« et , sheath WR iC irl Pet ‘44 pn wateeeey 4. mb) a4 7 at « Tass arent eB aL o" ent sen wey : dat reracet cab ro ‘ ry Je hee ae “ ”~ + ”™ F 4 thiibatt ~ Ww Wei M ook eda tae “ Pern) \ Wane 4) 9019) tr nS Lay ae ae Th eked eee Hi tee gietiats ts Hall de h ae Tetkuel ath te res ale We iemetiy Pont Cit Ay ur ished ey Po Peete de ded ‘eM PEAR if te i Higa ea re) a Hh and ¢ vee # ve iad tea ‘ ia myant aes \ 1 ny’ ‘i ROE wate Ta ak Nery erre we Wane head Vika ted Me Be 1 OA Eig Re ered’ Mahe Ae 4 way. ere wee Ap ey TW Ms tas Pda was joie < Fe yi * Bea Niele a ea oT IE ed eat ae cane def Pathol ee ay ail 44 eae Oekaki ‘ Lidl = ay ae Ua ae aia ry swe }4 ant sae ‘ vie Ee dee tite a erway i »! na vist ! prep ig 4a A qe at 4 we ath vd \ i iy Habit alae i aa a4 HN Nah Wied tsa Par on ee set a Das wate ‘bia Mdshided te ttl arabe ioe aes we” BS a Tegra th att yaa sbi jaa 1 tah WAY 4} aay aa alae ‘ia a Ad ee aah FP OWAP HE IR Kt He CUA E Pear nia 0 deel 4. aa * rie ak hard i ly a MORK, i Ale Lee WRI} ae ey q - iyo eh a \ fae aie aR AS) Yea tats td LALA SUES aod VR de lanes : 4 Wea dada’ Sete ee tiara eA all, yy ee Te de Yaad att aeae ge Jig a Le a Oo pods Ve Oat Fay sth Gade hh ses dd ‘ect a waht tapaag en a teed aA 44 4 j “ df ‘oa, 4 ‘ ‘ \ we ht ZV eh: < a ‘ ee PA day dei Deny pe EL te Ded) bine Lee i) wad Chk PF A) AN ak aot ay ri iy a wait ue iy 4 Tee \ hae faite} hale Lede ia a Ay Neal vat iD eu Ay, at yaad ROL TAY Side a at Pac aks Heres ke ” a EIN ie Scuba ie aia 4 rf ho hats aon ¥ ‘ Ni Ao cond 4) yn vi odd Sa aalys iP MATS Wee oe cbt ‘eth es lt - v7 so) toi . 4 ea ae i aah We aoe steele do ih tT ig id ‘EF op veh ee ee OF Hon Wi vy ale i ie ae ‘ fh mi a tae if ab Weal a = } ANE eK Fah af a‘ tn etd ae 0 heed a a MAN vw site 4 nt 19) ao bi Ay ONY Ne ge atid Oe i) nit Re ait vate mats) we on lle i i efi ae Cer bahe he as He te sf a) mad ah ces x moet \ a tite Mt i Ne i cae Ni saa: s ne! ee a AN ite Pree ny rhe srk ne Nine \ fin ene afjaae ! A ay Rie ni fee i ; ate at aa tea att Wels lian we p! iT Late. pia, ne i vs 0 oe i iat hs ts i aN as ay ae if ne ey ee ye Ns Ay x ae ae ihe ia ta os Atel at E nae ti ae i saat a aN he Nf ity cs ae ieee oe i ive dol Maks Ak sai ha b Gated Lalas in hoe ih nd eat hele A dace Pi lankat ent 4 Mae daha me Be if ae wine i Se tates if moe rn aoe : 3 oe bee WO Tt gig ahaa) ? wlaed he y Pd LW Othe vai a eat ” i a et NWaithe) weary in iM ee ut i eel Sethian: aed abate AY COA Hatha 4 oo Widen guurees 4 ily tren ' tae (oped ed gd 8 ae, “| IN 4 ‘ eb 4 pid ded a boda tn vite Vayeuaeg ty 4 aot ! Shy gid alates iW eg de / Le 9 aod Paaeat dell ae er i ad eave AW? OND beat Ay ra ‘yah 4 A y's « Ga! ADs ae aay apres oy Hk ia dak te blr ‘ ae vif deta, pedi ik wedded kes th ay th By di vtadthedieee od Fiend 4 Wibide 6 Key LaNrVEY yh s Need o it f phy he Ha fh A har a ULL A Dh had atts “ihe 4 wh Hits ‘ad's. yi ih ry Baile ay a ify hg a 4 y 4 ee he i Fioatied a ey ed HH aint Seb 4 1 4 He hips Aten wes rang i as a rfeate de wes ait Sara ane COM eto Rk Tor PA AL Pepa Aad dnd set he ARN) fly aea anid ated feed ede) ide eae ad SUC ay ated ee ede HAI dicen Oinen EEN A eae ern etna ge tet Oe a 4 len Panini: aiiee y ae LI ND i rf wis se iN Bia Ps BUN Ay ores Wa ML a ed “ * fr Sha Na Ny ne Rona MOR ea Men Rig A eros dete ae Naat ao ten ba Saar ater | he aia Sa paved Madea baad a8 Araya RT AN MN RATIO wae eats “ Manat Avid tomb towel LOM i WYCAA betta dt Wao garde host dea 4 aoe AT Lets edt 1 daniel Reg HOMES kaw Brg ETS Hh wane kh thd tity UALS aI) ised tac det 44 Pay YC: i ly Orv hod pate, wae CEA BP “au od tte ib ry ye aril We arden des Rak A Vara Was dada ba eent ; COTE nt want HN t gl i a) a ne lava a ee oe bh Rs Aira te sain iy SS ren ih neess fe i hey te aitieate tr 4 my ray) forty aera howe i eth -tE he the eh darth, Mea eh! eae wav ar ait) iy a4, vite ys ## na ae be nia a, , eh fy deka + oe a ett sai Fg b , is : eee begins , . tia a Ry iat Aas ren oy re ™, aan ~ “4 asd We cia el a0 iM et ibe Ahaha Wh) GUE dade oe ata? Whee ue Ae te Bingtl? LRabra Saas fo itd rere pete i ie oe ya Ny te Abate , Stl Amt Ue bed bled , Hii eulia Py se, athe A fini iM trot wer recy Ne H A i sh Ara sMiCA tt wu the ah eee hh eo hy a *i y Hh 4 soak ate Nh hete By det erat 4) Cekutiptit get eng ys De ) Ph) 6 ait We hat nite @: Solty at Da. iL patna ah A cnet Fy) . isa ee ate mas arg " adel ‘ nial, ry) ny ie at ay a1 Mast wi i aa pds heal ea # Ny hoa oes Es eben autre ee ee er neces ras eae Sie ~“e<. “i athe ae = Ueee es = ees Sree OR Se, == Eege oe oA = aS ee 4 OOK uh iy aq. way we ph Se ti a Ae ae oo ‘hy Keng v Garda a ue * m Ae it ae ves eens Ai re eet pends ’ oy . Prt Avett fe Aras + 4 sca oe nie ue hee itor a “4 see i, ae i eae “ i us pe ae rid pen a ist se ya Roca ae ria iia neil ae we 4 SS w hd ee ee ee tea ue nea on ah bl! sag we any oe oe rt ws ay A ales “ ‘* wae py Ap Oe hers i eG qe gt POA I NT Pert vid peer rer tee ayia Foy eaireenaats Arde a ean i : abel Wi val * wee ia va i fF uae ae 48 ih d Bye aia ni soe iw Te taaniara wntalaga | eed deste dtl a8 a, ey iY ¢ Pn oe bs ide ded: acmeli vt : PS sa ais fai i Ge" er ate ar GN Mi Dh ted Tada e 4 Por er Oke 2 4 Y ainie bali Beets be at vals Haat eri ack BA A Wea Se Cnn i swash We (ha i Ta) aed WP de dy nod wath a ae ad wes sed sak oad tee sania a sce8 ib ads rac “« Ne ra sun wee hoy A ah ‘ tea eg ated ty ad ue ea Mie wad err oe a Lea Me ei haat ad APU a \y saved aay Cy Dealer id ki te Te, Ria 4 ee et 4 jeat aay Gd ae ACN IN ASL)S OF THE SOUTH AFRICAN MUSEUM VOL CME Oe I a al oder lite eign) : 7 i) MR nf fm v oh Te Le % Au * D Re ay he ve yi \ % nT v ANNALS OF THE BoOUTH AFRICAN MUSEUM VOLUME. XLII OMe Ms a yu an eee ate ‘o ail E il RTE } cS Qi> af his 4 e. Ul ee tat PRINTED FOR THE ; TRUSTEES OF THE SOUTH AFRICAN MUSEUM : AND FOR THE GEOLOGICAL SURVEY OF SOUTH AFRICA "1953 — 1956 BY THE NATIONAL COMMERCIAL PRINTERS LIMITED, JAN VAN RIEBEECK STREET, ELSIES RIVER. TRUSTEES OF THE SOUTH AFRICAN MUSEUM. Dr. C. F. ALBERTYN. Pron. 'G. S! DE VILLIERS. Dr. M. R. Drennan, M.B., Ch.B., F.R.S.A.Afr.- CouNCILLOR H. E. GEARING. Cale SIBBEeTr,, ].P: Mimeow id. SKAIPE, |.Pi, PR S:S.Air-. SENATOR D. H. van ZyYL. SCIENTIFIC STAFF OF THE SOUTH AFRICAN MUSEUM. KEPPEL HARCOURT BARNARD, M.A., D.Sc., F.L.S., Director. ALBERT JOHN HeEsSsE, B.Sc., Ph.D., Assistant in charge of the Entomological Department. . Lizuwe DrrK Boonstra, D.Sc., Assistant in charge of the Palaeontological: | Department. Miss E. MARGARET SHAW, B.A., Assistant in charge of the Ethnological and. - Numismatic Departments. Miss G. Joyce Lewis, Ph.D., Assistant in charge of the Botanical Depart-— ment. | A. J. H. Goopwin, M.A., Honorary Keeper of the Archaeological Collections. LIST OF CONTRIBUTORS. L. D. BOoOoNSTRA. HH. a Note on some Rhynchosaurian remains from Tanganyika Territory Report on a collection of Fossil Reptilian bones from Tanganyika Territory Suggested clarification or the axehonne satis of the South African Titanosuchians : 7 The Gorgonopsians: Aelurognathus don and ‘Hipnesaiee boonstrai reconstructed ; The cranial morphology and taxonomy a the Tapinocemlakel genus Struthiocephalus 2 I at OES Sa The Lower Jaw Articulatory region in some Pristerognathid Therocephalians 2 ie The Pristerognathid ne Occohalians: fro the Tapinoce naan zone in the South African Museum wae The cranial structure of the Titanosuchian Anieccnuee The smallest Titanosuchid yet recovered from the Karroo Paranteosaurus gen. nov.: a Titanosuchian Reptile Struthiocephalellus: a new Deinocephalian The Girdles and Limbs of the South African Debceplaln B. S. COOKE. Some Fossil Mammals in the South African Museum collections . BE, EWER & R. SINGER, Fossil Carnivora from MHopefield . H. HAUGHTON. Phosphatic-Glauconitic Deposits off the West Coast of South Africa . N. KEEN 6 R. SINGER. Further Fossil Suidae from Hopefield . K. MILLER @ W. M,. FURNISH. Tertiary Nautiloids dredged near Cape of Good Hope . SINGER 5 E, N. KEEN. Fossil Suiformes from Hopefield PAGE 108 149 157 180 185 Lor 335 329 350 327 169 NEW GENERIC NAMES PROPOSED IN THIS VOLUME. meosatsanius (iherocephalidace) Boonstra «900.0; . 0. 2k ke 62 Micranteosaurus (Anteosauridae) Boonstra . . . . . . Cy 150 Neomegacyclops nom.nov. for Megacyclops Broom 1931 preocc., Boonstra 7 Pearanteosaurus (Anteosatiridae) Boonstra 50. a ee 57 Parascapanodon (Titanosuchidae) Boonstra .. SEN AONE We full tecvedt as ge Pristerognathoides (Therocephalidae) Boonstra . . . . . . 94 Pristerosaurus (Therocephalidae) Boonstra . : : ; BN SO Reeuemestes| (imerocephatdae)) Boonstra Woo ee 8 Struthiocephalellus (Struthiocephalidae) Boonstra . . . . . . 184 Semupazonsen) (Lherapsidac) (Boonstra: ake 9 Therioides (Therocephalidae) Boonstra PIN a Pu atic AR Te NR oat! A CR DATE, OF JSSUE,.OF THE: PARTS. Part 1, April, 1953. Part 2, January, 1954. Part 3, January, 1955. Part 4, March, 1956. PLATE, I-V. Vii VEE VEL TX. X-XIV. DY RV. XVII. XVIII. XIX. XX-XXIV. XXV. XXXVI. XXVIT-XXIX. XXX, XXXII. AXXITI. XXXII. XXXIV. XXXV. LIST. OF , PLATES. Anteosaurus abeli. Anteosaurus minor. Anteosaurus vorsterl. Anteosaurus abeli. Aelurognathus microdon. Hipposaurus boonstrai. Struthiocephalus. Micranteosaurus parvus. | Homoioceras bainii. Mesochoerus lategani. Struthiocephalus whaitsi. Aturia lotzi, Crocuta spelaea. Lycaon pictus magnus, and Crocuta spelaea. Mellivora capensis. Mesochoerus lategani, and paiceae. Mesochoerus, juvenile. Tapinochoerus meadowsi. Aelurognathus Aepyceros Agnosaurus Alcelaphus Alopecognathus . Anteosaurus Anthodon Antidorcas Aonyx : Archaeosuchus Arctocephalus Aturia 5 oe Aulacephalodon Avenantia 23, Balaena Balaenoptera Bubalus Cadulus Canis Carcharias Carcharodon Cephalophus Chlamys Connochaetes Conus Crocuta Cuspidaria Cynariognathus Damaliscus Dentalium Diceros Dicynodon Dinartamus . Dinocynodon Dinopolus Dinosphageus Dinosuchus Eccasaurus Enobius Equus Fusus LIsh OF PAGE. 29 165 BANS 26, 108, 327, 329, 766. 24, 334 GENERA. Gazella Giraffa Glanosuchus Herpestes Hippopotamus Hipposaurus Hippotragus Histiophorus Homoioceras Hyaena Hylochoerus Isurus Jonkeria Kannemeyeria Keratocephalus Lamiasaurus Leptailurus Loxodonta Lycaenops Lycaon Lystrosaurus Maraisaurus Megacyclops Megaptera Mellivora Mesochoerus Mesoplodon Metridiochoerus Micranteosaurus Mormosaurus Moschognathus Moschoides Moschops Moschosaurus Natica Notochoerus Neomegacyclops Notohipparion M 23; 164, Cars ey 29, 62, 170, 353) 156, ai 177, PAGE. 164 166 75 335 170 ae 166 333 161 336 353 333 285 212 340 165 Omochoerus Orca Ostrea Papio vee: Paranteosaurus Parapapio Parascapanodon Pecten Pedetes Pelea Peloroceras Pelosuchus Phacochoerus Phocosaurus Pleurotoma Pnigalion Potamochoerus . Pristerognathoides . Pristerognathus Pristerosaurus Procavia Propalaeonyx Ptomalestes Pusionella Pycnodonta 157; 164, Raphicerus Redunca : Rhachiocephalus Scapanodon Scapanodon ES PAGE. 25 269 Scaphonyx Scullya Scymnosaurus Staganosuchus Stenaulorhynchus Strepsiceros bl hae Struthiocephalellus Struthiocephaloides Struthiocephalus Struthionops Suricata 54> Sus Sylvicapra Syncerus . Tangagorgon Tapinocephalus Tapinochoerus Taurocephalus Taurotragus Terebratula Terebratulina Thecodontosaurus Therioides AOS Higa Titanognathus Titanosuchus Tragelaphus Trirachodon Xenophora Ziphius PAGE. 334 333 ANNALS OF THE SOUTH AFRICAN MUSEUM VOLUME XLII \ Descriptions of the Palaeontological Material collected by the South African Museum and the Geological Survey of South Africa. PART I, containing :— I. A Note on some Rhynchosaurian Remains from Tanganyika Territory. By L. D. Boonstra, D. Sc. (With two text-figures.) : A Report on a collection of Fossil Reptilian Bones from Tanganyika Territory. By L. D. Boonsrra, D.Sc. (With seven text-figures.) A suggested Clarification of the Taxonomic Status of the South African Titanosuchians. By L. D. Boonstra, D.Sc. (With Plates I-IX.) The Gorgonopsians, Aelurognathus microdon and Hipposaurus boonstrat, reconstructed. By L. D. Boonsrra, D.Sc. (With Plates X-XVI.) The Cranial Morphology and Taxonomy of the Tapinocephalid genus Struthiocephalus. By L. D. Boonstra, D.Sc. (With Plate XVII and _ ‘six text-figures.) _ The Lower Jaw Articulatory Region in some Pristerognathid Thero- cephalians. By L. D. Boonstra, D.Sc. (With four text-figures.) ISSUED APRIL 1953 PRICE 12s. 6d. PRINTED FOR THE TRUSTEES OF THE SOUTH AFRICAN MUSEUM | AND THE , GEOLOGICAL SURVEY OF SOUTH AFRICA BY THE RUSTICA PRESS LIMITED, COURT ROAD, WYNBERG, CAPE PONINIACES OF THE SOUTH AFRICAN MUSEUM VOLUME XLII 1. A Note on some Rhynchosaurian Remains from Tanganyika Territory. By LizEUWE D. BoonstraA, D.Sc. (With 2 text-figures.) From Mr. G. M. Stockley I have received three tins containing fossil remains from near Msamara, in the Tunduru district of the Southern Province of Tanganyika Territory, for determination. This material ($7662) consists of weathered fragments of skull, limb-bones and vertebrae. The largest skull fragment consists of part of the left upper jaw. The limb-bones include a proximal third of a right femur, a weathered distal end of a femur, a bit of humerus, scapula and tibia. The vertebral material consists of the weathered centra of dorsal and caudal vertebrae showing few distinctive features. SKULL (fig. 1) This weathered and rolled fragment consists of part of the left jugal, maxilla and palatine. On the lateral surface two nutritive foramina pierce the maxilla. The dentigerous border of the maxilla is convex antero-posteriorly. ‘The ventral surface of the maxilla is studded with a large number of worn peg-like teeth varying in diameter from 2 to 4 mm. These teeth are somewhat indefi- nitely arranged, but there does appear to be some indication of an alignment in four rows. The teeth on the inner or lingual margin of the maxilla are aligned in a fairly definite row. On the outer or labial margin and anteriorly the teeth are larger than those situated medio-posteriorly. The width of the dentigerous surface is wider posteriorly than anteriorly. Medial to the maxillary teeth there lies a groove filled with matrix, and the maxillo-palatine suture would lie along this groove. Lingual to this groove there lies a row of teeth on the labial border of the palatine, with at least three more indefinite rows of teeth lingually. As is the case with the maxilla the width of the dentigerous surface is posteriorly wider than anteriorly. As preserved, the length of the maxillary dentigerous surface is 100 mm., the width anteriorly 7 mm. and posteriorly 20 mm. The nature of the dentigerous plate of the maxillo-palatine indicates that we have here a fragment of a Rhynchosaurian skull showing particular affinities Won. XLII. PART I. SUL 23 1952 2 ANNALS OF THE SOUTH AFRICAN MUSEUM to Scaphonyx australis described by Von Huene from South America, Hyperoda- pedon gordom described by Huxley from Scotland, and Hyperodapedon huxleyi described by Lydekker from India. The length of the dentigerous surface in this specimen from Tanganyika is, as preserved, greater than in these three known forms. Fic. 1.—Maxillo-palatine complex of Scaphonyx stockleyi. a, lateral view (x 4). 6, ventral view (x 4). S.A.M. 11704. In H. gordoni the teeth are placed in rows much more definitely than in the Tanganyika specimen; in the former there are two rows on the maxilla whereas in the latter there are possibly four rows; and the width of the dentigerous surface of the maxilla is also smaller in H. gordoni. In H. huxleyi the teeth are also in much more definite rows, and the number of rows on the maxilla exceed those on the palatine. In the Indian beast the crowns form triangular pyramids whereas in the Tanganyika beast the crowns are bluntly conical. In Scaphonyx australis the teeth are of very similar shape and have very nearly the same arrangement. But, whereas the anterior maxillary teeth are larger than the posterior ones in the Tanganyika specimen, the reverse is the case in the South American specimen. NOTE ON SOME RHYNCHOSAURIAN REMAINS 3 From the available evidence, admittedly scanty, it would thus appear that we have here a type distinct from all the hitherto known Rhynchosaurians, but apparently approaching fairly closely the skull fragment described by Von Huene from South America and by him referred to as Scaphonyx australis. I propose that the Tanganyika beast be included in Smith Woodward’s genus Scaphonyx, but to be distinguished from the hitherto known species and to be known under the name Scaphonyx stockleyi n.sp. Type.—Skull fragment in the South African Museum, 5.A.M. 11704. Femur (fig. 2) The proximal third of a weathered right femur is preserved. The post-axial corner with the external trochanter is missing. Noteworthy features of this fragment are: the caput femoris is bent sharply dorsally; the internal tro- Fig. 2.—Proximal end of femur of Scaphonyx africanus. a, ventral view (xX #). b, dorsal view (x4). c¢, pre-axial view (x4). d, articular surface (x }). S.A.M 11705. chanter forms a prominent bulbous structure and is abruptly demarcated from the caput, and is situated some distance distally from the proximal articulatory surface; the inter-trochanteric fossa is comparatively shallow, but, because of the curvature of the bone and the presence of a low ridge, is clearly separated from the more distal ventral surface of the bone; the proximal articulatory surface is wide dorso-ventrally; the external trochanter was probably not strongly developed and the ilio-femoralis flange not extensive. This femur is about half the size of that (S344) described by Haughton under the name Stenaulorhynchus stockleyi but is obviously of a related type. Apart from the difference in size it differs in the shape of the caput; the internal trochanter is situated further distally; is more bulbous and is abruptly separated from the caput without a connecting neck. This femur, although also only half the size, shows closer affinities to that of Scaphonyx fischeri and Scaphonyx australis as described and figured by Von Huene. But here also the nature of the internal trochanter is strikingly different, and the femur cannot be considered as being that of either of these two South American species. 4. ANNALS OF THE SOUTH AFRICAN MUSEUM If Von Huene is correct in assigning his skull fragment and femoral fragment to the same species (Scaphonyx australis), we have a dentigerous plate of 80 mm. associated with a femur proximally 80 mm. wide. In the Tanganyika material we have a dentigerous plate of at least 100 mm., and we would thus expect that the associated femur would have a width proximally of about 100 mm. Actually the femur under consideration cannot proximally be of a greater width than 50 mm. The femur can thus only be of a younger beast or of a smaller species. ‘The ossification of the femoral fragment does not appear to be that of an immature half-grown animal, and is thus in all probability that of a species of half the size of Scaphonyx australis. For this species I propose the name Scaphonyx africanus. Type.—Proximal third of a right femur, S.A.M. 11705. The discovery by Mr. Stockley of these definite Rhynchosaurian remains in Tanganyika, together with the Rhynchosaurian limb-bones of Stenaulorynchus stockleyi previously described by Haughton from Nyjalila, will enable the strati- graphists to attempt a closer correlation of these beds with those that have yielded the species of Hyperodapedon, Scaphonyx, and the other closely related genera. The discovery of better-preserved material may make it necessary to remove these species from the genus Scaphonyx to a new genus. Until such time I do not think it necessary to create a new genus for these African forms. 2. A Report on a collection of Fossil Reptilian Bones from Tanganyika Territory. By LizEuWE D. Boonstra, D.Sc. (With 7 text-figures) INTRODUCTION Recently I received from Mr. G. M. Stockley eight cases of fossil reptilian bones collected from the Ruhuhu Coalfield region. This collection was obtained from three distinct localities. ‘Those labelled $346 come from the Matomondo area, $559 from the Ngaka-Kingori Hill area, and $340 from the Nijalila- Mkongeleko area. The geological horizon of the first two localities is given as the Lower “Bone Bed’ of the Songea Series, and the third as the Upper ‘Bone Bed’ of the same series. Stockley states that the bones ‘were picked up from the surface and thus are much weathered’. It is clear that all the fragments collected from a locality were given the same number, and each number undoubtedly includes fragments derived from a number of different skeletons. I have no means, other than the state of preservation, of determining the degree of association of the various fragments. For the greater part the fragments thus have to be considered individually. No evidence is given by the collector that the fragments bearing the same number were obtained or derived from the same horizon within the bed. On the contrary, it is evident on morphological grounds that from the Upper ‘Bone Bed’ there are at least four groups of specimens and that these were derived from four different levels within the bed. It should be stressed that in future collecting attention should be paid to the relative levels within the bed from which each specimen is collected. A. MATERIAL FROM THE LOWER ‘BONE BED’ 1. Anomodont Skull Fragments Under the label $346 there are a small number of weathered pieces which include fragments identifiable as parts of a fairly large anomodont skull with a narrow parietal crest. ‘This feature apparently excludes reference to the Aulacephalodon group of genera, and these fragments must thus represent a large species of the genus Dicynodon. Under the label $559 there are a large number of weathered pieces—cranial as well as post-cranial. Of the recognizable skull fragments the following identifications have been possible, mainly on pieces of the intertemporal region bearing the pineal foramen and the structures immediately surrounding it. (a) Inone piece of the intertemporal region of a fairly large dicynodont there is a medium-sized oval foramen situated at the posterior end of a depression formed by the preparietal. The parietal crest is of medium width and height. The distinguishing feature in this fragment is the nature of the preparietal. 6 ANNALS OF THE SOUTH AFRICAN MUSEUM From the anterior border of the foramen the sides of the preparietal diverge in anterior direction. The preparietal thus meets the frontal in a long transverse suture. The whole of the bone lies in a depression. Direct comparison proved the identity of this specimen with a form from the Luangwa Valley of Northern Rhodesia which I described under the name Dicynodon roberti. This specimen now bears the South African Museum Cat. No. 11706. _ (5) In another frag- ment (fig. 1) the pineal foramen pierces a large oval boss which is almost entirely formed by the preparietal. The pos- terior third of the boss is formed by the parietal, but this bone only stretches two-thirds up the posterior surface of the boss and is thus excluded from the border of the pineal foramen. The : preparietal completely a surrounds the foramen, which is of medium size and nearly circular. The foramen is directed anteriorly, and the plane in which it lies makes an angle of 45° with the plane of the dorsal sur- face of the skull. Pos- teriorly, the surface of the boss, here formed mainly by the parietal, a, dorsal view of pineal seston (4). , lateral view (lef of OP V2°8i Maia pineal boss (x 4). The parietal, which has only a small exposure, is then covered by the two post-orbitals which meet on the median line. Anteriorly, the boss is abruptly marked off from the surface of the frontals by a semi- circular groove. A near approach to this condition of the pineal region, in the more fully known Karoo species, is shown by some of the species of the genus Platycyclops, where the foramen is also completely surrounded by the preparietal raised in the form of a boss. A nearer approach still is shown in the species of the genus Megacyclops, where the parietal participates in and forms the posterior part of the boss. In the present specimen the shape of the boss differs remarkably REPORT ON COLLECTION OF FOSSIL REPTILIAN BONES 7 from that of M. whaitsi of the South African Karoo and of M. rugosus described by Haughton from Tanganyika, but agrees in other features, particularly in being so definitely anteriorly directed and in the manner in which the parietal participates in its formation. It differs from M. usilzensis of Von Huene in the slope of the boss and in the fact that the parietal is excluded from the border of the foramen. It is thus evident that we have here a new species of Megacyclops* and this fragment can be designated as the type specimen of Neomegacyclops cyclops n. sp. This specimen now bears the South African Museum Cat. No. i 70'7- (c) In a third fragment where the intertemporal region is preserved, the nature of the pineal region is strongly reminiscent of that known in the genus Rachiocephalus of the Endothiodon zone of the South African Karoo and may be provisionally referred to that genus. This specimen now bears the South African Museum Cat. No. 11708. (d) Another identifiable fragment is of the central part of a massive occiput. The basisphenoidal tubera are exceptionally massive and are, furthermore, peculiar in that they lie practically horizontally instead of approaching the vertical, so that the foramen ovale is directed nearly completely posteriorly, whereas the more usual direction is nearly completely ventrally. This condition is not known to exist in the genera Megacyclops and Rachiocephalus. This fragment is thus not derived from either of the large skulls that yielded the pineal regions mentioned above. In none of the larger Anomodonts examined by me nor in the published descriptions have I seen basisphenoidal tubera of a similar nature. This fragment thus indicates the existence in Tanganyika of a new type of the larger Anomodonts, of which one hopes that a more complete skull may soon be found and be designated as the type of this new species. This specimen now bears the South African Museum Cat. No. 11709. 2. Anomodont Limb-bones Included in the collection are a number of weathered distal and proximal ends of some large humeri, femora and ulnae. ‘The former two are of about the same size and nature as the humerus and femur described by Haughton and referred to as Eocyclops? and Megacyclops respectively. In addition to these there are some proximal and distal ends of humeri and femora of Anomodonts of smaller size which could be associated with a skull of the size of a form like Dicynodon huenei—a species peculiar to Tanganyika. These specimens now bear the South African Museum Cat. Nos. 11710 and 11742. 3. Pareiasaurian Vertebrae Two medium-sized vertebrae are preserved in a weathered condition. These are typically Pareiasaurian, and are smaller than those of the larger genera which predominate in the Tapinocephalus zone of the Union. They are also * I find that the name Megacyclops which Broom (1931) proposed for this anomodont genus is preoccupied for a crustacean (Kiefer, 1927). I propose that this anomodont genus be known under the new name Neomegacyclops nom. nov. 8 ANNALS OF THE SOUTH AFRICAN MUSEUM smaller than, and do not show the peculiarly narrow elongated dorsal spine so characteristic of, the vertebrae described by Haughton from locality Big, west of Kingori. These vertebrae prove the presence in Tanganyika of Pareiasaurs probably closely related to the medium-sized types of the South African Karoo. These now bear the South African Museum Cat. No. 11743. 4. Therapsid Cranial Material Among the mass of weathered fragments from the Ngaka-Kingori Hill area there are two incomplete snouts much weathered, and one fairly complete but Fic. 2.—Tangagorgon tenuirostris n. g. et n. sp. S.A.M. Cat. No. 11744. Lateral view of skull (x 3). somewhat distorted and weathered skull. The more complete skull is of a Gorgonopsian and the two snouts appear to be Therocephalian. (a) The more complete specimen (figs. 2 and 3) has the outer surface con- siderably eroded, so that it is difficult to determine the sutures. The skull is of medium size. The chief measurements are: Maximum@length ora i ee Basioccipital—Premaxilla 72 te Premaxilla—-Pineal’foramen’ 2...) ie Premaxalla““Orbit <0" en es Widthacross'squamosals’)) "525 1 Imterorbital width" oe ee a ee AOS; Se Intertemporal“width® (o's ea ee Widthacross' canines) iS ee ce el a se Height from Mx. edge—median suture . . . . 50 ,, Height of mentum ene nner iO. Length of molar, series: Left,(4 teeth) )\...4..9 ee Right (@ teeth); 4.5) 29) eee Lengthyotuncisomseresien 5.) sane Dental formula: Right, i5, co+1, m3. Left, 15,.cr--1, m4. 25 » REPORT ON COLLECTION OF FOSSIL REPTILIAN BONES e) The snout is long, high and narrow. Anterior to the orbits the cross-section is squarish. The orbits are situated in the posterior half of the skull and are laterally directed. The temporal openings are of medium size, oval, and directed as much dorsally as laterally. The pineal foramen is small and posteriorly situated. The frontals apparently form only a small part of the orbital border. The snout is higher than wide. ‘The maxilla is deep. The interparietal is only slightly inclined downwards from the general dorsal surface and is more a bone of the dorsal surface than of the occiput. From the posterior edge of the inter- parietal the occiput descends practically vertically. The nature of the rest of the occiput is obscured by the presence of the proatlantal arches, atlas and axis. The teeth are poorly preserved. On the left there are preserved five medium-sized incisors, a fairly weak canine with the tip of a replacing? canine just emerging, and the roots of four small molars. On the right side the last of the five incisors is small (a replacing tooth ?), the canine has been lost and a replacing? one is emer- ging; the crowns of three small molars are preserved, number 3 having fallen out. The characters enumerated above do not occur together in any known Gorgo- nopsian, and I therefore designate this specimen as the type of a new genus and species— T angagorgon tenuirostris. ‘This type specimen now bears the South African Fic. 3.—Tangagorgon tenuirostris n. g. et \ \ ‘ \ 1 ’ <= eof Museum Cat. iy : n. sp. S.A.M. Cat. No. 11744. Dorsal Sutth (Cee NG: 744 . view of skull with proatlas, atlas and (6) The smaller snout fragment is axis. (x 4) nearly completely stripped of bone but shows the roots of five strong and large incisors. This specimen, which. is probably Therocephalian, now bears the South African Museum Cat. No. 11745. (c) The larger snout fragment (fig. 4) represents the anterior half of a medium-sized skull. The tip of the snout is weathered away and no upper incisors are preserved. The lower edge of the dentary is also weathered away. A moderately sized canine is represented by a root, and four irregularly spaced and directed molars are present. The molars are implanted on a flange of bone lying in a plane medial to the general lateral maxillary surface. The lacrymal is of small antero-posterior extent and its intra-orbital surface is pierced by two IO ANNALS OF THE SOUTH AFRICAN MUSEUM foramina. The accompanying sketch gives the general shape and proportions. The affinities of this Therocephalian are uncertain, but the nature of the lacry- mal indicates that it is not a very primitive form, and that its affinities would lie with the Therocephalian forms younger than those from the Tapinocephalus zone. This specimen now bears the South African Museum Cat. No. 11746. B. MATERIAL FROM THE UPPER ‘BONE BED’ Under the field number $340 there were six cases of weathered fragments. The majority of these, mostly small pieces, are so worn as to be indeterminable. In the workshop few contacts were found and only a small number could be fitted together. The determinable pieces consist of cranial fragments, a large number of vertebrae, parts of the pelvic and shoulder girdles, three complete Fic. 4.—Therocephalian sp. S.A.M. Cat. No. 11746. Lateral view of incomplete and weathered snout. (x $) limb-bones and a large number of ends of various limb-bones. The following reptile groups are represented: Pareiasaurians, Anomodonts, Rhynchosau- rians, Pseudosuchians, Cynodonts, and a Theropodous Dinosaur. 1. Pareiasaurians Of this group there is preserved a single weathered distal end of a small right humerus. It is much smaller than the specimen from Tanganyika figured by Haughton and described under the new specific name Anthodon minusculus. In Haughton’s specimen the greatest width across the condyles is 87 mm., whereas in this specimen it is only 63 mm. In all other determinable features the two specimens agree very closely. It would thus appear that we have here a humeral end of a smaller species or more probably of a juvenile specimen of Anthodon minusculus. In the material from Tanganyika described by Haughton some errors of labelling are mentioned by him. The type specimen bears the number $342 REPORT ON COLLECTION OF FOSSIL REPTILIAN BONES Ii and a second specimen the number $350; the former number being given to specimens from a locality (Njalila) in the Upper ‘Bone Bed’ and the latter number to specimens from a locality (below and west of Kingori) in the Lower ‘Bone Bed’. Haughton assumed the number $342 to be an error in the labelling. Now, the present specimen bears the number $340, which refers to a locality (Njalila-Mkongeleko) in the Upper “Bone Bed’. The weight of evidence in regard to the labelling is thus that all these specimens are derived from the Upper ‘Bone Bed’. Palaeontologically, however, the Lower ‘Bone Bed’ is the horizon indicated, unless we consider it probable that this species survived practically unchanged into the Upper “Bone Bed’. Until this uncertainty is removed these specimens must be cited with caution in drawing stratigraphical or faunistic conclusions. 2. Anomodonts Fragments of skulls, lower jaws and various limb-bones prove the presence in these beds of species of medium sized to large Anomodonts. (a) A well-preserved right humerus of medium size (fig. 5) can, despite the paucity of our knowledge of the post-cranial skeleton of the numerous species of Anomodonts, be referred to the genus Lystrosaurus. ‘The distal condyles are undeveloped, the antero-ventral line is only weakly indicated, the lateral median line undeveloped, and no definite scar for the medial humeral head of the triceps is present. These characters exclude the terrestrial genera of the Anomo- donts, and agree with the condition manifested in the species of the genus Lystrosaurus. In the absence of a skull no specific determination is possible, and it is advisable, until further data are available, to refer to this specimen as Lystrosaurus sp. ‘The chief measurements are: Greatest lencthen \ 4a ee ee, 4m Greatest;proximal width= 377 © ;, Greatest aistal.width =~.) 2 2/2 61... Shattw cs Gh oe bves 5 ram: This specimen now bears the South Atican Museum Cat. No. 11748. (b) Under the number $340 are included about a dozen ends of limb-bones. Among these are seven distal and three proximal ends of humeri, three distal and two proximal ends of femora which, with slight individual differences, agree well with the corresponding bones of Kannemeyeria. ‘The widths across the distal ends of the humeri are: 180, 160, 158, 154, 148, 139 and 125? mm. These humeral ends are manifestly different from the humerus figured by Haughton under the name Focyclops(?) sp. These specimens now bear the South African Museum Cat. No. 11749. Some fragments of the pelvic and pectoral girdle may also very well belong to this genus. In addition to these limb-bones there are also preserved the ends of much smaller Anomodont humeri and femora. So that, contrary to our experience in the South African Karoo, small Anomodonts survived comparatively late in Tanganyika. These bones now bear the South African Museum Cat. No. 11750. ANNALS OF THE SOUTH AFRICAN MUSEUM 12 "(¢ x) smoata Jeunxoid pure so1ojue “[esrop ‘TeVUSA UT snJoUINET ys “gPL11 ‘oN VED ‘Fry's ‘ds sninvsousdI—G oy e hy) Jniid Wie REPORT ON COLLECTION OF FOSSIL REPTILIAN BONES 13 (c) ‘The Anomodont cranial material includes parts of the temporal arches, a quadrate, and some pieces of occiput. One occiput, with its tripartite condyle and its characteristically shaped and ventrally directed basisphenoidal tubera nearly surrounding the foramen ovale, is strongly reminiscent of the same struc- tures in the skull of Kannemeyerta and may with confidence be referred to that genus. Some pieces of maxilla with the canine roots preserved may also be included in this determination. These fragments now bear the South African Museum Cat. No. 11751. (d) Three other occipital fragments are more massive. Here the condyle is rounded with no grooves tripartitioning it. ‘The basioccipital tubera, though massive, do not descend so far ventrally and are situated some distance apart. A very similar condition is shown by a number of the larger Aulacephalodon-like forms, but as in the large number of described forms this area is neither figured nor described, closer comparison is not possible, and for more specific deter- minations it is necessary that we have better descriptions of known forms or better-preserved skulls from this area. These specimens now bear the South African Museum Cat. No. 11752. 3. Cynodonts One very badly eroded fragment consists of the anterior two-thirds of a fairly small Cynodont skull. The outer surface is so badly weathered that the structure is indeterminable, and on the palatal surface little more than the teeth sockets are preserved. These indicate that we have here a specimen of the genus Trirachodon. This specimen now bears the South African Museum Cat. No. 11755. 4. Rhynchosaurians The collection includes the following bones, or parts of bones, determinable as Rhynchosaurian: a complete humerus, two proximal and seven distal ends of humeri, six proximal and four distal femoral ends, parts of the pelvic and pectoral girdles, a large number of disarticulated vertebrae, a jaw fragment and two incomplete occiputs. (a) The Humerus (fig. 6).—Direct comparison of these humeral elements to the type humeral end of Stenaulorhynchus stockleyt proves them to be specifically identical. In size and in the proportions there are considerable differences in the hitherto described material and that at present under consideration, as will be apparent from the following table: Type Para- Huene’s Spec. Spec. Spec. Specs. a b type Spec. c geen fe via, length. 22) se) E00 : ? 145 166 G 2 a vats fiat Max. width across prox.end 10! 77 92 110 120 ? OE niet nan Piatt ae (eee - ee AL X< 30) OTOL On KOO)" 39> 241 ? ? iii a Vi Proc. lat.—prox.end . . 83 57 60? 81 94. ? abidots sal ig Proc. med.—prox.end . 85 63 88? IOI 110 ? be sO Thickness at med. corner . 37 26 33 31 40 Y Ee ia aol Max. width across dist. end 2 ? 80 100 ? OSE SOr a7! AO, ANNALS OF THE SOUTH AFRICAN MUSEUM 4 I suimeiq *(# xX) smorA [ewrxoid puv solazue ‘Jessop “fen ‘assaH] ‘[ vy aq Aq oA Ul snuouinyy 1ysry “SSL 11 ‘ON VeD “Py'sS. “2py205 snysudsojnouajs—'g “O14 REPORT ON COLLECTION OF FOSSIL REPTILIAN BONES I5 In the light of our present limited knowledge it is advisable to consider these differences of size and proportions as age and/or sex variations and not as denoting any specific distinctness. (b) The Femur.—In the collection there are six proximal and four distal ends of femora. Direct comparison with the type material proves these bones to be specifically identical to Haughton’s Stenaulorhynchus stockley. As in the type material there is considerable variation in the various dimensions: , a b c d e ie Max. width over trochanter . 93 73 72 71 65 64 Max. thickness of prox.end . 58 y 44 53 46 52 Trochanter—prox.end. . . 37 2 Be: 15 22 BT There is, moreover, considerable variation in point of size, shape and position of the trochanter: In a the slightly bulbous trochanter is connected with the proximal surface by a somewhat constricted neck. In 6 the trochanter is less bulbous and the neck less constricted. In ¢ the trochanter is hardly thickened, and instead of a neck a sharp ridge connects it to the proximal surface. In d the trochanter lies nearly in the same plane as the proximal surface, and in proximal view appears as a tongue-like extension of the proximal surface, without any neck. In e the bulbous trochanter connects with a short neck to the side of the proximal end to form a distinct step. In f a thickened ridge attaches the trochanter to the proximal end. In the specimen recently described by me under the name Scaphonyx africanus the bulbous trochanter is separated from the proximal surface by a much greater step than in e. The amount of variation shown by the above specimens impels one to reconsider the position of Scaphonyx africanus. ‘This must now be considered as an extreme variant of a femur of Stenaulorhynchus stockley1, and the name given by me becomes a synonym of Stenaulorhynchus stockley: Haughton. The distal femoral ends of our material agree very well with that of the type material and the shaft of one specimen with that figured by Von Huene. (c) The Girdles and Vertebrae——The material here preserved agrees in all essentials with the corresponding bones figured by Von Huene in his paper on Stenaulorhynchus. , | (d) The Occiput.