both the

lower and middle subdivisions exposed.

Both the genera Bradysaurus and Embrithosaurus are thus fully developed

from near the base of the zone, with some stragglers possibly surviving into the

middle part of the zone.

We have thus at least five species living unchanged through about 3,000

ft of strata. The time lap, according to Broom’s figures would be:

3,000,000 — 5,000,000 3000

—_—_ x

years.

I 1900

As for myself I should not like to be implicated in this arithmetic exercise.

THE FAUNA OF THE TAPINOCEPHALUS ZONE 33

ANOMODONTIA

A suborder of therapsid reptiles. Without a coronoid process to the dentary.

Quadrate large, anteriorly situated and supported by a strong quadrate ramus

of the pterygoid. Carnivorous or herbivorous. It consists of the infra-orders

Dinocephalia and Dicynodontia.

DINOCEPHALIA

An infra-order of anomodonts in which the pterygo-quadrate complex is

greatly strengthened. Carnivorous or herbivorous. It contains in South Africa

the families Anteosauridae, Titanosuchidae, Tapinocephalidae and Styraco-

cephalidae.

Anteosauridae

A family of dinocephalians which are fairly primitive carnivores, with

long intermeshing incisors. A stage further advanced than the Russian

Brithopidae.

From the Tapinocephalus zone seven genera have hitherto been named,

viz. Eccasaurus, Anteosaurus, Titanognathus, Dinosuchus, Micranteosaurus, Paranteo-

saurus and Pseudanteosaurus.

Of these, Dinosuchus and Titanognathus have already been considered

synonyms of Anteosaurus (Boonstra, 1954).

Eccasaurus, with a holotype of which the cranial material consists of only

a few typical anteosaurid incisors, appears to be only determinable as to family.

The skull fragment forming the holotype of Pseudanteosaurus can best be

considered as an immature specimen of Anteosaurus.

Maucranteosaurus, the holotype of which contains a small snout, has been

considered a new genus only on account of its small size and I now propose

that it be regarded as a young specimen of Anteosaurus.

ANTEOSAURUS

A genus of anteosaurids in which the postfrontal forms a boss of variable

size overhanging the dorso-posterior border of the orbit.

We have 32 skulls of Anteosaurus, of which 16 are reasonably well preserved

and on them ten species have been named. To differentiate between the species

the following main characters have been used: the number, size and shape of

the teeth, skull size, shape and the nature of the pachyostosis.

On re-examination it has become clear that the crowns of the teeth are

seldom well preserved; basing the count for the dental formula on the pre-

served roots is unreliable as this is affected by age and tooth generation; size

of skull is a function of age and also possibly sex; skull-shape is greatly affected

by post-mortem deformation, and the variability in the pachyostosis, which

may be specific in some respects, can just as well be the result of normal or

morbid physiological processes. Specific diagnosis consisting of the enumera-

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. Anteosauridae represented by Anteosaurus. Skull in lateral,

dorsal and ventral view.

tion of differences of degree in features such as the above can hardly be con-

sidered as sufficient indication of the existence of discrete species.

I thus suggest that only the oldest trivial name be retained. A. magnificus

thus has the following synonyms: abeli, acutirostris, crassifrons, cruentus, laticeps,

levops, lotzi, minor, minusculus, parvus, priscus and vorstert. Motivation for the

synonymy can be given as follows: cruentus, levops, minor, minusculus and parvus,

because the holotypes are immature animals; lotzi and priscus because the

specimens are specifically indeterminable; abeli, acutirostris, crassifrons, laticeps,

major and vorsteri because the characters used appear to be either the results

of post-mortem deformation or, the degree of the pachyostosis and this as

far as the pachyostosis is concerned, would fall within the range possible for

individual variation and the age or sex.

THE FAUNA OF THE TAPINOCEPHALUS ZONE 35

PARANTEOSAURUS

A genus of anteosaurids in which the postfrontal is not developed to form

a boss.

Titanosuchidae

These are fairly primitive herbivorous Dinocephalia in which the canine

has been retained and the very strong incisors have a piercing talon and a

crushing heel and the long series of postcanines have serrated spatulate crowns.

There is very little pachyostosis. The family contains two groups, viz. the long-

legged Titanosuchus and the short-legged species of Fonkerva.

When Owen described the first form in 1879 all the material from the Karoo

was so obviously new to science that even the poorest specimen warranted

description. Titanosuchus ferox was based on the roots of an incomplete set of

teeth.

In the race to name as many new species as possible Broom added a

ballast of names on just as poor material and even misidentified an obvious

carnivorous anteosaurid skull as co-specific with the herbivorous Titanosuchus

Serox.

Subsequently it has become apparent that even well-preserved batteries

of teeth exhibit so much variation, even in the left and right sides of the same

skull, that dental features are a very unreliable criterion for distinguishing

between titanosuchids.

In total 24 names appear in the literature.

A critical evaluation of these names seriatim

1. Titanosuchus ferox Owen, 1879

The type material is such that a diagnosis based on the available features

of the dentition is that given for the family which is based on the condition in

Jonkeria. The associated limb-bones, however, enable us to formulate a generic

diagnosis to distinguish this genus from Jonkeria.

2. Titanosuchus cloetei (Broom, 1903)

The characters determinable in this jaw-piece only allow one to determine

the family.

3. Scapanodon duplessist Broom, 1904

The cranial features are indeterminate. The humerus cannot be dis-

tinguished from that of Tztanosuchus and Scapanodon thus becomes a synonym.

The skull referred by Broom to the genus Scapanodon has been shown to fall

within the limits set for the genus Jonkeria.

4. Archaeosuchus cairncrosst Broom, 1905

The type specimen is so poor that no diagnosis can be made. The specimen

being indeterminable, the name is a nomen dubium.

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 8. Titanosuchidae represented by Jonkeria. Skull

in lateral, dorsal and ventral view.

5. Lamiasaurus newtoni Watson, 1914

This jaw fragment cannot be identified even as to family—it may be

either titanosuchid or anteosaurid.

6. Titanosuchus dubius (Haughton, 1915)

The teeth roots preserved only allow one to identify the specimen as being

of the family Titanosuchidae.

7. fonkeria truculenta Van Hoepen, 1916

Besides the good holotype, which consists of a good skull and much of

the postcranial skeleton, there is another good skull with lower jaw known.

In the South African Museum collection there are a number of specimens of

which I have described the dentition. All these specimens show that the teeth

are too variable for use in distinguishing between species of the genus. The

genus Jonkeria is distinguishable from Titanosuchus on features of the limb-bones.

THE FAUNA OF THE TAPINOCEPHALUS ZONE 37

8. Dinophoneus ingens Broom, 1923

The holotype, together with its synonym, Jonkeria pugnax, and three other

known skulls constitute a distinct species of Jonkeria as the genus is defined by

me in 1953. The genera Dinosphageus and Dinopolus can also be included in the

genus Jonkerta.

g. Dinartamus vanderbyli Broom, 1923

The fragments on which this name is based have features too indefinite

for a generic diagnosis and can only be considered as titanosuchian indeter-

minate as to genus.

10. Enobius strubent Broom, 1923

These pieces of jaw can only be identified as titanosuchian.

11. Jonkeria vanderbylt Broom, 1929

The holotype is a good skull of Jonkeria easily distinguishable from the

other species of the genus.

12. Dinosphageus haughtoni Broom, 1929

This fairly good skull, with some limb-bones, shows no features that would

exclude it from the genus Jonkeria, but can specifically be distinguished from

the other species of the genus.

13. Jonkeria crassus Broom, 1929

In the holotype consisting of dentaries and postcranial bones, the humerus

cannot be distinguished from that of Fonkerta haughtoni, and as there are no

other distinctive features it should be considered a synonym of 7. haughtonzt.

14. Phoneosuchus angusticeps Broom, 1929

This good mandible has all the characters of the genus Jonkeria and I can

find nothing to exclude it from the species truculenta.

15. Jonkeria pugnax Broom, 1929

I have already in 1935 shown this to be a synonym of 7. ingens.