—tIncluded in the material there are two imperfect and weathered occipital fragments that appear to be identical to the occiput described by Von Huene and figured in fig. 4 in his paper on Stenaulorhynchus and in fig. 6 in his “Die Verwantschaftsgeschichte der Rhynchosauriden des Siidamerikanischen Gondwanalandes’. A very eroded fragment in a different type of matrix shows a part of the maxilla and dentary which also appears to be Rhynchosaurian. All these Rhynchosaurian bones now bear the South African Museum Cat. No. 11753. ANNALS OF THE SOUTH AFRICAN MUSEUM 16 ‘(| x ) smora yeuutxoid pue so1a}ue “fesiop ‘fenuaA UIs ‘assay [vy ‘aq Aq suimeiqgg nooumy wysry “PSL11 on VO “y's ‘ds ‘u sosuavyruvsuny snyonsouosnjg—'L, *o1.J REPORT ON COLLECTION OF FOSSIL REPTILIAN BONES a7) 5. Pseudosuchians (fig. 7) Also bearing the field number $340 is a well-preserved right humerus. This bone shows undoubted and close relationship to the corresponding bone in the material collected by Dr. Nowack in the Njalila area, and included in the type material described by Von Huene under the name Stagonosuchus nyassicus, and representing a new genus and species of Stagonolepid Pseudosuchians. Although closely resembling Von Huene’s form, the following comparative table of measurements in mm. will show that some noteworthy differences in size and proportions exist: St. nyassicus St. tanganytkaensis Masamanriencth, 2. 2 320 200 Max. width across prox.end . . 180 12 Max. width across dist.end. . . 130 93 Maxewidth ofshaft . 9. 2... 50 38 roe lat—prox.end.° . . . . 70 52 Ectepicondylar flange—dist. end. 30 23 These differences appear to be of a specific nature, and I propose that this new species be known under the name Stagonosuchus tanganyikaensis. The main points of difference between nyassicus and tanganyikaensis may be enumerated: in nyassicus the processus lateralis lies in the same plane as the median corner of the proximal surface, whereas in tanganyikaensis the medial corner is not deflected and thus not situated so far distally; in tanganyikaensis the bicipital fossa is deeper and circumscribed much more definitely; the angle between the planes in which the proximal and distal ends lie is less in tanganyikaensis than in nyassicus ; proportionally the distal end is wider, the shaft has a greater maximum width and is relatively shorter, the processus lateralis extends further distally in tanganyikaensis than in nyassicus. ‘This humerus now bears the South African Museum Cat. No. 11754. : 6. Dinosaurs Among the large number of Rhynchosaurian vertebral elements in this collection I found a fairly small caudal vertebra lacking the upper part of the neural spine, of a Theropodous Dinosaur. The chief measurements are: Greatest lensth ofcentrum ~~. . .. 40 mm. Grcatestaheicht of centrum) isc js o 34 5, Greatesu width, of centrum >.2;.., ..-. 20 I have compared this vertebra with the Thecodontosaurus caudal vertebrae in our Museum and find a fairly close agreement, especially with the anterior caudals of a specimen from the Red Beds of the Stormberg Series of the Union. This specimen now bears the South African Museum Cat. No. 11793. CONCLUSIONS From the above account it is thus evident that in this collection there are from the Lower “Bone Bed’ no forms showing any close relationship to the fauna of the TYapinocephalus zone. ‘The Anomodonts, Pareiasaurs and Therapsids it 18 ANNALS OF THE SOUTH AFRICAN MUSEUM contains are all manifestly akin to species from the Endothiodon and Cistecephalus zones of the Karoo of the Union. The beds represented in the Matomondo and Ngaka-Kingori Hill areas are thus homotaxial to the upper two zones of the Lower Beaufort of the Union. The assemblage from the Upper “Bone Bed’ of the Njalila-Mkongeleko area, containing as it does small to medium-sized Anomodonts, Anomodonts of the Aulacephalodon group, a Lystrosaur, a Kannemeyeria, a Gynodont, a Rhyncho- saurian, a Pseudosuchian and a Theropodous Dinosaur, is related to the fauna known from the beds of the Karoo ranging from the top of the Cistecephalus zone of the Lower Beaufort right up to the Red Beds of the Stormberg. A more detailed recording of the relative levels in which the various fossils occur will enable the stratigrapher to subdivide the Upper “Bone Bed’ of this area into beds respectively homotaxial to the Middle and the Upper Beaufort, the Mol- teno and the Red Beds. It is probable that in this area some of the Anomodonts were actually derived from the top of the Lower Beaufort. ACKNOWLEDGMENTS My thanks are due to Mr. G. M. Stockley and his collectors for the oppor- tunity of examining this interesting collection, and to Dr. A. J. Hesse for two of the illustrations. REFERENCES Boonstra, L. D. ‘A Report on some Karroo Reptiles from the Luangwa Valley, Northern Rhodesia’, Q.7.G.S., xciv. 1938. wate L. D. ‘A Note on Some Rhynchosaurian Remains from Tanganyika Territory’, Ann. S. Afr. Mus., xli, p. 1. 1952. Brot, F., and ScHRODER, J. ‘Beobachtungen an Wirbeltieren der Karrooformation, xxiv, and ‘Uber Theriodontier-Reste aus der Karrooformation Ostafrikas’, Sitzb. Bay. Akad. Wiss. 1936. Lceron S. H. ‘On Karroo Vertebrates from Nyasaland’, Trans. Geol. Soc. S. Afr., xxix. 1926. Haucuton, S. H. ‘On a Collection of Karroo Vertebrates from Tanganyika Territory’, Q.7.G.S., Ixxxviil. 1932. Huene, F. von. ‘Uber Rhynchosaurier und andere Reptilien aus den Gondwana Ablagerungen Stidamerikas’, Geol. Pal. Abh., 17/1/1929. ‘Kurzer Uberblick iiber die terrestrischen Wirbeltierfaunen der jungeren Gondwana- zeit’, Centr. f. Min. usw., Abt. B, No. 6. 1933. ‘Stenaulorhynchus, ein Rhynchisauride der Ostafrikanischen Obertrias’, Nova Acta Leo- poldina, vi, 36. 1938. ‘Ein grosser Stagonolepide aus der jungeren Trias Ostafrikas’, N. Jahr. f. Min. usw., Bd. 80, Abt. B. 1938. ‘Die Alterbeziehungen der siidamerikanischen Gondwana Fauna’, Physis., xiv. 1939. ‘Die Lebensweise der Rhynchosauriden’, Pal. Zeit., xxi, 3. 1939. ‘Die Karroofauna im ostafrikanischen Ruhuhu Gebiet’, Zenérbl. f. Min. usw., Abt. B NOs 2a ‘Ein bese Pseudosuchier und ein Saurischier aus den ostafrikanischen Mandaschichten’, N. Jahr. f. Min. usw., Bd. 81, Abt. B. 1939. ‘Die Verwantschaftsgeschichte der Rhynchosauriden des sitidamerikanischen Gondwana- landes’, Physis., xiv. 1939. ‘Die Saurier der Karroo-, Gondwana- und verwandten Ablagerungen in faunistischer, biologischer und phylogenetischer Hinsicht’, NV. Jahr. f. Min. usw., Bd. 83, Abt. B. 1940. ‘Die Anomodontier des Ruhuhu-Gebietes in der Tubinger Sammlung’, Aeon braphica, XClV. 1942. eee aus dem Ruhuhu-Gebiet’, Pal. Zeit., xxiii, 3/4. 1944. PARRINGTON, F. R. “On the ‘Tooth-Replacement in ’Theriodont Reptiles’, Phil. Trans. Roy. Soc., B, 226, 532. 1936. ‘On the Cranial Anatomy of Cynodonts’, P..Z.S., cxvi, 2. 1946. Youne, C. C. ‘The Triassic Vertebrate Remains of China’, Am. Mus. Nov., No. 1324. 1946. 3. A suggested clarification of the Taxonomic Status of the South African Titanosuchians. By Lizuwe D. Boonstra, D.Sc. (With Plates I-I1X) INTRODUCTION Being engaged on a morphological study of the Deinocephalians the need has arisen to establish some order in the systematics of the group. In an assessment of the importance of the various morphological features the existence of a number of generic and specific names, not associated with at least some of the main morphological characters, is merely confusing and an encumbrance. In the literature there are about half a dozen generic and a somewhat greater number of specific names that signify next to nothing. One is led to wonder why they were ever created at all. One realizes that due to the nature of the material dealt with in palaeontology, it is not always possible to describe only relatively well-preserved specimens. ‘There is some justification, although one doubts the wisdom, if a newly discovered fragment establishes the existence of a group hitherto unknown to science (as, for instance, Owen’s creation of Titanosuchus Jerox), but there is no justification for subsequent authors to create additional names for specimens that do little more than prove that there are more indivi- duals of the group besides the original individual. It is, of course, fully realized that the vertebrate palaeontologist generally deals with individuals and that his species mostly refer to single individuals and are hardly ever the norm of a good series. In the Titanosuchians sixteen generic names have been created up to date. Of these only five (Jonkerta, Dinosphageus, Anteosaurus, Dinophoneus, and Dino- suchus) are based on reasonably fully preserved skulls, one (Phoneosuchus) on a good lower jaw, and the other ten (Tztanosuchus, Archaeosuchus, Dinartamus, Scapanodon, Lamiasaurus, Dinocynodon, Enobius, Scullya, Dinopolus, and Titanogna- thus) are all based on either skull fragments or parts of dentaries. In two former papers, in 1935 and 1936, I have shown that two of the above names must fall away—Dzinophoneus and Phoneosuchus being both synonyms of Jonkeria. In the sequel a further reduction of generic names will be proposed. At this stage I am only considering the cranial material on which, in any case, the authors have mainly created their new genera and species. I have critically examined most of the described specimens and for the rest have extracted the pertinent points from the descriptions of the authors concerned. In addition there is at the South African Museum a large collection, mainly collected by myself in recent years, of undescribed material which includes a dozen or so really good skulls. I now wish to present tentatively, for criticism by colleagues, a solution of the taxonomic maze formed by the plethora of names. With due Hg) 20 ANNALS OF THE SOUTH AFRICAN MUSEUM regard to the fetish of the law of priority I have attempted to include as many of the fragmentary specimens as is possible instead of simply regarding them as generically or specifically indeterminate. SEX AND AGE IN THE TITANOSUCHIANS In addition to the fragmentary nature of many of the types there are other difficulties in arriving at a reasonable classification. With the small number of individuals known we have little to indicate sex in the Titanosuchians. It must, however, be kept in mind that a smaller size, lighter build and a lesser degree of pachyostosis, particularly with regard to bosses, may indicate the female of the species. Without a reasonably long series of specimens it is difficult to determine the effects of age in the Titanosuchian skull. In the Tapinocephalian, Moschops, Gregory found an increasing pachyostosis, especially of the postorbital bar, a character indicating increasing age. In some of the Titanosuchians, especially of the Anteosaurus group, there is a slight variation in the thickening of the post- orbital bar, but whether this is due to age or sex is uncertain. In the Jonkeria group there are marked differences in size unaccompanied by other differences of generic value. Here size is not considered a character of generic value and at most may be regarded as specific, but in some cases may very well be a character of full maturity. Thus Dinosphageus is considered to be a large species of Jonkeria, but may very well prove to be a fully grown specimen of one of the previously described species of Jonkerta. ‘The strength and size of the dentary, particularly of the mentum, have been used as a character of systematic value, but is it not probable that a moderately strong dentary with a sloping mentum may become massive with a fairly upright mentum with increasing age? This was one reason for regarding the name Enodius as unnecessary. THE DENTITION The dentition of the Deinocephalians presents some very interesting condi- tions. A detailed study of the modifications and their implications should be undertaken. Here I can only touch on some aspects. A study of the teeth is made very difficult because of the poor preservation in most specimens. In the majority of cases the crowns are not preserved and we are forced to study stumps and cross-sections at varying levels which are not directly comparable. The presence of a lingual crushing surface internal to a talon in many teeth very materially affects the nature of the cross-section at different levels. Where crowns are preserved the intractable nature of the matrix of the Tapinocephalus zone makes it difficult to free them from the matrix satisfactorily. The Titanosuchians have a heterodont dentition with incisors, canine and postcanines. In some forms the incisors are fairly short with a labial talon and a lingual crushing surface, in others the incisors are long to very long with a talon long to very long and the lingual crushing surface is poorly developed or represented only by a cingulum, or wholly absent. ‘The canines are strong to TAXONOMIC STATUS OF SOUTH AFRICAN TITANOSUCHIANS DON very strong, simple, pointed teeth of a typical carnivorous nature with, in some cases, a possibly serrated posterior cutting edge. What little is known of the postcanines indicates a considerable variation in the nature of the crowns— some have flattened crowns with serrations, others short or longish conical teeth. Now, is the heterodont Titanosuchian dentition to be morphologically derived from the homodont dentition of the Tapinocephalians? The Tapino- cephalian teeth all have a talon, pointed or somewhat flattened, and a lingual crushing surface varying in strength, from which the different Titanosuchian types of teeth could be morphologically derived. The medium-sized incisors of a form like Jonkeria, with its short talon and definite lingual crushing surface, would then represent a more primitive stage than the long incisors of some of the Anteosaurians which are all talon with no crushing surface. The stages in this transition could then be used for taxonomic purposes. But, in Dinartamus, Broom found a mixture of these two types of incisors and the question arises whether this may not represent a difference in the consecutive sets of incisors. We do not know how many times the Titanosuchians replaced their incisors. Hitherto only two sets have been described. On this subject the external nature of the teeth gives little information and we have to rely on fortuitous weathering and fracturing or sectioning. The material at my disposal affords no evidence as to whether the Jonkerian type of incisor is the deciduous type and the Anteo- Saurian type typical of a later set of teeth. Until we have evidence supporting Broom’s observations on Dinartamus I propose to consider the presence or absence of the lingual crushing surface in the incisors to be a specific character present in the consecutive sets of teeth, and not dependent on age. Besides having these two types of incisors the Titanosuchians also possess a variable number of incisors—as described, from 0 to 5. In those specimens where the premaxilla is edentulous the condition can be interpreted in a number of ways: the incisors may have been shed postmortemly, or being juvenile they have not yet erupted, or being gerontic they have been lost. The nature of the material being what it is, it is difficult to decide which explanation best fits each case. Sectioning through the alveolar region is not always possible in order to base a count on the nature of the alveoli. The usual number of incisors is 5 but in some forms it is 4 or even 3. Can the number be considered a character of systematic value? If the juvenile and mature incisors are of the same size then obviously the premaxilla of a young animal could only house a smaller number than that of the mature animal. In using the number of incisor teeth as a taxonomic character the age of the animal must thus be taken into account. The number of postcanine teeth in the Titanosuchians varies greatly, from 1 to 19, and is of doubtful systematic value. In some, these teeth are small; in others, strong; in some specimens they are regularly spaced and in others quite irregular. The two sides are often dissimilar, e.g. in Enobius there are, according to Broom, in the left dentary ‘. .. portions of three molars. There certainly have been four, and very probably there have been five’, and on the right side 22 ANNALS OF THE SOUTH AFRICAN MUSEUM Broom found only one tooth behind the canine. In general it can be stated that in the Jonkerias the postcanines are regularly spaced and form a long series, whereas in the Anteosaurians they are irregular and the series is short. With the material at my disposal I can state that the dentary usually has one incisor less than the corresponding premaxilla. It would also appear that there are fewer postcanines in the dentary than in the opposing maxilla. Teeth on the palatines occur in the Anteosaurians and in the fragment called Scullya, but have as yet not been recorded in the Jonkerias although I found some indication in Jonkeria ingens (A.M.N.H. No. 5608).* These teeth are mostly seen only in section but in $S.A.M. No. 11592 there is preserved a small pointed tooth slightly recurved. In the Anteosaurians the palatine teeth are situated on a prominent elevation, semilunar or reniform in shape. Their function seems clear—the incisors and canines tear out a lump of flesh from the victim and this is then held by the raised palatine teeth as an intermediate stage in the swallowing process. The palatine teeth situated on this characteris- tic boss constitute a diagnostic character of value. KEY FOR THE GENERA Bearing the above remarks in mind we may now proceed with our attempt to arrange the Titanosuchians in some reasonable order. As a preliminary step, and in practice of considerable value, to facilitate the process of a rough-and- ready sorting out of the 14 genera, I propose making use of the following key: A. Forms without bosses on postorbital bar and angular, incisors not long and with step: 7 1. With many regular postcanines, no palatine boss—fonkeria, Dinospha- geus, Dinopolus. 2. With few postcanines—Dzinariamus. B. Forms with bosses on postorbital bar and angular, incisors long and with- out step: 3. With variable irregular postcanines, prominent palatine boss—Antéeo- saurus, Dinosuchus. 4. With variable irregular postcanines, probably with palatine boss— Titanosuchus, Scapanodon, Archeosuchus, Lamiasaurus (snout), Dinocynodon, Scullya, Titanognathus, Enobuus. REDEFINITION OF GENERA With our increased knowledge of the Titanosuchians it has become advisable to redefine the valid genera: * For the institutions housing the specimens here referred to the following abbreviations are used: A.M.N.H., American Museum of Natural History, New York. S.A.M., South African Museum, Cape Town. B.M., British Museum (Natural History), London. T.M., Trans- vaal Museum, Pretoria. A.K., Alte Akademie, Munich. K.M., Kimberley Museum, Kimberley. TAXONOMIC STATUS OF SOUTH AFRICAN TITANOSUCHIANS 23 I. Jonkeria van Hoepen 1916. se The genotype 7. truculenta is based upon an excellent skull and lower jaw. Io. It. cs T3. Skull size—this is medium to large, not massive. Bosses—there are no prominent bosses. Additional pachyostosis—the parietals form a prominent ridge thickened round the pineal foramen but do not form a well-demar- cated boss. Palatine—there is no prominent boss, but the palatine is possibly dentigerous. Premaxilla—the dentigerous border does not curve upwards and there is also no corresponding upward sweep of the dentary. Incisors—are of medium length with labial talon pointed and with well-developed lingual crushing surface. Pineal foramen—penetrates anterior part of the parietals and is thus some distance from the occipital border. Snout—is relatively long, broader than high, the frontals are only slightly swollen, the premaxilla is fairly flat. Squamosal—does not extend far ventrally and does not sweep far posteriorly, posterior to the interparietal. There is also little lateral sweep of the squamosal. Temporal fossa—this is fairly large, with the antero-posterior and dorso-ventral diameters moderate and approximately equal, not extending much laterally, as the squamosal does not sweep much outwards. Intertemporal width—this is small to moderate; the parietals are laterally pinched in to form a fairly high and narrow parietal crest. Dentary—this is strong but not massive, with sloping mentum. - 4—5 1 14—19 Memeaormiula ie, C7.) P-€ aaaqe Synonym. As thus defined Dinosphageus and Dinopolus become congeneric with Jonkeria. Anteosaurus Watson 1921. The genotype, A. magnificus, is based upon the major portion of a skull somewhat weathered. Tee Die 3. Skull size—this is small to very large, slightly to very massive. Bosses—a medium to very prominent boss is present on the dorsal part of the postorbital bar, and there is a prominent oval boss on the angular, the boss on the jugal is absent or low and moundlike to very prominent. Additional pachyostosis—the parietals are much thickened but do not form a crest, greatly thickened around the pineal foramen to form a mound or well-demarcated circular boss; the frontals are sreatly thickened to produce a medium to very prominent swelling. 24 HII. ANNALS OF THE SOUTH AFRICAN MUSEUM 4. Palatine—there is a prominent semilunar ridge-like or reniform boss- like eminence carrying irregular small pointed teeth. 5. Premaxilla—the dentigerous border curves antero-dorsally to form an obtuse angle with the maxillary border and there is little corre- sponding upward sweep of the anterior part of the dentary. 6. Incisors—are long to very long, the labial talon forming most or all of the tooth, with the lingual crushing surface greatly reduced to form little more than a cingulum, or are altogether absent. 7. Pineal foramen—penetrates the posterior part of the parietals and is thus near the occipital border. 8. Snout—is of medium length, higher than broad, the frontals are greatly swollen, the premaxilla is dorsally swollen and demarcated from the maxilla by a groove. 9g. Squamosal—extends moderately to far ventrally and sweeps far posteriorly, i.e. much posterior to the occipital surface of the inter- parietal, and sweeps far to very far outwards. 10. Temporal fossa—this is large, with the dorso-ventral diameter ereater than the antero-posterior, extending much laterally due to the outward sweep of the squamosal. 11. Intertemporal width—this is moderate to large, the parietals are laterally somewhat pinched in, but do not form a high and narrow crest. 12. Dentary—this is strong and fairly to very massive, with fairly upright mentum. 13. Dental formula—i.3=; ct pee Synonyms. As thus defined Dinosuchus becomes congeneric with Anteosaurus. Dinartamus Broom 1923. The genotype, D. vanderbyli, is based upon portions of a skull, probably associated. Skull size—this is probably large, not massive. Bosses—no evidence is preserved. Additional pachyostosis—no evidence of this is preserved. Palatine—no evidence of a palatine boss is preserved. Premaxilla—the border does not curve upwards. Incisors—according to Broom, ‘the first is of Deinocephalian type, but the other three incisors may have had pointed crowns. There is some little indication that this may have been so.’ Pineal foramen—this area is not preserved. Snout—this is weathered. Squamosal—this is not preserved. Temporal fossa—this region is not preserved. Intertemporal width—this region is not preserved. Soe ee eae ee ee Ce pes = a rv. TAXONOMIC STATUS OF SOUTH AFRICAN TITANOSUCHIANS 25 12. Dentary—in the associated specimen the dentary is apparently fairly massive with a fairly upright mentum. Ay a | 5 13. Dental formula—according to Broom, 1.3, c.;, p.c.3. Titanosuchus Owen. 1879. The genotype, 7. ferox, is based upon upper and lower jaw fragments showing sections of the teeth roots. The only diagnostic features that can be determined in the genotype are: dentary strong and massive with mentum apparently fairly upright, and dental formula: i3 : C4 s pc. Other forms considered here are: Titanosuchus cloete: (Broom 1903) is based on a piece of massive dentary, with the incisors lacking a lingual crushing surface and with the dental fonmaulas i. 4. °C. 1,0 p.c. 4+. Scapanodon (Broom 1904). ‘The type species is based upon two imperfect, badly preserved jaws showing a series of teeth: 1.2+,c. 1, p.c. 11+. Archaeosuchus (Broom 1905). ‘The type of the type species is a partial maxilla with some teeth; according to Broom c. 1, p.c. 8 (in 1932 Broom gives 7 postcanines). | Lamiasaurus (Watson 1914). Only the snout is considered here. The premaxillary border is not dorsally directed and the dental formula is MMPOrn. (C1, p.c. 3-4. Dinocynodon (Broom 1929). For the type dentary Haughton (1915) states that the symphysis is massive and square, and the dental formula is 1.4, C. I, p.c. 11+, and Broom gives as the only generic character ‘the extreme flattening of the large canine’. Now, in a specimen of Anteo- saurus, there is a canine on the point of being shed which is also flattened, and this character can thus hardly be considered of generic value. Scullya (Broom 1929). The type species is based upon ‘a very badly crushed snout’ with the dental formula i. 5, c. 1, p.c. 12, and the only other characters are the possible presence of teeth on the palatine and the dentary massive. Titanognathus (Broili and Schroder 1935). The type of the type species consists of skull fragments showing, “Schadel mit schmaler und steil vom pramaxillaren Kieferrand aufsteigender Schnauze, sehr gross. Prae- maxillarer Kieferrand gegeniiber dem maxillarer stark in die Hohe gezogen, Symphysenregion des Unterkiefers entsprechend erhoht gegenii- ber dem riickwartigen Abschnitt des Dentale. Zahnformel: 1. Ho Guts p-c: aoe Enobius (Broom 1923). The type of the type species consists of two dentaries; mentum massive and squarish, with the dental formula c. 1, DCI Be pcre From the above it is quite evident that this series of fragmentary ‘types’ affords no very trustworthy bases on which they can be distinguished from 26 ANNALS OF THE SOUTH AFRICAN MUSEUM each other generically. I propose lumping all these forms together and to redefine the genus Tztanosuchus compositely as follows: Cary Bodo Skull size—this is very probably large and massive. Bosses—are probably present. Additional pachyostosis—of this no evidence is preserved. Palatine—this is not preserved. Premaxilla—the dentigerous border curves antero-dorsally with an associated step-up of the anterior part of the alveolar border of the dentary. Incisors—these are apparently long, with no lingual crushing surface. Pineal foramen—no evidence is preserved. Snout—is probably as in Anteosaurus. Squamosal—is not preserved. Temporal fossa—this area is unknown. Intertemporal width—this area is not preserved. . Dentary—this is strong and massive with a squarish and upright mentum. ii el eS 1 (il =154: Dentaliormulas We Gs Ps eee Synonyms: Scapanodon, Archaeosuchus, Lamiasaurus (snout), Dinocynodon, Scullya, Enobius, and Titanognathus. THE SpPEcIFIC NAMES OF THE ABOVE FORMS Until a detailed study is completed it is impossible to make any statement on the validity of the specific names given by authors. Meanwhile a list is appended of the names as they stand at the present moment in the literature, but under the genera—Anteosaurus, Dinartamus, Jonkerta and Titanosuchus—as defined above. Genus Anteosaurus : A. abeli Boonstra 1952. Plates I-V and IX. Type. A good skull and lower jaw. S.A.M. No. 11296. A. magnificus Watson 1921. Type. Major part of skull. B.M. No. 3595. A. minor (Broom) 1929. Type. Fragment of skull. B.M. No. 5742. Referred specimens: S.A.M. No. 11492. A somewhat weathered skull. S.A.M. No. 11694. A good skull. (Plate V1, figs. 1, 2.) Additional diagnostic features revealed by these two skulls are: In S.A.M. No. 11492, the left premaxilla has no incisors preserved, but on the right side there are three incisors, fairly long, with no indication of a lingual step; in S.A.M. No. 11694 no incisors are present, the denti- gerous boss on the palatine is prominent and semilunar in outline; there is no indication of a jugal boss. The dental formula is i. 0-3, c. I, Picai7=e: TAXONOMIC STATUS OF SOUTH AFRICAN TITANOSUCHIANS 27 A. vorstert (Broom) 1936. Type. A good skull. T.M. 265, Broom’s Dinosuchus vorstert. Referred specimen: a good skull. S.A.M. No. 11577. (Plates VII and VIII.) Genus Dinartamus : D. vanderbyli Broom 1923. Type. Skull fragments. Coll. Broom. Genus Jonkeria: J. angusticeps (Broom) 1929. Type. Good lower jaw. A.M.N.H. No. 5633. Broom’s Phoneosuchus angusticeps. F. haughton (Broom) 1939. 3 ‘Type. Fairly good skull. S.A.M. No. 4343. Broom’s Dinosphageus haugh- tont. J. ingens (Broom) 1923. Type. Fair skull. A.M.N.H. No. 5634. Broom’s Dinophoneus ingens. Synonym. 7. pugnax (Broom) 1929. Fairly good skull and lower jaw. A.M.N.H. No. 5608. 7. truculenta van Hoepen 1916. Type. Good skull and lower jaw. T.M. 212. f. vanderbylt (Broom) 1929. Type. Good skull. A.M.N.H. No. 5620. J. spp. The lack of identifiable cranial elements makes the specific status of J. crassus (Broom) 1929, Type A.M.N.H. No. 5577, uncertain. The skull referred to Scapanodon duplessist by Broom 1923 is undoubtedly a Jonkeria species. Further study may prove synonomy with one of the named species and until then it is best left unnamed. Dzénopolus atrox Broom 1923, Type T.M. 274, consists of a snout. The incisors are short, the anterior three with ledge, the outer without ledge, according to Broom. The lower canine peculiar, but otherwise it falls under Jonkeria as here defined. ‘These characters may indicate that it is a distinct species—onkeria atrox Broom, bridging the gap between Dinartamus and Jonkeria. Genus Titanosuchus : T. cloetei (Broom) 1903. Type. Piece of dentary. S.A.M. No. 731. IT. dubius (Haughton) 1915. Type. Major part of dentaries. S.A.M. No. 2759. Broom’s Dinocynodon dubius. T. ferox Owen 1879. Type. Upper and lower jaw fragments. B.M. No. 49370. T. gigas (Broom) 19209. Type. A very badly crushed snout. Coll. Broom. Broom’s Scullya gigas. 28 ANNALS OF THE SOUTH AFRICAN MUSEUM T. lotzi (Broili and Schréder) 1936. Type. Skull fragments. A.K. (no number given). Broili and Schréder’s Titanognathus lotzi. T. strubent (Broom) 1923. Type. Partial dentaries. K.M. (no number given). Broom’s Enobius stru- bent. Incertae sedis. ‘The following forms do not merit separate generic rank and may best be regarded as species of Yitanosuchus, whose specific validity is questionable. T. duplessist (Broom) 1904. Type. Imperfect badly preserved jaws. S.A.M. No. 769. Broom’s Scapano- don duplessist. T. cairncrosst (Broom) 1905. Type. Part of maxilla. S.A.M. No. 916. Broom’s Archaeosuchus cairncrosst. T. newtont (Watson) 1914. Type. Snout. B.M. No. 49385. Watson’s Lamiasaurus newtoni. REFERENCES Boonstra, L. D. ‘A note on the synonomy of the two deinocephalians, Dinophoneus ingens Broom and Jonkeria pugnax (Broom).’ S. Afr. J. Sci., vol. 32, pp. 329-31. 1935. “The cranial morphology of some titanosuchid deinocephalians.’ Bull. Amer. Mus. Nat. Hist., vol. 72, pp. 99-116, pls. 9-17. 1936. Miljoene Jare Gelede in die Karoo. Johannesburg, Voortrekkerpers, 125 pp. 1948. °’n Nuwe Titanosuchiérsoort (Anteosaurus abeli).’ Tydsk. Wet. en Kuns, xu, bls. 142-9. 1952. Broom, R. ‘On evidence of a new species of Titanosuchus (T. cloetei).’ Ann. S. Afr. Mus., vol. 4, pp. 142-3. 1903. ‘Notice of a new fossil reptile (Scapanodon Duplessisi) from the lower Karroo beds of Prince Albert, Cape Colony.’ Rec. Albany Mus., vol. 1, pp. 182-3. 1904. ‘Notice of some new fossil reptiles from the Karroo beds of South Africa.’ ibid., vol. 1, [2]0e ee eh Soe comparison of the Permian reptiles of North America with those of South Africa.’ Bull. Amer. Mus. Nat. Hist., vol. 28, pp. 197-234. 1910. ‘On the structure of the skull in the carnivorous deinocephalian reptiles.’ Proc. Zool. Soc. London, pp. 661-84. 1923. ‘On the carnivorous mammal-like reptiles of the family Titanosuchidae.’ Ann. Transvaal Mus., vol. 13, pp. 9-36, pl. 4. 1929. The mammal-like reptiles of South Africa and the origin of mammals. London, H. G. and G. Witherby, xvi. pp. 376. 1932. ‘On some new genera and species of Karroo fossil reptiles, with notes on some others.’ Ann. Transvaal Mus., vol. 18, pp. 349-86. 1936a. ‘On the structure of the skull in a new type of dinocephalian reptile.’ Proc. Zool. Soc. London, pp. 733-42. 19360. Bro, F., and ScHRODER, J. ‘Ein Dinocephalen-Rest aus den unteren Beaufort-Schichten.’ Sitzber. Bayrischen Akad. Wiss., pp. 93-114. 1935. Grecory, W. K. ‘The skeleton of Moschops capensis Broom, a dinocephalian reptile from the Permian of South Africa.’ Bull. Amer. Mus. Nat. Hist., vol. 56, pp. 179-251, pls. 1-21. 1926. HavucutTon, S. H. ‘On two new therocephalians from the Gouph.’ Ann. S. Afr. Mus., vol. 12, PP- 55-7: 1915. Hoepen, E. C. N. van. ‘A new Karroo reptile.’ Ann. Transvaal Mus., vol. 5, suppl. 3, no. 3, I p. 1916. Owen, R. ‘Description of fragmentary indications of a huge kind of theriodont reptile ( Titano- suchus ferox Ow.) from Beaufort West, Gough Tract, Cape of Good Hope.’ Quart. Four. Geol. Soc., vol. 35, pp. 189-99, pl. 11. 1879. Watson, D. M. S. ‘The Deinocephalia, an order of mammal-like reptiles.’ Proc. Zool. Soc. London, pp: 749-86, pls. 4, 5. 1914. ‘The bases of classification of the Theriodontia.’ ibid., pp. 35-98. 1921. -azis Teanjeu + ynoqe ‘MoIA [ese “Q6SI1 “ON “W'V'S ‘uouttoods odAyT, “vaysuoog 1aqv sninvsoajup snp POW YD) 2 80104 | Plate mn. 5. Afr. Mus., Vol. XLII Peer Cas) Plate u Ann. S. Afr. Mus., Vol. XLII *pasojsaa MUL JAMO] JO oLSuUy Plate yr fr. Mus., Vol. XLII Ama. S. “OZIS jeanjyeu 1 | Node ‘MOIA [R191e'T ‘obSP CoN CIW'V'S ‘9dA1-o1Fr ‘Bsuo0g 773qD SNANDSOIJU’ Plate iv Ann. S. Afr. Mus., Vol. XLII ‘posojsod JNOUS Jo Jed JOIQJUW ‘9ZIS [Vanjeu F ynoqe ‘Mola vszjVT “GSOS1I “ON “WYy'S ‘edAl-o1-) ‘vajsuoog ijaqp snunpsoajuyy Ann. S. Afr. Mus., Vol. XLII Plate v Anteosaurus abeli Boonstra. Co-type. S.A.M. No. 11293. Dorsal view, about } natural size. Anterior part of snout restored. 2 4 Ann. S. Afr. Mus., Vol. XLII Plate 4 } « Anteosaurus minor (Broom). Referred specimen. S.A.M. No. 11694. About } natural size. Fig. 1, lateral view. Fig. 2, ventral view. Plate vu Ann. S. Afr. Mus., Vol. XLII va ‘OZIS [VANVU fF INOGV SMOTIA [V19]U'T ei _ I I ON W V S uguil ) ads PotTIJo YY ‘(wool ) 14948400 SNANDSOAU Ann. S. Afr. Mus., Vol. XLII | aie som | cm Anteosaurus vorsteri (Broom). Referred specimen. S.A.M. No. 11577. Dorsal view, about 4 natural size. Plate 1x Ann. S. Afr. Mus., Vol. XLII (-ojoyd siag ayeuoiseNy) ‘sayeyd SuroSei0j ay} ul poansy sed4j-o9 pure odA} 9y} JO sT[Nys 9y} UO paseq ‘peoy ay} Jo Joyseyd ut uoNoNssuOdaI ozIs-oyt] JO Ydersojoyd “esuoOg 179q) siinDsoajUp 4. The Gorgonopsians, Aelurognathus microdon and Hipposaurus boonstrai, reconstructed. By L. D. Boonstra, D.Sc. (With Plates X—-XVI) In 1934 I described a new species of Gorgonopsian under the name Aelurogna- thus microdon. ‘The specimen consisted of a skull, much of the vertebral column, an excellent pectoral girdle, a good fore-limb with most of the carpus, a good pelvis and most of the hind-limb. (Plate X.) The fragile nature of the preserved bones has made it impossible to attempt a free mount of the skeleton as preserved. During 1937 Mr. J. Drury, then modeller to the South African Museum, modelled in plaster of paris all the elements preserved, two-thirds natural size. Utilizing our knowledge derived from other gorgonopsians species (see op. cit.) the missing bones were restored. The resulting mount could be considered fairly accurate and a photograph of it was published in the ‘Report of the South African Museum for 1937’ and republished in a popular booklet Miljoene Fare Gelede in die Karoo. Apparently both these have passed unnoticed by overseas colleagues, and Colbert in his study of Lycaenops ornatus does not mention either of these publications. An augmented set of new photographs of the modelled skeleton (Plates XI- XIII) and of the animal reconstructed ‘in the flesh’ (Plate XIV), also by Drury, is here presented for comparison with the excellent set of photographs published by Colbert of the remarkable free-mount of Lycaenops ornatus as mounted by Charles Lamb after preparation by Jeremiah Walsh. Aelurognathus microdon has been reconstructed with 29 presacral vertebrae (including the proatlas) measuring 750 mm.; 3 sacrals (90 mm.); 29 caudals (585 mm.). All these measurements are projections. The total projected length of the skeleton is 1,530 mm. and measured over the curvature of the back is 1,650 mm. The shoulder height is 570 mm. and at the hips the height is 368 mm. THe STANCE OF AELUROGNATHUS The following remarks should be read in conjunction with the section in Colbert’s paper, ‘The Skeleton as a Whole’, as, after his masterly account, written with the help derived from Schaeffer’s movie-film of the alligator, I can here be brief and confine myself to the conditions in Aelurognathus without repetition of comparisons with other Therapsids. Drury’s model is mounted showing Aelurognathus in the standing position based on views held in 1937, some of which have since been modified necessita- ting some alterations to the original mount. (Plate XIV.) The curvature of the spine, as mounted, is probably over-accentuated with the apex of the curve too far forward and I now think that in life the back 29 30 ANNALS OF THE SOUTH AFRICAN MUSEUM would have been much straighter with the curve nearer the sacrum. In this I would be in agreement with Colbert. The ribs form a complete presacral set. On this point I had in 1937 corrected the erroneous view expressed in 1934 as to the probable absence of lumbar ribs in Lycaenops, thereby anticipating Colbert’s criticism of 1948 and thus adeeane its validity in advance. The skull hangs downward, dog-fashion, and, in harmony with this, I would now prefer the neck to show a more pronounced downward curve. The pectoral girdle was preserved in approximately natural articulation and ‘this position is retained with little correction in the model. As mounted Aelurognathus has a deep chest—much deeper than Lycaenops as shown in Colbert’s figure 22. In lateral view the scapula has a slight backward tilt when the animal is in the standing position. When executing a stride, the forward side would show the scapula tilting further backwards with its posterior edge everted, i.e. there would be some rotation of the scapula on its long axis. In anterior view the scapular girdle is V-shaped, but, with the scapular blade somewhat curved, the top edge with the cartilaginous suprascapula would not stand excessively away from the ribs. This I believe to be the natural position of the pectoral girdle, for in this position the glenoid articulation would appear to function properly. When executing a stride the left humerus would, in the forward position, be directed somewhat laterally, and the right humerus be directed backwards close in to the body. To keep both the humeri in articula- tion the direction in which the glenoids face must be in keeping. This would be achieved by a lateral sigmoidal curvature of the spine accompanied by a slight movement of the scapular girdle as a whole, and also of its two halves indivi- dually, in relation to the clavicular girdle. Thus the left half of the scapular girdle would move forwards, sag slightly and rotate on its long axis so that the glenoid is directed more outwards and, at the same time, the right half would concomitantly rotate so that the right glenoid is directed slightly inwards from the backwardly facing position. In the mount, with the animal in the standing position, i.e. nearly half-way through the stride, the humerus is directed obliquely outwards with the distal end appreciably lower than the proximal end, and the elbow is thus everted but less anteriorly and downwardly directed than would be the case at the commencement of the stride. The femur, in the 1937 mount, is directed obliquely forward and outward with the distal end much lower than the acetabulum. I now think that the femur was too much everted and that in life the knee would be closer in to the body. This correction has now been made to the mount as shown in the accompanying new photographs—on the right side the femur is brought in closer to the body than the left side. In the stride of the back limbs the spine would be flexed in the lumbar region to complete the sigmoid curve which the movement of the fore-limbs initiated. Plate x Ne See Air. Muss Vol; SIT bh 6 6 ‘ON (IW V'S ‘921s Teanyeu 9 L - jNOGqV uo S od nyis Ut YY ul yno prey UOPOLILUL snyJDUsOINJap jo U0}9] IAS ‘azis Jeanjyeu $ ynoqe ‘MoIA [eJOJV YT “UoposINU SNYDUTOMNJayy JO UOJI[BYS PeT[Ppoyy —_= —_ s vs 2 > Ss a Afr. S. Ann. Plate xu ‘ozis Jeanyeu # ynoqe ‘MOIA [BS 1 LOCL “UOposdiut SNYIDUDOANIIYT jo UO)JI[IAS PYTPP POA AW is, ‘ . & ‘: : . — as as ei ee ad a ae ~AA A eS s Ol, DULIL! Ty / Ann. S. Afr. Mus., V onbiqQ “UOPOLITU SNY TOL a0 njap jo UOJo[I zs P2T[2PPN Plate xin — — : > BS 2) = a a y) eI a Plate x1v ol. XLII Ann. S. Afr. Mus., \ ‘(uunasnyy Uvolapy YING) Aanaqy *[ Aq Ysop otf} Ur, UoposnU SNYJVUTOANJAP JO |PPOW Plate xv Ann. S. Afr. Mus., Vol. XLII ‘ozIs [eanyeu ? ynoqe ojoyg “sJoyNe oY} Aq stred-jo-19}se]d ur ozIs [enyeuU poy[apour wysu00g sninvsodquyy Plate xvt Wolk OOIUL cf) Ann. S. Afr. Mus uno f, ado’ ‘slag apouoispyy aq sYydvadojoyd I]t uoog § ‘UOTONASUOIIT JIAO IU} JO MolA onbyqo WwW. Ws nN ia / imosodquyy as i i S ————_— AELUROGNATHUS MICRODON AND HIPPOSAURUS BOONSTRAI 31 In the reconstruction the feet are probably mounted slightly too much in the semi-digitigrade position. They should, in the standing position, be more plantigrade. At the commencement of the stride they would be wholly planti- grade, and, at the completion of the stride, the digitigrade position would be obtained just before the foot is lifted from the ground. The fore-feet are correctly shown as forwardly directed, and, with the knees drawn in to the body in the corrected mount, the hind-feet also assume a more natural forward position than was the case in the 1937 mount. For comparison I have included here some hitherto unpublished photographs of a life-sized reconstruction of Hipposaurus boonstrat which I have myself modelled in plaster, with due acknowledgment to Colbert for ideas derived from his Lycaenops paper. (Plates XV and XVI.) Eipposaurus is here represented in the walking position and the greater straightness of the back is noteworthy. The right fore-foot, at the commence- ment of the stride, is flat on the ground; the left, nearly at the completion of the move, is semi-digitigrade. The left hind-foot, just after the commencement of the stride, is semi-plantigrade with the knee probably everted a little too far; the right hind-foot is just about to be lifted and swung forwards after having completed the stride. REFERENCES BoonstrA, L. D. ‘A Contribution to the Morphology of the Gorgonopsia.’ Ann. S. Afr. Mus. 31, P- 137- 1934. Report of the South African Museum for 1937, p. 9 and plate facing p. 18. 1938. Muljoene Fare Gelede in die Karoo. Voortrekkerpers, Johannesburg. p. 51. 1948. ‘Bewoners van die Oer-Karoo’, Die Huisgenoot, Cape Town, 6 Aug., p. 23. 1948. “‘Wereld van 250 Miljoen Jaar Gelede.’ Die Burger, Cape Town, 11 May, p. 3. 1950. Bystrow, A. P. ‘Rekonstuktionversuche einiger Vertreter der Nord-Dwina Fauna.’ Akad. Nauck, U.S.S.R., 4, p. 289. 1935. Co.Bert, E. H. “The Mammal-like Reptile, Lycaenops.’ Bull. Am. Mus. Nat. Hist., 89, 6, p. 357. 1948. 5. The cranial morphology and taxonomy of the Tapinocephalid genus Struthiocephalus. By L. D. Boonstra, D.Sc. (With Plate XVII and 6 text-figures) The one important genus of the Tapinocephalid Deinocephalians! of which the cranial structure has hitherto not been adequately described is Struthio- cephalus. Up to date five species of Struthiocephalus have been created. Few details of the cranial structure were published before I re-examined the type skull? and described the two new species, duplessisi® (since transferred to the new genus Struthtocephaloides*) and akraalensis.4 Of Broom’s rheedert® and Olson and Broom’s milleri® little more is known than can be gathered from Broom’s sketches of the outlines of the skull and the main skull openings—temporal fossae, orbits, nostrils and the pineal foramen. In the collection of the South African Museum there is, besides the type skulls of whatis: (S.A.M. 2678), and akraalensis (S.A.M. 3719), an excellently preserved skull of the species whaitst (S.A.M. 11591) showing nearly all the features of the dorsal, lateral, occipital and palatal surfaces and the right side of the brain-case, and a juvenile skull (S.A.M. 11493) of the same species. The skull of the mounted specimen which has associated with it much of the postcranial skeleton (S.A.M. 3012) is crushed and shows little of the detailed structure.’ The structural details determined in these five specimens are here pooled to give an account of the cranial morphology of the genus but the figures are all of the one specimen of whatisi (S.A.M. 11591). THE GENERAL SHAPE AND FORM The generic name very aptly describes the shape of the skull as being very similar to the shape of the head of an ostrich. Anterior to the orbital border the snout is anteriorly directed, long, shallow but fairly broad and, in relation to the part of the skull posterior to the orbits, appears weak. With the lower jaw occluded the muzzle appears much less weak as the mentum is quite massive. The skull is low—the width over the quadratojugals being much greater than the height in this plane. The bones of the posterior part of the skull are all strongly pachyostosed, whereas those of the snout are only relatively moderately thickened with the outer bone surface smooth. The transition from the posterior part of the dorsal surface to the dorsal and lateral surface of the snout is thus abrupt but less abrupt than in Mormosaurus. As the strong pachyostosis does not include the lacrimal and only affects the posterior part of the prefrontal the transition from the smooth snout to the rough posterior part lies further back than in Taurocephalus and Mormosaurus. The pachyostosis in Struthiocephalus is not general over the posterior part of the outer surface—the ‘cheek’ being still 32 CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 33 fairly light and its surface smooth, and on the dorsal surface the strong thicken- ing is still localized, emanating from distinct centres. The pachyostosis forms a strong rugose, dorsally somewhat bulbous, postorbital bar; a prominent rugose boss surrounding the pineal foramen and a peculiar naso-frontal boss. (The first is reminiscent of the condition in the Titanosuchid Anteosaurus, _where it is very strongly developed, and the last a feature which it has in common with the Tapinocephalid Keratocephalus.) The orbits, situated in the posterior half of the skull, are large, round and directed forwards and outwards with the thickened postero-dorsal half of the orbital border strongly overhanging. The nostrils are large and elongated, situated on the dorsal surface mainly dorsally, and only slightly laterally directed and are well back from the anterior edge of the snout, close to each other and separated by a strong internasal bar. The temporal fossa is fairly large, higher than long, and the pachyostosis of the postorbital bar and the posttemporal arch has not reduced its size much; its anterio-posterior diameter is still greater than in Mormosaurus and much greater than in the slitlike fossa of Tapznocephalus. Dorsally, it extends medially to form a bay encroaching into the parietal region (in akraalensis the fossa approaches the condition in Mormosaurus). The interparietal width is moderate, due to a pinching-in laterally of the parietals to form the dorso-median bay of the temporal fossa. Due to the forward position of the quadrate, which lies anterior to the plane of the orbit, the lower jaw is short and the maximum gape of the jaws is com- paratively small. : The anterior teeth of the upper jaw are directed much anteriorly but the intermeshing teeth of the dentary are directed dorsally. THE BONES OF THE DoRSAL AND LATERAL SURFACES (figs. I and 2) The matrix of the Tapinocephalus-zone being notoriously intractable, the determination of sutures in most specimens from this horizon is extremely difficult. ‘Thus Haughton in his description of S. whaitst and Broom in the case of S. rheedert and Olson and Broom in 'S. millert have, together, only figured parts of two sutures. It has only been through a laborious process of ‘artificial weathering’ by dilute hydrochloric acid that I have been able to determine most of the sutures, but some still remain indeterminable and others uncertain. The premaxillaries (P.Mx.) together form a large part of the snout. From the anterior border they stretch posteriorly to past the middle of the skull. From its anterior border each premaxilla narrows, where it forms the inner border of the nostril, then it stretches as a long tapering bone posteriorly, where it lies in a groove of the nasals. The nasals (N.) are long, narrow bones, which in their postero-median part are grooved to house the posterior tongue of the premaxillaries; posterior to the limits of the premaxillaries they meet on the dorsal surface in the median 34 ANNALS OF THE SOUTH AFRICAN MUSEUM line and are here thickened to form the anterior and major part of the naso- frontal boss. The septomaxillaries (S.Mx.) appear to be small splint-like bones forming the outer border of the nostrils, but their limits are uncertain in most specimens. The maxillaries (Mx.) are the largest bones of the snout, being long but shallow. Posteriorly a dorsal prong just meets the prefrontal, but in some speci- mens the lacrimal is intercalated, and a ventral prong extends far posteriorly with its upper edge applied to the lower border of the jugal. In between these two prongs lie the anterior ends of the lacrimal and jugal. Fic. 1. Struthiocephalus whatisi. Lateral view of the skull (S.A.M. 11591) (x4). All the figures are not perspective drawings but pro- jections drawn with the aid of a pantograph. The lateral view of the skull and that of the braincase are projected on to the median (sagittal) plane, the dorsal and ventral views are projected on to the plane of the alveolar border, and the occipital view at right angles to the plane of the alveolar border. The lacrimal (L.) ventrally meets the upper edge of the jugal in a straight suture and these two bones together extend anteriorly into the posterior fork of the maxilla. The lacrimal is in its anterior extent as in Mormosaurus and thus stretches much further anteriorly than it does in Taurocephalus. It forms only a small part of the relatively unthickened anterior orbital border. Dorsally the prefrontal does not, in S.A.M. 11591, altogether exclude the lacrimal from contact with the nasal; in the other specimens a tongue of the prefrontal is intercalated between the nasal and lacrimal. (In Taurocephalus the lacrimal does not meet the nasal, but in Mormosaurus it does.) The jugal (J.) is a strong bone; like the bones of the snout it is not greatly thickened and its surface is not rugose but smooth. It forms the antero-ventral comparatively unthickened border of the orbit. It extends far ventrally as a tapering element to be separated (in S.A.M. 11591) from the quadratojugal by a narrow incisure. Anteriorly it stretches far as a prolongation, with the lacrimal, into the posterior fork of the maxilla. The posterior border forms a shallow curve and is thus not deeply indented by an anterior wedge of the squamosal as in Mormosaurus and Taurocephalus. CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 35 The prefrontal (Pr.F.) is much thickened along its lateral edge to form the antero-dorsal thick, rugose and rounded portion of the orbital border. In all the specimens, except S.A.M. 11591, it has an anterior tongue which separates the lacrimal from the nasal. It does not extend much posteriorly, but is thickest here. Medially it is thinner and there is thus a hollow between the thickened orbital border and the naso-frontal boss. The frontal (F.) is a large element of the dorsal skull roof. Its median part is roughly rectangular and from here three tongues extend: one, anteriorly inter- Fic. 2.—Struthiocephalus whaitsi. Dorsal view of the skull (S.A.M. 11591) (x4). calated between the nasal and prefrontal; the second, entering the thickened rugose upper orbital border and, the third, is a wedge between the postfrontal and the parietal. Anteriorly, the frontals meet the nasals and here form the posterior minor part of the naso-frontal boss. For the rest the frontals are not greatly pachyostosed and in whaitsi form a shallow saddle between the naso- frontal and parietal bosses, whereas in akraalensis the frontals are thickened in the median line with laterally a deep depression which is further laterally bounded by a strong ridge formed by the postfrontal and postorbital. Pos- teriorly, the frontals meet the parietals in a nearly straight frontal suture just anterior to the parietal boss surrounding the pineal foramen. 26 ANNALS OF THE SOUTH AFRICAN MUSEUM The postfrontal (Po.F.) forms the dorso-posterior corner of the orbital border which is here greatly thickened and very rugose. Its suture with the postorbital is in Most specimens uncertain but in whazts: (S.A.M. 11591) it is a large bone forming the dorsal swollen part of the postorbital bar, and in akraalensis it forms part of the ridge lying lateral to the depression in the surface of the frontal. This identification of the large size of the postfrontal in Struthiocephalus leads one to suspect that this element is a much larger bone than it has hitherto been thought to be in most Tapinocephalia. The pachyostosis of the postfrontal has resulted in the postorbital being practically excluded from the dorsal surface of the skull and thus forms only the lower part of the postorbital bar. Another result of the thickening of the postfrontal has been that together with the enlarged and thickened prefrontal the frontal tends to become excluded from the orbital border. The parietals (P.) together form the greater part of the cranial roof. In their antero-median portion a large elevated and rugose boss is developed and is pierced by a large round pineal foramen. In their posterior half the parietals form a narrowed dorsal surface as they are here laterally pinched in. This pinching-in is less evident in akraalensis. Here a sharp edge separates the dorsal surface from a lateral surface, which forms the dorsal part of the median or inner face of the temporal fossa. ‘This lateral parietal surface, extending on to the posttemporal arch to meet the squamosal, effectively prevents the post- orbital from meeting the squamosal at this level. The medio-dorsal bay of the temporal fossa thus formed by the pinching-in of the parietals laterally is clearly shown in whaitsi, but in akraalensis the temporal fossa is more a continuous slit with this bay not clearly demarcated. Posteriorly, the parietals are buttressed by the interparietal in their median part and, more laterally, by the strong tabulars. The postorbital (P.O.) is a massive element forming the lower part of the thickened postorbital bar which in its upper postfrontal part is bulbous on a scale just less than in the Titanosuchid Anteosaurus. ‘The postorbital bar 1s strong, wide and fairly rugose in whaitsi, and very strong, very wide and strongly rugose in akraalensis. Posteriorly, the postero-lateral flange of the parietal lies between the posterior process of the postorbital and the squamosal. In a juvenile specimen of whaits1, S.A.M. 11493, the posterior process of the post- orbital still stretches far posteriorly, but even here does not meet the squamosal. Ventrally, the postorbital forms an overlapping suture with the squamosal. The squamosal (Sq.) is the main constituent bone of the ‘cheek’. It is a strong thickened element but, as in the jugal, its outer surface is not rugose but smooth in whaztsi, but with pits and rugae in akraalensis. Anteriorly, it meets the jugal in a long curved suture with no anterior wedge-shaped process as in Mormosaurus and Taurocephalus. Its postero-ventral corner overlaps on the outer surface of the quadratojugal. Dorsally, it is overlapped by the ventral edge of the postorbital, and, further, posteriorly, it forms the thickened lower border of the temporal fossa. From here it sweeps upwards to form most of the anterior CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 37 face of the posttemporal arch and here its dorsal end overlaps the postero- lateral flange of the parietal, where this bone forms the inner upper margin of the temporal fossa. Postero-ventrally, the squamosal forms the thickened rounded postero-ventral edge of the skull. This rounded border forms the lateral wall of the wide and deep auditory groove, which groove lies mostly in the squamosal. Medially to this groove, the squamosal forms a strong and prominent ridge, medio-ventrally buttressed by the paroccipital and further dorsally wholly formed by the tabular. This ridge, thus composed of squamosal, tabular and paroccipital, forms the median wall of the auditory groove and from it originated the strong depressor muscle of the mandible. The tabular (Tab.) in dorsal view is seen to form the posterior half of the dorsal part of the posttemporal arch, supporting the anterior half formed by the flange of the parietal and the up-sweeping flange of the squamosal. In lateral view, the tabular is seen to form the lateral part of the posterior edge of the skull. The interparietal (I.P.) in dorsal view, shows its upper edge where it forms the posterior buttress to the parietal in the median part of the posterior margin of the skull. The quadratojugal (Q.J.) in dorsal and lateral views is seen to form the antero-ventral corner of the ‘cheek’. Along its posterior border it is clasped by the squamosal and its inner surface supports the quadrate. Its dorsal margin does not abut against the jugal but is separated from it by a narrow incisure. THE OccrpuT (fig. 3) The occiput in Struthiocephalus forms a large surface, much broader than high and nearly semicircular in outline. It is shallowly concave from side to side. In the median line it is nearly ver- tical with its dorsal edge slightly further posteriorly and here it lies in a plane nearly at right angles to the plane of the maxillary alveolar bor- der. In the median line there is a ridge, which runs from the foramen magnum to the upper edge of the occipital surface. In whats: this ridge is wedge-shaped, broad dor- sally and tapering to the upper edge of the foramen magnum, a ee an eM eo whereas in akraalensis this ridge is straight and narrow with a sharp edge forming the median line with a deep depression lateral to it. In this species the occiput has its ventral part situated much anteriorly so that it is no longer nearly at right angles to the alveolar border but forms an obtuse angle with the alveolar plane and an acute angle with the dorsal surface. This may be due to post-mortem dorso-ventral pressure. 38 ANNALS OF THE SOUTH AFRICAN MUSEUM The condyle is directed postero-ventrally so that the skull would normally hang somewhat downwards. It forms a stout rounded knob, dorsally excavated by a groove leading into the foramen magnum. ‘The foramen magnum is large and oval. The posttemporal fossae are small slits, bounded dorsally by the supra- occipital and ventrally by the paroccipital. The lateral outer border of the occipital plate is formed by the squamosal and median to this lies the deep auditory groove, whose inner wall is formed by a strong and prominent ridge to whose formation the tabular, squamosal and paroccipital contribute. Ven- trally the condyles of the quadrates lie far anteriorly to the plane of the occiput. In occipital view the basioccipital condyle forms the median part of the ventral edge, and laterally the quadrate rami of the pterygoids together form a third of the ventral edge of the skull. In only one specimen (S.A.M. 11591) is the occiput well preserved, and even here the sutures between the basioccipital and exoccipital and between the supraoccipital and interparietal cannot be determined. : The basioccipital (B.Oc.) apparently forms the whole of the condyle with no participation by the exoccipital, which appears to be a small element lying dorso-laterally in a plane anterior to that of the condyle. A groove on the dorsal surface of the basioccipital leads into the foramen magnum. A notochordal pit lies in the centre of a shallow concavity in the postero-ventral surface of the condyle. The supraoccipital (S.Oc.) appears to form the major part of the median portion of the occipital plate. Laterally it stretches to the inner base of the prominent ridge composed of the tabular, squamosal and paroccipital and forming the inner rampart of the auditory groove, and here it meets the tabular in a long vertical suture. In its ventro-lateral part the supraoccipital forms the upper border of the slit-like posttemporal fossa. Medially and laterally to the posttemporal fossa it meets the paroccipital and enters the upper edge of the jugular foramen. Dorsally its junction with the interparietal cannot be deter- mined, but with this bone it forms the median occipital ridge to which it contributes the lower, narrower part. The interparietal (I.P.) forms the dorso-median part of the occipit but it is uncertain how far it stretches ventrally before meeting the supraoccipital. It forms the dorsal part of the median ridge, which, in whaitst, is broad in its inter- parietal part but in akraalensis is sharp and narrow throughout. The tabular (Tab.) as I have determined its limits, forms only a small part of the occipital surface as it appears not to stretch far medially. It forms the greater part of the prominent lateral ridge lying medially to the auditory groove. Dorsally the tabular flanks the postero-lateral parietal flange and the upsweeping dorsal squamosal flange and thus forms the postero-dorsal part of the temporal arch. This part of the tabular is especially strong in akraalensis, and here its dorsal edge is wide, strong and rugose. The paroccipital (P.Oc.) in occipital view is seen to be a strong bar medially abutting against the basioccipital and stretching laterally to the lateral ridge CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 39 where its dorso-lateral corner forms the most prominent part of this ridge. Its ventro-lateral edge overlaps and supports the quadrate and dorsally it meets the supraoccipital and forms the lower border of the small slit-like posttemporal fossa and that of the small foramen jugale. Medially to the jugular foramen it meets the ventral edge of the exoccipital. The quadrate (Q) in occipital view shows a squarish posterior surface with, ventrally, paired strong rounded knobs separated by a broad groove together forming the ginglymoid articulatory surface for the articular. Laterally the quadrate is overlapped by the squamosal and flanked by the quadratojugal. Dorsally the posterior face of the quadrate is overlapped by the paroccipital. Medially the long quadrate ramus of the pterygoid is applied to its inner surface ventral to where the expanded distal end of the stapes abuts against the quadrate. The quadratojugal (Q.J.) in posterior view is seen to form the latero-ventral corner of the skull. Dorsally its posterior surface is overlapped by the squamosal. The stapes (St.) is only partly exposed in occipital view. It is a stout rod with its proximal end obscured by the paroccipital and its distal expanded end is seen to be applied to the inner face of the quadrate. The pterygoid (Pt.) only shows its long quadrate ramus in occipital view. This is seen to extend very far posteriorly with its dorsal edge overlapping the distal end of the stapes and its postero-lateral end applied to the inner face of the quadrate. | If my interpretation of the relations of the interparietal, supraoccipital, tabular and paroccipital is correct the structure of the occiput in Struthiocephalus differs greatly from the condition in Mormosaurus, Taurocephalus, Tapinocephalus and Moschops. In these forms the tabular has a much greater occipital surface and the supraoccipital is a much smaller bone. In Taurocephalus and Moschops the tabular forms the whole of the dorsal border of the posttemporal fossa, but in Tapinocephalus the tabular, as in Struthiocephalus, is excluded from the post- temporal fossa. THE VENTRAL SURFACE OF THE SKULL (fig. 4) The palate and the basis crani lie in the same plane, with the strong lateral pterygoidal rami extending ventral to this plane and the suspensorium lying still further ventrally. Striking is the very anterior position of the articulatory surfaces of the quadrates, which lie anterior to the posterior third of the skull. The suborbital fossae are small, the choanae large and oval and the interpterygoid vacuity is a narrow slit not extending between the prevomers. The basioccipital (B.Oc.), in ventral view, is seen to carry a strong condyle pear-shaped in outline. Postero-ventrally the condyle is circularly excavated round the notochordal pit. Anteriorly to the condyle the basioccipital forms a squarish plate of bone directed antero-ventrally to meet the surface of the basisphenoid at an obtuse angle in a not very secure ankylosis. This surface carries a low median ridge flanked by shallow oval depressions and the anterior and lateral edges are rounded. Laterally the basioccipital is flanked by the 40 ANNALS OF THE SOUTH AFRICAN MUSEUM small exoccipital, whose limits are uncertain. Anterior to the jugular foramen the basioccipital is strongly supported by the paroccipital which it meets in a firm curved suture. The basisphenoid (B.Sph.) posteriorly meets the basioccipital at an angle so that the two ventral surfaces subtend an obtuse angle. The postero-lateral corner of the basisphenoid forms the anterior border of the foramen ovale. Anteriorly the basisphenoid extends as a blunt wedge in between the pterygoids but the exact position of the suture is uncertain and its probable position is eceywe Fic. 4.—Struthiocephalus whaitsi. Ventral view of the skull (S.A.M. 11591) (x §). given in broken lines. In the median line the basisphenoid carries a low, sharp keel, lateral to whose anterior end lie the carotid foramina. The pterygoids (Pt.) form a large part of the ventral surface. In the middle of the skull each pterygoid consists of a thin plate of bone meeting the basi- sphenoid obliquely along a long edge whose exact position has not been deter- mined but which I believe stretches in anterior direction to the median line from the notch leading into the pituitary fossa. In the median line the pterygoid meets its fellow to form a sharp median keel. Anterior to this keel lies the inter- pterygoid slit. Lateral to the median keel the pterygoid is deeply excavated and lateral to this wide and deep groove lies the quadrate ramus of the ptery- goid. This is a deep sheet of bone lying obliquely in the skull nearly at right CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 41 angles to the plane of the palate. The quadrate ramus extends far posteriorly and here its outer surface is applied to the inner face of the quadrate, well behind the plane of the condyle of the quadrate, and there is thus no anterior process of the quadrate to meet the pterygoid. The distal end of the stapes thus passes over the upper edge of the quadrate ramus of the pterygoid to reach the medial face of the quadrate. In no other Deinocephalian is the quadrate ramus of the pterygoid known to extend so far posterior to the quadrate condyle, which fact emphasizes how far the quadrate has shifted in an anterior direction. The lateral ramus of the pterygoid is only moderately strong and has no great ventral extent. This is in strong contrast to the condition in the Titano- suchids where the ramus is very strongly developed. In Struthiocephalus the ramus does not extend far laterally as it does in Taurocephalus and all the ‘Titanosuchids. The lateral edge of the ramus is supported by a descending process of the ecto- pterygoid, which, on account of the narrowness of the lateral pterygoid ramus, has a greater palatal face than in all other Deinocephalians. As in all Deino- cephalians the lateral ramus is connected with the quadrate ramus by a web of bone reducing the size of the suborbital vacuity of which it forms the antero- median border. Anteriorly the exact limits of the pterygoid are indetermined but are probably as indicated by broken lines in the figures. The ectopterygoid (Ec.Pt.) has a larger palatal surface than in other Deino- cephalians. Its anterior and median limits are not clearly shown but it appears to descend along the lateral edge of the lateral pterygoid ramus which it buttresses. Its posterior edge forms the antero-lateral border of the suborbital fossa. Postero-laterally it abuts against the jugal in a sigmoid suture and laterally against the maxilla. The palatine (Pal.), from where the suture with the pterygoid and ecto- pterygoid appears to lie, stretches antero-laterally as a thickened bone to form the rounded lateral two-thirds of the choanal border, and has its lateral edge applied to the inner maxillary surface where it flanks the alveolar border. Near the median line the palatine ends with a short sharp ridge running parallel to the median line. The two palatines thus do not meet each other, as median to their inner ridged borders a posterior tongue of the prevomers intervenes. Just lateral to the ridged inner edge there lies a rounded mound on which there are indications of the roots of a small number of small palatine teeth. The prevomers (P.V.) are strong elements together forming a massive inter- choanal bar. Anteriorly they underlie the inner surface of the premaxillaries with their anterior edges bevelled. Posteriorly they widen and overlie the palatines postero-laterally and in the median line send a tongue posteriorly in between the ridged inner edges of the palatines. Anteriorly the median suture is open and the interchoanal bar is here grooved, whereas in the posterior half of the bar a keel is developed along the median line. The premaxillary (P.Mx.) alveolar border is very massive. Each premaxilla carries three strong teeth which are directed antero-ventrally. Posterior to the functional teeth there are indications of crowns which may be either replacing 42 ANNALS OF THE SOUTH AFRICAN MUSEUM or replaced teeth. In a juvenile specimen of whaitsi (S.A.M. 11493) where the crowns of the teeth are just erupting the labial edge of the premaxillaries is sharp and appears to form a cutting edge functioning as such until the teeth are sufficiently developed. . The maxilla (Mx.) has its alveolar border anteriorly massive and wide but then it tapers rapidly in posterior direction, and behind the last tooth a sharp edge is continued by the jugal, sweeping down towards the quadrato-jugal. Stumps of teeth and infilled alveoli in most specimens show that there were ten to eleven maxillary teeth. The anterior four are large, the fifth appreciably smaller, the sixth abruptly smaller and the series then decreases gradually in size in posterior direction. In the juvenile specimen of whaits: (S.A.M. 11493) the labial edge of the maxilla, as is the case in the premaxilla, is sharp and during immaturity apparently forms a cutting edge. The jugal (J) in ventral view has a narrow and deep flange of bone sweeping from the sharp outer edge of the maxilla down towards the quadratojugal. Internally and dorsally to this sharp edge the jugal is thickened and extending internally forms the lateral border of the suborbital fossa, and anteriorly it meets the ectopterygoid in a sigmoid suture. The quadrate (Q.) has its articulatory condyle very prominent in ventral view as this forms the most ventral part of the skull. The articulatory surface has a median trochlear surface bounded internally and externally by longitudinally oval condyles. ‘The rounded articulatory surfaces are sharply demarcated from both the posterior and the anterior face of the upper part of the bone. Laterally a ridge bounded medially and laterally by a groove lies lateral to the condyle in the plane of the ventral edge of the quadratojugal. Dorsally the posterior surface of the quadrate is seen wedged in between the paroccipital internally and the quadratojugal and squamosal externally. Internally the long quadrate ramus of the pterygoid is applied to the medio-posterior face of the quadrate. The extremity of the ramus extends far posteriorly of the plane of the quadrate condyle and there is no anterior process of the quadrate to meet the quadrate ramus of the pterygoid. No foramen quadrati could be located with certainty but it may be small and may lie just medio-dorsally of the ridge on the quadrate lying laterally to the condyle. The stapes (St.) is in position in S.A.M. 11591 on both sides. It is a stout rod with expanded distal and proximal ends. The distal end applied to the inner face of the quadrate is underlain by the long quadrate ramus of the pterygoid and has its postero-distal corner much expanded. No foramen can be determined. The quadratojugal (Q.J.) in ventral view is seen to form the angle of the ‘cheek’. Its ventral edge lies in a plane dorsal to the condyle of the quadrate from which it is separated by a ridge bounded on both sides by a groove. The posterior surface of the quadratojugal is seen to be overlapped by the descending posterior squamosal process. CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 43 The squamosal (Sq.) in ventral view forms the outer edge of the posterior third of the skull. Median to this edge lies the auditory groove whose inner wall is formed by a ridge formed by the squamosal, paroccipital and tabular. Further ventrally the squamosal supports the posterior face of the quadratojugal and quadrate. The paroccipital (P.Oc.) shows a much greater face in ventral view than it does in occipital view. This rotation of the paroccipital from the occipital plane on to the ventral surface is one of the results of the forward shift of the quadrates and is a character distinguishing all Deinocephalians from the other Therapsids. Fic. 5.—Struthiocephalus whaitsi. The right side of the braincase in lateral view (S.A.M. 11591) (x4). The suborbital bar, postorbital bar and occipital plate fractured and here seen in section. The right quadrate ramus of the pterygoid and the right stapes are also seen in section where they have been fractured. The paroccipital is a strong massive element acting as a very firm connecting link between the bones of the ‘cheek’ and the suspensorium lying laterally and the basioccipital of the cranial base. Posteriorly the thickening of the paroccipi- tal and supraoccipital have all but obliterated the posttemporal fossa which is only preserved as a narrow slit. The postero-lateral corner of the paroccipital forms the most prominent part of the strong ridge from which the depressor mandibulae originates. Antero-medially the paroccipital forms the posterior half of the border of the foramen ovale. In ventral view the tabulars, interparietal and supraoccipitals are seen to lie well posterior to the basioccipital condyle with the first two forming the posterior edge of the skull which, as is the case in most Tapinocephalids, is nearly a straight line, whereas in the Titanosuchids this edge is concave. In akraalensis more of the occiput is seen in ventral view than is the case in whaitst. THE BRAINCASE IN LATERAL VIEW (fig. 5) In a specimen of whaitsi (S.A.M. 11591) a fracture through the posttemporal arch, postorbital and suborbital bars has enabled me to prepare the lateral surface of the braincase on the right side. But after the removal of the intractable 44 ANNALS OF THE SOUTH AFRICAN MUSEUM matrix from the temporal fossa and orbit the surface of the internal bones thus exposed is not sufficiently clear so as to determine the limits of the constituent bones with any great degree of certainty. The accompanying figure shows what structural details have been determined. The parasphenoid (P.S.) is the largest element forming a large part of the fenestrated septum. A dorsal sheet of bone supports the ventral edge of the sphenethmoid and a well-developed anterior process is directed obliquely forwards. Of this part of the parasphenoid Efremof says that in many cases, ‘den vordere Forsatz des Parasphenoid knorpelig blieb’, whereas in fact the parasphenoid is an os investitiens and thus not an element preformed in cartilage. Of the sphenethmoid (S.E.) only the lower part can be seen where it rests on the upper edge of the dorsal parasphenoidal process. The prootic (P.O.) is seen wedged in the postero-dorsal corner and its rela- tions with the sphenethmoid are uncertain. The opening for the trigeminus and the fenestra of the fossa hypophyseos are situated as shown in the figure. THE Lower JAw In S.A.M. 11693 most of the dentaries are preserved and in 8.A.M. 11493 the crushed posterior half of the right mandibulary ramus is present, but in both only the outer surface could be prepared. What could be determined of the structure I have included in the composite figure accompanying the description of Struthtocephaloides duplessist. The hinge of the lower jaw lies very far forward, in the plane of the orbit. The dentary forms nearly two-thirds of the ramus and its mentum is massive and fairly upright. The teeth are directed upwards and only slightly outwards to intermesh with the labially directed teeth of the upper jaw. The angular has a large outer flange and the surangular has a strong rounded dorsal border curving upwards and forwards from the articular. THE DeEntTiTION (fig. 6) In all specimens stumps, imperfect crowns and empty alveoli are all that are preserved. In fragments of the Fic. 6.—Struthiocepha- lower jaw of S.A.M. 11591 a few crowns of the teeth lus whaitsi. A tooth i : of the lower jaw of the lower jaw are preserved and one is figured here. (S.A.M. 11591) All the teeth have a labial talon or pointed cusp at (x4). a, lingual } i : view. 6, lateral view. whose base there lies lingually a cup-shaped base with a serrated edge. The three teeth in the premaxilla and the first four maxillary teeth are large with long labial cusps. Then abruptly the fifth tooth is much smaller with a short labial talon and then the rest of the teeth decrease still further in size posteriorly. CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 45 A tooth just erupting in the lower jaw (S.A.M. 11591), which is thus as yet unabraded, shows three longitudinal ridges on the lingual surface of the talon. With use the talon is abraded and the ridges disappear. Through use the serrations on the labial edge of the cup are also worn down and lost. There are indications that in the anterior teeth there can be at least three successive sets of teeth. TAXONOMIC DIAGNOSES As Mormosaurids I have grouped together (1936) the genera Mormosaurus, Struthiocephalus and Taurocephalus. The following amended diagnosis for this group is suggested: Skull large, long and moderately wide; snout long or fairly long and shallow; cranial bones strongly thickened with parietal, naso- frontal and postorbital bosses undeveloped, distinctly developed or tending to coalesce in the general pachyostosis; postorbital bar moderately wide to wide and massive; the facial surface fairly or very abruptly demarcated from the thickened cranial surface; temporal fossae fairly small with the dorso-ventral diameter appreciably or not very much greater than the antero-posterior; intertemporal region fairly narrow to moderately wide and laterally distinctly or only slightly bayed; parietals entering supratemporal border; quadrate ramus of the pterygoid extending only up to or well posterior to the quadratic condyle, which is situated far forward; tabular entering or not entering the post- temporal fossa; no differentiation of teeth into incisors, canines and postcanines. GENERIC DIAGNOSES A. Mormosaurus 1. Snout short, very shallow, facial surface not extending posterior to the anterior orbital border, very abrupt transition from the facial to the cranial surface. 2. Dorsal cranial surface very strongly pachyostosed, with the centres of thickening coalesced and transition on to face very abrupt along a very definite transverse line forming a transverse wall. 3. Additional pachyostosis: a. Nasal boss laterally confluent with the thickening of the postorbital bar and the prefrontal and thus forming a transverse wall from orbit to orbit. 6. Parietal boss not very distinct, confluent with the general machyostasts of the parietal and frontal. c. Postorbital bar wide and massive but without distinct bulbous boss and thus flowing evenly on to the general dorsally thickened surface. d. Posttemporal arch greatly thickened and rugose. e. Orbits not visible in dorsal view and in dorsal view the postorbitals do not form the lateral border of the skull. f. Antero-posterior diameter of temporal fossa small and fossa transversely oval. g. Dorsal parietal surface fairly wide, laterally indistinctly bayed. 