16. Dinocynodon dubius (Broom, 1929)

The oval outline of the cross section of the canine can hardly be a character

sO important as to justify a new generic name. It can only be identified as

titanosuchian.

17. Scullya gigas Broom, 1929

This poorly preserved snout shows no definite titanosuchian characters.

The possible presence of teeth on the palatine may be an anteosaurid character.

The specimen must be considered indeterminable.

18. Dinopolus atrox Broom, 1936

The features exhibited in this snout are those known as characters of the

genus Jonkeria and cannot be used for a specific diagnosis. Dinopolus atrox is

thus an indeterminable species of the genus Jonkeria.

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

19. Scapanodon septemfontis Boonstra, 1955

Only postcranial bones are known. As the imperfect humerus cannot be

distinguished from that of Scapanodon duplessisi, septemfontis represents a synonym

of the former species and thus of the genus Titanosuchus.

20. Parascapanodon avifontis Boonstra, 1955

The humerus and femur are so similar to that of Titanosuchus and

Scapanodon that this form should be included in the genus Tztanosuchus.

A number of specimens, described by me under the name Parascapanodon,

together give a full picture of the dentition, but nothing determined in these

specimens is incompatible with the inadequately preserved dentition of the

holotype of Titanosuchus ferox. I thus suggest that the characters—cranial,

dental and postcranial—described under the names Titanosuchus, Scapanodon

and Parascapanodon be considered as diagnostic of the form Titanosuchus ferox.

21. JFonkeria koupensis Boonstra, 1955

The holotype is a good pelvis readily distinguishable from that of any

other known species of Yonkeria.

22. Fonkeria parva Boonstra, 1955

A small humerus is quite distinct from that of the other known species of

Jonkeria.

23. Jonkeria rossouwi Boonstra, 1955

The holotype consists of postcranial bones readily distinguishable from

those of the other species of the genus. Moreover, two other specimens are

known that show the same distinctive features.

24. Fonkerta boonstrai Janensch, 1959

Janensch has given a convincing diagnosis of the specific features of the

holotype skull. He also stresses the herbivorous nature of the dentition.

The family Titanosuchidae is thus composed of two genera and nine

species. The genera Titanosuchus and Jonkeria cannot in the present state of

our knowledge be distinguished from one another on either cranial or dental

characters. But in Titanosuchus the limb-bones are long, whereas in all the

species of Jonkerta they are short and squat.

In Jonkeria some of the species, where good cranial material is known, can

be distinguished on differences in cranial structure and the others only on

postcranial features.

Tapinocephalidae

This is an advanced family of herbivorous Dinocephalia without a ‘canine’ ;

all the teeth in maturity have a talon and a crushing heel and the upper and

lower teeth of the whole battery intermesh. The pachyostosis is moderately

to very greatly developed.

The family can be subdivided into a number of sub-families, viz.

Struthiocephalinae, Tapinocephalinae, Moschopinae and Riebeeckosaurinae.

THE FAUNA OF THE TAPINOCEPHALUS ZONE 39

Fig. 9. Tapinocephalidae represented by Tapinocephalus. Skull in lateral, dorsal and ventral view.

Struthiocephalinae

These are tapinocephalids with a long fairly strong snout and with mode-

rate pachyostosis. Six genera have been named. Two of these, Struthiocephalellus

and Moschosaurus, appear to be growth stages of Struthiocephalus and are thus

synonyms.

Struthiocephalus is a genus with a naso-frontal boss in maturity. The seven

described species—parvus, longiceps, millert, whaitsi, rheederi, akraalensis and

kitchingt— have been shown by me to represent a growth series and six of these

trivial names are thus synonyms of the first described species, viz. whaitst.

Struthiocephaloides is a genus without a naso-frontal boss in maturity. The

species duplessisi is wide across the tabulars whereas cavifrons is narrow across

the tabulars.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 10. Struthiocephalinae represented by

A. Struthiocephalus

oides

ocephalus Skulls in lateral view.

ops

id hosaurus

F. Struthiocephalus

THE FAUNA OF THE TAPINOCEPHALUS ZONE 41

Struthionops, with one species, intermedius, is a genus without a naso-frontal

boss, with moderate pachyostosis and a snout only moderately long.

Taurocephalus, with one species, lerouxt, has a long and strong snout and

with a long series of teeth — twenty in the upper jaw.

Fig. 11. Moschopinae represented by:

. Criocephalus

Moschops ]

. Delphinognathus P Skulls in lateral view.

Avenantia J

Moschops

. Criocephalus _)

: es (ae in dorsal view.

Delphinognathus

Skull in ventral view.

STamsgan>

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

Moschopinae

Medium-sized tapinocephalids with a short snout running up to the frons

in an even curve. The cranial roof is moderately to very greatly thickened by

pachyostosis.

Delphinognathus, a monotypic genus with species conocephalus, is known

from a single skull which has a conical boss on the parietal surrounding the

pineal foramen. There is a notch on the ventral edge between the jugal and

quadratojugal. The pachyostosis is moderate. It may possibly be a young skull

of a Moschops species.

Avenantia kruisvleiensis is only moderately pachyostosed and the intertem-

poral region is narrow and the temporal fossa large.

Moschops (with synonyms Moschoides, Agnosaurus, Moschognathus and

Pnigalion) is a genus strongly pachyostosed with a broad intertemporal region

and greatly reduced temporal fossae. There are two species—capensis and

koupensis —known from good material, and two species—whaitst and oweni—of

doubtful validity.

Criocephalus: in addition to the poor holotype specimen a further half-

dozen skulls are now known. In this genus the intertemporal region is very

broad, overhanging the greatly reduced temporal fossae. The cranial roof is

very greatly thickened by pachyostosis to make the parietal canal enormous.

The species —vanderbyli—is South African and gunyankaensis is Rhodesian.

Riebeeckosaurinae

This is a subfamily of the tapinocephalids in which the only species,

Riebeeckosaurus longirostris, known from two skulls, has a skull with a very long

and slender snout and a narrow intertemporal region which forms a narrow

sagittal crest. The pachyostosis is moderate.

Tapinocephalinae

The tapinocephalines are large and massive tapinocephalids in which

the snout is moderate to short and weak; they have either a prominent naso-

frontal boss or a greatly swollen frons. They are greatly to very greatly

pachyostosed.

Of the described genera Pelosuchus, without cranial parts known, is, on

the postcranial features, a synonym of Keratocephalus and the snout of Taurops

is so similar to that of Tapinocephalus that it should be included in that genus.

The greatly variable skulls known of the genera Mormosaurus and Keratocephalus

form a link between the subfamilies Struthiocephalinae and Tapinocephalinae.

Tapinocephalus: the one species —atherstonei—is known from a number of

skulls and postcranial bones. The skull is large with a massive swollen frons and

a short weak snout. The skull roof is greatly pachyostotic.

Phocosaurus megischion differs from Tapinocephalus in that the transition from

the frons to the snout is not abrupt.

THE FAUNA OF THE TAPINOCEPHALUS ZONE

Fig. 12. Tapinocephalinae represented by:

Phocosaurus

. Mormosaurus »Skulls in lateral view.

. Keratocephalus

. Phocosaurus

Mormosaurus }>Skulls in dorsal view.

Keratocephalus

rE Ob >

44. ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 13. Riebeeckosaurinae repre-

sented by skull of Riebeeckosaurus in

lateral and dorsal view.

Mormosaurus: in the holotype skull of the species, seeleyz, there is a naso-

frontal boss but in another skull there is a moderately strong frons. The snout is

strong and the pachyostosis moderate.

Keratocephalus: a number of skulls described and referred to the type,

moloch, show considerable variability in the pachyostotic development. There

is a prominent naso-frontal boss and the strength of the snout varies con-

siderably.

Styracocephalidae

In this family a number of imperfect skulls are known to constitute a

family of small advanced herbivorous dinocephalians with the retention of a

moderate canine. Characteristic are the prominent posteriorly directed tabular

bosses.