46 = oo ANNALS OF THE SOUTH AFRICAN MUSEUM Dentition feeble, uniform. Quadrate moderately far forward with the quadrate ramus of the pterygoid not extending much posterior to the plane of the quadratic condyle, not underlying the distal end of the stapes. ; The lateral ramus of the pterygoid does not form a prominent transverse bar and the width across the transverse rami is small. The intersquamosal width is large. The basioccipital condyle is directed posteriorly. The frontal appears to be small and is excluded from the orbital border, the supraoccipital is wide and low, the tabular with a moderate occipital face and apparently enters the posttemporal fossa, the lacrimal meets the nasal, the squamosal with an anterior wedge into the jugal. (This diagnosis is based on Watson’s description and figures.) B. Struthiocephalus Snout long, shallow, facial surface extending far back, posterior to the anterior orbital border, fairly abrupt transition on to the cranial surface. Dorsal cranial surface strongly pachyostosed, but centres of greatest thicken- ing still distinct and the transition on to the face fairly abrupt but not along a definite line and thus not forming a transverse wall. Additional pachyostosis : a. Naso-frontal boss not laterally confluent with the pachyostosis of the postorbital bar and the prefrontal and thus not forming a transverse wall from orbit to orbit. b. Parietal boss distinct. Postorbital bar moderately wide to wide, fairly massive to massive, with or without distinct bulbous boss and with or without a ridge de aneotne it from the dorsal surface. d. Posttemporal arch moderately to shea thickened, fairly smooth to rugose. e. Orbits just visible in dorsal view, and in dorsal view the postorbitals do not form the lateral border of the skull. f. Antero-posterior diameter of temporal fossa moderate to fairly small and fossa oval to slitlike. g. Dorsal parietal surface narrow to moderately wide, laterally distinctly or indistinctly bayed. Dentition well developed with 14 teeth of which 3 are on the premaxilla; the fifth maxillary tooth is abruptly smaller than those anterior to it and then posteriorly the series gradually decreases in size. Quadrate far forward with the quadrate ramus of the pterygoid extending much posterior to the plane of the quadratic condyles, underlying the distal end of the stapes. The lateral ramus of the pterygoid forms a strong and prominent bar but the width across the transverse rami is small. {olny Ay CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 47 . The intersquamosal width is moderate to fairly moderate. . The basioccipital condyle is directed postero-ventrally. . The frontal is of medium size and enters the orbital border, the supra- occipital is wide but apparently fairly high, the tabular with a small occipi- tal face and excluded from the posttemporal fossa, the lacrimal meets or does not meet the nasal, the squamosal without anterior wedge into the jugal. C. Taurocephalus . Snout of medium length, fairly high, facial surface not extending posterior to the anterior orbital border, transition on to cranial surface not abrupt but through a gentle curve. . Dorsal cranial surface fairly strongly pachyostosed, but centres of greatest thickening still distinct and the transition on to the face through a gentle curve and no transverse wall is thus formed. . Additional pachyostosis: a. No distinct naso-frontal boss present, and from orbit to orbit the dowel surface is evenly convex. __b, The parietal boss forms a rounded mound, highest round the pineal __ border but with its edges running into the general parietal surface. c. Postorbital bar fairly wide and massive, though extending very far laterally it is dorsally not bulbous and flows evenly on to the generally thickened dorsal surface; the width over the postorbitals is very great and in dorsal view they form the lateral edge of the skull. d. Posttemporal arch moderately thick and smooth. e. Orbits just visible in dorsal view anterior to the postorbital bar which here forms the lateral border of the skull in dorsal view. jf. Antero-posterior diameter of the temporal fossa fairly great and thus broadly oval. - g. Dorsal parietal surface wide and laterally disenctly bayed. . -Dentition well developed with 21 teeth of which 4 are in the premaxilla; there is an evenly graded decrease in size in posterior direction. Quadrate not very far forward; the quadrate ramus of the pterygoid not extending pésterior to the plane of the quadratic condyles, only underlying the anterior corner of the distal end of the stapes; quadrate with anterior process meeting the quadrate ramus of the pterygoid. . The lateral ramus of the pterygoid forms a strong and prominent bar and the width across the transverse rami is large. . The intersquamosal width is moderate, just more than the inter-postorbital width. . The basioccipital condyle is directed posteriorly. . The frontal is large and enters the orbital border, the supraoccipital is narrow and high, the tabular has a large occipital face and enters the post- temporal fossa, the lacrimal does not meet the nasal, the squamosal with a sharp anterior wedge into the jugal. 48 ANNALS OF THE SOUTH AFRICAN MUSEUM D. Struthiocephaloides® Snout long, shallow, facial surface extending far back, posterior to the anterior orbital border, transition on to the cranial surface not abrupt. Dorsal cranial surface strongly pachyostosed, but centres of greatest thicken- ing still fairly distinct, but the transition on to the face not abrupt, with no indication of a transverse interorbital step. Additional pachyostosis: a. No naso-frontal boss (but with a swelling just anterior to the posterior end of the premaxilla in cavifrons). b. Parietal boss distinct in cavifrons but in duplessist undeveloped or indistinct due to confluence with the general pachyostosis of the parietal and frontal. c. Postorbital bar very wide, massive, but without a bulbous boss in its dorsal part. d. Posttemporal arch moderately to strongly thickened, fairly smooth. e. Orbits just to plainly visible in dorsal view, and in dorsal view the post- orbitals do not form the lateral border of the skull. f. Antero-posterior diameter of temporal fossa moderate to fairly large, fossa oval or narrow (in cavifrons the fossa lies obliquely in the skull). g. Dorsal, parietal surface wide, laterally not pinched in. Dentition well developed with 14 teeth of which 3 are in the premaxilla, and from the 5th the teeth gradually decrease in size. Quadrate far forward, with the quadrate ramus of the pterygoid extending much posterior to the plane of the quadratic condyles. The lateral ramus of the pterygoid forms a fairly strong and prominent bar but the width across the transverse rami is small. The intersquamosal width is moderate to large. The basioccipital condyle is directed much ventrally. The frontal is fairly small, entering the orbital border in duplessist but not in cavifrons, the tabular excluded from the temporal fossa, the lacrimal does not meet the nasal, the squamosal without anterior wedge in to the jugal. E. Struthionops Snout fairly short and high, facial surface not extending posterior to anterior orbital border, transition to cranial surface not abrupt but through a gentle curve. Dorsal cranial surface moderately pachyostosed, centres of greatest thickening distinct, no interorbital step. Additional pachyostosis: a. Naso-frontal boss very low and not confluent with the thickening on the prefrontals. b. The parietal boss is a fairly prominent mound, highest round the pineal foramen and with its outer edges running into the general parietal surface. Ann. S. Afr. Mus., Vol. XLII Plate XVII eee RECONSTRUCTION OF STRUTHIOCEPHALUS 1 as exhibited in the South laster of the whole anima ion in p . Modelled by the author. ife-sized reconstruct from the | African Museum Photograph of the head of Struthiocephalus taken CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 49 3 Postorbital bar fairly narrow and slender with dorsal swelling. d. Posttemporal arch fairly weak and smooth. e. Orbits large and well visible in dorsal view, postorbitals form the lateral border of the skull as seen in dorsal view. f. Antero-posterior diameter of temporal fossa fairly large and broadly oval. g. Dorsal parietal surface wide, but laterally slightly, pinched in. 4. Dentition with probably 14-15 teeth evenly decreasing in size in posterior direction. 5. Quadrate not far forward. 6. The lateral ramus of the pterygoid strong and prominent but transverse width across the rami moderate. The intersquamosal width is large, just more than the inter-postorbital width. The basioccipital condyle is not preserved. g. The frontal is large, but is just excluded from the orbital border, the lacrimal just meets the nasal, the squamosal without a wedge into the jugal. oo DIAGNOSES FOR THE SPECIES OF STRUTHIOCEPHALUS S. whaitsi 1. Naso-frontal, parietal and bulbous boss on postorbital prominent. 2. Snout moderately long, shallow and fairly weak. 3. Moderately broad over squamosals. 4. Pineal foramen moderately far from occipital border. 5. Postorbital bar wide. 6. Temporal fossa fairly rounded. 7. Median occipital ridge wedge-shaped with rounded edges. 8. Nares well back. 9g. Quadrate well forward. 10. Interorbital width large. 11. Interparietal width small. 12, Posttemporal arch moderately strong. S. rheederi Naso-frontal, parietal and bulbous boss on postorbital prominent. Snout fairly long, moderately shallow but broad. Broad over squamosals. Pineal foramen very near occipital border. Postorbital bar probably wide (not evident from Broom’s account). Temporal fossa slit-like. Median occipital ridge not figured or described by Broom. Nares very far back. Quadrate not figured or described by Broom. 10. Interorbital width very large. 11. Interparietal width large. 12. Posttemporal arch moderately strong. SOS A Pee a es S. milleri 1. Naso-frontal and parietal bosses very prominent and bulbous boss on postorbital bar moderately prominent. Snout very long and very shallow. Relatively narrow over squamosals. Pineal foramen near occipital border. Postorbital bar narrow. Temporal fossa fairly rounded. Median occipital ridge not figured or described by Olson and Broom, Nares fairly near premaxillary edge. © OW HAP Quadrate not figured or described by Olson and Broom. 5O ANNALS OF THE SOUTH AFRICAN MUSEUM 10. Interorbital width small. 11. Interparietal width fairly small. 12. Posttemporal arch robust. ‘S. akraalensis 1. Naso-frontal boss large but low, parietal boss prominent, postorbital bar dorsally greatly thickened with the development of a longitudinal ridge raised above the surface of the frontal which thus appears excavated. 2. Snout very long and shallow. 3. Broad over the squamosals. 4. Pineal foramen far from occipital border. 5. Postorbital bar very wide and strongly rugose. 6. ‘Temporal fossa slit-lke. 7. Median occipital ridge straight with sharp median edge. 8. Nares far back. g. Quadrates very far forward. 10. Interorbital width very large. 11. Interparietal width very large. 12. Posttemporal arch very robust. THe Curer MEASUREMENTS COMPARED whaitst rheedert milleri akraalensts Length 558 624 480 655 Width : 345 432 300 390 © Interorbital width . 170 180 130 175 Interparietal width. : Sete ste OO 140 105 167 Pineal foramen to occipital edge shine ti 65 48 - 44 go Narisito Bo Mixedge uae 85 114 52 95 Keys To FActnirATE Rapip TAxonomic IDENTIFICATION Deinocephalia. Large therapsids, with pachyostosed skulls, with large quadrates, quadratojugals forming the corner of the ‘cheek’, premaxillaries with long facial exposure. Deinocephalians with differentiated carnivorous dentition, cranial bones not greatly pachyostosed, snout long, low parietal crest, ‘quadrates not situated far anteriorly. Medium-sized to large Deinocephalians with undifferentiated herbi- vorous dentition, cranial bones slightly to very greatly pachyostosed, in all but the Moschosaurids the snout is weakened, there is no parietal crest (except in Riebeeckosaurus and Avenantia); the quadrates are situated moderately to very far anteriorly. ‘Titanosuchia. Tapinocephalia. GROUPS IN THE TAPINOCEPHALIA Little pachyostosis Great pachyostosis Facial bones thickened to run evenly on to the cranial surface Facial bones not so thickened as to run 1 evenly on to the cranial surface . Snout very much shortened and ‘weakened Snout not very much shortened and weakened oF BP Po GENERA OF THE MORMOSAURUS-GROUP Skull with abrupt step from face to cranial surface Skull without abrupt step : Distinct naso-frontal boss present Distinct naso-frontal boss absent ‘ Width over postorbitals not very great and postorbitals not forming lateral edge of skull in dorsal view . Width over parietals great and ieee forming lateral edge of skull in dorsal view : Width across jugals much less than across postorbitals Width across jugals not much less than across postorbitals D RYN so ox Moschosaurus-group 3 Moschops-group 5 T apinocephalus-group . | Mormosaurus-group Mormosaurus y) Struthiocephalus 5 Struthiocephaloides 7 T aurocephalus Struthionops re CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS Hy THE SPECIES OF STRUTHIOCEPHALUS ie onout very Jong iand, very, shallow) 33%) .) G4. mallert 2. Snout not very long and very shallow . Ae aie : 3. With prominent ridge median to ee boss Ne 12) akraalensis 4. Without this ridge ' hs ey ed) 8 5. Pineal foramen very near occipital border . Bian NOs bi oy WIMECUENE 6. Pineal foramen not near occipital border . . . . . = whaitsi DIscussIoN It would be unwise to attempt a discussion of the relationships of the Deino- cephalians with other orders and inter se until all the material in the South African Museum has been studied. At this stage I am confining my remarks to some points which the study of Struthiocephalus has brought to the fore. Age in the Struthiocephalus skull A specimen of whavist (S.A.M. 11493) has a skull just as long as that of the type (S.A.M. 2678) and another specimen referred to this species (S.A.M. 11591), but in it the upper teeth are just commencing to erupt and the outer edge of both premaxillaries and maxillaries is sharp and appears to form a functioning cutting edge until the teeth are fully erupted. All the cranial bones are still little affected by any pachyostosis. The postorbital bar is narrow and lightly built with only its dorsal part slightly expanded and here the postfrontal is large and has not yet overgrown the postorbital. ‘The posterior flange of the postorbital forming the dorsal border of the temporal fossa is little reduced and though not meeting the squamosal extends far posteriorly and is not encroached on or overhung by the parietal. ‘The antero-posterior diameter of the temporal fossa is nearly equal to the dorso-ventral diameter. The posttemporal arch is not thickened and does not encroach into the fossa. ‘The quadrates still lie in a plane posterior to the anterior orbital border. The whole outer surface is still smooth and free from rugosities. Unfortunately the state of preservation does not reveal the structure of the occiput and the ventral aspect of the skull. What could be determined in this juvenile skull is however sufficient to show that many of the characters peculiar to the adult Tapinocephalian skull are mainly due to the pachyostosis which increasing age brings about and which obscures its essential therapsid nature. The Cranial Pachyostosis Apart from the fact that in both Moschops and Struthiocephalus an intensifica- tion of the thickening of the cranial bones takes place during the life of the individual, it also appears to be a process that can be traced phyletically. In Moschosaurus, which is undoubtedly the most primitive of the South African Tapinocephalians, there is very little pachyostosis and in Agnosaurus it is also slight, whereas in all the other known forms a lesser or greater degree of bone- thickening is apparent. Less of the skull is affected in the Mormosaurus-group than in the Moschops- and Tapinocephalus-groups. The thickening chiefly affects the supra- and interorbital region, the intertemporal skull roof, the post- orbital bar; the posttemporal arch and the face and ‘cheek’ are relatively little 52 ANNALS OF THE SOUTH AFRICAN MUSEUM affected. The thickening in these regions very materially affects the size and shape of the temporal fossa, the orbits, the braincase and the mechanism of the lower jaw. The Supra- and Interorbital region In the Mormosaurus-group the moderate to fairly strong pachyostosis lies posterior to the plane of the anterior orbital border. It is least in Struthiocephalus, Struthiocephaloides, and Struthionops, and here the relatively unaffected snout extends posteriorly to the plane of the anterior orbital border where the transi- tion on to the supraorbital thickened part is not very abrupt. This transition is not very abrupt chiefly because the pachyostosis has not affected the lacrimal, prefrontal, nasal and frontal to any great extent, only the posterior part of the prefrontal and nasal and the medio-anterior part of the frontal being affected. The thickening here does not extend transversely across the skull from orbit to orbit, but is in evidence at three separate centres, viz. the tops of the two post- orbital bars and in the naso-frontal boss. In Taurocephalus more of the prefrontal and the posterior part of the nasals are affected and the above-mentioned three centres tend to coalesce and the transition from the face on to the cranium is by a gentle curve. In Mormosaurus the process has extended further and the pachyos- tosis extends anterior to the plane of the anterior orbital border with confluence of the three centres to form a very distinct transverse step raised high above the surface of the snout. In the Tapinocephalus-group this process has progressed much further in Tapinocephalus to include all the prefrontal, frontal and the posterior part of the nasal to a plane well in advance of the anterior orbital border and the descent on to the surface of the snout is very abrupt from the very high transverse rampart. (But in Keratocephalus this region has remained essentially Stéruthio- cephalus-like with an accentuation of the naso-frontal boss.) In the Moschops-group the pachyostosis is continued into the bones of the snout but is so graded that instead of forming a step there is an even curve from the tip of the snout on to the supra- and interorbital region. The Intertemporal Skull Roof In Moschosaurus and Agnosaurus the intertemporal width is small with the sides pinched in and with little thickening of the parietal. In the Mormosaurus-group this width is increased with the stages in this process shown by the five genera Struthiocephalus, Struthtocephaloides, Struthionops, Taurocephalus and Mormosaurus, in this order. In Struthiocephalus this widening has advanced least but the direction of this process is evident in the series formed by its constituent species. The loss of the dorsal bay to the temporal fossa can be traced through the species of Struthiocephalus to Mormosaurus. In the Tapinocephalus-group the intertemporal width attains its greatest dimensions and there is no sign of the bay. In Moschops a parallel process of widening of the intertemporal region has taken place but there is still evidence of the bay. CRANIAL MORPHOLOGY AND TAXONOMY OF STRUTHIOCEPHALUS 53 The thickening of the parietal is at first mainly in evidence round the pineal foramen. In Struthiocephalus the thickness of the parietal at the pineal foramen varies from 40 to 80 mm. and in a Tapinocephalid species from Gunyanka’s Kraal in Southern Rhodesia it reaches the enormous thickness of 310 mm. With the increasing pachyostosis of the parietal the boss round the pineal foramen tends to become engulfed by the general thickening and in the Rhodesian specimen there is little evidence of a separate pineal boss. The Postorbital Bar The postorbital bar is light and slender in Moschosaurus and Agnosaurus, moderate in Taurocephalus and Struthionops, strong to very strong in Struthio- cephalus and Struthiocephaloides, very strong and massive in Mormosaurus, and very wide, strong and very massive in Tapinocephalus. In Struthiocephalus the dorsal end of the bar formed by the postfrontal has a tendency to form a bulbous boss reminiscent of the condition in the Titanosuchid Anteosaurus, but in Tapino- cephalus especially this is incorporated in the general extensive pachyostosis. An increasing massiveness of the posttemporal arch parallels that of the post- orbital bar, and together they have the effect of reducing the antero-posterior diameter of the temporal fossa. Adverse Effects of the Pachyostosis The downward growth of the roof-bones, together with the reduction of the angle between skull roof and occiput, boxes in the braincase in the dorso- posterior corner of the skull. The encroachment of the posttemporal arch and postorbital bar very greatly reduces the size of the temporal fossa and conse- quently of the adductor mandibulae. The thickened bone overhanging the orbits very materially restricts the field of vision. No doubt the increasing pachyostosis indicates the road which led to extinction. REFERENCES 14 Watson, D. M.S. P.Z.S., p. 749. 1914. 1b Haucuton, S. H. Ann. S. Afr. Mus., xii, 2, p. 52. 1915. 1¢ Grecory, W. K. Bull. Am. Mus. Nat. Hist., 61, p. 179. 1926. id Broom, R. Mammal-like Reptiles of South Africa, p. 18. 1932. 1€ BoonstRA, L. D. Bull. Am. Mus. Nat. Hist., 72, p. 75. 1936. if Erremov, J. A. Nova Acta Leopoldina, N.F. 9, 39, p. 155. 1940. * Boonstra, L. D. Ann. Mag. Nat. Hist., xii, 5, p. 455. 1952. 3 Boonstra, L. D. ibid., p. 509. 1952. 4 Boonstra, L. D. S. Afr. Four. Sci., 48, 8, p. 247. 1952. 5 Broom, R. P.Z.S., p. 302. 1937. § Otson, E. C., and Broom, R. Journ. Pal., 4, 7, p. 615. 1937. 7 Havucuton, S. H. Report of the South African Museum for 1915, p. 4. 1916. 8 Boonstra, L. D. Tydskr. Wet. Kuns, xii, 2, p. 237. 1952. ® Boonstra, L. D. ibid. xii, 2, p. 246. 10 BoonstrA, L. D. Ann. Mag. Nat. Hist. xii, 5, p. 988. 1952. 11 Boonstra, L. D. Tydskr. Wet. Kuns. xii, 2, p. 242. 1952. 6. The Lower Faw Articulatory Region in some Pristerognathid Therocephalians. By LiEuwE D. Boonstra, D.Sc. (With 4 text-figures) In the course of the preparation of a paper, mainly of a taxonomic nature, for the Annals of the South African Museum on the hundred-odd specimens of Therocephalians from the Tapinocephalus-zone preserved in the South African Museum, I have determined certain points in the structure of the posterior part of the lower jaw and the relations of the quadrate, quadratojugal and squamosal. A short account of the structure in some of the better-preserved specimens is presented here. New genera and species mentioned here will be fully described in the forthcoming taxonomic paper referred to above. SCYMNOSAURUS FEROX (fig. 1) A specimen collected by me on the farm Rietkuil in the district of Beaufort West (S.A.M. 9084) consists of a fairly good skull and parts of some limb-bones. By direct comparison with the type in our collection I have identified it with Scymnosaurus ferox. Natural weathering has partly exposed the mandibular articulatory region, but also unfortunately removed the lateral surface of the quadrate, quadratojugal and the lateral edge of the squamosal. The dentary has a concave posterior edge overlapping the angular and its dorso-posterior coronoid process does not extend above the subtemporal arch but is directed more posteriorly in the direction of the squamosal. The angular hasa fairly large lateral face with a prominent dorso-ventral ridge and a lesser ridge roughly at right angles. The anterior half of its ventral edge is fairly thick, but the posterior half has a double thin edge—the outer being the reflected flange. Posteriorly the outer (reflected) flange extends to the surangular and ventrally it underlaps the articular. Dorsally there is a small notch, anterior to which the angular apparently overlapped the outer surangular face. This dorsal notch, not exposing the outer surface of the inner angular sheet, but instead the external surangular surface, thus has different relations than in other Pristerognathids where this region has been described. A notch in the ventral edge indicates the level to which the reflected flange usually extends posteriorly in other Pristerognathids. Externally the surangular shows a thickened rounded and curved dorsal and postero-dorsal girder forming the edge of the mandible. Further ventrally the surangular curves firmly round the lateral surface of the articular and, with the internal prearticular, transmits the stresses arising anteriorly to the articular bone. The articular is a comparatively small bone of peculiar shape; externally a tongue extending dorsally is anteriorly clasped by the surangular and posteriorly o4 ARTICULATORY REGION IN PRISTEROGNATHID THEROCEPHALIANS I b. Fic. I.—Scymnosaurus ferox. S.A.M. 9084, Rietkuil, Beaufort West. (x # nat. size.) a. Lateral view. The lateral surface of the quadrate, quadratojugal and squamosal has been weathered away so that these elements are seen in section. b. Cross-section of above at A.B. c. Oblique postero-lateral view. The lateral edge of the squamosal, quadrate and quadratojugal weathered and seen in section. These and subsequent figures are all ortho-projections taken with a pantograph. An.—angular. Art.—articular. B.Oc.—basioccipital. B.Sph.—basisphenoid. Cor.—coronoid. D.—dentary. Ep.Pt.—epipterygoid. Ex.Oc.—exoccipital. F.J.—foramen jugale. I.P.—inter- parietal. J.—jugal. Pa.—parietal. P.O.—postorbital. P.Oc.—paroccipital. Pr.Art.—prearticular. Pr.Ot.—prootic. P.T.F.—posttemporal fenestra. Pt.—pterygoid. Q.—quadrate. Q.J.—quadrato- jugal. S.A.(n)—surangular. S.Oc.—supra-occipital. Sq.—squamosal. St.—stapes. Tab.—tabular. 39 56 ANNALS OF THE SOUTH AFRICAN MUSEUM forms the front face for the articulation with the quadrate condyle; it then extends posteriorly to form the ventral part of the articulatory surface, but no retro-articular process is developed. In posterior view it is seen that the articular has a dorsal spur, which, if it extends anteriorly, must form a ridge dividing the articulatory surface into a smaller lateral concavity and a larger median concavity. The quadratic condyle would thus be bipartite with a smaller rounded outer and an inner larger roller-shaped condyle. Medio-ventrally the under-surface of the articular is clasped by the prearticular, which transmits the greater part of the thrust from the anterior part of the mandible to the articular. Only part of the quadrate is exposed in lateral and in posterior view. Its lateral surface has been somewhat weathered away so that the drawing (fig. 14) really shows a section face. The apparent bipartite nature of the quadratic condyle has been noted above. In section the outer condyle has a circular outline. From the antero-dorsal corner of this condyle the lateral edge of the quadrate is seen extending dorsally to meet the squamosal. Further medially a flange of the squamosal clasps the quadrate from above. From the antero- median corner of the quadrate a process stretches antero-medially to meet the quadrate ramus of the pterygoid. In posterior view it is seen that from the external condyle the quadrate extends medially, forming a transverse mass forming the major part of the articulatory surface. Dorsal to this transverse body the quadrate extends dorsally as a strong pillar of which the posterior | surface is overlapped by a descending sheet of the squamosal. The medial face of the quadrate abuts against the latero-anterior corner of the strong paroccipital. On the median surface of the internal condyle there appears to be a shallow depression for the reception of the distal end of the stapes. The tympanic membrane must have been situated in this region, but except for the ventral squamosal edge no other points of attachment can be determined. The quadratojugal is only seen in section, where it is seen as a triangular element with its base resting on the postero-dorsal surface of the rounded external condyle and its apical part lying posterior to the ascending column of the quadrate and in the hollow between this and the descending flange of the squamosal. ‘The median limits of the quadratojugal and the quadrate foramen are in posterior view hidden by the descending flange of the squamosal. The squamosal has its lower antero-lateral edge weathered away and a section medial to this is shown in the figure. There is thus exposed much of the quadratojugal, the upper part of the quadrate and the internal forwardly directed domed sheet of the squamosal. All of which would be covered laterally by the squamosal when fully present. In posterior view the descending sheet of the squamosal covers the quadratojugal and most of the quadrate above the condyles. Its ventral edge is concave and probably served as an edge of attach- ment for the tympanic membrane. The median edge of the descending sheet of the squamosal forms a backwardly directed ridge where it abuts against the paroccipital. This ridge forms the median wall of the auditory groove, which is thus situated directly dorsal of the probable location of the tympanic membrane. ARTICULATORY REGION IN PRISTEROGNATHID THEROCEPHALIANS 57 ll b. Fic. I1.—Therioides cyniscus Gen. et Sp. Nov. S.A.M. 11888, Vindraers- fontein, Beaufort West (x #3). a, lateral view. 6, occipital view. 53. : ANNALS OF THE SOUTH AFRICAN MUSEUM Anterior to the descending sheet of the squamosal the paroccipital appears to abut against the median surface of the ascending pillar of the quadrate. TTHERIOIDES CyNiscus Gen. et Sp. Nov. (fig. IT) On the farm Vindraersfontein, Beaufort West, I collected a fair skull and fore-limb which I believe to be new and for which I propose the name T heriozdes cyniscus gen. et sp. nov. and the genotype bears the S.A.M. No. 11888. This new genus may preliminarily be described as a Pristerognathid with dental formula i. 6, c. 1, p.c. 6-7, postcanines slender and small, mentum fairly strong and squarish, quadrate situated far ventrally, maximum length about 270 mm., snout as high as broad (55 mm.), orbit well in posterior half of skull. In Therioides the posterior edge of the dentary sweeps sharply in posterior direction and is directed more in the direction of the squamosal than towards the dorsally situated temporal opening. The angular has a smooth surface in its dorsal part; antero-ventrally there is a groove and another groove is directed postero-ventrally. On its lower border a thin sheet of bone lying medially projects below the edge of the outer surface. The reflected lamina is strongly developed but does not extend to the surangular. A deep notch lies dorsal to the reflected lamina and the bone surface in the notch is formed by the angular. In lateral view the surangular shows as a curved girder, anteriorly lying medially of the upper angular edge. Postero- ventrally it firmly clasps the articular. The articular is small with a slight indication of a retro-articular process. Medio-ventrally the articular is under- lapped by the prearticular. The quadrate in lateral view shows little more than the circular outline of the outer condyle above which the quadratojugal lies. In posterior view the trans- versely situated condylar part is partly exposed and the stout dorsal pillar-like body of the bone is seen to be overlapped by the squamosal. Medio-ventrally the rounded corner of the quadrate appears to form a surface to receive the concave distal end of the stapes. The quadratojugal is not clearly exposed but the visible bone immediately above the external rounded condyle of the quadrate is apparently the lateral part of the quadratojugal. The postero-lateral corner of the squamosal is indentured immediately above what I believe to be part of the quadratojugal. The ventral border of the posterior descending flange of the squamosal is shallowly concave and this edge probably served for the attachment of the tympanic membrane. Lateral to where the squamosal abuts against the distal end of the paroccipital, the squa- mosal has a high, thin sigmoid rampart forming the median wall of the auditory groove. The ventro-distal corner of the paroccipital has a sharp edge to which the tympanic membrane was in part attached. The stapes is a slender rod in which I have not been able to detect a foramen. Its distal end has its dorsal corner prolonged and this, tipped with an extrastapedial cartilage, was probably ARTICULATORY .REGION IN PRISTEROGNATHID THEROCEPHALIANS 59 2 ‘ = @& @& a ® 4 om. a & 5f Py OP Sl OOM . @ “lg. @ . . 2 us & we bed - = . ° . PS NCA tena a Ss is CE aC = @ eof. ae Oe - 2 oe ey we ° #l3e07 2 ° CA oY EO ewan a a SG ds a ? Bo 22" ve wifeey BS Sere phd tes Seat 5 . & Oars 2s. 3 e r | P 5 ella erie toe ae . Gia F a ie ie FO Oa Fy Sats Ar we fou Cre We F 2 Seen S50 BI S780 = - Cig Sette ce y 4 ° : >” PRSAS ZOD Ce ec LL we heat ae 4 5 BSS ep ene Od ee 6 oes = . Ra eA Oe eS = OREO Ores} ° < . os See RON Oe so e ‘ i J By Ore CSS Sh film ele wee e See . | zs 5 =s ‘ cs ee Ae AS ee Sh, Mite ig it cl oa a ch ne = TaN hai! we Tatu demisess 2h. itr. boewohansesiey vl baicuelci) ob een Owe Sci ¥ . P 7 ZSHENGIoO I Lect ae teoe Svoo gem boritiahaltaan wi 7 % 4 4 : [ ) > REN et Pe ee el 2 The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at iene intervals as material becomes available. As far as possible each volume is devoted exclusively to a particular subject (Zoology, Botany, etc.). Two or more volumes may be i in course of ss publication concurrently. Most of the Geological and Palaeontological papers are issued in conjunction ‘with the — Geological Survey of the Union of South Africa. *\ Some volumes and parts are out of print, and others are only sold as parts of a set, or volume, _ respectively. The prices of parts published prior to 1940 have been increased. Out of print: Vols. I, II, V (Parts 1, 2, 9), VU, VIII, Ix ee I), XII (Part 7), XXII, XXIV (Part 2), XXXI (Parts 1, 2, 3). Vol. III. 1903-1905 Zoology IV. 1903-1908 Palaeontology ; V. 1906-1910 Geology, Palaeontology, Zoology, Anthropology : VI. 1908-1910 Zoology. IX. 1911-1918. --Botany | =. ed (excl. ‘Part 1) X. sig911-1914 Zoclogy XI. s911-1918 Zoology oe ah olen XII. 1913-1924 Palaeontology anal Geateey .. (exclo Part 9) XIII. 1913-1923 eas y and oes a é Rees 8 XIV. 1915-1924 Zoology XV. 1914-1916 Zoology XVI. 1917-1933 Botany : XVII. 1917-1920 Zoology .. XVIII. 1921 Zoology a XIX. 1924-1925 Zoology .. Bs ot XX. 1924-1926 Zoology .. oo aN ‘< XXII. 1925-1927 Zoology — XXIII. 1925-1926 Zoology ied) 9 eee XXIV. 1929-1938 Anthropology nae Biktolany . (excl. Part 2) XXV. 1927-1928 Zoology .. Ns XXVI. “1928 Zoology . XXVIII. 1929 Anthropology XXVIII. 1929-1932 . Palaeontology XXIX. 1929-1931 Zoology XXX. 1931-1935 Zoology . INDEX of papers, authors, and subjects, hed 4 in Vols; LXXX . XXXI. 1934-1950 Palaeontology : (Par 4 ae XXXII. 1935-1940 Zoology ‘ a XXXITI. 1939 Zoology XXXIV. 1938 Zoology XXXV. - Reserved for conclusion of siaherph i in Vol. XXXIV. XXXVI. 1942-1948 Zoology as i XXXVIT.. 1947-1952 Archaeology — XXXVITII. 1950 Zoology a oe XXXIX. 1952 Zoology .. a ae Ns a at XL. 1952- Botany %)) .) ae oe maa! (Part 1) XLI.. 1952- Zoology (Part 1) Copies may be obtained from— The LIBRARIAN, Soutn Arrican Museum, CAPE Tose except the Geological and Palaeontological parts, which are obtainable from the ~ GOVERNMENT PRINTER, PRETORIA. irae OCW ND HH HRD HWNORWORWOWOR NO Hw HH oO NW —= N <* oak ce s COMOMMODFOCAOMIIVD090ROZORAMROMTOMNSDOO® mDomo00sd ANNALS : 3 eo aa ee * ee se ee ee = Weer oe Se a tt eaters a SOUTH AFRICAN MUSEUM : VOLUME XLII _ Descriptions of the Palaeontological Material collected by the South African Museum and the Geological Survey of South Africa. PART II, containing: — ‘i 7, The Pristerognathid Therocephalians from the Tapinocephalus-zone ff in the South African Museum. By L. D. Boonstra, D.Sc. (With ae 22 text-figures.) : _ 8. The Cranial Structure of the Titanosuchian: Anteosaurus. By L. D. iy Boonstra, D.Sc. (With 22 text-figures.) sy 9g. Ihe smallest Titanosuchid yet recovered from the Karroo. By L. D. jm Boonstra, D.Sc. (With Plate XVIII and 5 text-figures.) yi 10. Paranteosaurus Gen. Nov.: A Titanosuchian Reptile. By L. D. P _ Boonstra, D.Sc. (With 2 ‘text-figures.) oa £. ey hipt Ting 2 is i { REN FAN 4 Viger ATL g cif ER bs ISSUED JANUARY 1954. PRICE 155. od. — :-t TE ape a PRINTED FOR THE ae TRUSTEES OF THE SOUTH AFRICAN MUSEUM AND THE GEOLOGICAL SURVEY OF SOUTH AFRICA + > :< - he ae POY grids PEK? BY THE NATIONAL COMMERCIAL PRINTERS LIMITED, VAN RIEBEECK STREET, ELSIES RIVER tae | | | | | ht AN Pi MC a, ys 7. The Pnsterognathid Therocephalians from the Tapinocephalus-zone in the South African Museum. By L. D. Boonstra, D.Sc. (With 22 text-figures) In the course of my study of the fauna of the TAPINOCEPHALUS-zone the collection of Therocephalians in the collection of the South African Museum | has grown to number nearly a hundred specimens. The majority belong to the family Pristerognathidae. Unfortunately, well preserved specimens are rare and, although we have some good skulls, most of the specimens are weathered snouts and in only a few cases have we postcranial elements associated with the cranial material. In this paper I propose to deal only | with the cranial material. The Pristerognathids represent the bulk of the small to fairly large carnivores in the fauna of the Tapinocephalus-zone. With them occur members of some other Therocephalian families viz. Lycosuchidae and Scylacosauridae and as a lesser element some Scaloposauridae, which were _ probably insectivorous. The balance of the carnivorous section in the fauna is formed by a few small Gorgonopsians and then we have the large and massive Titanosuchid Deinocephalians. The food source of these carnivores consisted mainly of the number of small herbivorous Dicynodonts and the abundant large Pareiasaurs and | Tapinocephalid Deinocephalians. Some of the Pristerognathids were | probably carrion eaters. Family Pristerognathidae D1aGnosIs Fairly small to large early carnivorous Therocephalians from the Tapinocephalus-zone with a dental formula varying within the limits, a, C.—, pee The incisors and canines usually well developed and the posi-canines, with few exceptions, relatively feebly developed. Skull long, fairly high and narrow; snout long, fairly narrow; sagittal crest high and sharp; temporal fossa moderately long and broad; low to fairly low over post-orbital arches; lower jaw strong, dentigerous ramus of dentary not curved and relatively long, post-dentary part of mandible strong, strong coronoid process. Prefrontal and _ postorbital well developed; and postirontal well to fairly well developed; jugal arch moderate to strong; frontal with moderate entry into orbital border or excluded from it; septomaxilla with well developed facial surface; pineal foramen medium to 65 WOL. XLII. PART II. 66 ANNALS OF THE SOUTH AFRICAN MUSEUM fairly large; parietal narrow with sharp crest; paroccipital strong; occiput fairly high and broad, deeply concave; epipterygoid narrow; anterior palatal openings medium to long; suborbital vacuities large; palate flat; with only the anterior — ramus of the pterygoid bearing teeth; basisphenoidal tubera of medium size, parasphenoidal keel deep; suspensorium posteriorly situated high up and only moderately laterally displaced. Within the family two groups can be recognised which may yet have to be considered as subfamilies: a. Those with 6 incisors and with skulls where the length is double or more the breadth. b. Those with 5 incisors or with an inconstant 6th and where the skull length is less than double the breadth. GENERAL DESCRIPTION Incisors, canines and post-canines have a sharp posterior cutting edge which is serrated — the serrations vary from fine to fairly coarse. The incisors are moderately curved teeth with a more or less flattened extremity forming a curved cutting edge. They vary from fairly weak to very strong seizing and tearing teeth. The first incisor with its fellow form a pair of teeth closely set to each other, with the root rounded in cross section and always smaller than the second incisor. The second to the 5th incisor are subequal or decrease in size backwards. In the forms with six incisors the 6th tooth is inconstant in some genera and in the others it is mostly much smaller and weaker than its predecessors. There is always only one canine; except in one genus the canine is always a strong curved dog-tooth with its extremity more pointed than is the case in the incisors; the upper canine is situated far forward in the maxilla (except in one species) and to house its long strong root the maxillary edge has grown downwards so that the premaxillary edge is situated at a higher level; usually the alveolar border curves evenly upwards anterior tc the canine, but in some the transition is by a distinct step. The diastema between last incisor and canine varies considerably in length — sometimes the canine follows immediately after the last incisor. Behind the canine the diastema to the first postcanine also varies considerably, but only in one specimen does the first postcanine lie right up against the canine. The number of postcanines varies considerably in the Pristerognathids — from 2 to 9. The postcanines of the family, considered as a group, must be described as weakly developed. They are usually small slightly curved teeth either closely set or well spaced. In only one genus (Cynarniognathus) do they form a set of well developed teeth. They are often irregular and of little functional importance. In the dentary there are apparently always 3 incisors, which in the closed jaws are overlapped by the upper incisors. The lower canine lies lingually THE PRISTEROGNATHID THEROCEPHALIANS 67 and anteriorly to the upper canine and its point in the closed jaws is housed in the anterior section of the “‘choanae’’. Little is known of the postcanines of the dentary — they lie lingually to those of the maxilla. It is thus evident that the Pristerognathids in their feeding relied on the anterior part of the jaws; the large canine acting as piercing instrument to effect the kill and these together with the incisors were used to hold the victim and then acted as the instruments for tearing away pieces of flesh. Those with weaker incisors were probably carrion eaters. In most Pristerog- nathids the postcanines acted chiefly as protruding points preventing flesh from slipping out of the mouth. The transverse pterygoidal rami limited the action of the lower jaw to movement in a vertical plane and there was thus no chewing or cutting by the postcanines. The small set of recurved teeth on the pterygoidal ridge served to hold the flesh in the stage preliminary to deglutition. The postfrontal is well developed in some species, whereas in others it is only a fairly small splintlike bone; it has only a small posterior extent, in some forms extending up to the plane of the fronto-parietal suture, in others a little further and at most to the level of the pineal foramen. It seldom extends further posteriorly than the postorbital. The posterior tongue is usually confined to the dorsal surface but sometimes lies on the lateral face of the parietal crest. The postorbital varies in the extent of its participation in the postorbital bar, which is slender to fairly slender, never widened; in some forms it has a small entry into the orbital border and the temporal border; in others the entry into the orbital border is small, but the entry into the temporal border is large; this depends on how high the dorsal jugal ramus extends and to what extent the jugal is confined to the posterior half of the postorbital bar. A postero-dorsal tongue of the postorbital sometimes just enters the supratemporal border where it is applied to the lateral face of the parietal or it sometimes develops quite a fair-sized sheet of bone flanking the parietal, but seldom extends to just behind the plane of the pineal foramen. The jugal is always well developed; it is triradiate with a dorsal, anterior and posterior ramus. The anterior ramus forms the lower part of the orbital border and a varying portion of the lower part of the posterior orbital border; it mostly forms a sharp pointed wedge in between the lacrimal and maxilla, but this is sometimes truncated. The dorsal ramus sometimes forms the whole posterior half of the postorbital bar and thus this part of the temporal border; in other forms its dorsal extent is much smaller and this part of the temporal border is formed by the postorbital. The posterior ramus always extends very far posteriorly as a long pointed wedge underlying the anterior ramus of the squamosal, together forming the temporal arch. Usually the jugal forms nothing or very little of the upper border of the temporal arch, but in those forms where the anterior ramus of the squamosal is short it forms the anterior 68 ANNALS OF THE SOUTH AFRICAN MUSEUM portion of the upper border of the temporal arch. (This long posterior ramus of the jugal in the Pristerognathids resembles the condition seen in Cynodonts much more than does the jugal of the Gorgonopsians. ) The squamosal is always well developed; it has an anterior ramus extending far anteriorly to form most of the upper border of the lower temporal arch where it overlies the jugal; its ventral ramus extends far ventrally, laterally largely covering the quadratojugal and posteriorly covering most of the posterior face of the quadrate, which is housed in a recess in the antero-ventral face of the descending ramus of the squamosal; the medially directed ramus of the squamosal forms the posterior border of the temporal fossa and is here applied to the outer face ot the parietal; this overlap of the squamosal on the parietal is fairly small so that the postorbital and squamosal only flank the lateral face of the parietal in its extreme anterior and posterior part respectively, leaving the parietal to form nearly all of the upper border of the temporal fossa. Where the squamosal meets the strong paroccipital the edge of the squamosal is everted to form the auditory groove. The sweep of the squamosal posteriorly carries the posterior border of the temporal arch far back in the long narrow headed forms and thus forms a deeply concave occiput, whereas in the shorter broader-headed forms there is only a moderate backward sweep with a shallowly concave occiput. There is only a moderate lateral sweep of the squamosal so that the temporal fossa never becomes very wide. There is more lateral sweep in those forms where the posterior sweep is least. In dorsal view the Pristerognathids show little more of the upper occipital bones than their dorsal edges; the occiput is thus upright. In many forms the condyle is clearly visible in dorsal view, but in the longheaded forms it is either just visible or just hidden by the backward sweep of the parietals. The quadratic condyles lie well below the level of the occipital condyle and in a plane well below that of the alveolar border of the maxilla; they have migrated very little in lateral direction so that the outer edge of the quadrate ramus of the pterygoid is fairly straight; there has also been no forward migration and they still occupy a posterior position. The quadratojugal is much reduced and is a small element not entering the condylar surface but resting with its base on the ledge of the quadrate above the outer part of the condyle and with two weak dorsal processes fitting into recesses in the overlapping squamosal. The large prefrontal has a rounded ridge separating a dorsal from a lateral face, but farther anteriorly the snout is rounded with no abrupt transition from dorsal to lateral surface. The lacrimal is small and has little anterior extension due to the upward growth of the maxilla; it lies in the preorbital depression which is usually fairly shallow and extending as a shallow groove in the direction of the canines; but in some forms this depression is deep with sharp margins and THE PRISTEROGNATHID THEROCEPHALIANS 69 anteriorly ends abruptly. The homology of the gland occupying the preorbital depression is uncertain. The long and strong root ot the canine is housed in a strong rounded bulge of the maxilla, which is consequently a high bone. The outer surface of the maxillary bulge often has an ornamentation of sub-crocodilian pitting, but in some forms it appears to be fairly smooth. It is pierced by a number of nutritive foramina. The nasals are long and fairly narrow; together they are hourglass-shaped; the constricted waist being due to the upward growth of the maxilla overlapping the lateral edge. In the lower jaw the symphysis is always weak and unankylosed, with the splenial just entering the symphysis and forming a very subordinate part and not visible ventrally. The angular has a large outer face with a notch and a reflected lamina. Systematic Descriptions Genus Scymnosaurus Broom. Broom, 1903, p. 152. Genotype, S. ferox Broom. Large Pristerognathids with dental formula i.5, c.1I, p.c.2-4; incisors and canines large and strong, postcanines fairly weak to medium, varying in number — 2, 3 or 4. (In one specimen a small 6th incisor has been seen, but only on the one side.) Skull very large (Max. Length 375 to 475 mm.); preorbital hollow fairly shallow, not sharply demarcated and really not more than a groove stretching from the orbit in the direction of the canine; septomaxilla and septomaxillary foramen well developed; frontal either excluded or possibly just entering the supraorbital border; snout probably always slightly broader than high, slightly narrowed between orbits and canines; orbits partly in anterior half of skull. Mandibular symphysis, formed solely by dentaries, weak, unankylosed, mentum sloping little or moderately. Scymnosaurus ferox Broom. (Figs. I, Ila.) Broom, 1903, p. 152. Holotype. S.A.M. 632. Locality unknown. Coll. unknown. A well preserved but imperfect snout. There are 3 to 4 postcanines; mentum moderately sloping and symphysis weak; maximum length of skull 375 to 381 mm. (as reconstructed); frontal possibly still entering orbital border. S.A.M. 632 (Fig. Ia). Broom’s holotype consists of the anterior third of the mandibular ramus, the dentigerous border of the premaxillaries and the maxillaries up to just behind the 3rd postcanine on the right and the 2nd postcanine on the left side. The horizontal fracture through the snout shows the dorsal surface of the anterior third of the palate, which is as figured by Broom moos, Pl. 18, fig, 9). 70 ANNALS OF THE SOUTH AFRICAN MUSEUM The first four incisors are strong, whereas the 5th is much shorter and this tooth shows fine serrations on its sharp posterior edge; the robust canine has fine serrations on its sharp posterior edge; only 3 postcanines are preserved and these decrease rapidly in size in posterior direction; the postcanines bear coarser serra- tions on the posterior edge. REFERRED SPECIMENS S.A.M. 3430 (Fig. 10). Janwillemsfontein, Prince Albert, ,,,,Coll. Haughton & Whaits. This is an imperfect snout in which is_ preserved the symphysis and the major part of the premaxillaries and maxillames’ In’) "both pre- maxillaries 5 incisors are preserved and here the 5th is only slightly shorter than the anterior 4 incisors. But on the left side there is a stump of what appears to be a small 6th incisor. On the right side a stump of the canine is preserved, as also stumps of the 2nd and 4th postcanines and empty alveoli of the rst and 3rd. On the left side the canine has dropped out and so has the Ist postcanine, but stumps of the 2nd and 3rd are preserved. The maxilla overlaps the premaxilla right up to the level of the anterior edge of e “ ay SAM. 632. e ©. ¢ nes. e te tts ee ty Oe wn sn ee oreo eese a tL Lene 7S ios is Se ee ‘ r) 7 = ci “Sr-. — US > ny Ee os . . mlis CS x we © ET Sere rn Be HY = pais e ‘: ‘ ‘ s Tea ot e te! , - Or em en ah ied Bye . ‘ PIR OS cid . CE BO Os zs v PR Ae 7s ’ ~ od : : Se oe ‘. his . « Ce x ss . ante ‘ Soi eee XN Or y a ~ D. - Og oe Oa eee ’ Senne tem eee ect tte S ae, AM 31,30. e ¢ se e Mn ee A = se TF Me CL we me He ec eeaese we ee ae ) o-ce S a s d- = , rc > \ ies ‘ ‘ ’ A dl ind ~ =. = ~ a ‘ e ah Mh NERY is mated : , OB SS ee ee eee $ me 7 — aof ac ‘ one’ Sn ‘ aR aes : Bea Ae Se Hi Be - —, “ ‘ > ert : 4 Gk -cre Sings » a: e \ . . oot Leoitie) ) me cae e me en neces SAM. U3LI. Fic. 1.—a, Scymnosauvus ferox. ype: S.A.M. 632. (x %-) ~ Right sidetverssnenc fragment. 6b, Scymnosaurus ferox. S.A.M. 3430. (x 2.) Right side of snout fragment. c, Scymnosaurus ferox. S.A.M. 4341. (x #¢.) Right side of snout. Lettering of Figures: A.—angular; Art.—articular; B.O..—basioc- cipital; B.S.—basisphenoid; D.—dentary; Ep. Pt.—epipterygoid; Ex.—exoccipital; F.— frontal; F.J.—foramen jugale; F.O. fenestra ovalis; I.P.—interparietal (postparietal); J.— jugal; L.—lacrimal; Mx.—maxilla; N.—nasal; P.—parietal; Pal.—palatine; P.F.—pretrontal; P.Mx.—premaxilla; P.O.—postorbital; P.Oc.— paroccipital; Po.F.—postfrontal; Pr.Ot.— proédtic; P.S.—parasphenoid; Pt.—pterygoid; P.V.—prevomer Q.—dquadrate; Q.J.—quad- ratojugal; S.A.—surangular; S.Mx.—septo- maxilla; S.Oc.—supraoccipital; Sq.—squa- mosal; St.—stapes; Tab.—tabular; Tr.— transversum. The figures are orthoprojections taken with a pantograph and, except where specifically stated, they are of the specimens as preserved without correction of any distortion. Recon- structed parts and uncertain features are shown in broken line. All figures drawn by the author. THE PRISTEROGNATHID THEROCEPHALIANS We the 4th incisor. The bulge of the maxilla above the canine is rugosely pitted, but no nutritive foramina can be seen. The septomaxillary foramen is of moderate size (6 mm.). S.A.M. 4341 (Fig. 1c). Stinkfontein, Prince Albert. Coll. Haughton. This is a weathered snout bleached white. Each premaxilla has the stumps of 5 incisors preserved; in cross section the 5th is only slightly smaller than the 4th; in the right maxilla imperfect crowns of 4 postcanines are preserved; the 3rd and 4th are the smallest and the 2nd the largest maxillary tooth. The septomaxilla has a large facial exposure and a stout spur directed anteriorly into the nostril. There is a fairly large oval septomaxillary foramen and in the maxillary edge, greatly overlapping the premaxilla, there is a nutritive foramen. The maxillary bulge is rugosely sculptured. The nasals broaden as they approach the nostril. S.A.M. 9084 (Fig. 2a). Rietkuil, Beaufort West. Coll. Boonstra. This specimen consists of the major part of a distorted skull, lacking the intertemporal region, together with the ends of a humerus, radius and ulna. The articulatory region has recently been described and figured (Boonstra, HSS, lig.) 1). ‘ «Yale deoran . mC S| gs cl) ese mm, ee ~ ee? “Ms ecee* Fic. 2.—a, Scymnosaurus ferox. S.A.M. 9084. (x #.) Left side of skull and lower jaw with correction of the distortion. b, Scymnosaurus major. S.A.M. 10556. (x ¢-) Right side of preorbital part of skull. 72 ANNALS OF THE SOUTH AFRICAN MUSEUM With the full length of the lower jaw preserved it is possible to estimate the maximum length of the Scymnosaurus ferox skull viz. 375 mm. The incisors are large and robust with the 5th only slightly smailer than its predecessor and the sharp posterior edge bears moderately fine serrations. The 3 postcanines are short stout teeth with the 3rd much smaller than the anterior ones and the serrations on the posterior edge are as in the incisors and the canine. The 5th incisor lies close to the canine, whereas in the specimens mentioned above there is an appreciable diastema. The preorbital depression has clearly demarcated borders. The maxillary bulge is pitted and is perforated by a number of nutritive foramina. The septomaxillary foramen is fairly large (6 mm.) and the septomaxilla well developed but with less facial exposure than in S.A.M. 4341. 7 SAM. 9005. Fig. 3:') Seymnosaurus major. Type: | S.A NE.) Coen: (x 4.) .@, Dorsal view. 0, Left side. THE PRISTEROGNATHID THEROCEPHALIANS TS The orbit lies partly in the anterior half of the skull. The dentary is strong, with a straight ventral edge, a sloping mentum and a weak symphysis. See: 4347. SCYMNOSAURUS MAJOR. Sp. Nov. Holotype. S.A.M. 9005. Klein-Koedoeskop, Beaufort West. Coll. Boonstra. The well preserved anterior two thirds of a skull together with parts of the pectoral and pelvic girdles, ends of humerus and radius and ulna. There are 2 to 3 postcanines; mentum fairly upright and symphysis fairly strong; maximum length of skull 450 to 474 mm. (as reconstructed); frontal excluded from orbital border. S.A.M. 9005 (Fig. 3 a and 0). The estimated maximum length of the skull is 450 mm. The incisors are large and strong (length of 3rd incisor 55 mm.); the length of the canine is 80 mm. (as reconstructed), whereas the two postcanines are small (15 mm.). All the teeth have a finely serrated sharp posterior edge. The 5th incisor lies close up to the canine, but the distance between the canine and the first postcanine is large (30 mm. on the right side and 50 mm. on the left). The preorbital depression is deep but with rounded margins. The maxillary bulge has pronounced pits and radiating grooves. The septomaxilla is well developed and the foramen fairly large (11 mm.). The maxilla is high but short. The nasals widen considerably in their anterior half. The frontals are small, very narrow and well removed from the orbital border. The prefrontals are large and the postfrontals fairly large. The orbits are small and round (45 mm.) and lie partly in the anterior half of the skull. The mentum is fairly upright but the two rami are loosely joined at the symphysis. S.A.M. 10556 (Fig. 2b). Knoffelfontein, Beaufort West. Coll Boonstra. This specimen consists of a weathered and bleached preorbital part of a large skull (474 mm. as reconstructed). Only cross sections of the roots of the teeth are preserved. The five incisors are practically of equal size. 74 ANNALS OF THE SOUTH AFRICAN MUSEUM Anterior to the canine there is a distinct diastema and the edge of the maxilla curves sharply downwards (the “‘step’’ of Watson). Posterior to the canine there is a diastema of 27 mm., then follow three closely packed small postcanines. The preorbital depression forms an oblique groove from the orbit in the direction of the canine. The maxillary bulge is strongly pitted. The septomaxilla is well developed with a strong intra-nostril spine. The septomaxillary foramen is very large (19 mm.) and is kidney shaped with a spur from the septomaxilla dividing it into two, in very much the same way as the medially directed spine divides the nostril. The maxilla is low and fairly long. The mentum is upright and the symphysis quite strong. SCYMNOSAURUS Sp. INCERTAE SEDIS The following fragmentary specimens can be referred to the genus but specific determination is uncertain, S.A.M. 8999. Locality unknown. Coll. unknown. A weathered snout fractured cleanly just behind the canines and from which little more can be determined than that there were 5 large subequal incisors and a fairly strong canine. S.A.M. 11459. Buffelsvle1, Beaufort West. Coll. Boonstra and Marais. A weathered snout in pieces, with the dental formula 1.5, c.1I., p.c.3+? together with parts of a carpus and tarsus. S.A.M. 11833. Lammerkraal, Prince Albert. Coll. Boonstra and Pienaar. A weathered snout with I postcanine on the right side and 2 on the: lett: S.A.M. 11961. Dikbome, Laingsburg. Coll. Boonstra. A weathered snout with dental formula 1.5, c.1, p.m ee S.A.M. 9126. Voélfontein, Prince Albert. Coll. Boonstra. This consists of a fairly well preserved snout with the teeth of the left side preserved i.5, C.I, p.c.3-+. The teeth are very similar to those of the genotype. Being only 2 the size of the genotype this snout may represent a small species of Scymnosaurus. THE PRISTEROGNATHID THEROCEPHALIANS 75 Genus Scymnosaurus: Measurements in Mms. Sey ELOX.: S. major. 632 3430 434% 9084 9005 10556 prvee— Post.sq. edge’ 4 -. 3752 375? 38% 375 4560? 474? Biwi Ant.Orb. border =. - —— — eT eMOg | 215 216? Width of Snout over Canines . 105? 106 105 #00? I15 100 ici tmterorbital 9... — — Corr 72. = Height of Snout at Post. edge Gigeaminers...... eee OO LOO 4 TOs LOOn. pT XO go Length of Upper Incisor Series (65 75 65 70 85 60 ea, #1010) 70 60 70 75 60 Diastema C.-T... ] 9 ae 15 3 6 18 r a Oe 16 5 9 16 DMiastemay C2—P.C. . 14 = == 20 51 = Ts, h ed aes 24 20 27 25 Length of P.C. Series oo == = 42 18 = Bey 20 — AI 36 21 30 Ant.-Post.Diam. of Canine 1 22 a 25 24 3311 25 ig 22 20 20 a 26 25 Genus Glanosuchus Broom. Broom, 1904, p. 85. Genotype G. macrops Broom. Large Pristerognathids with dental formula 1.5+1, c.1, p.c.5; the first 5 incisors are large but the 6th is inconstant and when present very small; the canine is large and strong; the postcanines are moderately to weakly developed. Skull large (Max. length 315-321 mm.); preorbital hollow fairly shallow, not sharply demarcated and really not more than a groove stretching from the orbit in the direction of the canine; septomaxilla and septomaxillary foramen well developed; frontal enters orbital border; postfrontal quite well developed; snout broader than high, slightly narrowed between orbit and canines; orbits well in posterior half of skull. Mandibular symphysis formed solely by dentaries, weak, unankylosed, mentum sloping. | Epipterygoid only slightly widened. Sagittal crest of parietals only moderately high but fairly sharp. Glanosuchus macrops Broom. (Figs. 4 and 5.) Broom, 1904, p. 85. Holotype. S.A.M. 637. Knofloksfontein, Beaufort West. Coll. Snyman. A well preserved skull lacking only the posterolateral corners of the skull and thus the suspensorial region. 76 ANNALS OF THE SOUTH AFRICAN MUSEUM Broom’s description was based on the only partially cleaned skull. I have cleaned up the outer surface, thus revealing the sutures and prepared the palate and basicranium so that these features can now be described. In Fig. 4 a, b and c the structural, features Of (ihe lateral and dorsal surfaces are shown. The _ septo- maxilla has a large facial exposure, the spur project- ing into the nostril is well developed and the foramen fairly large (6 mm.). The maxilla is fairly high and long with its dentigerous border curving upwards anterior to the posterior edge (ot) \the jcanine: yiihe preorbital hollow has a fairly definite dorsal and posterior rim but anteriorly it flows into the general maxillary surface in the direction of the canine. The maxillary bulge above the canine is fairly smooth. The nasals are widest anteriorly. The frontal is of medium size apparently just entering the orbital f border. In dorsal view the Fic. 4.—Glanosuchus macrops. Type. S.A.M. occiput is seen to be deeply 637- (X4%-) @, Right side. 6, Left side. 5 c, Dorsal view. d, Ventral view. concave. In outline the epipterygoid is hourglass-shaped and although not much widened it is not a columnar columella crani. Its base, resting on the quadrate ramus of the pterygoid, has no great posterior process. Although the edge of the sagittal crest is not preserved it is clear that the crest was not very high and moderately sharp. The temporal fossa is short but roomy. The ventral surface (Fig. 4d). With the dentaries in occlusion it has not been possible to expose the choanal region. The prevomers posteriorly form a truncated spatulate sheet of bone underlying the median edges of the palatine and the anterior edge of the anterior pterygoidal ramus. The lateral edge of the prevomer lies medial to an oblique ridge on the palatine. This ridge is continued on the pterygoid where it is dentigerous (a cross section of THE PRISTEROGNATHID THEROCEPHALIANS 77 at least one tooth is clearly visible). The palatine is a relatively small bone, anteriorly bounding the choana and posteriorly forming the front margin of the oval shaped large suborbital vacuity. Two oblique sutures, meeting in an obtuse angle, separate the palatine from the prevomer and pterygoid. The pterygoid is a typically tetraradiate bone; the anterior ramus with its fellow, forms the lozenge-shaped middle part of the palate; anteriorly, between the two diverging dentigerous ridges, a V shaped hollow extends on to the prevomerine surface where it shallows and fades out; laterally the anterior ramus forms the inner border of the suborbital vacuity; the posterior part of the anterior ramus carries a prominent oblique ridge stretching from near the median line to near the edge of the suborbital vacuity; this ridge is dentigerous, carrying small sharp teeth irregularly arranged in two rows; in the median line just posterior to these dentigerous ridges there appears to be a small interpterygoidal slit. The transverse pterygoidal ramus forms a strong bar which laterally descends to the level of the lower dentary border and this distal end lies just median to the angle of the dentary. The movement of the lower jaw is thus suided and confined with little lateral play allowed. Against the lateral part of the anterior face of the transverse pterygoidal bar lies the transversum; from here the transversum ascends rapidly and curving forwards as a thin process forms the lateral edge of the suborbital vacuity and meets the palatine. The posterior ramus of the pterygoid with its fellow forms a median keel which clasps the anterior end of the parasphenoidal rostrum. The quadrate ramus of the pterygoid is not completely preserved and one does thus not know whether it met the quadrate or not. On its upper surface the quadrate ramus carries the base of the epipterygoid. I have not been able to determine whether the epipterygoid has a posteriorly directed process running along the quadrate ramus. In ventral view it is seen that the edge of the quadrate ramus is only slightly concave and the epipterygoid has a roomy cavum epiptericum median to it and the anterior part of the prootic. The parasphenoid is in ventral view seen as a sheet of bone underlying the basisphenoidal tubera and narrowing anteriorly is clasped by the pterygoids where these bones form a fairly deep median keel. Ventrally the occipital condyle is formed by the strong basioccipital. Postero-laterally to the basisphenoidal tuber lies the fenestra ovalis with the proximal end of the stapes tm situ on both sides. The rim of the fenestra is formed postero-laterally by the opisthotic, postero-medially by the basioccipital, antero-medially by the basisphenoid and antero-dorsally by the prodtic. Only part of the occiput is preserved; above the foramen magnum the occiput is deeply excavated; the posttemporal fenestrae are large; the 78 ANNALS OF THE SOUTH AFRICAN MUSEUM paroccipital (opisthotic) strong and only the dorsal edge of the interparietal and tabular seen on the dorsal surface. S.A.M. 11843. (Fig. 5a@ and b.) Lammerkraal, Prince Albert. Coll. Boonstra. : This good specimen consists of the major part of a skull lacking only the suspensorial region and the lower edge of the dentary. It is complementary to the holotype skull in that it shows that the frontal has a small entry into the orbital border, | the «postiremtal ame postorbital flank the parietal for a short distance within the temporal fossa; the sagittal crest of the parietals is preserved as a not very high but fairly sharp crista; the temporal fossa is short but roomy. But the postcanines are smaller and more slender although in both skulls they occupy the same space (48) ;mm.).. ihe; yothameieen) 1s preserved on the right side and here also it is very much smaller Fic. 5.—Glanosuchus macrops. S.A.M. than the anterior incisors. As in 11843. (X¢-)_ 4 Lateral view. b, the type all the teeth have a finely Dorsal view. Left side incorporating . features determined of the right side. serrated posterior border. The occiput is less deeply concave than in the type skull. The preorbital depressien is very shallow and the maxillary surface smooth over the bulge where the long canine root is housed. S.A.M. 11964. Locality unknown. Coll. unknown. This is a weathered snout showing the upper dentition to be 1.5+1, C.I, p.c.2+?, with the 6th incisor very small.as in the two foregoing specimens. Although appreciably smaller than the type the dentition agrees well with that of the type so that I am referring it to the species macrops. S.A‘: M. 903: *Seekoeigat, Prince’ /Albert)))Colly Mu Bless A very much weathered snout, which on account of the presence of a small 6th incisor can be included in the genus Glanosuchus. Genus Ptomalestes. Gen. Nov. Fairly large Pristerognathids with the dental formula i.5, c.1, p.c.6; the incisors are subequal and fairly weak, the canine strong and the postcanines small slender teeth well spaced. THE PRISTEROGNATHID THEROCEPHALIANS 79 Skull fairly large (Max. Length 258 mm.); preorbital depression fairly deep but without abrupt margins and shallowing in the direction of the canine; frontal just entering orbital border; postfrontal quite well developed; snout broader than high, only slightly narrowed between orbit and canine; orbits well in posterior half of skull. Mandibular symphysis formed solely by dentaries, weak, unankylosed, mentum sloping. Sagittal crest of parietals only moderately high and narrow. Ptomalestes avidus Sp. Nov. (Fig. 6a—d.) Genotype. S.A.M. 11942. Steenboksfontein, Laingsburg. Coll. Boonstra. A fairly well preserved skull, but it has been subjected to a slight shear and some weathering, together with some cervical vertebrae, part of the pectoral girdle, humeri, radii and ulnae. In lateral view (Fig. 6a) the maxilla is seen to be low and long and has a long overlap on the premaxilla; the small slender _post- canines are _ irregularly spaced and occupy 51 mm. with a diastema of 12 mm. to the canine; anterior to the canine the maxillary edge curves upwards (‘‘step’’ of Watson) and the diastema is 13 mm. on _ the left and 15 mm. on the Hight side. On the pre- maxilla there are 5 incisors, fairly small and all more or less the same size. All the teeth have serrated sharp posterior edges. The prefrontal is large with a dorsal and a lateral face. The jugal has a stout dorsal ramus forming the Fic. 6.—Ptomalestes avidus. ype: wovAeML: . LEO4Z)) (SC 4) a, a. Wateral view.’ b;, Dorsal lower and hind part of view. (G7) Vellital view. id, Occipitaly view: the postorbital bar; the posterior ramus extends moderately far backwards underlying the anterior 80 ANNALS OF THE SOUTH AFRICAN MUSEUM part of the anterior squamosal ramus forming the infratemporal arch; the anterior jugal ramus is short. , The postorbital is of moderate size forming the postero-dorsal corner of the orbital border and has only a short posterior tongue applied to the lateral face of the parietal. The squamosal has a deep descending process to hold and support the quadrate and quadratojugal, but I have not been able to determine the relations of these three bones as seen in lateral view. The temporal fossa is short but roomy. The angular has a fairly large external face with radiating ridges and posteriorly a deep notch and a well developed reflected lamina with a thin ventral edge. The surangular forms, in lateral view, the dorsal girderlike border of the jaw between the dentary and the articular. The lower edge of the dentary is straight and the posterior edge concave between the angle and the strong coronoid process, which, extending into the temporal fossa, stands out well dorsal to the surangular and the dentigerous border of the dentary. In dorsal view (Fig. 6b) the parietals are seen to form a moderately high, fairly sharp sagittal crest. The frontal is fairly short, narrow and is practically excluded from the orbital border. The postfrontal is quite well developed and the prefrontal has a large dorsal surface. The occipital edge is fairly deeply concave with only the dorsal edges of the interparietal and tabulars seen in dorsal view, but the posttemporal arch does not sweep very much in posterior direction. The temporal fossa is short but wide. In ventral view (Fig. 6c) the dentaries are in position so that the anterior and lateral portions of the palate are not exposed. The choanal openings are short and the interchoanal bar formed by the prevomers stout. Posteriorly the prevomers form a truncated spatulate sheet of bone underlying the palatines and pterygoids. The palatine is of fair size with its outline roughly that of a parallelogram; it carries an oblique ridge which is continued on the pterygoid. The anterior pterygoidal rami form a median flat surface laterally bounded by the above mentioned oblique ridges; no teeth can be seen on these ridges; laterally the pterygoid bounds the large suborbital vacuity; posteriorly the anterior ramus carries an eminence in which a number of small teeth are implanted mainly in a curved row. The transverse pterygoidal ramus forms a moderately strong bar inclined ventrally in lateral direction; its corner does not descend so far as the ventral edge of the dentary where this forms its obtuse angle. Posteriorly the pterygoid clasps the anterior process of the parasphenoid and forms the lateral sheets of bone of the fairly deep median keel. The quadrate ramus of the pterygoid has a fairly straight lateral edge, well removed from the braincase, and meets the quadrate. THE PRISTEROGNATHID THEROCEPHALIANS 81 I have not been able to determine the relations of the epipterygoid to the quadrate ramus of the pterygoid. The transversum is biramic; a transverse ramus is applied, as a vertical sheet of bone, to the anterior face of the transverse pterygoidal ramus; a longitudinal ramus forms a girder, ascending in the skull in anterior direction to meet the palatine at its junction with the maxilla, and laterally bounding the large suborbital vacuity. The parasphenoid forms a fairly deep median keel anteriorly clasped by the pterygoids, and presumably forms the underface of the basisphenoidal tubera. The basisphenoidal tubera are situated well below the level of the basioccipital condyle; their posterior borders are developed as a sharp ridge forming the anterior part of the deep rim bounding the fenestra ovalis; the medial part of the rim of the fenestra is formed by an equally sharp ridge formed by the basioccipital confluent with that of the basisphenoid. Medially to the deep rim of the fenestra ovalis the under surface of the basisphenoid and basioccipital appears as a wide deep groove. The strong opisthotic abutting against the basisphenoid and basioccipital forms the outer and more dorsally situated part of the border of the fenestra ovalis. Laterally the paroccipital bar abuts against the everted sheet of the squamosal which forms the inner face of the auditory groove. The lower jaw, being in articulation, covers most of the quadrate in ventral view, but medially the inner rounded edge of the median condyle can be seen and laterally the posterior edge of the lateral quadratic condyle. The sharp ventral edge of the reflected lamina of the angular is shown in fig. 6c as is also the unankylosed mandibular symphysis. In occipital view (Fig. 6d) it is evident that the occiput is low and that the quadrate complex lies well below the basis cranu and that the squamosal bulging outwards lateral to the roomy temporal fossa has not carried the quadrate laterally with it. The limits of the interparietal, supraoccipital and tabulars cannot all be determined, but where uncertain are probably as shown in broken lines. The posttemporal fenestrae are large, the auditory groove fairly shallow but above the foramen magnum there is a very deep circular depression. The paroccipital bar is strong and the descending sheet of the squamosal covers the greater part of the face of the quadrate. The deep rms of the fenestra ovalis are clearly seen and are formed by the basisphenoid and basioccipital. S.A.M. 11460. Buffelsvlei, Beaufort West. Coll. Boonstra and Marais. This specimen consists of an imperfect anterior half of a skull smaller than that of the type. There are 5 incisors and 5 postcanines; the incisors are fairly weak, the first 4 subequal and the 5th much smaller; the postcanines 2—Annals 82 ANNALS OF THE SOUTH AFRICAN MUSEUM are small and slender; on both sides the 3rd is missing; they occupy 20 mm. on the left and 22 mm. on the right side; the diastema anterior to the canine as well as that posterior to it measures 12 mm. On the assumption that its smaller size and the differences in the dentition may be due to its being a juvenile I am including this specimen in the species avidus, but with corroborative evidence from a more fully preserved skull it may well prove to be a smaller species specifically distinct from the genotype. S.A.M. go12a. Klein-Koedoeskop, Beaufort West. Coll. Boonstra. This specimen is an anterior weathered part of a snout in which the incisors agree with those of the type. Genus Pnsterosaurus Gen. Nov. Fairly large Pristerognathids with the dental formula i.?, c.I, p.c.3 (in all probability there were 6 incisors); the canine small and weak; the postcanines also small and weak and closely packed; in all probability the incisors were also small and weak. Skull fairly large (Max. Length 225? to 255? mm.); preorbital hollow shallow to very shallow without definite margins; frontal with small entry into the orbital border; postfrontal well developed; snout broader than high, not narrowed between orbit and canine; orbits in all probability in anterior half of skull. Sagittal crest of parietals high and sharp. Pristerosaurus microdon Sp. Nov. (Fig. 7.) Genotype. S.A.M. 9083. Rietkuil, Beaufort West. Coll. Boonstra. A well preserved skull lacking only the precanine portion of the snout. In lateral view (Fig. 7a) the maxilla is seen to be short but high. The canine is small, probably not more than 15 mm. long and antero-posterior diameter 6 mm; the three small postcanines occupy 13 mm. on the right side and 15 mm. on the left. The prefrontal is large with a rounded ridge separating a dorsal from a lateral face. The jugal is a large triradiate bone; the dorsal ramus forms the major part of the postorbital bar; the posterior ramus extends far posteriorly and with the anterior ramus of the squamosal which overlies it forms the infratemporal arch. (In this feature this early Therocephalian exhibits a condition found in the later Cynodonts and not found in the Gorgonopsians.) The anterior jugal ramus forms the infraorbital bar with its extremity wedged in between the lacrimal and maxilla. The postorbital forms the postero-dorsal part of the orbital margin and curving towards the median line is applied to the temporal face of the sagittal crest formed by the parietal, but does not extend far in posterior direction. THE PRISI!EROGNATHID THEROCEPHALIANS 83 The squamosal has a deep descending process supporting the quadratojugal and quadrate, but the displacement of the lower jaw has disturbed the articulatory region so that the relations here are difficult to determine. SAM.4083. Fic. 7.—Pristerosaurus microdon. Type. S.A.M. 9083. (x 4.) a, Lateral view. b, Dorsal view. Left temporal arch corrected on basis of the right side. The angular has a large external face with a notch situated above the reflected lamina; a curved ridge separates a dorsal hollow from the lower part of the external surface. The temporal fossa is long and roomy. In dorsal view (Fig. 7b) the high sharp parietal crest is evident. The frontal is long with a small entry into the orbital border and the prefrontal has a large dorsal face separated from the lateral face by a rounded ridge. The occiput is deeply concave with the posttemporal arch sweeping far posteriorly. Above the foramen magnum the occiput is very deeply excavated; the posttemporal fossae are small; the paroccipital is strong, with its distal end everted where it meets the squamosal to form the inner boundary of the deep auditory groove. Little of the ventral surface could be prepared; the condyle is moderately large and single; the basisphenoidal tubera are very strong (as in Trochosaurus) and the fenestra ovalis has a prominent sharp ventral rim; the parasphenoidal keel is deep and narrow. The quadrate complex is posteriorly overlapped by a sheet of the squamosal descending far ventrally. 84 ANNALS OF THE SOUTH AFRICAN MUSEUM The specimen in the British Museum B.M.N.H. R.4100 which has been referred to Lycosuchus and then to Scymnosaurus can obviously not be included in either of these two genera and undoubtedly falls within the limits set for the genus instituted here and constitutes a second species — Pristerosaurus watsom (Broom). Measurements. Glanosuchus, Ptomalestes and Pmsterosaurus Pristero- Glanosuchus. Ptomalestes. saurus. 637 11843 I1964 903 I1942 11460 9083 Pr. Mx:==B.Ovscondyle .: inet 310. .200)0) 70) tn See ee ER LON Pr: Ni Post! Squsedge..° 8. 32m? 32 heya ah ee eee Save Pr. Mx.2-Ant: Orb, bonder 9.) 44.172 \ a0 — i | aero Gm Pr.Mx.—Pin.For. AND) SDAA EN N21 dm Z20%) eee = be Re Sel Width of Snout over Canines 82 80 i 65 70 48? Width of Snout over last P.C. 85 O57) nee 95 75 58 Width *Tmteronbitale 2) ts 52 54 a 44 — 40 Width Intertemporal over Pinon) SiMe arc i OF ae 23 a 24 a 20 Height of Snout at Post. Edge Of) Camimen Wye en ee ae 61 Se 55 47 55? Height)atiP Ow bam © se 5.5 5. eS 527 — — Joven 45 Length of Upper Incisor series |. 