DICYNODONTIA

In the Tapinocephalus zone the Dicynodontia are represented by small

advanced anomodonts with edentulous beaks covered by horn and with the

maxillary and dentary teeth greatly reduced or absent. They differ from the

Dinocephalia in that, although the quadrate is strong, the quadrate ramus of

the pterygoid is not greatly strengthened and by the characteristically hori-

zontally turned squamosal bar.

Two families—Endothiodontidae and Dicynodontidae—occur in the

Tapinocephalus zone.

THE FAUNA OF THE TAPINOCEPHALUS ZONE 45

Fig. 14. Styracocephalidae represented by skull of Styracocephalus in lateral, dorsal

and ventral view.

Endothiodontidae

A family of dicynodonts with postcanine (molar) teeth in the maxilla.

Seven genera have been named of which two have been found to be synonyms,

viz. Opisthoctenodon of Pristerodon and Brachyprosopus of Brachyuraniscus.

Brachyuraniscus is a genus with three species with pineal boss, in which the

molars are in a row and the maxilla reaches the choana.

Broilius, with one species without pineal boss, in which the molars are irregu-

larly spaced and the maxilla reaches the choana.

Koupia, with one species; the molars are irregularly spaced and the relations

of the maxilla to the choanal border not determined.

Robertia broomiana: the molars are irregular, there is no pineal boss and the

maxilla does not enter the anterior choanal border.

Pristerodon brachyops: the molars are in a row, without a pineal boss and

the maxilla reaches the choana.

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 15. Endothiodontidae represented by composite figures of the

skull in lateral, dorsal and ventral view.

Dicynodontidae

A family which in the Tapinocephalus zone consists of two genera of small

dicynodonts without postcanine (molar) teeth.

Dicynodon, with eight described species from this zone, in which the

maxilla does not reach the choana.

Oudenodon, with one species, where the maxilla reaches the choanal border.

THERIODONTIA

A suborder of the Therapsida in which the South African forms have a

coronoid process to the dentary; the quadrate has not moved anteriorly, it is

small and the quadrate ramus of the pterygoid is weak. Carnivorous or insecti-

vorous. In the TYapinocephalus zone the sub-order is represented by the two

infra-orders Gorgonopsia and Therocephalia.

THE FAUNA OF THE TAPINOCEPHALUS ZONE 47

Fig. 16. Dicynodontidae represented by the skull of Dicynodon

in lateral, dorsal and ventral view.

Fig. 17. Galeopsid represented by the

skull of Galepus in lateral view.

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

GORGONOPSIA

This infra-order, as represented in the Tapinocephalus zone, consists of

fairly primitive theriodonts, in which the intertemporal region is still very wide

with the postorbital in its dorsal part lying horizontally, with the jaw adductors

originating from the under surface of the parietal and postorbital.

Of the nine described genera I regard two (Cyniscodon and Eriphostoma)

as without diagnosable generic characters and the two poor specimens as

indeterminate gorgonopsians. Of the remaining seven a further two (Aeluro-

sauroides and Broomisaurus) are represented only by snouts.

The described genera have been placed in five families by Watson &

Romer (1956). I recognize only two, viz. Hipposauridae and Galesuchidae.

Fig. 18. Galesuchid gorgonopsian, composite figures based on

the known genera. Skull in lateral, dorsal and ventral view.

THE FAUNA OF THE TAPINOCEPHALUS ZONE 49

Hipposauridae

The representatives of this family in the Tapinocephalus zone consist of

three fairly good skulls and three partial skulls forming a single genus Hippo-

saurus with two species — boonstrai and major. These are primitive gorgonopsians

only little advanced beyond the Russian Phthinosuchidae, which are the most

primitive gorgonopsians known. The hipposaurids are characterized by a

very broad intertemporal region, a long tooth row, a deep suspensorium and

by the fact that the snout is curved down strongly in relation to the skull roof.

Fig. 19. Hipposaurid gorgonopsian represented by the

skull of Hipposaurus in lateral and dorsal view.

50 ANNALS OF THE SOUTH AFRICAN MUSEUM

Galesuchidae

Primitive gorgonopsians with a broad intertemporal region, a reduced

postcanine series, a shallow suspensorium with the quadrate extending to

just below the level of the alveolar border and the snout is not bent down.

Pachyrhinos has a fairly large skull, with a dental formula I5, C1, PC4.

The other genera are fairly small and appear fairly closely related.

Eoarctops has for the upper jaw a dental formula 14, C1, PC3.

Galesuchus has for the upper jaw a dental formula I?, C1, PQ4.

Scylacognathus has for the upper jaw a dental formula I5, C1, PC5.

THEROCEPHALIA

This infra-order, as represented in the Tapinocephalus zone, consists of

small to large primitive theriodonts in which the intertemporal region is

narrowed with usually a sagittal crest and the posterior process of the post-

orbital lies vertically so that the jaw adductors arise from the lateral face of

the postorbital and parietal.

A large number of genera (24) have been described and these can be

arranged in four families— Pristerognathidae, Lycosuchidae, Alopecodontidae

and Scaloposauridae.

Pristerognathidae

Primitive Therocephalia with five to six incisors and a single canine and

two to nine variable postcanines. The epipterygoid is narrow. Two subfamilies

have been established —Scymnosaurinae and Pristerognathinae.

Scymnosaurinae

Fairly large to large pristerognathids with five incisors and two to six

postcanines. Five genera with seven species have been assigned to this subfamily,

which appears to be very variable.

Scymnosaurus, with three species, is known from six large but incomplete

skulls. Postcanines fairly weak to medium varying in number —gQ, 3 or 4.

Glanosuchus, with one species, is known from two good skulls and two

snouts. Large, with five postcanines and a variable small sixth incisor.

Ptomalestes: this monotypic genus has five to six fairly weak postcanines.

The holotype is a well-preserved skull.

Pristerosaurus with a fairly large skull with weak canines and three small

postcanines.

Kinnosaurus is a medium-sized scymnosaurine with five widely spaced

weak incisors and two rather feeble postcanines.

Pristerognathinae

In 1895 Seeley described the first therocephalian under the name

Pristerognathus polyodon—the holotype specimen being a poor weathered snout.

Seeley had no difficulty in deciding that he had a fragment of an animal new

THE FAUNA OF THE TAPINOCEPHALUS ZONE 51

Fig. 20. Pristerognathid therocephalian represented by

the skull of Glanosuchus in lateral, dorsal and ventral view.

to science. Why? Because hitherto no animal was known that had six upper

and three lower incisor teeth and this was the only feature shown by his type.

Broom, in 1904, described an even poorer snout as Pristerognathus baini because

his specimen was somewhat smaller than Seeley’s.

We now know that six upper incisors occur in a number of therocephalian

species but this is not even a subordinal character, so that this feature by itself

has no diagnostic value.

Pristerognathus polyodon is thus a nomen dubium and so is Broom’s name—

P. baini. I have transferred some of the later specimens which have been referred

to the genus Pristerognathus to the genus Pristerognathoides of which a valid

generic diagnosis can be given.

Two other described genera — Hyorhynchus and Ictidopareia—are both only

identifiable as to family. My genus Maraisaurus appears to be based on a juvenile

and is best included in the genus Pristerognathoides, and Brink’s Karowallteria is

based on a snout which falls within the range of the genus. Alopognathus, with

52 ANNALS OF THE SOUTH AFRICAN MUSEUM

four species, contains moderate to fairly large pristerognathines with upper

jaw dental formula 16 C1 PC4-6. The squamosal has an everted lateral edge.

Cynariognathus with four species, is medium-sized with upper jaw dental formula

I5-6, C1, PC6-g; the posterior incisors are smaller than the anterior ones,

the canine is long and strong and the postcanines are fairly strong and form a

closely set series. Lycedops is monotypic and has only four postcanines and the

temporal fossa is short and broad.

Pristerognathoides, with six described species, has a dental formula 16, C1,

PC5-6; the postcanines small, weak, and well spaced and the lateral edge of

the squamosal is not everted.

Therioides is monotypic and known only from a single skull with some of

the postcranial skeleton. Dental formula 16, C1, PC 6; the postcanines are

small and slender and well spaced. The descending process of the squamosal is

very deep so that the lower jaw articulation is situated far ventrally.