55 47 AD he 40 35 = Lee No) 50 45 45 40 35 i Diastema C.-T a 10 Ty th wae 13 r2 a r 7 ke TO nO 15 13 sos Diastema C= Pee: hy a he ee 12 12 2 r 16 18 15, 8 be 12 1 2 Meneth).or ue © Series Wy ee — SS = 20 16 Te Ao WGP get aie 51 22 15 Ant:-Post, Diam, of) .Canine 11) 20 18 Sp) aS 13 12 af 1 Me 17 TO Lh) as 15 12 = Genus Thernoides Boonstra. Boonstra, 1953, p. 58. Genotype. TZ. cyniscus Boonstra. DIAGNOSIS Fairly large Pristerognathids with the dental formula 1.6, c.1, p.c.6; the incisors are subequal, moderately long and slender, the canine long and strong and the postcanines well spaced, small, slender teeth. THE PRISTEROGNATHID THEROCEPHALIANS 85 Skull fairly large (Max. Length 275 mm., as reconstructed); preorbital depression fairly deep but without abrupt margins and shallowing in the direction of the canine; frontal probably just entering orbital border; snout as broad as high, only slightly narrowed behind canines; orbit well in posterior half of skull. Mandibular symphysis fairly weak, but mentum fairly upright. Quadrate situated very low down in skull, far ventral of the occipital condyle. Therioides cyniscus Boonstra. (Fig. 8.) Beonsird, 1953, Pp. 56. Holotype. S.A.M. 11883." Vindraersfontein, Beaufort West. Coll. Boonstra. A slightly distorted and somewhat weathered skull, together with part of the pectoral girdle, humerus, radius, ulna and part of manus. In lateral view (Fig. 8) the maxilla is seen to be fairly low but long with a moderate overlap on the premaxilla; the small and slender postcanines are evenly spaced, occupying 39 mm. on the left and 40 mm. on the right side with a diastema of 10 mm. on the left and 9 mm. on the right side to the canine; the canine is robust and 48 mm. long; no “‘step’’ anterior to canine, and the diastema to the last incisor is small (10 mm. on left and 9 mm. on right). On the premaxilla there are 6 incisors occupying 33 mm. on the right and 5 incisors occupying 30 mm. on the left side; they are slender with a mean length of 15 mm.; the 6th is weaker than the anterior teeth. All the teeth have serrated sharp posterior edges. The septomaxilla has a fairly large facial exposure and the septomaxillary foramen is large. SAM. 96S. iy ae : Fic. 8.—Therioides cyniscus. Type. S.A.M. 11888. (x 4.) Lateral view. Incorporating features of the right side. The prefrontal is fairly large with a dorsal and a smaller lateral face. The jugal bar is strong. The squamosal has a very deep descending process 86 ANNALS OF THE SOUTH AFRICAN MUSEUM carrying the quadrate very far down in the skull with the result that the articulation is situated on the same level as the ventral border of the dentary. I have recently described this area in detail (Boonstra, 1953). The occiput is only partly preserved. The paroccipital is strong and its ventro-median corner well developed and this, together with the basioccipital and basisphenoid processes, forms the strong rim of the fenestra ovalis, which is thus situated low down in the skull, well below the level of the occipital condyle. The basisphenoidal tubera are well developed and the parasphenoidal keel deep. The posttemporal fenestra is fairly large and the auditory groove - well developed. The lateral sweep of the squamosals has not carried the quadrates far laterally. Genus Pristerognathus Seeley. Seeley, 1895, p. 994. Genotype. P. polyodon Seeley. DIAGNOSIS : Laer . 6 3? Pristerognathids with the dental formula oc €.2, eae Pristerognathus polyodon, Seeley. The holotype is in the British Museum (Natural History), B.M.N.H. R. 2581, Tamboerfontein, Beaufort West, and consists of a poor weathered snout. This genus has only historical importance since the poor genotype shows only the following diagnostic characters: dental formula, ae Ci, pod, In the premaxilla the first incisor’s root has a circular cross section and it lies close to its fellow; incisors 2-5 are ovate in section and the 6th is smaller than the anterior teeth; they occupy 35 mm. In the dentary there are three incisors decreasing in size in posterior direction; they are long and curved with finely serrated posterior edges; they occupy 17 mm. The upper canine has a diameter of 14 mm. and a serrated posterior edge. There is a step in the maxillary border anterior to the canine. The symphysis, formed chiefly if not entirely by the dentary, is unankylosed and the mentum is sloping. The width of the snout is 55 mm. and the height 46 mm. The bulge of the maxilla above the canine has a sub-crocodilian pitted surface. Pristerognathus baim (Broom). Broom, 1904, p. 87. Holotype. S.A.M. 583. Locality unknown. Coll. unknown. THE PRISTEROGNATHID THEROCEPHALIANS 87 A very poor weathered snout fragment. There are roots of 6 upper and 3 lower incisors. The upper incisors occupy 34 mm. (measured round the curve); the 6th is smaller than the preceeding teeth; the first smaller than the succeeding 3 and lies close to its fellow. There is no reason for this fragment to bear a distinct specific name and I regard bain; as a synonym of polyodon. I propose that the generic name Pristerognathus be retained for Seeley’s historic specimen and be used as a suitable label for all Pristerognathid Therocephalian specimens in which no other diagnostic characters are shown except the presence of 6 upper incisors occupying a space of 35 mm. For specimens showing other additional characters other generic names should be used as has already been done viz. Alopecognathus, Cynariognathus, etc. The following list of specimens comprises a number of fragmentary and weathered snouts which on the nature of the incisors can only be referred to the genus Pristerognathus as characterised above. Se ven 63r Koup, Coll. Joubert: Ly 751 Seekoeigat, P.A. Coll. Du Plessis. A 752 Seekoeigat, P.A. Coll. Du Plessis. pews iiettontem, P.A. Coll. J. H. Whaits. ,, 1213 Unknown locality. », 3432 Janwillemsfontein, P.A. Coll. Haughton & Whaits. », 9015 Klein-Koedoeskop, B.W. Coll. Boonstra. ,, 9084a Rietkuil, B.W. Coll. Boonstra. peerir Ore Vier, P2A: Coll. Boonstra. », 11456 Buffelsvlei, B.W. Coll. Boonstra & Marais. », 11586 Koedoeskop, B.W. Coll. Boonstra. Menrrea2 7 Lammerkraal, P)A.~ Coll. Boonstra. feito |, Veldmansnvier, P2A. Coll: Boonstra. Pete a berdewater,. PA. Coll. Boonstra, ge L872 is - ite WORSE) Ws 4 ee ke 74 : in - if PATIO TS % A Y 3 yt o76 ‘3 ,, 11936 Bosluiskraal, Laingsburg. Coll. Boonstra, ,, 11950 Klein-Koedoeskop, B.W. Coll. Boonstra. », 11957 Abrahamskraal, P.A. Coll. Le Roux. ,, 11960 Dikbome, Laingsburg. Coll. Boonstra. ,, 11963 Dikbome, Laingsburg. Coll. Boonstra. ,, 11965 Skoppelmaaikraal, Laingsburg. Coll. Botes. #2210906... Seekoeigat,,,P.A...Coll. Bz) Bain. py. EIGQO7), »Seekoecigat,..PAn,Colly io Baim: 88 ANNALS OF THE SOUTH AFRICAN MUSEUM Genus Alopecognathus Broom. Broom, I9I5, p. 116. Genotype. A. angusticeps Broom. The holotype of this species :s in the American Museum of Natural History. A.M.N.H.5559. Coll. Whaits. DIAGNOSIS Moderate to fairly large Pristerognathids with the dental formula 1.6, C.I, p.c.4-5; the anterior incisors are fairly long, slender teeth (the ist incisors, as in all Pristerognathids, are smaller than the 2nd to 5th) but the 6th is appreciably to very much smaller; the canine is long and strong; the postcanines are small and fairly weak, not close set. Skull moderate to fairly large (Max. Length 240? mm. to 276 mm.); preorbital hollow varies from a fairly shallow hollow shallowing evenly in the direction of the canines and without abrupt borders to a deep depression with sharply demarcated borders especially anteriorly; septomaxilla with well developed facial exposure and septomaxillary foramen well developed; frontal with small entry into orbital border or just excluded from it; prefrontal large with well marked dorsal and lateral face; postfrontal moderately to well developed; snout wider than high, broader over last postcanines than over canines; orbits just entering anterior half of skull. Mandibular symphysis weak and mentum sloping very much. Sagittal crest of parietals moderately high but with sharp edge. Quadrate low down, but well above the level of the lower border of the dentary. Temporal fossa fairly long and wide. Squamosal with everted lateral edge. Alopecognathus angusticeps Broom. The holotype in the American Museum has been redescribed by me some years ago when studying the South African specimens sold by Dr. Broom to the American Museum (Boonstra, 1935; p. 2). DIAGNOSIS There are 5-6 postcanines; maximum length of skull 275 mm.; preorbital depression fairly shallow to deep, extending in direction of canine; postfrontal well developed; squamosal laterally everted. S.A.M. 9112 (Fig. 9). Stinkfontein, Prince Albert. Coll. Boonstra. A fairly well preserved anterior three quarters of a skull. From the figures and from the accompanying list of measurements it is clear that this specimen from Stinkfontein is co-specific with the genotype, THE PRISTEROGNATHID THEROCEPHALIANS 89 notwithstanding certain less important differences e.g. in our specimen there are 6 postcanines occupying 40 mm., and the preorbital depression is deep. Pit | es th SAM 4112. Fic. 9.—Alopecognathus angusticeps. S.A.M. QIi2. (x a, Dorsal view. b, Lateral view. Cols F This specimen allows us to correct two points in my description (Boonstra, 1935) of the genotype viz. with the anterior part of the sagittal crest preserved it is now clear that in my reconstruction the height of the crest should have been much higher; the preserved frontonasal suture in this specimen shows that the broken line indication in my figure of 1935 should have been further forward so that the frontal is a long narrow bone and the nasal shorter than I thought in 1935. Alopecognathus angustioriceps (Boonstra). Boonstra, 1953, p. 63. Holotype. S.A.M. 9342. (Fig. 10.) Kroonplaas, Beaufort West. Coll. Boonstra. A very good undistorted and practically complete skull. go ANNALS OF THE SOUTH AFRICAN MUSEUM DIAGNOSIS There are 5 postcanines; maximum length of skull 252 mm.; preorbital depression shallow, continued anteriorly as a shallow groove; postfrontal small; squamosal with lateral bulge. age ~~ ~ x” any we Ws C Fic. 10.—Alopecognathus angustioriceps. Type. S.A.M. 9342. (x 4.) a, Lateral view. 0b, Dorsal view. c, Ventral view. In lateral view (Fig. 10a) the maxilla is seen to be fairly low and of moderate length with a moderate overlap on the premaxilla; it carries 5 small THE PRISTEROGNATHID THEROCEPHALIANS Qi (1st very small) postcanines occupying 27 mm. on the left and 29 mm. on the right side. The diastema between canine and the first postcanine is 10 mm. on the left and rr mm. on the right. The canine is well developed with a length of 30? mm. and antero-posterior diameter of 14 mm. Anterior to the canine is a diastema of 12 mm. on the left and 14 mm. on the right. The premaxilla carries 6 teeth, the anterior 5 are long and slender and the 6th small and slender occupying 33 mm. on both sides. The septomaxilla has a well developed facial surface and intra-nostril spur. The prefrontal has a small lateral face. The jugal has a short dorsal ramus and a well developed posterior ramus overlain by a moderately long anterior ramus of the squamosal. The postorbital forms most of the postorbital bar. The squamosal has a deep descending ramus supporting the quadrate complex. I have recently figured and described the suspensorial region (Boonstra, 1953). The dentary is large with a strong coronoid process; the symphysis is weak and the mentum sloping. In dorsal view (Fig. 10b) the long slender shape of the skull is evident; the orbits just enter the anterior half of the skull; the temporal fossa is long and roomy; the everted anterior ramus of the squamosal forming most of the lower temporal arch is noteworthy (seen also in the genotype). The frontal is just excluded from the orbital margin; both the frontal and nasal are narrow. The prefrontal has a large dorsal face bounded laterally by a sharp ridge. The postfrontal is small and the postorbital well developed but extending little along the lateral face of the parietal. The parietals are narrow and form a sharp sagittal crest which is, however, not high. The upper occipital edge is very deeply concave. In ventral view (Fig. 10c) it is seen that with the dentaries in position it has not been found feasible to clean the anterior part of the palate. Just anterior to the transverse rami of the pterygoids the pterygoids bear two dentigerous ridges. The lateral corner of the transverse pterygoidal ramus lies in the plane of the angle of the dentary but does not descend to the level of the ventral edge of the dentary. Posteriorly the pterygoid clasps the parasphenoidal rostrum which lies in the median line and the quadrate ramus, with a slightly curved lateral edge, extends to the quadrate. The parasphenoidal keel is deep. The basisphenoidal tuber, the medio-ventral corner of the opisthotic and the antero-lateral corner of the basioccipital form three distinct protuberances which constitute the tripartite rim bounding the fenestra ovalis. The paroccipital bar is strong; the basioccipital condyle fairly weak. The squamosal extends down far to cover the posterior face of the quadrate complex. The squamosal sweeps far posteriorly to make the occiput deeply concave, and the occipital condyle and the quadratic condyles are situated far anterior to the posterior limits of the skull. The quadratic condyles also lie in a plane well ventral to that of the occipital condyle. Q2 ANNALS OF THE SOUTH AFRICAN MUSEUM Measurements. Alopecognathus A.M.N.H. S.A.M. S.A.M. 5299 OTT 739344 Prbix.—-B.O., condyle}, Lib Avapeeah Me iebe 234 — 215 Pr. Mx. Pesto! | etlge my man. amare Bee be: 275 260 252 Py. Mx¢ 2 AntOrbs: (border ie ee ne annie 140 138 [NG PG. Mix POT. eal ok UN cage mer en ka 210 210 180 Width ‘of -Snout vover ‘Camimes in) 04) sa 55 56 50 Width vot ‘Snomitover jlasth biG. ja. ao ae 68 70 53 Width, TnterorbitalQe\ oe Weck se. i a ane 35 45 40 Width, Intertemporallover sransHor.. 10? 15 17, Height of Snout at Post. edge of Canine . . 50 51 48 Height at PostiOrp. ban) iii. saurwai\ Sepia 45 Ae 45 Length of Upper Incisor Series Le 33 34? 33 I. a SE OS Diastema “C2er IF r2 1 2 I. = = 14 Diastemra (CEC. Ih. 12 16 10 ite =o = II Peneth, of vb ©) Series lig 31 40 27 r. wax 39 29 Ant.-Post.Diam. of Canine I. 12 15 13 ie = — 13 Genus Cynariognathus Broom. Broom, 1931, p. 161. Genotype. C. platyrhinus Broom. The holotype of this species is in the American Museum of Natural History. A.M.N.H. 5502. Collected by J. H. Whaits and sold to the American Museum by Dr. Broom. DIAGNOSIS Medium sized Pristerognathids with the dental formula i.5-6, c.1I, p.c.6-9; the posterior incisors are smaller than the anterior ones; the canine is long and strong; the postcanines are fairly strong and closely set. Skull of medium size (Max. Length 260-290 mm.); preorbital depression shallow; septomaxilla fairly small; frontal with a moderate entry into the orbital border; prefrontal large with well marked dorsal and lateral face; postfrontal moderately developed; snout broader than high; broader over last postcanines than over canines; orbit in posterior half of skull; skull low over postorbital bar; orbits high up in the skull. Premaxilla weak. Angle THE PRISTEROGNATHID THEROCEPHALIANS 93 of dentary obtuse with posterior edge of dentary not deeply concave. Dentary not shallow behind the canine. Cynariognathus pauciondens. Sp. Nov. (Fig. II.) Holotype. S.A.M. 11560a. Kroonplaas, Beaufort West. Coll. Boonstra. A weathered snout with some parts of the postcranial skeleton. DIAGNOSIS Six closely packed postcanines occupying 27 mm. on the left and 28 mm. | on the right side; breadth of snout over the canines 50 mm.; height of snout at posterior edge of canine 40 mm.; preorbital length of skull 99 mm. -ee 7? See . Fic. 11.—Cynariognathus paucioridens. Type. S.A.M. 11560a. (x 4.) a, Dorsal view. b, Lateral view. This skull has a long low maxilla, the lacrimal is large and the prefrontal fairly small; the dentary is fairly lightly built; the preorbital groove is fairly deep. Little more can be determined from this specimen and I would have hesitated to name it were it not that a second specimen, also with 6 closely packed postcanines, proves that we have here a definitely new species of Cynariognathus. Q4 ANNALS OF THE SOUTH AFRICAN MUSEUM S.A.M. 11586. Koedoeskop, Beaufort West. Coll. Boonstra. The antorbital weathered half of a skull. Here there are 6 closely packed, fairly weak postcanines occupying 28 mm. and there are apparently 5 incisors occupying 30 mm.; the height of the snout is 40 mm. and the width 53 mm.; the antorbital length is 100? mm. 5.A.M. 1080. Fraserburg Rd., Prince Albert. Coll. J: H. Whaite: A weathered antorbital part of a skull together with a complete hindfoot. In this specimen there are 7 postcanines; they are closely set and robust and occupy 26 mm. The height of the snout is 35 mm. and its breadth 48 mm.; the antorbital length as estimated is only 75 mm. The shorter snout and the larger number of postcanines make inclusion in the above species provisional. Cynariognathus spp. S.A.M. 3713. Bloukrans, Prince Albert. Coll. Haughton. S.A.M. g088a. Klein-Koedoeskop, Beaufort West. Coll. Boonstra. S.A.M. 11968. Locality and Collector unknown. These three incomplete snouts are included in the genus Cynanognathus as the height of the snouts is much smaller than the breadth and because in the postcanine series the teeth are closely packed. But as the posterior postcanines are not preserved the determination of the species remains uncertain. Genus Pristerognathotdes Gen. Nov. Genotype. Alopecognathus minor Haughton. DIAGNOSIS Medium sized Pristerognathids with the dental formula 1.6, c.1, p.c.5-6; the incisors are small to fairly strong teeth, with the 6th incisor only slightly to much smaller than its predecessors; the canine is fairly long; the postcanines are small, weak and well spaced. Skull of medium size (Max. Length 222-287 mm.); preorbital depression shallow to fairly shallow continuing as a groove in the direction of the canine; septomaxilla fairly small; frontals with a fairly small entry into the orbital border; prefrontal large with well marked dorsal and lateral face; postfrontal moderately well developed; snout wider than high; broader over last postcanine than over canines; orbits in posterior half of skull or just entering anterior half; skull very low over postorbital bar; orbits high up in skull. THE PRISTEROGNATHID THEROCEPHALIANS 95 Sagittal crest high and with sharp edge. Quadrate fairly low down, but well above the level of the lower border of the dentary. Temporal fossa fairly long and moderately wide. Premaxilla weak. Squamosal with lateral edge not everted. Angle of dentary squarish with fairly deep concave posterior edge sweeping | far posteriorly towards the coronoid process. Dentary shallow behind _ lower canine. Pristerognathoides minor (Haughton). (Fig. 12.) Haughton, 1918, p. 180. Holotpye. S.A.M. 3415. Klipbank, Beaufort West. Coll. Whaits. A good skull lacking only the temporal arches. | DIAGNOSIS | There are 4 postcanines; maximum length of skull 240? mm.; preorbital | depression deep with very definite and abrupt margins; postfrontal large. Since Haughton’s description I have carried the development of the skull _ further, exposing more of the palate, and removed the matrix from the | preorbital depression. In lateral view (Fig. 12a) the maxilla is seen to be low and long with _a long overlap on the premaxilla; on the right 4 postcanines are preserved but there is a space between the rst and 2nd which may have housed an additional tooth; the length of the series is 28 mm.; on the left 3 postcanines are preserved with a gap between the 1st and 2nd; the length of the series is 26 mm.; the postcanines are short but stout. Between the Ist postcanine and the canine there is a diastema of 8 mm. on the left and 12 mm. on the right side. |The canine is long and strong (diameter 11-12 and length 24? mm.). Anterior to the canine there is a diastema of 10 mm. to the last incisor. There are 6 incisors of which the 5th and 6th are shorter than the anterior ones and the series occupies 32 mm. on both sides. All the teeth have finely serrated posterior borders. The septomaxilla has a fairly large facial exposure and a well developed intra-nostril spur; the septomaxillary foramen is fairly large. The preorbital depression is a deep hollow, especially on the right side, with a sharp rim, especially along its dorsal and ventral borders; a low longitudinal ridge divides the hollow into two equal parts on the right side, but on the left the lower part is the larger; anteriorly the hollow ends abruptly and is not continued in the direction of the canine. The bulge of the maxilla above the canine is not rugosely pitted. The anterior orbital border just enters the anterior half of the skull. The 96 ANNALS OF THE SOUTH AFRICAN MUSEUM postfrontal is very well developed with a large entry into the orbital border; whereas the postorbital appears to have only a very small entry into the orbital border, forming the posterior part of the postorbital bar. The edge of the sagittal crest is not preserved but was probably sharp and fairly high with little flanking of the parietal by the postorbital. S.AM.3LI5. ’ « & tos SA ee ey oe SD) oer Ce f . vetted Got ee: frp ao Se > = or Pola a Fic. 12.—Pristerognathoides minor. Type. S.A.M. 3415. (x #4.) a, Lateral view. Incorporating features of the left side. b, Dorsal view. c, Ventral view. The articulatory region is not well shown since the bone has been stripped. The angular has a deep notch and a small bone above the articular I have THE PRISTEROGNATHID THEROCEPHALIANS 07 ident:fied as the quadratojugal. The squamosal has a deep descending process posteriorly supporting the quadrate complex. In dorsal view (Fig. 12b) the width across the temporal arches (as restored) is great making a roomy although short temporal fossa, whereas the snout, unconstricted behind the canines, is relatively narrow. The frontal is fairly small and probably just excluded from the orbital border. The postfrontal is very large and the postorbital weak, both with little posterior prolongation along the lateral face of the parietals. The prefrontal has a well developed | dorsal face separated from the smaller lateral face by a sharp ridge which _ also forms the sharp dorsal rim of the preorbital depression. The nasals are large, expanded anteriorly and posteriorly. The dorsal occipital edge is deeply concave. In ventral view (Fig. 12c) development has exposed all but the choanal region of the palate. The posterior ends of the prevomers form a shovel-shaped sheet of bone underlying the palatines and pterygoids. The _ pterygoids have long anterior rami, which bear, in their posterior part, a dentigerous ridge; the transverse pterygoidal ramus is not very strong or _ wide and its anterior face is lined by a descending sheet of the transversum which extends anteriorly as a girderlike ramus to form the outer border of the large suborbital vacuity. There is a small interpterygoid vacuity. The outer edge of the quadrate ramus of the pterygoid is slightly concave, thus increasing the size of the temporal space. In the median line the parasphenoid underlies the basisphenoid and anteriorly forms a deep keel. Surrounding the fenestra ovalis there is a fairly strong rim formed by the basioccipital, opisthotic and basisphenoid-parasphenoid. The paroccipital bar is strong and its ventro-anterior corner abuts against the quadrate. Lateral to the quadrate lies a small guadratojugal which does not enter the condylar surface. The squamosal covers most of the posterior face of the quadrate complex. S.A.M. 4332 (Fig. 13.) Wilgerbos, Prince Albert. Coll. Haughton. A good skull of which some aspects of the internal structure have been described by Haughton (Haughton, 1918). The skull is very similar to that of the holotype of Pristerognathoides minor. On the left side there are 5 postcanines occupying 29 mm. The distance between canine and postcanines is 15 mm. on the left side and between canine and postcanines is 15 mm. on the left side and between canine and incisor the diastema is 9 mm. and the 6 incisors measure 30 mm. The preorbital depression is fairly deep, but shallows in the direction of the canine. As is evident from the figures there are several other minor differences in the proportions of some of the surface bones, but notwithstanding these I am referring this skull to P. minor. 8—Annals ANNALS OF THE SOUTH AFRICAN MUSEUM SAM 4332. oan WN. : ysis = ie ee oe eS Win se mi sae SO xP at [! = Vj ‘ Zs ” se Fic. 13.—Pristerognathoides minor. S.A.M. 4332. (x a, Dorsal view. 0b, Lateral view. 3+) S.A.M. 3435 (Fig. 14). Jakkalsfontein, Prince Albert. Coll. Rogers. A well preserved antorbital part of a skull laterally compressed. SAM. 3135. Fic. 14.—Pnisterognathoides minor. S.A.M. 3435. (x 4.) Lateral view. Although the snout as preserved appears to be shorter and the ma il consequently relatively higher and shorter than in the type, the dentition is” very similar and the clearly demarcated preorbital depression has an abrupt anterior border as in the type specimen. I am thus referring this snout to. ny P. minor. a 4 S.A.M. 11891 (Fig. 15). Lammerkraal, Prince Albert. Coll. Boonstra & Pienaar. : ki i i: td THE PRISTEROGNATHID THEROCEPHALIANS 99 A good skull lacking only the right postorbital and temporal arches. In this skull there are on the right side roots of six postcanines. The snout is broader than in the type and the preorbital depression is fairly shallow and although there are other minor differences as can be seen from the figures and the table of measurements I am referring it to the species minor. SAM. 11541. + ee ey Canes aus oes - Soe n8 abe oe = 7 & SOSA Soe ee 5 o> tA pe ES, B a ®*eae” Fic. 15.—Pristerognathoides minor. S.A.M. 11891. (x 4.) a, Dorsal view. 0b, Lateral view. _ Pnisterognathoides roggeveldensis (Boonstra). (Fig. 16.) Boonstra, 1953, p. 60. Holotype. S.A.M. 9356a. Roggekloof, Sutherland. Coll. Walker. A fair though somewhat distorted skull lacking the posterior part of the skull roof. DIAGNOSIS Skull long and narrow; with five weak, well spaced postcanines; incisors fairly strong with the 6th only slightly smaller than the 5th; orbit well in posterior half of skull; squamosal extending well forward on lateral face of parietal; prefrontal narrow. In lateral view (Fig. 16b) the maxilla is seen to be long and high; the septomaxilla has a moderate facial exposure; the orbit is small and situated high up in the skull; the lacrimal large and the preorbital depression fairly shallow. I have recently (Boonstra, 1953) described the articulatory region. I00 ANNALS OF THE SOUTH AFRICAN MUSEUM SAM. G35bQa. Fic. 16.—Pristerognathoides roggeveldensis. Type. -S.A.M. 9356a. (x #) a, Dorsal view. b, Lateral view. In dorsal view (Fig. 16a) the snout appears long and narrow and the temporal fossa long and narrow. Pristerognathoides vanwyki (Broom). (Fig. 7) Broom, 1925, p. 318. Holotype. S.A.M. 6533. Bloukrans, Prince Albert. ‘Coll, Le“ Roux: A weathered and distorted skull. DIAGNOSIS Skull moderately long and narrow; with six weak well spaced postcanines; incisors fairly weak, with the sixth very slender; orbit just in posterior half of skull; prefrontal large. In lateral view (Fig. 17b) the maxilla is seen to be fairly short and high; the septomaxilla has a good facial exposure; the orbit is small and situated high up in the skull; the lacrimal is large and the preorbital depression shallow. In dorsal view (Fig. 17a) the skull is fairly long and fairly narrow and the temporal fossa fairly short and narrow. S.A.M. 11893 (Fig. 18). Lammerkraal, Prince Albert. Coll. Pienaar. THE PRISTEROGNATHID THEROCEPHALIANS TO SAM 6533... oe 4 “eeee?” -c° oe” e M@weooer’ Fic. 17.—Prsterognathoides vanwyki. Type. S.A.M. a, Dorsal view. b, Lateral view. 6533= (Xx Incorporating features of the left si ) uk sue de SAM. 11893. Fic. 18.—Pristerognathoides vanwyki. S.A.M. 11893. a, Dorsal view. (OS aya) 6, Lateral view. 102 ANNALS OF THE SOUTH AFRICAN MUSEUM This snout, presented to the Museum by Mr. J. Pienaar of Lammerkraal, has only two postcanines preserved, but as this is probably only due to © postmortem loss and the other characters preserved agree fairly well with those of vanwyki I am referring it to that species. S.A. M: 11689, (Fig. 19). Prince Albert Road. Coll. Hesse: SAM Hb $4. This dorso-ventrally crushed skull has the postcanines badly preserved but there were probably 6. Although the snout is longer and there are differences of actual size and proportions | am referring it to vanwyki. Fic. 19.—Pmnsterognathoides vanwyki. S.A.M. 11689. (x +4.) a, Dorsal view. b, Lateral view. Dorso-ventral crushing corrected. Pristerognathoides paruus Sp. Nov. (Fig. 20.) Holotype. S.A.M. 3611. -Bloukrans, Prince Albert. © Coll. Scholtz A weathered anterior two thirds of the skull. DIAGNOSIS Small low and fairly narrow skull; with 5 small well spaced postcanines; incisors weak with the 5th weaker than the anterior ones and the 6th very feeble, the diastema between incisors and canine great (15 mm.), and canine > slender; prefrontal large; very low over postorbital arch. The skull is small with the maximum length probably not more than 195 mm., the maxilla is long and low; the suborbital arch slender and the dentary weak; the preorbital depression quite deep. THE PRISTEROGNATHID THEROCEPHALIANS 5.AM, 360). Fic. 20.—Pysterognathoides parvus Sp. Nov. Type. MEASUREMENTS—Pristerognathoides SE Aci 3 Om. a, Dorsal view. 6b, Lateral view. Incorporating some features of the left side. 103 (X 3+) 3415 | 4332 | 3435 | 11891 | 9356a| 6533 | 11893 | 11689) 3611 Pr. Mx.—B.O. condyle oe ae Bo) 25) 200 — 200 —_ — = —= ee Pr. Mx.—Post. Sq. edge St af <. |_240 218 — 215 255 2227 | 213'2-|\ 235 — Pr. Mx.—Ant. Orb. border .. te Se |) GIS) 110 90 113 117 106 | 108? | 118 97 Pr. Mx.—Pin. For. .. &. we «|| 165 155 — | 160 — | 158 — | 1772) 148 Width of Snout over Canines as Bie 50 47 _ 55 52? | 45 50 52 45 Width of Snout over last P.C. me So 55 60 -- 60 — 52 =if/ — 59 Width Interorbital .. ope ae Ae 33 35 — 30 — 25 2} — 30 28 Width Intertemporal over Pin. For. nie 20 — — 16 _ 14 — 17 25 Height of Snout at Post. edge of Canine .. 40 45 40 39 53? 40 51 STi? 40 Height at P. Orb. bar =% “ aos Zor) +30 a 20 — — — 302) |) 30)? Length of Upper Incisor Series ate E 32 33 _— — 35 30? 35 30 35 r 32 _ 30 35 — — 34 — 35 Diastema C.—I. an as ze 1. il 8 5 — 9 9 13 9 10 r il 11 6 5 — — 13 ~ 9 Diastema C.—P.C. .. ae ae 1. 8 15 12 15 16 10 11 12 10 Tt 11 Ui 9 14 10 10 10 11 8 Length of P.C. series .. he oh jail 33% 30 22 26 27 27 — _— 25 r 29 — 27 27 DON 135 a 3325) 21 Ant.-Post. Diam. of Canine .. Ns i 12 10 10 11 12 13 12 12 5 ; Tr 11 10 9 7 12 12 12 11 6 I04 ANNALS OF THE SOUTH AFRICAN MUSEUM Genus Maraisaurus Boonstra. Boonstra, 1953, p. 62. Genotype. M. parvus Boonstra. DIAGNOSIS Small Pristerognathids with dental formula unknown but probably 1.02 feelhar p.c.4% Skull very small [Max. Length (as reconstructed) 162? mm.]; preorbital depression very deep with sharp rim; frontal with moderate entry into orbital border; postfrontal well developed; orbits large and just entering anterior half of skull. Sagittal crest of parietals fairly low, but with sharp edge. Maraisaurus parvus Boonstra (Fig. 21). Genotype. S.A.M. 11944. Buffelsvlei, Beaufort West. Coll. Marais. The posterior two thirds of a somewhat weathered skull lacking the snout. Lateral view (Fig. 216). The articulatory region of the lower jaw has recently been figured and described (Boonstra, 1953, p. 62). The prefrontal has a sharp dorsal edge separating } a small lateral from a larger dorsal ‘ face. The jugal is well developed with an anterior ramus forming the ai stout suborbital bar; a weaker SAMIIM44UL. vit dorsal ramus forming the lower : od. half of the orbital bar and a fairly long posterior ramus, overlain by an anterior ramus of the squamosal, forming the anterior and much of the ventral part of the lower temporal arch. The postorbital is small forming only a small part of the orbital margin and posteriorly extending as a weak tongue on to the lateral face of the parietal. The squamosal has a moderate descending flange carrying the quadrate complex but not extending far ventrally so that the quadratic condyle still lies well above the level of the lower border of the dentary. In dorsal view (Fig. 21a) it is seen that the temporal fossa is fairly long and wide, the inter- Fic. 21.—Maraisaurus parvus. Type. temporal region narrow and the >”: cenit Boe ae et sagittal crest low but sharp. The frontal is long and has a fairly large entry into the orbital border: the Vee tet ay at THE PRISTEROGNATHID THEROCEPHALIANS 105 postfrontal is well developed; the postorbital has only a small entry into the orbital border and its posterior tongue weak and flanks the parietal for only a short distance; the ridge on the prefrontal sharply separates the dorsal from the lateral surface. An Unidentified Pristerognathid. S.A.M. 11959 (Fig. 22). Dikbome, Laingsburg. Coll. Boonstra. Fragmentary weathered skull pieces. The dental formula is 1.6, c.I, p-C-5. The incisors are robust, with the 6th strong, occupying 40 mm. on the left and 43 mm. on the right side. There is no diastema between incisors and canine and the diastema between canine and postcanines is very small (5 mm. on the right and 6 mm. on the left side). The postcanines are irregular with the first, fourth and fifth fairly weak but the third quite robust and occupying 36 mm. on the left and 45 mm. on the right side. The articulatory region has recently been described (Boonstra, 1953)- This form appears to occupy a position intermediate between genera like Scymnosaurus and Cynariognathus. SAM, 11459. Fic. 22.—An Unidentified Pristerognathid. S.A.M. 11959 a, Lateral view (x 4.) b, Oblique view of the articular region. (x 3.) The quadratojugal is not preserved; when present it rested on the ledge on the posterior surface of the quadrate. DISCUSSION As all recent authors derive the Therapsids from the Pelycosaurs it will be of interest to compare the Pristerognathidae — the most primitive Therocephalian family — with the Pelycosaurs. From Romer’s monograph I have extracted a list of morphological features mentioned by him and am comparing the same structures as identified in the Pristerognathids. The Pristerognathids agree with the Pelycosaurs: in most forms the frontal enters the supraorbital border, but in some this entry is small and in 106 ANNALS OF THE SOUTH AFRICAN MUSEUM some the frontal is secondarily excluded; a ridge separates a dorsal from a lateral surface in the prefrontal, but the general dorsal surface is not sharply separated from the lateral surface; otic notch closed at junction of squamosal and tabular; step anterior to canine as in predaceous Pelycosaurs; quadrate situated below plane of maxillary teeth; maxilla with its large canine crowds out lacrimal to form a junction with the nasal as in the advanced Pelycosaurs; squamosal covers most of posterior face of the quadrate; choanae well forward; expansion of posterior end of prevomer; quadrate ramus of pterygoid reaching quadrate; epipterygoid narrow, but not a typical columella cranu and reaches parietal; quadrate in contact with paroccipital; angular notch as in Sphenacodontidae. The Pristerognathids differ from the Pelycosaurs: the parietal separates the postorbital from the squamosal; there is no basal movable articulation; the palate is not Rhynchocephalian-like; supratemporal absent; dorsal surface not sharply demarcated from lateral surface and sides not steep; premaxilla not extending posteriorly between nasals; parietal short but narrow; upper edge of occiput not moved forward and occiput is thus not slanting and the surface of the interparietal and tabular are not visible in dorsal view; condyle and quadrates not in posterior position; lacrimal always with little anterior extent; the postorbital bar is always slender, never developed as a plate of bone between orbit and temporal fossa; maxilla never meets quadratojugal; jugal though extending far posteriorly never meets the quadratojugal; quadratojugal always very small; septomaxilla with large facial exposure; only the anterior ramus of the pterygoid bearing teeth and no teeth on the lateral pterygoidal flange nor any on the palatine and ectopterygoid; lateral pterygoid flange never meeting the jugal; suborbital vacuity always present and large; epipterygoid somewhat broader and reaching the parietal; angular notch present and dentary always with a strong coronoid process. It will furthermore be of interest to note in what characters the Pristerognathids agree with the Cynodonts viz. dentary with large coronoid process; sloping mentum; interparietal region narrow and sagittal crest; postorbital does not meet squamosal; preparietal absent; posterior end of prevomer spatulate and underlying palatines and pterygoids; jugal stretching far posteriorly ventral to the anterior ramus of the overlying squamosal; frontal sometimes excluded from orbital border as in Cynodonts. REFERENCES Boonstra, L. D. 1934. A Contribution to the Morphology of the Mammal-like Reptiles of the Suborder Therocephalia. Ann. S. Afr. Mus. XXXI, pp. 215-267. Boonstra, L. D. 1935. On Some South African Reptiles of the Suborder Therocephalia preserved in the American Museum of Natural History. Am. Mus, (Noy. (77 2.:) Pps ais b2: Boonstra, L. D. 1953. The Lower Jaw Articulatory Region in some Pristerognathid Therocephalians. Ann. S. Afr. Mus. XLII, 1, pp. 54-63. _THE PRISTEROGNATHID THEROCEPHALIANS 107 Broom, R. 1903. On Some New Primitive Theriodonts in the South African Museum. Ann. S. Afr. Mus. IV, pp. 147-158. Broom, R. 1904. On Two New Therocephalian Reptiles (Glanosuchus macrops and Pnsievopnatnus baini)..~ Trans. 5. Afr.’ Phil’ Soc. 15, pp. 85-88. Broom, R. 1915. On Some New Carnivorous Therapsids in the Collection of the British Museum. Proc. Zool. Soc., pp. 163-173. Broom, R. 1915. Permian, Triassic and Jurassic Reptiles of South Africa. Bull. ames vas. Nat. Hist. XXV, Il, pp. 105-164. Broom, R. 1925. On some Carnivorous Therapsida. Rec. Alb. Mus. III, pp. 309-326. Broom, R. 1931. Notices of some new Genera and Species of Karroo Fossil Reptiles. Rec. Alb. Mus. IV, pp. 161-166. HauGutTon, S. H. 1918. Some New Carnivorous Therapsida, with Notes upon the Brain-Case in Certain Species. Ann. S. Afr. Mus. XII, p. 175-216. SEELEY, H.G. 1895. On the Therosuchia. Phil. Trans. Roy. Soc. 185, pp. 987-1018. 8. The Cranial Structure of the Titanosuchian: Anteosaurus. By L. D. Boonstra, D.Sc. (With 22 text-figures.) HISTORICAL Although the Anteosaurus skull was first described by Broom as long ago as 1910, the details of the cranial structure are still very inadequately known. Watson in 1914 gave some details of the structure of the incomplete skull in the British Museum and in 1921 the same author, giving details of the snout, instituted the name Anteosaurus for the specimen hitherto thought to be a Titanosuchus. Broom in 1929 founded an additional species — A. minor — on a piece of the skull roof. Broili and Schréder in 1935 described certain skull fragments under the name TJitanognathus lotzt. In 1936 the present author figured and described a distorted skull of A. minor that had been sold to the American Museum by Dr. Broom. In the same year Broom described a good skull and lower jaw under the name Dinosuchus vorstenr. In 1948 the present author published a figure of a skull, which in 1952 was named A. abeli, and in 1953 a taxonomic account of the Titanosuchians included a number of photos of Anteosaurus skulls in the South African Museum. MATERIAL The present paper is based mainly on the large number of specimens in the collection of the South African Museum, viz. S.A.M. 2752. Vivier Siding, Beaufort West. Coll. Haughton & Whaits. Posterior two thirds of the skull without basis cranii. S.A.M. 4340. Leeurivier, Beaufort West. Coll. Haughton. A good skull, though distorted by a simple shear, with part of the lower jaw. S.A.M. 5621. Leeurivier, Beaufort West. Coll. Haughton. A snout and part of the skull roof. S.A.M. 9123. Voélfontein, Prince Albert. Coll. Boonstra. A weathered skull. S.A.M. 9139. Voélfontein, Prince Albert. Coll. Boonstra. A weathered skull fragment. 108 CRANIAL STRUCTURE OF THE TITANOSUCHIAN I0Q S.A.M. 9140. Voélfontein, Prince Albert. Coll. Boonstra. A partial disarticulated skull. S.A.M. 9329. Kruisvlei, Beaufort West. Coll. Boonstra. A good skull and much of the lower jaw. S.A.M. 11293. Boesmansrivier, Beaufort West. Coll. Boonstra. A good weathered skull, slightly dorso-ventrally compressed and distorted, with some bones of the occiput disarticulated. S.A.M. 11296. Kruisrivier, Sutherland. Coll. Boonstra & Laurenson. A very good skull and lower jaw, though somewhat distorted by a simple shear. S.A.M. 11302. Buffelsvlei, Beaufort West. Coll. Boonstra & Marais. A fair, weathered skull and lower jaw. S.A.M. 11492. Mynhardtskraal, Beaufort West. Coll. Boonstra. A fairly complete skull without lower jaw. S.A.M. 11576. Klein-Koedoeskop, Beaufort West. Coll. Boonstra. A snout with fairly well preserved teeth. S.A.M. 11577. Bulwater, Beaufort West. Coll. Boonstra & Truter. A good practically undistorted three quarters of a well preserved skull. S.A.M. 11592. Dikbome, Laingsburg. Coll. Boonstra & Du Plessis. A weathered skull, but with a good palatal region. S.A.M. 11694. Koringplaas, Laingsburg. Coll. Boonstra & Du Plessis. A very good undistorted skull without the lower jaw. S.A.M. 11929. Kruisvlei, Beaufort West. Coll. Boonstra. The greater part of a skull in intractable matrix. S.A.M. 11946. Buffelsvlei, Beaufort West. Coll. Boonstra & Marais. A nearly complete, good skull, slightly distorted by a simple shear. S.A.M. 11949. Nuwefontein, Fraserburg. Coll. Boonstra & Jooste. A partial skull, snout end occiput not in contact. GENERAL SKULL FoRM There is some difficulty in determining the correct skull form in Anteosaurus, due to the post-mortem deformation the available material has usually undergone. In only one skull (S.A.M. 11694) there appears to be little disturbance of the original symmetry. In 8 the deformation is chiefly due to dorso-ventral pressure, but accompanied by some measure of distortion. In 5 specimens the skulls have been subjected to a shearing action — mostly in the form of a simple shear. In these sheared specimens it is of interest to note that they were all lying on their left sides. In only 1 specimen, lying on its right side, the deformation was due almost wholly to compression from side to side. IIO ANNALS OF THE SOUTH AFRICAN MUSEUM In the figures the legend indicates where an attempt has been made to correct the effects of the deformation. The resulting correction made from the dorsal and ventral aspect has in some cases not produced the same result and a mean between the two results may indicate the correct condition, but not necessarily so. The Anteosaurus skull is large to very large (480-800 mm. max. length; 222-612 mm. max. width). The snout (with the mandible) is much higher than broad and thus, notwithstanding the width over the temporal region, the skull gives the impression of being high and narrow. This is in sharp contrast to the other Titanosuchians and the Tapinocephalia. In those specimens where the incisors are present the carnivorous nature is strongly evident. The prominent boss-like development of the upper part of the postorbital bar is striking, and in some specimens this rugged appearance is further strengthened by the presence of bosses on the jugal and angular and around the pineal foramen with a lesser or greater amount of swelling of the frons. The great lateral and posterior sweep of the temporal arches is characteristic; the temporal fossa is large and extends far ventrally with a relatively narrow infra-temporal arch. The ventral postero-lateral corner of the skull is not formed by the quadratojugal lying on the surface, but, lying medially, gives an un-Deinocephalian-like appearance to the Anteosaurus skull. The orbits are of medium size and face anterolaterally; the nostrils are non-terminal and lie laterally, whereas in all other Deinocephalians they are directed much dorsally. The intertemporal region is fairly narrow, but without any suggestion of a sagittal crest. The mentum of the lower jaw is high and squarish. | The anterior part of the dentigerous border of the upper jaw sweeps sharply upwards, exposing the long intermeshing incisor teeth. The lower jaw is hinged fairly far posteriorly. a. The Skull in Lateral View (Figs. 1, 6, 8, 11, 1) In lateral view the skull is roughly pearshaped in outline. The preorbital portion is much longer than the postorbital part. The lateral direction of the nostril, orbit and temporal opening is apparent. The temporal opening extends far ventrally and the infratemporal bar is narrow. The quadrato- jugal is not a bone of the surface but lies medially of the postero-lateral corner of the skull. The side of the snout is fairly vertical. The Premaxilla (P.M.) forms the anterior } of the upper edge of tne skull; anteriorly it has a rounded curve to the dentigerous border, which curves CRANIAL STRUCTURE OF THE TITANOSUCHIAN IIl sharply upwards in antero-posterior direction; from the nostril a groove in the surface of the bone’ runs anteriorly, lying above the curved premaxilla-maxillary suture; on the lower border of the nostril the premaxilla is separated from the septomaxilla by a narrow tongue of the maxilla; the anterior 4 of the internarial bar is formed by a strong girder of the premaxilla, but in the posterior half the nasal helps in forming the internarial bar; ‘ Bis ay ‘ 9 yO et 2° * Ar. = o . . -~e- $¢.) fairly strong and prominent. No jugal boss, angular boss unknown. Fronto-naso-prefrontal swelling small, passing evenly on to the anterior nasal surface. Pineal boss prominent, with sharp circular border, situated very near the occipital edge. Sharp upward inclination of the premaxillary edge. The occiput is high and fairly broad, very deeply concave with a great posterior sweep of the temporal arch and the upper part of the temporal fossa roomy antereo-posteriorly; the temporal arch not rising above the very I40 ANNALS OF THE SOUTH AFRICAN MUSEUM Fic. 18.—Anteosaurus cruentus. S. Nov. Fic. 19.—Anteosaurus cruentus. Sp. Nov. Holotype. S.A.M. 11694, Koringplaas, Holotype. S.A.M. 11694, Koringplaas, Laingsburg. Dorsal view. (x %.-) Laingsburg. Ventral view. (x ¢-) narrow intertemporal surface. Palate long, with fairly robust lateral pterygoidal flanges. Basis cranu long, but the basisphenoid is short. The exoccipitals form- ing a large part of the dorso- lateral corners of the condyle. No trace of incisors is preserved; the alveolar face of the pre- maxillaries shows a matrix filled groove divided in its posterior part into distinct adveoli; there Po.F. i Fic. 20.—Anteosaurus cruentus. Sp. Nov. appears to be room for 5 incisors Holotype. S.A.M. 11694, Koringplaas, when developed. On the right Taingshurg. Occipital var" aaanae: the canine root is followed by roots of 6 postcanines, but on the left only four roots with a possible fifth can be seen. | CRANIAL STRUCTURE OF THE TITANOSUCHIAN I4I S.A.M. 9140 from Voélfontein, Prince Albert, is an imperfect disarticulated skull agreeing fairly well with the type, but is of some interest in that it shows a number of the roofbones as separate elements with exposed sutural faces. Anteosaurus levops Sp. Nov. (Fig. 21) Holotype. A weathered skull, without the lower jaw. S.A.M. 11492. Mynhardtskraal, Beaufort West. Coll. Boonstra. Skull fairly small, maximum length 485 mm. Snout fairly short, lightly built, narrow and low. Intersquamosal width relatively large (415 mm.). Postfrontal boss strong and prominent. No jugal boss; angular boss unknown. Fronto-naso-prefrontal swelling strong with a distinct step onto the anterior nasal surface and laterally slightly overhanging the sides of the skull. Pineal boss apparently prominent, reaching the occipital edge. Upward inclination Fic. 21.—Anteosaurus levops. Sp. Nov. Holotype. S.A.M. 11492, Mynhardtskraal, Beaufort West. Dorsal view. Temporal fossae not cleared of matrix. | (X &-) of the premaxillary edge moderate. The occiput is fairly low and broad; deeply concave, not vertical; strong postero-lateral sweep of the temporal ‘arches and the upper part of the temporal fossa shortened in antero-posterior direction; the temporal arch rising above the plane of the narrow intertemporal 142 ANNALS OF THE SOUTH AFRICAN MUSEUM surface. Palate long and narrow. JBasis cranu fairly short. Condyle unknown. In the right premaxilla parts of the crowns of 3 incisors are preserved, but on the left there is a matrix filled groove with no sign of any teeth. Anteosaurus minusculus Sp. Nov. Boonstra, L. D. 1936. Anteosaurus minor (Broom) in errore. Holotype. A distorted skull with the greater part of the arches and most of the palate missing. American Museum of Natural History, No. 2224. Vanderbylskraal?, Beaufort West. Coll. Broom. This specimen in the American Museum is stated by Broom to be the topotype of the skull fragment in the British Museum (Natural History) R.5742. In my paper on the Titanosuchids in the American Museum I attempted a reconstruction of the dorsal aspect of the skull and showed the premaxilla as a long posteriorly tapering bone. With our present knowledge of the truncated posterior end of the premaxilla in all species of Anteousaurus this was obviously an error in observation. As this skull has well developed postfrontal bosses it cannot beiong to Pseudanteosaurus minor (Broom) and I propose to regard it as a new species of Anteosaurus under the specific name — minusculus sp. nov. The skull is fairly small with a maximum length of 480 mm.; the snout is long, broad and fairly high; the intersquamosal width was probably small (225? mm.); the postfrontal boss quite strong and prominent; there is no jugal boss; the fronto-naso-prefrontal swelling is weak; the occiput is vertical, very deeply concave with a great posterior sweep of the temporal arches and the upper part of the temporal fossa is roomy in antero-posterior direction. Anteosaurus sp. (Fig. 22) S.A.M. 2752, Viviers Siding, Beaufort West. Coll. Hanghton and Whaits. In this specimen we have only the posterior two thirds of the upper surface of a skull. Although it cannot be included in any of the above described species I am not naming it. The skull differs from all the described forms in that the prominent mound-like pineal boss overhangs the occipital surface; the occiput is very deeply concave from side to side with the two horns of the parietal directed nearly wholly in posterior direction; in the posterior CRANIAL STRUCTURE OF THE TITANOSUCHIAN 143 ~ : ~ ° OI SORE) ’ Py sere io as yy Se, } ~=—.Iy 4 . e _ ? ae HW - id > . ‘ € Le) ‘ t ' ‘ ‘ al ! ‘ z Jf T. WAL Fic. 22.—Anteosaurus Sim SACMS 2752; Viviers Siding, Beaufort West. Dorsal View. (x #4.) part of the frontals there is a step bringing this part of the surface down to a lower level than that of the anterior part of the frontals. The postorbitals form a small part of the posterior surface of the boss, whose surface is in other species wholly formed by the postfrontal. These bosses are only moderately strong and prominent. There is very little swelling in the fronto-naso-prefrontal region, but notwithstanding this the prefrontal is so developed to exclude the frontal from entering the supraorbital border. DISCUSSION From the above account it is clear that Anteosaurus is a genus of the Deinocephalia quite distinct from all the known South African genera. It may be characterised as follows: skull large, with prominent postfrontal bosses; temporal fossa with large dorso-ventral diameter and deep antero-ventral bay; infratemporal bar narrow; quadratojugal no longer lying ANNALS OF THE SOUTH AFRICAN MUSEUM 144 ‘soundy poysiqnd 3194) Wosy Uayxe} syusWOINsvou Aq poyuoUINsne wey) Aq U9AIS s;UoUTOINseoW OY) PopNjou! savy | SIOYINE Jo4I0 Aq poquosop SWIJOJ OY} JO BSED OY} UY ésol 6097 €cl 97 69 TOT 0s £0€ O€e Tvl OIZ 8P 877 Is Cvse é Tes $87 Lyl S149SO411ND 077 OIZ 608 €L7 Sty O17 OLT 18 Sm = i nae OFC | L0S7 = og re. — papain WC} Fe = a a O9€ | cS8E = “aa ra — | é19$ | 9€9 8) | 4e= OO@ | SOZT | SZ | ZOIZ O8% | €67 | SL@ | $97 | GOE | ELIZ Pelt eal ee lea | Pe < S SS S > S|! Risa. ts [ota | es. | Ra ‘WW NI SLNAWAYNSVAW AAIHO SOSULLT “Iq “JET JOAO YIPIA, ‘S'a JO y3u07 aur] Id ‘eT JO o3pe jsogq—" Wd ‘O'a—"W'd SOJBN] JZAO JNOUS JO YIPIAA SOSSOq ‘J ‘Od I9AO UIPIAA . TIPIAA [e10du19}103;07 TIPIAA [e31Qs1019;UT *‘sDg JOAO UIPIM “Xe ‘dT — 10g “Uld WqIO— W'd ‘dI—Wd “qQei—W'd CRANIAL STRUCTURE OF THE TITANOSUCHIAN I45 on the surface on the latero-postero-ventral corner of the skull but shifted medially as in the higher Therapsids; alveolar border of the premaxilla inclined upwards to give working space to the long, simple, pointed incisors; a short series of postcanine teeth, rather small and irregular; deeply concave occiput; ridge forming the outer border of the occiput proper formed solely by the tabular and squamosal; postorbital covering the lateral face of the parietal and with a long contact with the squamosal; fronto-naso-prefrontal region swollen to a greater or lesser extent; with bosses on the jugal and angular in the larger species; number of incisors variable; great posterior overlap of the maxilla over the jugal. If the Titanosachia are defined as Deinocephalians with a carnivorous dentition, with quadrate not displaced far anteriorly, with concave occiput, with spacious temporal fossa, then Anteosaurus is undoubtedly a Titanosuchian. The only other S.A. Titanosuchian genus in which the skull is adequately known is Jonkena. Anteosaurus differs from Jonkeria in a number of important points: in Anteosaurus the quadratojugal is no longer a surface bone, but has shifted medially (in one species of Jonkeria viz. vanderbyli, there is an indication that here also there is a similar tendency); in Anteosaurus the infratemporal bar is narrow (again J. vanderbyli shows a similar tendency); in Anteosaurus the temporal fossa extends far ventrally and has in addition an anterior bay of the fossa extending to under the postorbital bar; the postfrontal develops a large boss in Anteosaurus, whereas this bone is a small element in Jonkeria; large jugal and angular bosses are unknown in Jonkerna and _ the fronto-naso-prefrontal region remains unswollen; the pineal foramen is situated near the occipital edge in Anteosaurus, whereas in Jonkeria it is situated in the plane of the postorbital bar; in Jonkeria the premaxillary edge does not curve sharply upwards; no reniform palatine boss bearing teeth is known in Jonkeria; the lateral pterygoidal flanges are much weaker in Jonkena; in Jonkerna the posterior sheet of the postorbital does not cover the whole lateral face of the parietal; in Jonkeria the jugal is a much smaller bone and does not flare out laterally as it does so characteristically in Anteosaurus, nor does it extend so far posteriorly lying along the inner face of the squamosal; in Jonkeria the postcanines form a long series, whereas this is short in Anteosaurus; the posterior process of the premaxilla in Jonkeria, as in the Tapinocephalians, is much longer than in Anteosaurus, in Jonkeria the ridge on the occiput is formed by the paroccipital, squamosal and tabular, whereas in Anteosaurus the paroccipital does not enter into it at all; in Anteosaurus the quadrate rami of the pterygoid curve much outwards as they approach the quadrate, whereas in Jonkeria they lie nearly parallel to the median line; in Anteosaurus the stapes is a much longer bone; in 6—Annals 146 ANNALS OF THE SOUTH AFRICAN MUSEUM Anteosaurus the maxilla has a much greater overlap over the jugal. These differences, together with others not listed here, to my mind show that these two genera lie on lines of development sufficiently divergent to warrant our placing them in different families, which I propose to name the Anteosauridae and the Jonkeridae. It is thus clear that the Anteosauridae, although retaining a number of primitive characters, have advanced farther in some points of structure than their contemporaries the Jonkeridae. COMPARISON WITH OTHER THERAPSIDS In South Africa the Therapsids are first encountered in_ the Tapinocephalus-zone. With its complex of monoclinal folds it has up to the present not been possible to establish from what level within the zone the various known Therapsid finds have come. Until then we are forced to consider the assemblage of forms from this zone as being contemporaneous. The thus contemporary Therapsids from the Tapinocephalus-zone are: Anningiamorpha, Dromasauria, Anomodontia, Gorgonopsia, Therocephalia and Deinocephalia. These are all present as well established groups clearly distinct from each other. The Anningiamorphs are not very well known, but appear to be a group in which a large number of primitive characters have persisted. The Dromasaurians are only known from four specimens and combine a number of primitive characters with some rather specialised. The Anomodonts, although but poorly represented in this zone compared to the great diversity developed in the younger rocks of the Karroo, are already quite specialised when they are first encountered. No group with such a remarkable edentulous premaxilla can be anything but firmly set on an independent line of development. The Gorgonopsians, with but a few forms present in the Tapinocephalus- zone, blossomed exceedingly in later ages, but the oldest known species are already definite Gorgonopsians with a habitus well established and clearly their own and the subsequent developments in no way exceeded the limitations inherent in these early forms from the Tapimocephalus-zone. The Therocephalians of the Tapinocephalus-zone are a virile suborder of the Therapsids already represented by a large number of species, which, representing different lines of development, can be placed in a number of different families. Having already, by the beginning of the Tapinocephalus- zone times, split into a number of families it is clear that the tempo of development within the suborder during the antecedent ages must have been greater than in the Gorgonopsians. This was probably due to a greater lability in the original stock. As their successors — not only the higher Therocephalian families but also their off-spring the Cynodonts, Bauriamorphs CRANIAL STRUCTURE OF THE TITANOSUCHIAN 147 and Ictidosaurians, of the later Karroo Beds — show, this lability or greater potentiality for further development beyond the confines of the suborder was maintained. The Deinocephalians, when first encountered in the Tapinocephaius-zone, were already at the end of their tether. They are represented by the end-products of their particular line of development which culminated in the 3 specialised groups — Titanosuchia, Tapinocephalia and Styracocephalia. CHARACTERS OF A POSTULATED PRIMITIVE THERAPSID ANCESTOR If we postulate a primitive Therapsid ancestor, common to all the Therapsids, situated on a morphological level somewhere between that of the early Pelycosaurs and the first Therapsids we would expect it to have the following characters: intertemporal region broad and flat; preparietal absent; premaxilla without long posterior process intercalated between the nasals; the postfrontal well developed; the postorbital covering the lateral face of the parietal and meeting the squamosal; the pineal foramen well in advance of the occipital edge i.e. the upper occipital edge has not yet migrated anteriorly; the premaxilla below the nostril would still be shallow; the zygomatic arch shallow; the temporal fossa relatively small with the squamosal not bowing out laterally or posteriorly; the quadratojugal still lying on the outer surface i.e. not yet migrated internal to the squamosal; the occiput low, fairly narrow, vertical, with its upper border not moved anteriorly, not deeply concave from side to side; the paroccipital just beginning to strengthen and to support the quadrate; the quadrate would still be large with its condyle lying fairly far ventrally, but still in the posterior position i.e. in line with the occipital condyle; the premaxilla would be dentigerous and with hardly any palatal face; the maxillary teeth would form a long series and there would probably already be a specialised canine; choanae and nares anteriorly situated; no suborbital opening; the lateral pterygoidal processes would still be situated fairly far back and be fairly weak; the basisphenoid-pterygoid joint would no longer be movable and the contact of the basisphenoid with the pterygoid shifted posteriorly; the basisphenoid processes flattening; the quadrate ramus of the pterygoid reduced in height; the parasphenoidal rostrum would still be visible in the posterior part of the interpterygoidal slit; there would be no coronoid process to the dentary; the lower jaw long and the angular notch and reflected lamina would be beginning to develop. Tabulating the characters of the Therapsids of the Tapinocephalus-zone on the basis of the above list one gets the following result: The Anningiamorpha are not sufficiently well known to make a count of the characters corresponding with those in the above list, but those preserved 148 ANNALS OF THE SOUTH AFRICAN MUSEUM point to this group as approaching the postulated primitive Therapsid condition most closely. | The Dromasauria, as far as they are known, appear to occupy the second place. Then come the three groups of the Deinocephalia — which, although clearly distinct from each other — due to particular specialisation — seem to stand on more or less the same morphological level. Somewhat further removed from the postulated primitive Therapsid condition are the Dicynodonts, and then come the Gorgonopsians, followed by the Therocephalians in which the rate of development away from the primitive Therapsid condition has been the greatest. Finally, Anteosaurus has advanced beyond the primitive Therapsid stage in the following characters: the premaxilla, below the nostril has increased in depth and then the alveolar edge has curved upwards; the temporal fossa has become very roomy and the squamosal is bowed out strongly both laterally and posteriorly; the quadratojugal has migrated from the outer surface to lie medial of the zygoma; the occiput has greatly increased its surface by becoming both broad and high; the paroccipital process has become very strong and supports the quadrate firmly; the length of the postcanine series has decreased and the long incisors have become intermeshing teeth; the lateral pterygoidal processes are situated well forward and are very strongly developed; the basisphenoid-pterygoid region has advanced as in higher Therapsids, but there are no strong basisphenoidal tubera and lastly we have the characteristic pachyostosis with the development of the peculiar postfrontal, jugal and angular besses. REFERENCES Boonstra, L. D. 1936. The Cranial Morphology of Some Titanosuchid Deino- cephalians. Bull. Am. Mus. Nat. Hist. 72, 3, 99-116. Boonstra, L. D. 1948. Miljoene Jare Gelede in die Karoo. Voortrekkerpers, Johannesburg. Boonstra, L. D. 1952. ’n Nuwe TJitanosuchiérsoort (Anteosaurus abelt). Tydskrif vir Wetenskap en Kumns. 12, 1, 150-151. Boonstra, L. D. 1953. A Suggested Clarification of the Taxonomic Status of the South Atrican’ Titanosuchians,, Anny S) Air Mus, 42)) a) meres: Broiri, F. AND SCHRGODER, J. 1935. Ein Dinocephalen-Rest aus den wunteren Beaufort-Schichten. Sitzb. Bay. Akad. Wiss. Sonderdruck. 93-114. Broom, R. 1910. Observations on some Specimens of South African Fossil Reptiles preserved in the British Museum. Trans. Roy. Soc. S. Afr. 2, 1, 19-25. Broom, R. 1929. On the Carnivorous Mammal-like Reptiles of the Family Titanosuchidae. Ann. Trans. Mus. 13, 1, 9-36. Broom, R. 1936. On some New Genera and Species of Karroo Fossil Reptiles, with Notes on some others. Ann. Trans. Mus. 18, 4, 349-386. Broom, R. 1936. On the Structure of the Skull in a New Type of Deinocephalian Reptile. Proc. Zool. Soc. 733-742. Watson, D. M. S. 1914. The Deinocephalia, an Order of Mammal-like Reptiles Proc. Zool. Soc. 749-786. , Watson, D. M.S. 1921. The Basis of Classification of the Theriodontia. Proc. Zool. Soc. 35-98. 9g. The Smallest Titanosuchid yet recovered from the Karroo. By oe Boonstra, D.Sc. (With Plate XVIII and 5 text-figures) In the collection of the South African Museum there is a specimen (S.A.M. 4323) collected by Haughton on the Merweville Commonage in 1917. This had been entered in the register as a Gorgonopsian, presumably because of its small size. The specimen as preserved consists of the anterior third of a small skull, the major part of a manus, a tarsus, a nearly complete femur, a radius, a fibula, part of the head of the humerus, a coracoid, a series of caudal vertebrae and some other fragments. This is the first specimen of a South African Deinocephalian in which most of the bones of the fore- and hindfoot have been found in articulation. THE SKULL (fig. 1) In the accompanying figure the lateral aspect of the snout is given, with the missing part of the skull indicated by broken lines. The snout is very similar in general build to that known in the large Anteosaurus, although in size it is less than 4 of Anteosaurus abel. The nostril is not terminal; the alveolar border, anterior to the canine, sweeps sharply upwards; this reduction of the premaxilla creates the space necessary for the large anteriorly directed anterior incisors, whose function has become that of snatching, piercing and tearing teeth (cf. mechanical grab); the five upper incisors, increasing rapidly 2e oer ee ee “ayec2 -- - ° %ace oe” o? Ph Se s a Er, ‘ ° : a 2 Anstey ou Fic. 1.—Micranteosaurus parvus Gen. et Sp. Nov. Lateral view of anterior third of the skull with the missing posterior two thirds in broken lines based on the structure of Anteosaurus. S.A.M. 4323. Commonage, Merweville, Beaufort West District. (x 4.) 149 150 ANNALS OF THE SOUTH AFRICAN MUSEUM in size from number 5 to number 1, intermesh with the four incisors of the dentary; the lower incisors also increase rapidly in size from number 4 to number 1, but they are directed upwards and very much less anteriorly than are the upper incisors; the posterior border of the incisors forms a sharp cutting edge with fairly fine serrations; the upper incisors occupy 55 mm. on the left and 57 mm. on the right side (measured over the curve). Between the last incisor and the canine there is a diastema of 5 mm. on the left and 4 mm. on the right side. The canine is a strong curved tooth with a length of 35 mm. and at the base of the crown the antero-posterior diameter is I4 mm.; its posterior edge, though sharp, is not serrated. Between the canine and the first postcanine there is a diastema of 27 mm. on the left and 23 mm. on the right side; on the right the first and second postcanines are in part preserved, whereas on the left the first, third and fourth are in part preserved; the postcanines are small stubby teeth apparently irregularly spaced and functionally unimportant. The dentary is strong with the alveolar border housing the incisors bent downwards to increase the space necessary for the very long anterior incisors; the symphysis is strong; the mentum squarish with ventrally a “‘gonial’’ or “‘digastric’’ tubercle. The premaxilla extends some distance posteriorly between the nasals. The limits of the septomaxilla are not very clear, but the bone is small and apparently forms the ventral border of the nostril as shown in the figure. The maxilla is not swollen above the canine so that it would appear that the root of the canine is not strong. Anteriorly the maxilla has a large overlap over the premaxilla. Femur (fig. 2a) Except for its distal end the femur is fairly well preserved. It is a long slender bone with both its proximal and distal end unexpanded and the shaft long and slender. A twist in the shaft places the distal end at night angles to the proximal end. The proximal facet is directed appreciably anteriorly. No external trochanter is differentiated on the posterior edge of the bone and the area for the insertion of the ilio-femoralis is narrow. Anteriorly a ridge separates this area from the area for the insertion of the pubo-ischio-femoralis internus. The posterior condyle has no epicondylar widening and the facet for the fibula is terminal. The intercondylar fossa is shallow and so is the popliteal fossa. The intertrochanteric fossa is shallow, and with no clearly differentiated external and internal trochanters developed, it is not clearly demarcated either anteriorly or posteriorly. There is thus little left of the primitive Y midges. It is thus evident that this femur differs greatly from that hitherto described in any South African Deinocephalian where the bone is usually a short stout SMALLEST TITANOSUCHID YET RECOVERED FROM KARROO I5I element with short wide shaft and greatly expanded distal and proximal ends. Although superficially resembling the femur of Therocephalians and Gorgonopsians, it differs strikingly in the absence of a differentiated external trochanter and in the rotation of the ends on the shaft. a b d Fic. 2.—Micranteosaurus parvus Gen. et Sp. Nov. S.A.M. 4323, Commonage, Merweville, Beaufort West District. a, Worsal view of left femur. (x 4.) 0, Ventral view of fibula. (x 4.) c, Dorsa view of left coracoid. (x 4.) 4d, Dorsal view of radius. (x #.) Fiputa (fig. 20) The fibula is a lightly built long and slender bone with its proximal end strongly expanded and with its articulating facet for the femur terminal. Whereas the proximal end is flattened, the distal facet is broadly oval in outline. Pes. (plate XVIIIa and fig. 3) When this specimen came under my notice it had already been partly cleared of matrix and the parts glued together. Before preparing it further I embedded the whole in plaster. In the accompanying plate I give a photograph of the pes after preparation and still in the plaster bed. From this it is evident that the two proximal tarsals joined by matrix were rotated as a unit through 180° when joined by glue to the distal part of the pes in 152 ANNALS OF THE SOUTH AFRICAN MUSEUM the original preparation. In Fig. 3 the proximal tarsal elements are shown right side up. | The intermedium is an ovoid bone with its outer border concave and facing a similar concavity of the fibulare, thus creating a passage for an artery. Fic. 3.—Micranteosaurus parvus Gen.,.,et) Spx j Nowy 1;S A. Meat 4g2ee Commonage, Merweville, Beaufort West District. Semi-diagrammatic restoration of the right pes seen in dorsal view > about 2?. C— centrale; F—fibulare; I—inter- medium. : The fibulare is a dorso-ventrally flattened bone, but thickened both | proximally and distally to form articulating facets for the fibula and centrale and 4th distale respectively. The single centrale is a small pebble-like bone. | Four distalia are preserved, but a fifth must also have been present. | Five metacarpals are preserved as shown in the illustrations; they are | dorso-ventrally flattened bones, constricted in the waist and with their distal _ ends more expanded than the proximal ends. No. 1 differs but little from No. 2, but the 4th and especially the 5th are much reduced. Phalanges. In the first two digits no phalanges are preserved, and in the 3rd and. 4th only the proximal end of the first phalanges are present; in the | little toe there is a fairly long phalanx and an ungual phalanx preserved. SMALLEST TITANOSUCHID YET RECOVERED FROM KARROO 153 The tarsal formula is thus 2, I, 5, and the phalangeal formula Bae 3°, 4? 2 The pes is thus still closely related to that of the Pelycosaurians. CoRACOID (fig. 2c) The left coracoid is preserved. It is roughly circular in outline; thickened laterally where it carries an articular facet to form the lower part of the glenoid articulation; medially it forms a fairly thin sheet of bone with a concave upper and a convex under surface. Anteriorly it has a free edge and is not suturally united to the procoracoid, as is the case in most Therapsids including the Deinocephalians such as Jonkeria and Moschops. This free coracoid is, hower, encountered in a number of Deinocephalians, e.g. Stvuthiocephalus, Tapinocephalus, Pelosuchus, etc. Rapius (fig. 2d) The, radius is much shorter than the fibula. It is a fairly slender bone somewhat flattened dorso-ventrally; its proximal end is expanded to about twice the width of the distal end, but the shaft has no waist-like constriction. Manus (plate XVIIIb and fig. 4) When the manus came to my notice it had also been partially prepared, but all the constituent bones were still joined to each other by matrix. I also embedded the whole in a block of plaster before continuing the preparation. As is evident from the photograph, the first digit lies extended, the second and third folded inwards, in the fourth digit the phalanges have been displaced and the fifth lies extended. In the proximal row of the tarsus the intermedium and ulnare have been displaced medially (anteriorly). . In Fig. 4 I give a restoration of the dorsal aspect of the manus. In the proximal row of the carpus there are a radiale, intermedium and ulnare. The radiale is roughly rectangular in outline; dorso-ventrally compressed; it is thickened distally and proximally to form facets for the distals (1 & 2) and radius respectively, with both the dorsal and ventral surfaces’ concave antero-posteriorly. The intermedium is a fairly small thin flat bone. The ulnare is the longest bone of the proximal row. It is a stout bone, proximally thick and knob-like, with a large convex proximal facet for the ulna; anteriorly it is weaker, with a flat distal articular facet for the distals (4 and 5); just behind the distal face the bone has a constricted waist. The centrale is not preserved, but was probably a fair sized bone lying anterior to the intermedium and between the distal ends of the radiale and ulnare, articulating distally with two distals (2 and 3). - There are 5 distal carpals. The first is a small pebble-like bone; the second is the largest of the distals, its dorsal surface is excavated with ridges 154 ANNALS OF THE SOUTH AFRICAN MUSEUM on the preaxial, distal and proximal edges; the third also has an excavated dorsal surface; the fourth and fifth are pebble-like. Fic. 4.—Micranteosaurus parvus Gens eto Sp Nov: /SiAGME ©4323; Commonage, Merweville, Beaufort - West District. Semi-diagrammatic restoration of right manus seen in dorsal view x plus-minus 4. C.— centrale; I—intermedium; Re— tadiale; Ue—ulnare. As a whole the carpus is very Pelycosaur-like, approaching that of Ophiacodon fairly closely, except that there is only one centrale. The metacarpals are somewhat dorso-ventrally flattened bones in general rod-like with slightly constricted waists; the decrease in size from the first to the fourth is fairly evenly graded, but the fifth is much reduced; the first metacarpal is distinguished by being much broader than the others and thus relatively more flattened with expanded ends. In the first digit the first phalanx is broad proximally, with a waist situated in the anterior half; the distal end is much narrower than the proximal; the second phalanx is much smaller, also broader proximally than distally, with the waist nearly in the middle; anterior to the second roughly hourglass-shaped phalanx there is preserved the proximal end of the ungual phalanx, so that in the first digit there are 3 segments. This is most unusual, but there is no doubt that such is the case. Even without the preserved proximal end of the ungual phalanx the count would also be 3, for the second phalanx with its hourglass-shape could not possibly be an ungual phalanx. SMALLEST TITANOSUCHID YET RECOVERED FROM KARROO 155 In the second digit there are also 3 segments with each of the constituent phalanges closely resembling those of the first digit. The third digit has four segments. The phalanges of the fourth digit have been displaced and all but one lost. What I believe to be the second phalanx lies above the displaced fourth metacarpal. The fourth digit probably had 4 segments as is the case in the third digit. The fifth digit is much reduced with a short hourglass-shaped first phalanx and a fairly high, narrow and long ungual phalanx. The carpal formula is thus 3, 1, 5, and the phalangeal formula 3, 3, 4, 4?, 2, and in structure thus near that of the Pelycosaurs. A MoscHopip CarpPus (fig. 5) For comparison I am including here a figure of a carpus, which, on the humerus, I have identified as a Moschopid fairly near Moschops. This specimen, S.A.M. 9157, collected by me at Wolwefontein, Prince Albert, consists of a good humerus, radius and ulna, with elements of the carpus still joined by matrix to the distal end of the epipodial in natural articulation. In the proximal row there are four bones, viz. radiale, intermedium, ulnare and pisiforme. Only the proximal part of the radiale is preserved, but it would appear that the complete bone was a strong rounded element. The intermedium is_ nearly circular in outline, with its dorso-ventra] diameter about equal to the antero-posterior diameter, and it is thus not a thin plate-like bone. The ulnare is a large bone, subcircular in outline; a longitudinal ridge divides its dorsal surface into two faces, of which the postaxial one is the larger; it is a much thinner bone than the radiale and intermedium, and its under surface is deeply concave. Postaxially there lies a pebble-like pisiforme. In between the ulnare and the radiale there was Fic. 5-—An_ undeter- mined Moschopid. S.A.M. 9157, Wolwe- fontein, Prince Albert District. Part of right fore-imb x 2. C— centrale; I—inter- medium; P—pisiforme; R—radius; Re— radiale; U—ulna; Ue— ulnare. a fair sized centrale; the actual bone has been lost, but an impression in the matrix on the preaxial surface of the ulnare indicates its position. None of the five distals is preserved. This Moschopid carpus is thus structurally close to that of the Pelycosaurs, but the shape of the individual bones is quite different, whereas in Micranteosaurus the individual bones very closely resemble those of the Pelycosaurs in shape. 156 ANNALS OF THE SOUTH AFRICAN MUSEUM TAXONOMIC Although we know nothing about the posterior two thirds of the skull, the snout is sufficiently characteristic to enable us to establish its affinities. The dorsally curving alveolar border of the premaxilla, the long simple intermeshing incisors, the short postcanine series; the fairly narrow, high snout, and the position of the nostril show that this small Titanosuchid is structurally nearly akin to the large Anteosaurus. Because of this great difference in size I propose that it be considered a new genus, under the name Micranteosaurus parvus Gen. et Sp. Nov. With Anteosaurus it is to be included in the Titanosuchid family Anteosauridae. RET LARLY Sie AD A EB) ORE We. See Rone eee WM TIAY ep Pate XVIII. Micranteosaurus parvus Gen. et Sp. Nov. S.A.M. 4323, Commonage, Merweville, Beaufort West District. a, Pes. The proximal segment shows the ventral surface of the intermedium and fibulare as these two bones have been turned upside-down in the original preparation. This error is corrected in the restoration (Fig. 3) x about 3 nat. size. b, Manus in dorsal view x about */ Daw.~SiZe. Ann. S. Afr. Mus., Vol. XLII Plate xvi a SY es : Shani es ee 10. Paranteosaurus, Gen. Nov.: A Titanosuchian Reptile. By L. D. Boonstra, D.Sc. (W:th 2 text-figures) In 1940 I collected parts of a skull, a proximal end of a femur and a vertebra of a Jitanosuchian on the farm Mynhardtskraal, Beaufort West (S.A.M. 11485). The cranial material consists of a weathered pre-orbital part and the dorsal cranial roof without actual contact. The snout on development yielded a good palatal exposure (Fig. 1), and in the other part the interorbital region and the structure of the postorbital bars could be determined (Fig. 2). Fic. 1.—Paranteosaurus primus Gen et Sp. Nov. Orthoprojection of the Ventral View of the skull of the holotype, S.A.M. 11485, Mynhardtskraal, Beaufort West. (x }.) 157 158 ANNALS OF THE SOUTH AFRICAN MUSEUM The palate as shown in Fig. 1 agrees very closely in structure to that of the Titanosuchian genus Anteosaurus. As in Anteosaurus the alveolar border of the premaxilla does not lie in the same plane as the maxillary border, but curving upwards makes an obtuse angle just anterior to the canine; on both sides only the root of the last incisor is preserved, with anteriorly a matrix-filled groove with little indication of a division into separate alveoli; in the left maxilla there is in this groove room for two incisors and in the right possibly for three; the incisor count thus falls within the limits set for Anteosaurus,; the canines have only the roots preserved, but these indicate that the canines were directed much more anteriorly than in Anteosaurus, as is usual in Anteosaurus, five irregularly spaced small postcanines are present; these are small labio-lingually flattened conical teeth; in a horizontal fracture through the right maxilla a replacing root is seen lying linguo-anteriorly to the third postcanine. The crescentic dentigerous boss on the palatine and the relations of the premaxilla, prevomer, maxilla, palatine, transversum and the pterygoid are as in Anteosaurus. a Os ®eseaee* . “e Fic, 2.