Fig. 21. Lycosuchid therocephalian represented by the skull

of Trochosaurus in lateral, dorsal and ventral view.

THE FAUNA OF THE TAPINOCEPHALUS ZONE 53

Lycosuchidae

Early, fairly large Therocephalia with fairly broad flattened skulls with

two functional canines in the maxilla, advanced broadened epipterygoid and

low sagittal crest. Otherwise very similar to the pristerognathids. With four

monotypic genera.

Hyaenasuchus —16 C2 PC4

Lycosuchus —I5 C2 PCr

Trochorhinus —15 C2 PQ3

Trochosaurus —15 C2 PC4

Alopecodontidae

Fairly small early Therocephalia of which the skulls are very inadequately

known. The characteristic feature of the family is the presence of two small

canines in the maxilla anterior to the single large canine. The family is com-

posed of three monotypic genera based on five incomplete to very poor skulls.

Alopecideops gracilis with a very slender snout; dental formula 16, C3, PC 7.

Alopecodon with dental formula I7 C3, PC 7. The species rugosus and minor

cannot be specifically distinguished from the holotype species priscus.

Pardosuchus whaitsi with dental formula 16, C3 PCs.

Scaloposauridae

From the Tapinocephalus zone a number of small forms are very inade-

quately known. They show the following scaloposaurid characters: small

skulls with a long snout and slender lower jaw; the intertemporal region is

fairly broad without a parietal crest; one to three canines and nine to ten post-

canines. But the postorbital bar is complete and the dentary has a coronoid

process.

Fig. 22. Scaloposaurid therocephalian repre-

sented by the skull of Blattoidealestes in lateral

view.

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

The two forms under the names Simorhinella baini and Ictidopareia brevirostris

should be considered nomina dubia, the former because it is based on an immature

animal and the latter because the type is lost and it has never been adequately

described. In Blattoidealestes gracilis some of the postcanines are triconodont.

The paratype of Icticephalus polycynodon is from this zone and Watson has

shown that there exists a close affinity between this genus and Scaloposaurus.

SysTEMATIC LisT OF THE TETRAPODS

AMPHIBIA

LABYRINTHODONTIA

RHACHITOMI

Rhinesuchidae

Rhinesuchus avenanti

Rhinesuchus whaitsi

Rhinesuchoides tenuiceps.

REPTILIA

ANAPSIDA

COTYLOSAURIA

DIADECTOMORPHA

Pareiasauridae

Bradysaurinae

Bradysaurus angusta

Bradysaurus baini

Bradysaurus seeleyt

Bradysaurus strubent

Embrithosaurus schwar zi

CHELONIA?

EUNOTOSAURIA

Eunotosauridae

Eunotosaurus africanus

LEPIDOSAURIA

EOSUCHIA

Millerettiformes

Millerettidae

Broomia perplexa

SYNAPSIDA

PELYCOSAURIA

SPHENACODONTIA

Varanopsidae

Anningia megalops

Elliotsmithia longiceps

THE FAUNA OF THE TAPINOCEPHALUS ZONE

THERAPSIDA

ANOMODONTIA

DINOCEPHALIA

Anteosauridae

Anteosaurus magnificus

Paranteosaurus primus

Titanosuchidae

Jonkeria boonstrat

Jonkeria haughtoni

Jonkeria ingens

Jonkeria koupensis

Jonkeria parva

Jonkeria rossouwt

Jonkeria truculenta

Jonkeria vanderbyli

Titanosuchus ferox

Tapinocephalidae

Struthiocephalinae

Struthiocephalus whaitsi

Struthiocephaloides cavifrons

Struthiocephaloides duplessisi

Struthionops intermedius

Taurocephalus lerouxt

Moschopinae

Avenantia kruisvleiensis

Criocephalus vanderbyli

Delphinognathus conocephalus

Moschops capensis

Moschops koupensis

Moschops owent

Moschops whaitsi

Riebeeckosaurinae

Riebeeckosaurus longirostris

Tapinocephalinae

Keratocephalus moloch

Mormosaurus seeleyt

Phocosaurus megischion

Tapinocephalus atherstoner

Styracocephalidae

Styracocephalus platyrhynchus

ANNALS OF THE SOUTH AFRICAN MUSEUM

DICYNODONTIA

Endothiodontidae

Brachyuraniscus broomi

Brachyuraniscus merwevillensis

Brachyuraniscus reuningi

Broilius antjiesfonteinensis

Koupia koupensis

Pristerodon brachyops

Robertia broomiana

Dicynodontidae

Dicynodon antjiesfonteinensis

Dicynodon gamkaensis

Dicynodon haughtonianus

Dicynodon jouberti

Dicynodon megalorhinus

Dicynodon pseudojouberti

Dicynodon schroedert

Dicynodon vanderhorsti

Oudenodon huener

Dromasauridae

Galeops whaitst

THERIODONTIA

GORGONOPSIA

Galesuchidae

Aelurosauroides watsoni

Broomiasaurus planiceps

Eoarctops vanderbyli

Galesuchus gracilis

Pachyrhinos kaiseri

Scylacognathus parvus

Hipposauridae

Hipposaurus boonstrai

Hipposaurus major

THEROCEPHALIA

Pristerognathidae

Scymnosaurinae

Glanosuchus macrops

Ptomalestes avidus

Pristerosaurus microdon

Scymnosaurus ferox

Scymnosaurus major

Scymnosaurus watsont

Kinnosaurus paucidens

THE FAUNA OF THE TAPINOCEPHALUS ZONE 57

Pristerognathinae

Alopecognathus angusticeps

Alopecognathus angustioriceps

Alopecognathus megalops

Alopecognathus skinnert

Cynariognathus gallowayi

Cynariognathus paucioridens

Cynariognathus platyrhinus

Cynariognathus seeleyi

Lycedops scholt zt

Pristerognathoides minor

Pristerognathoides parvus

Pristerognathoides peyert

Pristerognathoides roggeveldensis

Pristerognathoides vanderbyli

Pristerognathoides vanwykt

Theriodes cyniscus

Lycosuchidae

Hyaenasuchus whaitsi

Lycosuchus vanderrieti

Trochorhinus vanhoepent

Trochosaurus major

Alopecodontidae

Alopecideops gracilis

Alopecodon priscus

Pardosuchus whaitst

Scaloposauridae

Blattoidealestes gracilis

Icticephalus polycynodon.

Tapinocephalus ZONE FAUNAS OUTSIDE SOUTH AFRICA

Outside the main Karoo Basin in South Africa this fauna is known from

only one other area in Africa, viz. in southern Rhodesia, where a Criocephalus

and a therocephalian, probably pristerognathid, have been found.

From the rest of Gondwanaland no representatives of this fauna have as

yet been recorded.

The only country which has as yet yielded a comparable fauna, which

may be of the same age, is Russia. Here, the following families typical of the

Tapinocephalus zone, viz. Anteosauridae, Tapinocephalidae and Pristerognathi-

dae, have been recorded. No single genus occurs in both areas, but the Russian

Doliosauriscus is very close to our Anteosaurus and the Russian Ulemosaurus is very

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

similar to our Moschops. For the rest there are considerable differences between

the two faunas, the most important being the presence of more primitive forms

in Russia, which incline one to the view that the Russian fauna of Zone I and

possibly of Zone II are somewhat older.

CoMPARATIVE TABLE OF THE FAUNAS OF THE Tapinocephalus ZONE OF SOUTH

AFRICA AND ZONES I AND II OF THE RUSSIAN SUCCESSION

GENERA

T apinocephalus

zone Zone I Zone II

South Africa Russia Russia

Subclass:

Anapsida

Order:

Cotylosauria

Suborder:

Captorhinomorpha

Family:

Captorhinidae — — Hecatogomphius

Suborder:

Didectamorpha

Infra-order:

Pareiasuria

Family:

Rhipaeosauridae — — Rhipaeosaurus

— — Leptoropha

Family:

Pareiasauridae Bradysaurus — =

Embrithosaurus _— —

Infra-order:

Procolophonia

Family:

Nyctiphruretidae — — Nyctiphruretus

= = Nycteroleter

— — Nyctiboetus

Order:

Chelonia ?

Suborder:

Eunotosauria

Family:

Eunotosauridae Eunotosaurus — —

Subclass:

Lepidosauria

Order:

Eosuchia

Suborder:

Millerettiformes

THE FAUNA OF THE TAPINOCEPHALUS ZONE

Family:

Millerettidae

Family:

Mesenosauridae

Subclass:

Synapsida

Order:

Pelycosauria

Suborder:

Sphenacondontia

Family:

Varanopsidae

Suborder:

Edaphosauria

Family:

Phreatosauridae

Family:

Caseidae

Order:

Therapsida

Suborder:

Eotitanosuchia

Family:

Eotitanosuchidae

Suborder:

Anomodontia

Infra-order:

Dinocephalia

Family:

Brithopidae

Family:

Anteosauridae

T apinocephalus

Zone

South Africa

Broomia

Anningia

Elliotsmithia

Anterosaurus

Paranteosaurus

GENERA

Zone I

Russia

Phreatophasma

Phreatosuchus

Eotitanosuchus

Biarmosaurus

Biarmosuchus

Archaeosyodon

Chthomalopus

Zone II

Russia

Mesenosaurus

Phreatosaurus

Ennotosaurus

Phthinosaurus

Phthinosuchus

Brithopus

Notosyodon

Syodon

Titanophoneus

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family:

Titanosuchidae

Family:

Tapinocephalidae

Subfamily:

Struthiocephalinae

Subfamily:

Moschopinae

Subfamily:

Riebeeckesaurinae

Tapinocephalinae

Family:

Styracocephalidae

Family:

Estemmenosuchidae

Infraorder:

Dicynodontia

Family:

Otsheriidae

Family:

Venyukovidae

Family:

Endothiodontidae

T apinocephalus

zone

South Africa

Jonkeria

Titanosuchus

Struthiocephalus

Struthiocepha-

loides

Struthionops

Taurocephalus

Avenentia

Criocephalus

Delphinognathus

Moschops

Riebeeckosaurus

Keratocephalus

Mormosaurus

Phocosaurus

T apinocephalus

Styracocephalus

Brachyuraniscus

Broilius

Koupia

Robertia

GENERA

Zone I

Russia

Anoplosuchus

Estemmenosuchus

Otsheria

Zone II

Russia

Admetophoneus

Deuterosaurus

Doliosauriscus

Ulemosaurus

Mnemetosaurus

Molybdopygus

Venyukovia

THE FAUNA OF THE TAPINOCEPHALUS ZONE

Family:

Dicynodontidae

Family:

Dromasauridae

Suborder:

Theriodontia

Infra-order:

Gorgonopsia

Family:

Galesuchidae

Family:

Hipposauridae

Infra-order:

Therocephalia

Family:

Pristerognathidae

Family:

Lycosuchidae

Family:

Alopecodontidae

Family:

Scaloposauridae

T apinocephalus

ZONE

Dicynodon

Oudendon

Galeops

Aelurosauroides

Broomisaurus

Eoarctops

Pachyrhinos

Scylacognathus

Hipposaurus

Glanosuchus

Ptomalestes

Pristerosaurus

Scymnosaurus

Amnosaurus

Alopecognathus

Cynariognathus

Pristerognathoides

Theriodes

Hyaenosuchus

Lycosuchus

Trochorhinus

Trochosaurus

Alopecideops

Alopecodon

Pardosuchus

Blattoidealestes

Icticephalus

South Africa

GENERA

Zone I

Russia

Zone II

Russia

Unnamed

Porosteognathus

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

The (3) suborders Anapsida, Lepidosauria and Synapsida are common

to the three zones of South Africa and Russia.

The (4) orders Cotylosauria, Eosuchia, Pelycosauria and Therapsida are

common to the three zones but the order Chelonia? is unrepresented in Russia.

The (4) sub-orders Diadectamorpha, Millerettiformes, Anomodontia and

Theriodontia are common to the three zones, but the (2) sub-orders Eunoto-

sauria and Sphenacodontia are unrepresented in Russia and the (3) sub-orders

Captorhinomorpha, Edaphosauria and Eotitanosuchia are absent from

South Africa.

The (5) infra-orders Pareiasauria, Dinocephalia, Dicynodontia, Gorgonop-

sia and Therocephalia are present in South Africa as well as in Russia. The (1)

infra-order Procolophonia was unrepresented in South Africa during this

period.

Only the (3) families Anteosauridae, Tapinocephalidae and Pristerogna-

thidae are common to the two countries. Nine families occur only in Russia

and 14 only in South Africa.

Russia and South Africa have no genera in common. Of the 81 genera

here recognized as firmly established South Africa has 55 and Russia 26. But

Ulemosaurus is very near to Moschops and Doliosauriscus is very near to Anteosaurus.

ORIGIN OF THE Tapinocephalus ZONE REPTILIAN FAUNA

It is abundantly clear that the fossil remains at present known from the

rocks of the Tapinocephalus zone represent but a small part of the fauna which

must have lived during these times. Of the invertebrates we know next to

nothing and of the fish and amphibians we know little. Do the reptiles we

know fully represent this group? How many others lived in the Karoo at

this age of whom no remains have as yet been found or were not preserved as

fossils ?

The oldest reptiles of Carboniferous and Lower Permian age comprise the

orders Cotylosauria, Mesosauria, Protorosauria and Pelycosauria. Of these, the

Mesosauria did not survive beyond the Carboniferous. The Protorosauria,

which elsewhere survived to the Jurassic, have no representatives in the

Tapinocephalus zone. Of the stem group of the reptiles— the Cotylosauria — only

the bradysaurines occur in the Tapinocephalus zone and here form a well-

developed population of large herbivores. Of the pelycosaurs a few ‘Last of

the Mohicans’ lived into Tapinocephalus zone times. One of the oldest eosuchians

is present in this zone.

It is the newly evolved Therapsida which form the overwhelming, domi-

nant group in the fauna of the Tapinocephalus zone. Whence came these

therapsids — parvenus and immediately dominant?

In the Karoo, rocks of the Ecca Series underlie those of the Tapinocephalus

zone conformably and one would expect these to contain the immediate fore-

bears of the reptiles of the Tapinocephalus zone. Hitherto the Ecca has not yielded

a single reptilian specimen.

THE FAUNA OF THE TAPINOCEPHALUS ZONE 63

Why?

It is possible that during Ecca times the Karoo Basin was inhabited by an

ancestral fauna but that for some reason no remains of these animals were

preserved as fossils.

What were the factors that precluded preservation? Could it be that the

nature of the sedimentation during Ecca times was such that fossilization was

made impossible? In the western part of the Karoo Basin there is nothing to

indicate any radical difference in the sedimentation processes which formed

the rocks of the Ecca beds and those forming the rocks of the Tapinocephalus

zone. In fact the dividing line between these beds stipulated by stratigraphers

is an arbitrary one and lies below the first band of purple mudstones encoun-

tered in the column. Above and below this arbitrary line the constituent mud-

stones and sandstones were deposited in a similar way, viz. by fresh water

dropping its load of similar fine silt or coarser sand, according to the rate of

flow, in depressions in a general low-lying flood plain. If the hard parts of the

reptiles were included and subsequently mineralized above the arbitrary line

they should, if present, also have been entombed and preserved below this line.

The only reasonable conclusion is thus that during Ecca times the Karoo

Basin was not inhabited by a reptilian fauna.

The presence of purple mudstones above the arbitrary line and their

absence below the line may indicate that, although the mechanical nature of

the sedimentation was similar in both cases, there was in fact some other

difference. The green-blue colour of the Ecca mudstones is due to the presence

of the lower oxides of iron and the purplish colour in the purple bands of the

Tapinocephalus and higher zones is due to the presence of the higher red oxides

of iron. An increase in temperature is known to be conducive to the further

oxidation of iron salts. It is thus reasonable to assume that periods of higher

temperature occurred in Tapinocephalus zone times and that this made reptilian

life possible and that reptiles were absent in the Karoo Basin during Ecca times

because the temperature was too low for reptile life.

In the north-eastern part of the Karoo Basin the middle portion of the

Ecca beds has yielded a well-developed flora which was also the source of the

extensive and massive seams of coal. Did this flora also live in a moist and cold

climate? In the rest of Gondwanaland, beds of Ecca age have also proved

barren of reptiles.

In the lower beds of Dwyka age, the presence of Mesosaurus in various

parts of Gondwanaland shows that at this early age the reptiles had developed

at least one group probably closely related to the therapsids and far from

primitive. It would thus appear that the stem-reptiles of the Therapsida lived

well down in Carboniferous times.

With no ancestors in Gondwanaland, one turns to the northern hemisphere

in the search for the forebears of the Tapinocephalus zone fauna of the Karoo.

The presence of a comparable fauna in Zones I and II of the Russian

succession makes this imperative. The presence of some forms in Russia more

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

primitive than those of the Karoo indicates that the Russian beds may lie

nearer to the home of the ancestors.

However, the Lower Permian of the northern hemisphere has as yet

yielded no therapsids so that we still do not know any of the first forms of this

order.

Between the therapsids and the morphologically more primitive pelyco-

saurs and cotylosaurs of the northern hemisphere we still have a morphological

gap.

The pelycosaurs and cotylosaurs of the Lower Permian of Europe and

America would have had the first therapsids as contemporaries, and could thus

not have been their direct ancestors.

Moreover, we know of no Lower Permian pelycosaur that is unspecialised

enough to have been an ancestor of any known therapsid.

The pelycosaurs known can be considered as an offshoot of the direct

therapsid line.

This pushes the origin of the therapsids another step back, viz. to the more

primitive cotylosaurs of the Carboniferous.

DESCENDANTS OF THE FAUNA OF THE Tapinocephalus ZONE

In Africa, the conformably overlying Endothiodon and Cistecephalus zones,

comprising the Upper Permian, have the following direct descendants:

The bradysaurines gave rise to the pareiasaurines.

The varanopsids peter out.

The Dicynodontia really start to flourish and so do the Gorgonopsia.

The Therocephalia develop new families and give rise to the first cynodonts.

Of the eosuchians the millerettids continue and two other families arise:

All these groups spread out to East or Central Africa.

The comparable Russian faunas of Zones I and II evolved on similar

lines but less robustly. Some pareiasaurs and dicynodonts spread to western

Europe and dicynodonts to Indo-China.

ACKNOWLEDGEMENT

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

Scientific and Industrial Research for a grant towards the publication of this

paper.

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Kaapstad: Nasionale Boekhandel.

Boonstra, L. D. 1948a. Miljoene jare gelede in die Karoo. Johannesburg: Voortrekkerpers.

Boonstra, L. D. 19485. On the anomodont reptiles from the Tapinocephalus-zone of the Karoo

system. In RoyAL Society oF SoutH Arrica. Robert Broom commemorative volume: 57-64.

Cape Town: Royal Society of South Africa. (Special publication.)

Boonstra, L. D. 1949. Reptiele van die verre verlede. In Afrikaanse kinderensiklopedie: 2111-2114.

Kaapstad: Nasionale Boekhandel.

Boonstra, L. D. 19514. Kurze Notiz tiber den Schadel der Dinocephalen-Gattung Keratocephalus

F. v. Huene. Neues Fb. Geol. Palaont. Mh. 11: 341-343.

BoonstrRA, L. D. 1951. The museum—why have it? Lantern 1: 288-289.

Boonstra, L. D. 1952a. Agnosaurus gen. nov.: ’n nuwe geslag van die deinocephaliérs. Tydskr.

Wet. Kuns 12: 242-245.

Boonstra, L. D. 19525. Further observations on the type-skull of Struthiocephalus whaitsi. Ann.

Mag. nat. Hist. (12) 5: 455-459.

BoonstraA, L. D. 1952c. Die gorgonopsiér-geslag Hipposaurus, en die familie Ictidorhinidae.

Tydskr. Wet. Kuns 12: 142-149.

Boonstra, L. D. 1952d. A new deinocephalian from the Karroo. Ann. Mag. nat. Hist. (12) 5:

988-989.

Boonstra, L. D. 1952e. ’n Nuwe tapinocephalide, Riebeeckosaurus longirostris gen. et sp. nov.

Tydskr. Wet. Kuns 12: 246-2409.

Boonstra, L. D. 1952f. ’n Nuwe soort van tapinocephalide deinocephaliér: Struthiocephalus

akraalensis sp. nov. S. Afr. F. Sci. 48: 247-248.

Boonstra, L. D. 1952g. ’n Nuwe titanosuchiérsoort (Anteosaurus abeli). Tydskr. Wet. Kuns 12:

150-151.

Boonstra, L. D. 1952h. On a new tapinocephalid deinocephalian. Ann. Mag. nat. Hist. (12) 5:

50Q-51I.

Boonstra, L. D. 19527. Reconstructing life-sized models of prehistoric reptiles. SAMAB 5:

179-183.

Boonstra, L. D. 19527. Struthiocephaloides: ’n nuwe genus van mormosauride tapinocephaliérs.

Tydskr. Wet. Kuns 12: 237-241.

Boonstra, L. D. 19534. The cranial morphology and taxonomy of the tapinocephalid genus

Struthiocephalus. Ann. S. Afr. Mus. 42: 32-53.

Boonstra, L. D. 1953b. The gorgonopsians, Aelurognathus microdon and Hipposaurus boonstrai,

reconstructed. Ann. S. Afr. Mus. 42: 29-31.

Boonstra, L. D. 1953c. The lower jaw articulatory region in some pristerognathid therocepha-

lians. Ann. S. Afr. Mus. 42: 54-63.

Boonstra, L. D. 1953¢. A suggested clarification of the taxonomic status of the South African

titanosuchians. Ann. S. Afr. Mus. 42: 19-28.

Boonstra, L. D. 1954a. The cranial structure of the titanosuchian: Anteosaurus. Ann. S. Afr.

Mus. 42: 108-148.

Boonstra, L. D. 19545. Paranteosaurus gen. nov.: a titanosuchian reptile. Ann. S. Afr. Mus. 42:

157-159.

Boonstra, L. D. 1954c. The pristerognathid therocephalians from the Tapinocephalus zone in

the South African Museum. Ann. S. Afr. Mus. 42: 65-107.

Boonstra, L. D. 1954d. A scaloposaurid from the Tapinocephalus zone. Ann. Mag. nat. Hist.

(12) 7: 153-157.

Boonstra, L. D. 1954e. The smallest titanosuchid yet recovered from the Karroo. Ann. S. Afr.

Mus. 42: 149-156.

THE FAUNA OF THE TAPINOCEPHALUS ZONE 67

Boonstra, L. D. 19554. The girdles and limbs of the South African Deinocephalia. Ann. S.

Afr. Mus. 42: 185-326.

Boonstra, L. D. 19550. Struthiocephalellus: a new deinocephalian. Ann. S. Afr. Mus. 42: 180-184.

Boonstra, L. D. 1956a. Pareiasauriér-studies. Deel XII. ’n Lewensrekonstruksie van Bradysaurus

seeleyi. Ann. S. Afr. Mus. 43: 171-172.

Boonstra, L. D. 19566. The skull of Tapinocephalus and its near relatives. Ann. S. Afr. Mus. 43:

137-169.

Boonstra, L. D. 1957. The moschopid skulls in the South African Museum. Ann. S. Afr. Mus.

44: 15-38.

Boonstra, L. D. 1962. The dentition of the titanosuchian dinocephalians. Ann. S. Afr. Mus.

46: 57-112.

Boonstra, L. D. 19632. Diversity within the South African Dinocephalia. S. Afr. 7. Sci. 59:

196-206.

Boonstra, L. D. 19635. Early dichotomies in the therapsids. S. Afr. F. Sci. 59: 176-195.

Boonstra, L. D. 1965a. The girdles and limbs of the Gorgonopsia of the Tapinocephalus zone.

Ann. S. Afr. Mus. 48: 237-249.

Boonstra, L. D. 19655. The Russian dinocephalian Deuterosaurus. Ann. S. Afr. Mus. 48: 233-236.

Boonstra, L. D. 1965¢. The skull of Struthiocephalus kitchingi. Ann. S. Afr. Mus. 48: 251-265.

Boonstra, L. D. 1966a. The dinocephalian manus and pes. Ann. S. Afr. Mus. 50: 13-26.

Boonstra, L. D. 1966. The girdles and limbs of the Dicynodontia of the Tapinocephalus zone.

Ann. S. Afr. Mus. 50: 1-11.

Boonstra, L. D. 1967a. An early stage in the evolution of the mammalian quadrupedal walking

gait. Ann. S. Afr. Mus. 50: 27-42.

Boonstra, L. D. 19675. Langs verskillende weé (Pareiasauria en Dicynodontia). S. Afr. F. Sci.

63: 201-206.

Boonstra, L. D. 19684. The braincase, basicranial axis and median septum in the Dinocephalia.

Ann. S. Afr. Mus. 50: 195-273.

BoonstrA, L. D. 19685. The terrestrial reptile fauna of Tapinocephalus zone age and Gondwana-

land. S. Afr. F. Sci. 64: 199-204.

Brink, A. S. 1958. Struthiocephalus kitchingi sp. nov. Palaeont. afr. 5: 39-56.

Brink, A. S. & Kitcuinc, J. 1951. Some theriodonts in the collection of the Bernard Price

Institute. Ann. Mag. nat. Hist. (12) 4: 1218-1236.

Brorut, F. & ScHRODER, J. 1934. Beobachtungen an Wirbeltieren der Karrooformation. IV.

Ein neuer Gorgonopside aus den unteren Beaufort-Schichten. Sber. bayer. Akad. Wiss. 1934:

209-223.

Brom, F. & Scur6épER, J. 193542. Beobachtungen an Wirbeltieren der Karrooformation.

VIII. Ein Dinocephalen-Rest aus den unteren Beaufort-Schichten. Sher. bayer. Akad. Wiss.

1935: 93-114.

Brom, F. & Scur6pER, J. 19355. Beobachtungen an Wirbeltieren der Karrooformation. XII.

Uber einige primitive Anomodontier-Schadel aus den unteren Beaufort-Schichten. Sber.

bayer. Akad. Wiss. 1935: 223-278.

Broiu, F. & Scurdéper, J. 1936a. Beobachtungen an Wirbeltieren der Karrooformation. XV.

Ein Therocephalier aus den unteren Beaufort-Schichten. Sher. bayer. Akad. Wiss. 1936: 1-20.

Brom, F. & Scuréper, J. 1936). Beobachtungen an Wirbeltieren der Karrooformation.

XXIII. Ein weiterer Therocephalier aus den unteren Beaufort-Schichten. Sher. bayer.

Akad. Wiss. 1936: 283-310.

Brot, F. & Scuroper, J. 1937. Beobachtungen an Wirbeltieren der Karrooformation. XXVIII.

Uber einige neue Anomodontier aus der Tapinocephalus-Zone. Sher. bayer. Akad. Wiss.

1937: 118-168.

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

Broom, R. 19034. On an almost perfect skeleton of Pareiasaurus serridens Owen. Ann. S. Afr. Mus.

4: 123-138.

Broom, R. 19036. On an almost perfect skull of a new primitive theriodont (Lycosuchus vanderrieti).

Trans. S. Afr. phil. Soc. 14: 197-205.

Broom, R. 1903c. On evidence of a new species of Titanosuchus (T. cloetei). Ann. S. Afr. Mus. 4:

142-143.

Broom, R. 1903d. On the classification of the theriodonts and their allies. Rep. S. Afr. Ass.

Advmt Sci. 1: 1-9.

Broom, R. 1903e. On the presence of a pair of distinct prevomers in Tztanosuchus. Ann. S. Afr.

Mus. 4: 144-146.

Broom, R. 1903 f. On some new primitive theriodonts in the South African Museum. Ann. S.

Afr. Mus. 4: 147-158.

Broom, R. 1904a. Notice of a new fossil reptile (Scapanodon Duplessisi) from the lower Karoo

beds of Prince Albert, Cape Colony. Rec. Albany Mus. 1: 182-183.

Broom, R. 1904b. On a new species of Oudendon (O. megalorhinus) from the Gough, S. Africa.

Rec. Albany Mus. 1: 180-181.

Broom, R. 1904¢c. On two new therocephalian reptiles (Glanosuchus macrops and Pristerognathus

baini). Trans. S. Afr. phil. Soc. 15: 85-88.

Broom, R. 19054. Notice of some new fossil reptiles from the Karroo beds of South Africa.

Rec. Albany Mus. 1: 331-337.

Broom, R. 19056. On the classification of the Karroo system. Rep. S. Afr. Ass. Adumt Sci. 2: 38-44.

Broom, R. 1907. On the geological horizons of the vertebrate genera of the Karroo formation.

Rec. Albany Mus. 2: 156-163.

Broom, R. 1908a. On a new labyrinthodont Rhinesuchus Whaitsi from the Permian beds of

South Africa. Ann. S. Afr. Mus. 4: 373-376.

Broom, R. 1908). On some new therocephalian reptiles. Ann. S. Afr. Mus. 4: 361-367.

Broom, R. 1g08c. On the inter-relationships of the known therocephalian genera. Ann. S. Afr.

Mus. 4: 369-372.

Broom, R. 19094. An attempt to determine the horizons of the fossil vertebrates of the Karroo.

Ann. S. Afr. Mus. 7: 285-2809.

Broom, R. 1909). Notice of some new South African fossil amphibians and reptiles. Ann. S.

Afr. Mus. 7: 270-278.

Broom, R. 1goge. On the skull of Tapinocephalus. Geol. Mag. (n.s., 5) 6: 400-402.

Broom, R. 1g10a. A comparison of the Permian reptiles of North America with those of South

Africa. Bull. Am. Mus. nat. Hist. 28: 197-234.

Broom, R. 1910). Observations on some specimens of South African fossil reptiles preserved in

the British Museum. Trans. R. Soc. S. Afr. 2: 19-25.

Broom, R. 1911. On some new South African Permian reptiles. Proc. zool. Soc. Lond. 1911:

1073-1082.

Broom, R. 1912a. On a new species of Proppapus, and on the pose of the pareiasaurian limb.

Ann. S. Afr. Mus. 7: 323-331.

Broom, R. 1912. On some new fossil reptiles from the Permian and Triassic beds of South

Africa. Proc. zool. Soc. Lond. 1912: 859-876.

Broom, R. 1913a. On four new fossil reptiles from the Beaufort series, South Africa. Rec. Albany

Mus. 2: 397-401.

Broom, R. 19136. On the manus and pes of Pareiasaurus. Ann. S. Afr. Mus. 7: 353-357-

Broom, R. 1913c. A revision of the reptiles of the Karroo. Ann. S. Afr. Mus. 7: 361-366.

Broom, R. 19142. Croonian lecture: on the origin of mammals. Phil. Trans. R. Soc. (B) 206: 1-48.

THE FAUNA OF THE TAPINOCEPHALUS ZONE 69

Broom, R. 19145. A further comparison of the South African dinocephalians with American

pelycosaurs. Bull. Am. Mus. nat. Hist. 33: 135-141.

Broom, R. 1914c. Further observations on South African reptiles. Am. Mus. 7. 14: 139-145.

Broom, R. 19154. Catalogue of types and figured specimens of fossil vertebrates in the American

Museum of Natural History. II. Permian, Triassic and Jurassic reptiles of South Africa.

Bull. Am. Mus. nat. Hist. 25: 105-164.

Broom, R. 1915. On some new carnivorous therapsids in the collection of the British Museum.

Proc. zool. Soc. Lond. 1915: 163-173.

Broom, R. 1921. On the structure of the reptilian tarsus. Proc. zool. Soc. Lond. 1921: 143-155.

Broom, R. 1923. On the structure of the skull in the carnivorous dinocephalian reptiles. Proc.

zool. Soc. Lond. 1923: 661-684.

Broom, R. 1924. On some points in the structure of the pareiasaurian skull. Proc. zool. Soc. Lond.

1924: 499-508.

Broom, R. 1925. On some carnivorous therapsids. Rec. Albany Mus. 3: 309-326.

Broom, R. 1927. On a new type of mammal-like reptile from the South African Karroo beds

(Anningia megalops). Proc. zool. Soc. Lond. 1927: 227-232.

Broom, R. 1928. On Tapinocephalus and two other dinocephalians. Ann. S. Afr. Mus. 22: 427-438.

Broom, R. 19294. On some recent new light on the origin of mammals. Proc. Linn. Soc. N.uS.W.

54: 688-6094.

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

Trans. Mus. 13: 9-36.

Broom, R. 1930. On the structure of the mammal-like reptiles of the sub-order Gorgonopsia.

Phil. Trans. R. Soc. (B) 218: 345-371.

Broom, R. 1931. Notices of some new genera and species of Karroo fossil reptiles. Rec. Albany

Mus. 4: 161-166.

Broom, R. 1932. The mammal-like reptiles of South Africa and the origin of mammals. London:

Witherby.

Broom, R. 19354. A new genus and some new species of mammal-like reptiles. Ann. Transv.

Mus. 28: 1-12.

Broom, R. 1935). Notes on some species of pareiasaurian reptiles. Ann. Transv. Mus. 28: 37-51.

Broom, R. 1936a. On the structure of the skull in a new type of dinocephalian reptile. Proc.

zool. Soc. Lond. 1936: 733-742.

Broom, R. 19365. On the structure of the skull in the mammal-like reptiles of the suborder

Therocephalia. Phil Trans. R. Soc. (B) 226: 1-42.

Broom, R. 19374. A few more new fossil reptiles from the Karroo. Ann. Transv. Mus. 19: 141-146.

Broom, R. 19376. A further contribution to our knowledge of the fossil reptiles of the Karroo.

Proc. zool. Soc. Lond. 1937: 299-318.

Broom, R. 1940. Some new Karroo reptiles from the Graaff-Reinet district. Ann. Transv. Mus.

20: 71-87.

Broom, R. 1941. Some new Karroo reptiles, with notes on a few others. Ann. Transv. Mus. 20:

193-213.

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

South Africa. Trans. Kans. Acad. Sci. 40: 221-231.

Byrne, F. 1940. Notes on the evolution of the mammal-like reptiles. Trans. Kans. Acad. Sci. 43:

291-206.

Camp, C. L., Taytor, D. N. & WELLES, S. P. 1942. Bibliography of fossil vertebrates 1934-1938.

Spec. Pap. geol. Soc. Am. 42: i-v, 1-663.

Carp, B. 1952. An interesting discovery. Afr. wild Life 6: 137.

70 ANNALS OF THE SOUTH AFRICAN MUSEUM

Case, E. C. 1926. Environment of tetrapod life in the late Paleozoic of regions other than North

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Cys, J. M. 1967. Osteology of the pristerognathid Cynariognathus platyrhinus (Reptilia: Therio-

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Du Tort, A. L. 1948. The climatic setting of the vertebrate faunas of the Karroo system and

its significance. In Roya Society oF SoutH Arrica. Robert Broom commemorative volume:

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Fenton, C. L. & Fenton, M. A. 1958. The fossil book: a record of prehistoric life. New York:

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Grecory, W. K. 1920. Mounted skeleton of Moschops capensis Broom. Bull. geol. Soc. Am. 31: 223.

Grecory, W. K. 1926. The skeleton of Moschops capensis Broom, a dinocephalian reptile from

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

Grecory, W. K. 1946. Pareiasaurs versus placodonts as near ancestors to the turtles. Bull.

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Grecory, W. K. & Camp, C. L. 1918. Studies in comparative myology and osteology. No. III.

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HaucutTon, S. H. 1913. Ona skull of Tapinocephalus atherstoni, Owen. Ann. S. Afr. Mus. 12: 40-42.

Haucuton, 8. H. 19154. On a new dinocephalian from the Gouph. Ann. S. Afr. Mus. 12: 52-54.

Haucuton, S. H. 19155. On a new type of Dinocephalia (Moschosaurus longiceps). Ann. S. Afr.

Mus. 12: 78-81.

HaucutTon, 8S. H. 1915c. On some new gorgonopsians. Ann. S. Afr. Mus. 12: 82-90.

Haucuton, S. H. 1915d. On two new therocephalians from the Gouph. Ann. S. Afr. Mus. 12:

55-57:

Haucuron, S. H. 1917. Descriptive catalogue of the Anomodontia, with special reference to the

examples in the South African Museum. Ann. S. Afr. Mus. 12: 127-174.

Haucuron, S. H. 1918. Some new carnivorous Therapsida, with notes upon the brain-case in

certain species. Ann. S. Afr. Mus. 12: 175-216.

Haucuton, S. H. 191g. A review of the reptilian fauna of the Karroo system of South Africa.

Trans. geol. Soc. S. Afr. 22: 1-25.

Haucuton, S. H. 19244. On some gorgonopsian skulls in the collection of the South African

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HaucurTon, S. H. 19246. A bibliographic list of the pre-Stormberg Karroo Reptilia, with a

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Haucuton, S. H. 1925. Investigations in South African fossil reptiles and Amphibia. Part 13:

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Haucuton, S. H. 1953. Gondwanaland and the distribution of early reptiles. Trans. geol. Soc.

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THE FAUNA OF THE TAPINOCEPHALUS ZONE 71

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SEELEY, H. G. 1889. Researches on the structure, organization, and classification of the fossil

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215-296.

SEELEY, H. G. 1890. Some scientific results of a mission to South Africa. Trans. S. Afr. phil.

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SEELEY, H. G. 1892a. On Delphinognathus conocephalus (Seeley) from the middle Karroo beds,

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INSTRUCTIONS TO AUTHORS

Based on

CONFERENCE OF BIOLOGICAL EDITORS, COMMITTEE ON FORM AND STYLE. 1960.

Style manual for biological journals. Washington: American Institute of Biological Sciences.

MANUSCRIPT

To be typewritten, double spaced, with good margins, arranged in the following order:

(1) Heading, consisting of informative but brief title, name(s) of author(s), address(es) of

author(s), number of illustrations (plates, figures, enumerated maps and tables) in the article.

(2) Contents. (3) The main text, divided into principal divisions with major headings; sub-

headings to be used sparingly and enumeration of headings to be avoided. (4) Summary.

(5) Acknowledgements. (6) References, as below. (7) Key to lettering of figures. (8) Explana-

tion to plates.

ILLUSTRATIONS

To be reducible to 4? in. x 7 in. (74 in. including caption). A metric scale to appear with

all photographs.

REFERENCES

Harvard system (name and year) to be used: author’s name and year of publication given

in text; full references at the end of the article, arranged alphabetically by names, chronologi-

cally within each name, with suffixes a, b, etc. to the year for more than one paper by the

same author in that year.

For books give title in italics, edition, volume number, place of publication, publisher.

For journal articles give title of article, title of journal in italics (abbreviated according to

the World list of scientific periodicals. 4th ed. London: Butterworths, 1963), series in

parentheses, volume number, part number (only if independently paged) in parentheses,

pagination.

Examples (note capitalization and punctuation)

BuLtoucu, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FiscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. 7. Conch., Paris

88: 100-140.

Fiscuer, P.-H., Duvat, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines.

Archs Zool. exp. gén. 74: 627-634.

Koun, A. J. 19604. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee

region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19605. Spawning behaviour, egg masses and larval development in Conus from

the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In Schultze, L.

KLoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-

Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

ZOOLOGICAL NOMENCLATURE

To be governed by the rulings of the latest International code of zoological nomenclature issued

by the International Trust for Zoological Nomenclature (particularly articles 22 and 51).

The Harvard system of reference to be used in the synonymy lists, with the full references

incorporated in the list at the end of the article, and not given in contracted form in the

synonymy list.

Example

Scalaria coronata Lamarck, 1816: pl. 451, figs 5 a, 6; Liste: 11. Turton, 1932: 80.