—Paranteosaurus primus Gen. et Sp. Nov. Orthoprojection of the dorsal view of the skull of the holotype, S.A.M. 11485, Mynhardtskraal, Beaufort West. (x }.) PARANTEOSAURUS I59 On the basis of the palate alone there is nothing to exclude the specimen from the genus Anteosaurus. Similarly the outer surface of the snout (Fig. 2) | is very similar to that of Anteosaurus, although the snout is relatively lower. ! But in the structure of the skull roof and particularly of the postorbital bar our specimen differs very markedly from Anteosaurus. This marked difference is due to the small size of the postfrontal, which is here a small bone forming the dorso-posterior orbital margin, whereas in Anteosaurus the strong pachyostotic development of the postfrontal has resulted in this element overflowing on to the surface of the frontal and overgrowing much of the postorbital and forming the prominent boss so characteristic of Anteosaurus. An intermediary stage between the condition of the postfrontal in this specimen and in Anteosaurus is shown by the Russian genus Titanophoneus. The postfrontal in Pseudanteosaurus with its posterior tongue-like prolongation is of quite a different nature. | No features in this skull, which may be thought to indicate youthfulness, are not also encountered in some species of Anteosaurus. I thus conclude that the smallness of the postfrontal and the light build of the postorbital bar do not represent a growth-stage in the individual, but are in fact a retention of an older morphological stage of the family Anteosauridae. TAXONOMIC For this new form of the Anteosauridae I propose the name Paranteosaurus primus Gen. et Sp. Nov. It may be characterised as follows: A medium-sized Anteosaurid (max. length probably about 570 mm.), with small postfrontal not extending posteriorly, lightly built postorbital bar, without any sign of a boss-like development in the upper part of the postorbital bar, dental formula i.3?-4?, c.1, p.c.5. REFERENCES Boonstra, L. D. 1954. The Cranial Structure of the Titanosuchian: Anteosaurus. Ann. S. Afr. Mus. 42, 2, 106-148. EFREMOV, J. A. 1940. Preliminary Description of the New Permian and Triassic etrapodatrom U'S,S.R- Acad. Se. U'S:S.R., 10;)2, 1-140. N WEP: aie: ANNALS OF THE SOUTH AFRICAN MUSEUM VOLUME XIII Descriptions of the Palaeontological Material acquired by the South African Museum and the Geological Survey of South Africa. PART III, containing:— Ld: 12. 13. 14. Some fossil Mammals in the South African Museum collections. By i. B.S. Cooke, D.Sc: (With map and Plate XIX.) Fossil Suiformes from Hopefield. By R. SINGER and E. N. KEEN. (With Plates XX—XXIV and one text-figure.) Struthiocephalellus: A new Deinocephalian. By L. D. Boonstra, D.Sc. (With 3 text-figures.) The Girdles and Limbs of the South African Deinocephalia. By L. D. Boonstra, D.Sc. (With 6 diagrams, 108 text-figures and Plate XXV.) . ISSUED JANUARY 1955. PRICE {1 tos. od. PRINTED FOR THE TRUSTEES OF THE SOUTH AFRICAN MUSEUM AND THE CEOLGGICAL SURVEY. OF. SOUTH AFRICA BY THE NATIONAL COMMERCIAL PRINTERS’ LIMITED, JAN VAN RIEBEECK STREET, ELSIES RIVER Ir. Some Fossil Mammals in the South African Museum Collections.* By H B25. Cooks, D.Sc., F.ReS:S-Atr. Johannesburg. (With map and Plate XIX.) ABSTRACT The collections of the South African Museum at Cape Town include the first fossil mammal to be discovered in Southern Africa, Homotoceras bainit. There are also some two hundred fossil teeth and bones of Quaternary mammals from twenty-four sites, eleven of which have not previously been recorded. Few of the specimens have been mentioned in published literature unless they represented new species, and lists of faunal assemblages exist only for two of the sites— Taung and Florisbad. The present account records all the species identified in the collections and lists the assemblages for each locality. INTRODUCTION The fossil collections of the South African Museum at Cape Town include some two hundred bones and teeth of Quaternary mammals of which only very few specimens have been mentioned in published literature. Twenty- four sites are represented, eleven of these being new records. Of the thirteen known localities, faunal assemblages have been listed only for two — Taung and Florisbad — while the remaining eleven sites have been recorded merely by the mention or description of isolated specimens. It is the purpose of this account to provide as complete a record as possible for the assemblages for each locality. The accompanying map shows the wide distribution of the sites. I. Modder River, O.F.S. One of the most striking of the Quaternary mammalian specimens dis- played in the South African Museum is the frontlet and gigantic horn cores of an extinct buffalo, “‘Bubalus’’ bainii. The remains were recovered from a depth of forty feet in alluvial deposits of the Modder River, in the Orange * Read before Section D of the South African Association for the Advancement of Science at its Jubilee Congress in Cape Town, July, 1952. VOL. XLII. PART III. ANNALS OF THE SOUTH AFRICAN MUSEUM 162 MTD 140d Po YoIYsV HLNOS . JO NOINN NOGNO? LSVI A “ aNwi ¢- } : : _ OLasva NIFLNO4NI079 | cles =P) Q , = @ Vv : 3! iS LVLS @&laa HONVUO FOSSIL MAMMALS IN S.A. MUSEUM COLLECTIONS 163 Free State in 1839. The remarkable civil engineer and naturalist, Andrew Geddes Bain, saw the specimen and persuaded the actual finder, Mr. Martin Smith, to send it to the Geological Society of London. Its subsequent history is uncertain but it was displayed in the South African Museum and was seen by the eminent English palaeontologist, Dr. Seeley, who described it in r89r under the name Bubalus bainu.* Several other specimens of this giant buffalo have since been found in South Africa and three related species have been described from North Africa (1951), East Africa (1933) and the Anglo-Egyptian Sudan (1949). In describing the Sudan specimen, the late Miss D. M. A. Bate (1949, 1951) created a new genus Homovoceras for the African long-horned buffaloes and showed that this genus is allied to the African Syncerus and is very distinct from the Asiatic Bubalus. The South African form is accordingly now Homotoceras bain (Seeley). Another fossil is recorded as coming from the banks of the Modder River “half way between Kimberley and Bloemfontein’’. It comprises an imperfect frontlet and partial horn core of a large hartebeest, described by Broom in 1909 and named Bubalis [sic] priscus. If Broom’s generic reference is correct it should now be placed in Alcelaphus as A. priscus, but it is possible that it might fall within the genus Peloroceras. , 2. Delport’s Hope, Vaal River The fauna from this site has been listed by the writer (1949) from material in the McGregor Memorial Museum, Kimberley. The South African Museum collection includes a number of bone fragments from which only the proximal end of a right metacarpal can be identified as belonging to a large bovid, possibly a kudu. 3. Sheppard Island, Vaal River When material from this site was listed by the writer in 1949, it was thought that some of the specimens collected by van Riet Lowe in 1928 had been destroyed in a fire at the University of the Witwatersrand in 1931. Six of these have now been found in the S.A. Museum collections. The species represented are Equus sanduithi, E. capensis, E. burchel, Syncerus caffer, and possibly, a kudu. 4. Taungt In 1920 Haughton described seven small skulls of baboons from the limestone deposit at Taung but the full paper was not published. The name * See Appendix, p. 168. t Union of S. Afr. Rep. Form and Spelling of Geographical Proper Names, 1939, recom- mended ‘‘Taung’’’ instead of the previous ‘‘Taungs’’. 164 ANNALS OF THE SOUTH AFRICAN MUSEUM Papio antiquus was proposed and was printed in an abstract (1924). In 1926 J. H. S. Gear collected additional material and distinguished two species, the larger of which he named Papio africanus and the smaller P. izodi, presumably regarding Haughton’s name as a nomen nudum. In 1940 the generic reference was transferred by Broom to Parapapio and Gear’s views were upheld. In 1948, however, Broom re-examined Haughton’s original specimens (which are in the S.A. Museum) and concluded that only the larger species was represented and that the original specific name antiquus must be held to ante-date africanus. The neotype skull of the species is numbered S.A.M. 5356 and the best jaw is S.A.M. 5357. The generic reference remains amended and the correct designation is now Parapapio antiquus (Haughton). 5. Florisbad (Hagenstad Salt Pan) The S.A. Museum collections include a selection of specimens from the spring deposit at Florisbad first described briefly by Broom (1913) and later by Dreyer and Lyle (1931), whose list is complete but subject to several specific amendments. The assemblage includes the following species: Pedetes hagenstadi, Aonyx robustus, Diceros bicornis, Equus quagga, E. lylet, E. capensis, Hippopotamus amphibius, Phacochoerus helmei, P. compactus, P. aethiopicus, Homotoceras bainu, Pelea capreolus, Pelorocerus helmei, Conno- chaetes antiquus (2), Damaliscus albifrons, Taurotragus oryx, Strepsiceros strepsiceros, Gazella bondi (?), Antidorcas marsupialis, Cephalophus sp., Syluicapra grimmia. 6. Hoogstede, Tarkastad District Five specimens from the farm Hoogstede, west of Queenstown, are lightly mineralised and are probably surface finds. One specimen is a tooth of a. domestic ox. The remainder represent a lion, Equus burchelh and Pelorocerus sp., possibly P. broomi. 7. Lwartkops, Sundays River A small fragment of an upper molar of Hippopotamus sp. is the only fossil mammal from this locality, which has yielded much invertebrate material to other collections. 8. Victoria West From the farm Jakkalsfontein near Victoria West comes an upper molar of Equus burcheli. There is an upper premolar of Equus capensis from an unknown locality in the same area. FOSSIL MAMMALS IN S.A. MUSEUM COLLECTIONS 165 9. Brakfontein, Three Sisters A single specimen from the farm Brakfontein, near Three Sisters, is a fragment of a left lower jaw of Diceros bicornis. to. Beaufort West A lightly mineralised milk molar of Equus cf. burchelli comes from the farm Little England near Beaufort West. II. Cango Caves, Oudtshoorn Six specimens from the Cango Caves are referable to Procavia capensis, Connochaetes sp. and Tragelaphus cf. scriptus. 12. Linkerhandsgat and Noottgedacht, Stanford A few miles north-east of Stanford is an unusual occurrence of chalky limestone and calcified sands containing bones and teeth of mammals. Three specimens from Linkerhandsgat are identified as Crocuta sp., Thos mesomelas and Alcelaphus cf. caama. From the adjoining farm Nooitgedacht, there are three fragments representing Redunca arundium and Aepyceros melampus. A larger private collection and material collected by the writer will be dealt with at some length in a separate paper. 13. Hawston The sand-dunes along the coast near Stanford and Hawston have yielded almost unmineralised teeth of the African elephant and of the black rhinoceros. 14. Skildegat Cave, Fish Hoek This cave, also known as Peers’ Cave, was excavated twenty-five years ago by B. and V. Peers and it is probable that the eight teeth from this locality which are in the S.A. Museum collections were recovered during the excavations. A bovid incisor cannot be determined generically but the remaining specimens belong to Equus zebra and E. capensis. 15. Kalk Bay This area yielded three molars of Hippopotamus MR and part of a lower molar of Loxodonta africana. 16. Yzerplaats, Maitland District The type series of teeth of Equus capensis, described by Broom in 1909 and figured in 1928, was contained in a block of limestone washed up on the beach at Yzerplaats. : 166 ANNALS OF THE SOUTH AFRICAN MUSEUM 17. Bloembosch, Darling District The dune-covered farm Bloembosch, north of Yzerplaats, has furnished nearly fifty specimens, including bones, imperfect jaws and isolated teeth. Equus capensis is abundant, the collection including the plesiotype upper left second molar referred to Broom’s species by Haughton (1932). Other equine material is not specifically identifiable except for an unmineralised lower jaw of a young Equus zebra. Diceros bicornis occurs. The only carnivore is Cyvrocuta crocuta. Artiodactyls include Hippopotamus cf. amphibius, Guwiraffa camelopardalis, Homotoceras bainu, Syncerus caffer, Connochaetes sp., Hippotragus cf. niger. The two lower mandibles which the writer (Cooke 1947) named Hippotragus problematicus, are also in the collection; it was suggested that these jaws might represent the almost unknown H. leucophaeus but Dr. Broom informed the writer shortly before his death that he had found material of the Blue Buck during his last European-American tour and that the Bloembosch fossil was clearly distinct. His notes and drawings have not so far been found. The recently discovered site at Elandsfontein, near Hopefield, which is being studied by the University of Cape Town, has yielded abundant fossil material of a character similar to the Bloembosch specimens;* it has also provided a fossil human cranium. 18. Saldanha Bay The only species represented by the five isolated teeth from this locality is Equus capensis. One of the specimens is the isolated upper fourth premolar described by Broom (1913) as ‘‘from Darling’’, and another is the plesiotype left lower premolar described by Haughton (1932). 19. Hoedpjesbaa The limestone quarries at Hoedjiesbaai, near Saldanha Bay, have furnished sixteen very nice specimens, mostly partial jaws with teeth. The five species are Procavia cf. capensis, Thos mesomelas, Arctocephalus pusillus, Suricata sp. (or possibly Cynictis), and Raphicerus campesiris. This is believed to be the first record of the Cape sea lion in the fossil state. 20. Geelwal Karoo, Van Rhynsdorp District This site has provided a partial lower jaw of Thos mesomelas. 21. Near Springbok, Namaqualand A site 40 miles east of Springbok yielded the type series of cheek teeth described by Haughton (1932) as Notohipparion namaquense. The original illustration exaggerates the breadth of the crowns as the plane of drawing is parallel to the rather oblique grinding surface. * See: Following article by R. Singer and E. N. Keen. FOSSIL MAMMALS IN S.A. MUSEUM COLLECTIONS 167 22. Bogenfels, S.W.A. The collections include an incomplete lower jaw from Bogenfels, S.W.A., labelled “‘“Propalaeonyx africanus Stromer’’. It comes, presumably, from the so-called Miocene beds. 23. Kalk Plateau, S.W.A. Two equine teeth, apparently of Equus burchelli, are recorded as coming from a well on the Kalk Plateau east of Marienthal in S.W.A. fa Wsaros, S.W.A. The type series of upper and lower cheek teeth of Equus sandwithi (Haughton 1932), was associated with a number of other specimens not previously mentioned. It is now possible to reconstruct all the cheek teeth, though not of a single individual. One tooth of Equus capensis is also represented in the assemblage, and the bovid genera Connochaetes and Strepsiceros are present but the species cannot be determined. 25. Other Locahties The collection includes a lime-encrusted upper premolar of Equus cf. zebra from Broken Hill, Northern Rhodesia. There is a piece of. grey limestone from the vicinity of Mt. Lemagrut (near the famous Olduvai gorge), with part of the left lower jaw of a rhinoceros exposed. 26. Unknown Localities Twenty specimens are without locality records. The species represented are Equus burcheli, E. kuhni, E. capensis, Hippopotamus sp., Damaliscus sp., Connochaetes sp. and Phacochoerus sp. REFERENCES Bate, D. M. A. 1949. A new African fossil long-horned buffalo. Ann. Mag. Nat. Hist. (12), Il, pp. 396-308. Bate, D. M. A. 1951. The mammals from Singa and Abu Hugar. In Fossil Mammals of Africa, II, pp. 1-28. British Museum (Nat. Hist.), London. Broom, R. 1909. On a large extinct species of Bubalis. Ann. S. Afr. Mus. VII, PP: 27, 280. Broom, R. 1909. On the evidence of a large horse recently extinct in South Africa. IOI: BP. Zo... .2o2. Broom, R. 1913. Man contemporaneous with extinct animals. Ann. S. Afr. Mus. 1, pp. 13-16: Broom, R. 1928. On some new mammals from the Diamond gravels of the Kimberley District. Ann. S. Afr. Mus. XXII, pp. 439-444. . Broom, R. 1940. The South African Pleistocene Cercopithecid Apes. Ann. Transv. Mus. XX, pp. 89-100. Broom, R. 1948. Some South African Pliocene and Pleistocene Mammals. Ann. Transv. Mus. XXI, pp. 1-38. 168 ANNALS OF THE SOUTH AFRICAN MUSEUM Broom, R. and ScuHepers, G. W. H. 1946. The South African Fossil Ape-men; the Australopithecinae. Tvansv. Mus. Mem. No. 2, pp. 1-272. Cooke, H. B. S. 1947. Some fossil hippotragine antelopes from South Africa. So. Aly. J. Sct. SLIT, pp. i226-237. Cooke, H. B. S. 1949. Fossil mammals of the Vaal River basin. Geol. Surv. Mem. XXXV (3), pp. I-109g. DREYER, T. F. and Lyre, A. 1931. New fossil mammals and man from South Africa. Grey Univ. College Dept. Zool., Bloemfontein, pp. 1-60. Gear, J. H. S. 1926. A preliminary account of the baboon remains from Taungs. S. Agr oY set: OL pp ig3aa-7a9. HaucutTon, S. H. 1924. Papio antiquus n.sp. Trans. Roy. Soc. S. Afr. XII. Min. & Proce!" May)1920)"'p. (EX VEE. Haucuton, S. H. 1932. The fossil Equidae of South Africa. Ann. S. Afr. Mus. XXVIII, pp. 407-427. SEELEY, H. G. 1891. On Bubalus bainii (Seeley). Geol. Mag. VIII, pp. 199-202. APPENDIX Bubalus bainii The date when the specimen was received at the Museum is not recorded. Two entries in the Account Book for 1856 may refer to this specimen: ‘‘March. To Ford repairing horns, £1 15s.’’ Ford had already repaired the models of natives, presumably made of plaster, thus having a knowledge of plaster-work, which would be suitable for the repair of fossils. ‘‘September. To G. West, paint for fossil head, 3s. 6d.”’ The skull was on exhibition over the entrance to the old Museum (now the west wing of the South African Library) in 1860, when Layard compiled his Catalogue: “Immediately overhead is a magnificent fossil frontlet of an extinct bovine from Thaba ’Nchu.’’ (Layard, E. L. Catalogue of the South African Museum, Part 1, Mammals, p. 7. Cape Town, 1862. Preface dated 1st January, 1861, publication delayed until 1862. See Annual Report S.A. Mus. for 1862.) It was in the same position when Seeley saw it. The locality Thaba ’Nchu is not mentioned in Bain’s references to this fossil Bovine. Bain did not visit the eastern part of the Orange Free State until 1845. Thaba ’Nchu, however, does lie within the catchment area of the sources of the Modder River. lLayard’s entry seems to be the only record of the locality, unless possibly a more precise site can be traced by research in the Deeds Office for Mr. Martin Smith’s farm —dif he possessed one. In Dr. Cooke’s map the site of the locality (No. 1) is placed south of Kimberley in the western part of the Orange Free State; but if Layard was correct it should be placed in the eastern part between Bloemfontein and the border of Basutoland. Seeley (p. 201) gave two reasons for considering this type specimen as the one referred to by Bain in 1839. The second reason seems to be acceptable. In his letter to Sir H. de la Beche in 1844, read to the Geological Society in 1845, and printed (abridged) in Trans. Geol. Soc. Lond. (2), VII, 4, 1856, Bain said that Martin Smith’s fossil was ‘‘in Cape Town’’. The South African Museum possesses Bain’s own copy of this part of the Transactions, in which two words have been added by Bain (p. 59) to make the sentence read: ‘‘This fossil is now in the Cape Town Museum.’’ Seeley’s first reason is less acceptable because there may be two interpretations. Thomas Bain’s words ‘‘his father’s fossil’? may mean either the fossil which his father induced Martin Smith to send to Cape Town, or an example which A. G. Bain himself found. It is, therefore, appropriate to reproduce here (pl. XIX) Bain’s MS. drawing and description (from memory) of a specimen of this bovine which he himself found on Mr. G. Southey’s farm near Graaff-Reinet. The original sketch is in the S. Afr. Museum library — but what happened to the specimen? See: Lister, M. H. Journals of Andrew Geddes Bain. Van Riebeeck Soc. Publ. No. 30, p. 230. Cape Town, 1949. Also Seeley, H. G., 1891 (op. te PVATE GES MiSs, Wl SIUIUE Annes. Afr. 12. Fossil Suiformes from Hopefield.* By R. SinGER and E. N. Keen. Anatomy Department, University of Cape Town. (With Plates XX—XXIV and 1 text-figure.) INTRODUCTION The fossil site on the farm ‘‘Elandsfontein’’ situated 10 miles south-west of Hopefield, 90 miles north of Cape Town, was located by one of us (R.S.) in May, 1951.f On numerous subsequent visits various members of the University of Cape Town staff, Professor M. R. Drennan, Messrs. J. A. Mabbutt, K. Jolly and the authors, have collected highly fossilized bones and stone implements from the surface of the site. The range of specimens already identified includes a large number of extinct and existing mammals, the proportions of which suggest an early Upper Pleistocene period, with the possibility of extension into the late Middle Pleistocene. In January, 1953, Jolly, on a field trip with Singer, discovered pieces of the greater portion of the calvarium of an hominid. On three subsequent expeditions Jolly and Singer retrieved additional portions of the cranium which has since been reconstructed (Drennan, 1953; Singer, 1954). Fluorine tests, carried out through the courtesy of Dr. K. P. Oakley of the British Museum, reveal that this Saldanha Skull and the Mesochoerus fossils (described in this paper) were apparently contemporaneous. Elandsfontein lies in the sandy veld between the Sout River and the Langebaan-Saldanha Lagoon. The site is 300 feet above sea-level and is divided into wind-scoured kloofs or depressions by sand-dunes which are either drifting or, where covered by vegetation, stationary. Ridges of ferricrete cut diagonally across the length of the site, and in places the dunes are capped by massive calcrete mounds or flat boulders of partly silicified surface limestone. Softer, cellular calcretes are found in certain places at ~ * Simpson (1945) has given reasons for placing the Hippopotamidae and the Smidae into the separate infraorders ANCODONTA and SUINA; both are, however, included in the suborder SUIFORMES. + The existence of fossil-bearing sites in the Darling-Hopefield district has been known for years (Broom, R., Annals S. Afr. Mus. VII, p. 281, 1909, and XII, p. 13, 1913; Cooke, H. B. S., XLII, p. 161, 1955), but hitherto no thorough investigation of the area had been made. 279 ANNALS GF THE SOUTH AFRICAN MUSEUM the lowest parts of the depressions. The tortuous courses of the ferricrete ridges indicate that they are the indurated lower flanks of old sand-dunes now stripped bare of the sand walls. This ferruginization is usually associated with moist ground conditions, a fairly high stable water-table and an abundance of vegetable acids in the soil. It seems that this fossil site was at one time a large vlei or lagoon along the edge of which animals roamed. The rich collections of stone implements indicate the presence of Man on the site from a late stage of the Chelles-Acheul (Stellenbosch) Culture until the period when the Bush races were developing their culture. The occupation is not a continuous one. A large number of the implements exemplify the transition from the South African Earlier to the Middle Stone Age. DESCRIPTION OF MATERIAL Fam. HIPPOPOTAMIDAE Hippopotamus amphibius Linnaeus 1758 The collection of fossil mammalian remains from the Hopefield site includes several fragments of jaws and teeth which are identified as hippo- potamus remains. Many of the specimens derive from young animals and show deciduous or unerupted permanent teeth. Six of the permanent molar teeth are complete and can be compared in size with the limits stated in Cooke’s (1949) survey of all the South African fossil material recovered up to that date. None of the measurements fall outside the wide limits which are characteristic of this species. There are therefore no grounds for separating the remains from the existing species Hippopotamus amphibius, fossil specimens of which have been found in several other South African localities. Fam. SUIDAE Genus Mesochoerus, Shaw and Cooke, 1941 Mesochoerus lategant, sp. nov. Dragnosis: A Mesochoerus resembling M. olduvaiensis (Leakey, 1942) in the structure of the third lower molars. The second lower molars have a well developed anterior median pillar. The crowns of the third lower molars are longer by 10% and narrower by 10%, and the height of the unwom pillars is greater by 10% than in M. olduvaiensis. The pillars of the upper molars are obliquely arranged, those on the labial side being placed slightly in front of their lingual partners. The fourth premolars are molariform. Type: The complete upper and lower cheek dentition of a single animal, recovered in seven fragments numbered S6, S7, S8, S14, S15, $16, S17 in the Hopefield collection in the Anatomy Dept., University of Cape Town. (Plate XX.) Specimens $7, S14, S16 and S17, constituting the right dentition. FOSSIL SUIFORMES FROM HOPEFIELD rE have been donated to the S.A. Museum where they have received catalogue number I1I712. Among the fossil specimens collected from the Hopefield site are several upper and lower teeth and partial jaws which appear to belong to a single species of large suid. All the fossils at Hopefield have been more or less damaged by the movement of the sand in which they lie. For this reason recovery of an intact jaw is uncommon, and the study is necessarily confined to the characters of the individual teeth, which are often found lying free and not close to any other teeth or suid remains. The collection consists of teeth and jaws from ten individuals and includes a complete upper and lower molar dentition of a single animal (S6: left M: Mz M:;; S15 and S8: left M° M* M’; S16 and S17: right M: Mz M, P, P;; S14 and $7: right M* M* M’ P*). Three other pairs of left and nght lower third molars (S2 and $3, S12 and S13, Sg and Sga) appeared to derive from a single animal in each case, and one pair of right and left upper third molars (Str and S1o) similarly resembled each other. These upper molars could not, however, be made to occlude satisfactorily with any of the lowers. Five individuals are represented by isolated teeth (Sr, mght M:; S4, nght M’; S5, left M° and M’; S2o, left M* and M’; and Sat, right M,). The various teeth resemble each other sufficiently closely to support the conclusion that only one species is represented. LOWER CHEEK TEETH Intervals: 4/5 3/4 D3 V2 Lateral Sth 4th 3rd 2nd Ist Pillars: pair pair pair pair pair Fic. 1. Third lower molars: ten specimens from six individuals. A typical specimen in moderate wear is illustrated in Fig. 1, which also shows the convention by which the lateral and median pillars are identified. The crowns of these 172 ANNALS OF THE SOUTH AFRICAN MUSEUM teeth are each made up of five pairs of lateral pillars arranged symmetrically (Si, S2, S6, Pl. XXI). Each pair appears to meet in the median line of the tooth early in wear, but becomes separated by median pillars in later wear. The usual number of median pillars is nine. A single median pillar is found in front of the first lateral pair, sometimes in the median line, sometimes leaning to the labial side of the tooth. There are two median pillars in the 1/2, 2/3 and 3/4 intervals. These may be of equal size, or the posterior one may be larger than the anterior. A single median pillar is usual in the 4/5 interval, and another in the posterior median position, i.e. at the posterior extremity of the tooth. A few of the specimens show duplication of the median pillars in the 4/5 and posterior positions. The tooth erupts in such a way that the first three pairs of lateral pillars rapidly come into wear one after another, beginning in front (Pl. XX, B). Wear commences in these anterior pillars before the fourth and fifth pairs erupt. Later on growth and subsequent wear proceed at a faster rate in the posterior part of the tooth, so that in late wear the tooth may present an appearance of relatively even attrition. The upper third molars erupt in a similar way. In general the pattern of the worn surface of the lower tooth is simpler, less complex and less folded than that of the corresponding upper tooth. There is a cingulum at the crown-root junction in all specimens, but in some it is very poorly developed (S21, Pl. XXII). In many of the teeth the crown is partially covered with a thick layer of cement (S2r, Pl. XXII). Roots are very well developed along the whole length of the tooth, even at the stage when the posterior pillars are neither fully developed nor erupted (S21, Pl. XXII); at this stage the length of the root may exceed the height of the crown. The roots supporting the anterior three pairs of lateral pillars are more or less separated from their neighbours, and straddle a wide dental canal (S16, Pl. XXII, D). In some cases the tips of these roots are distorted in quite irregular ways (So, Pl. XXIII, B). Extra median rootlets are occasionally present in both upper and lower third molars, and vary from a minute nodule to a spicule as long as the neighbouring lateral roots (Sq, S14, Pl. XXIV). The curvature of the roots is backwards and to the labial side; the labial roots are always larger than those on the lingual side of the tooth. The roots of the posterior two pairs of lateral pillars tend to be fused together to form flat plates arranged antero-posteriorly. Towards the back of the tooth these plates converge, and fuse together in the median line of the tooth, under the posterior median pillar (cf. Pl. XXIV, D, S4). 173 ‘podojaaop ATyUoTOTJNsUL JO ‘UJOM ATJUETOTFFNsSUL ‘ueyoIq st usuTTOeds oY} STEYA\ poz}TWIO oI SOINSI ‘apaowd sndaoyr0sayy pue sisuawanpjo snaaoyoosayy JO SUOISUSWIP pozJoder oy} YM Uostreduloo pue ‘sIe[ofT Jomo'T pPsIIY], JO suotsuounqd :][ ATavL A ol iz] (=, Ay — VI (ors G1 Te ze ve ve — ad 89 ad A09NT a oe — — — o£ — — — (Ke Ez ol ed A,0[0O}{ s apaawd SnséaoyIo0sa O. es — — a 4 I't Lt — gz — — Lt vz G9 odAyereg an — — Q'z o'r Le —— Qz — — Li Cz C-Lo ad A010}, =| sisuawwaAnNp1O SnsaoY4I0Sa S Q'z Wet Ve z'6z bgt I'ZZ LEEL ueoy 5 +91 — — ver — — Qz 6z gz — Iz — (y)1zs 0 — QZ QZ ean gf —- — Lz — gI IZ CL (W)EIS = —_ o£ beg ex QE Cz Qz Qz — gl IZ CL (q)zISs wa of —- — — — — — — — IZ €z — (x1) 86s am) o gz — — — — — — — — ZZ bz — (7) 6s ra or +26 Ez Gr G-€ ee st €€ = Gr zz LL Qt) 9rs Re €€ — — ‘Sas — = — — ol €z CL (1) 9S oF = a8 Re ort a — _- — Oz Ze Lo Qt) €s =< ao a = = = — — — oz ZZ — (q) zs +0z a — = ous Zz — —- a oz Zz EL (pais ‘reid = -repid -seqrd = “req yyS yy p1e puz ‘Iol10}s0g ‘Ioojuy ‘“WseyT ‘“YIpeolg ‘“Yy}peomg ‘yoo 9Aoqe ‘Yypeorg ‘“YypeoIg *Yyy3ue'T yy} suo] WSO = s-UBU’T sreyid uromun jo }ysIOF] [esnjI99 -}se}e9IN) S90} POID) ‘WIUI UI S}OOI Jo yysUeT ‘“UMOID Jo suoTyJodo1g ‘WIUI UI UMOID JO SUOTSUSUITC 174 ANNALS OF THE SOUTH AFRICAN MUSEUM Second lower molars: three specimens from two individuals. The crowns of these teeth are made up of two symmetrically placed pairs of lateral pillars, a single well developed anterior median pillar, two median pillars in the interval between the lateral pairs, and a fairly large posterior median pillar. The anterior median pillar is eccentrically placed on the labial side of the tooth, and the posterior median pillar on the lingual side of the median line (S6, Pl. XXI, B). In the two specimens in which the pattern is clear there are small accessory enamel nodules behind each labial lateral pillar and a small row of nodules at the extreme anterior part of the tooth, in front of the anterior median pillar (S17, Pl. XXII, E). There are four well developed separate roots, which appear to diverge whether the tooth is viewed end-on or from the side. The general curvature of the root is backwards and to the labial side (S17, Pl. XXII, F). The posterior roots are larger than the anterior. Dimensions of Crown. Greatest Greatest Occlusal Length of Length. Breadth. Breadth. Roots. $3 (R) 24 20 — _ S6 (L) 29 18 16 — $17 (R) 30 20 14 29 Mean 28 19 15 Mesochoerus olduvaiensis Holotype 28 20 17 —_— Paratype 29 ZU 18 — Mesochoerus paiceae Neotype 32 19 18 20-25 TaBLE II: Dimensions of Second Lower Molars in mm., and comparison with the dimensions reported for M. olduvaiensis and M. paiceae. First lower molars: two specimens from different individuals. Both these teeth are extremely worn (S6, Pl. XXI, B; S17, Pl. XXII, E). The second and third molars belonging to each of these first molars have been found, and it can be seen that at the time when the first molar was worn down to the roots, the posterior part of the third molar was incompletely developed and unerupted (S6, Pl. XXI, B). The degree of wear of these teeth precludes a determination of the enamel pattern in the earlier stages of wear. In each there remain two fairly large convoluted pillars, one in front of the other, with small islands of enamel in the centre of the outlines, indicating an originally more complex pattern from which the present appearances were derived. | There are four roots, as in the second molar, diverging in both views, the posterior roots being more massive than the anterior (S17, Pl. XXII, F; 56) Pl x) FOSSIL SUIFORMES FROM HOPEFIELD 175 The greatest length, greatest breadth, and occlusal breadth of the crowns of the two teeth are 20, 14 and 12 mm. (S17); 20, 14 and 14 mm. (S6), respectively. One of the roots exceeded 20 mm. in length. Fourth lower premolar. There is one partly broken representative of this tooth, and it is molariform in type (S17, Pl. XXII, E, F). The crown structure is poorly defined, but seems to be made up largely of two pillars arranged antero-posteriorly. The greatest length of the crown is 16 mm., and the greatest breadth 12 mm. Two roots are disposed antero-posteriorly, and appear to have exceeded 14 mm. in length. The posterior root is partially divided into labial and lingual parts near its tip. Third lower premolar. Only the remains of the roots of one of these teeth were found, embedded in the fragment of jaw bearing a second and first molar (S17, Pl. XXII, E, F). Two roots appear to be arranged antero- posteriorly, and one of these is estimated to have exceeded 22 mm. in length. The length of the lower molar series in the type specimen was 120 mm. on the left side and 121 mm. on the right side. UPPER CHEEK TEETH When the upper teeth are compared with the lower teeth, certain differences are sufficiently marked to be summarised as follows: The upper cheek teeth are broader than the lower, and show more complex and folded enamel patterns; the structure of the third upper molars is less regular, and there is a tendency in all the teeth for the arrangement of the columns to be oblique, the columns on the labial side of the tooth being placed slightly in front of the level of their lingual companions (Pls. XXIII and XXIV). Third upper molars: seven specimens from five individuals. As already mentioned, the crown structure of these teeth is less regular than in the case of the lower third molars (S20, Pl. XXIV, A). The first two, and in some cases three, pairs of lateral pillars can be compared with the corresponding structures in the lower teeth, although they are obliquely placed, with the labial pillars ahead of the lingual partners. At least three regular lateral pillars are found on the lingual side of all the specimens, but there may be only two such regular lateral pillars on the labial side. Behind this regular part of the tooth is found a variable number (5—7) of smaller pillars, irregularly arranged in such a way as to defy division into median and lateral pillars. The posterior extremity of the tooth may consist of a single pillar, or of two pillars of approximately equal size. In the anterior median position is a pillar either frankly divided into a central triangular mass with two 176 ANNALS OF THE SOUTH AFRICAN MUSEUM small outriders, one on each side, or showing a clear tendency to be divided in this way. In the 1/2 and 2/3 intervals there is usually a single smaller median pillar; occasionally this is duplicated. A curious feature of the lateral pillars is a groove which appears on the outer side of the pillars low down, near the crown/root junction. This is well seen on the lingual side of the specimen S15 (Pl. XX, B). The third upper molars erupt in much the same way as the corresponding lower teeth (see p. 172). As in the case of the corresponding lower teeth, the upper third molars have well developed roots along their whole length. The roots supporting the first two lateral pillars are usually separate, and may be divided so that two rootlets support one pillar. Behind this the roots are more or less fused (S4, Pl. XXIV, D), sometimes into plates similar to those seen in the lower teeth. The roots diverge widely when seen end-on, and their general curvature is backwards (S5, Pl. XXIII, B). Second upper molars: four specimens from three individuals. The funda- mental structure of the crown is two pairs of lateral pillars, an anterior median pillar, a single central pillar, and a posterior median pillar. The enamel bounding these pillars is considerably folded and the picture is complicated by the presence of additional small enamel nodules. The maximum number of these is eight, one on each side of the anterior and posterior median pillars and two between each pair of lateral pillars. The disposition of the pillars shows the same slant as is seen in the third molars, with the labial side in advance. The anterior median pillar is situated more or less on the lingual side of the tooth (S20, Pl. XXIV, A). In later wear the posterior median pillar tends to fuse with the posterior lateral pillar on the lingual side (S20, Pl. XXIV, A). Four roots are present, the posterior pair being larger than the anterior. The general curvature of the roots is backwards (S5, Pl. XXIII, B). Dimensions of Crown. Greatest Greatest Occlusal Height of unworn Length of Length. Breadth. Breadth. pillars above root. Roots. 3rd Posterior Anterior. Posterior. pillar. group. Third Upper Molars S4 (R) 65 25 22 29 22 34 25 $5 (L) 64 25 20 36 30 34 2g S10(R) 67 26 23 — 26 — — S1r1(L) 70 26 Sie. — 27 — — S14(R) 70 24 3" 7 ary SYSTEMATIC DESCRIPTIONS Deinocephaha Sub-ordinal Characters of the Girdles and Limbs. The pectoral girdle is structurally Therapsid, but comparatively large and massive and without an acromion; the cleithrum is splintlike or well developed; the interclavicle with its anterior spatulate end turned upwards; the three scapulo-coracoid elements not firmly ankylosed; there is no ossified sternum. a a = GIRDLES AND LIMBS OF DEINOCEPHALIA 205 The humerus is structurally Therapsid but comparatively massive and with either one or two epicondylar foramina; the radial facet is either much distally or much ventrally situated. The ulna has a massive olecranon. The pelvic girdle is structurally Therapsid but large and massive; the pubo-ischiadic plate is short; the acetabulum large; there is only a simple pubic foramen; in most groups there is developed a distinctive ridge on the posterior edge of the posterior iliac process for the origin of the m. ilio- fibularis; the anterior pubic edge is everted; the symphisis is weak. The femur, of typically Therapsid structure, is either short and massive or long and slender. Tapinocephaha Infra-ordinal Characters of the Girdles and Limbs. The pectoral girdle is fairly light to moderately massive; the scapula is fairly low to low (310-530 mm.) with the upper part of its blade fairly narrow to fairly broad (120-270 mm.); the scapular head of the triceps originates from a low mound; the internal opening of the foramen supra- coracoideus does not open into or is connected by a groove to the subscapular groove; the glenoidal facet of the scapula is small or of medium size and faces little or not at all externally. — The coracoidal plate is fairly short or long antero-posteriorly; the pre- coracoid is fairly large, but fairly weak with a thin anterior border and a fairly strong dorso-posterior apex; the precoracoid excluded from the glenoid or just enters the rim; its outer face is only moderately convex. The coracoid is small to medium, light to fairly strong with a fairly large glenoidal facet facing dorso-posteriorly and also much externally. Except near the glenoid the three bones of the scapulo-coracoid are weakly united suturally. The clavicle is a medium-sized flattened bone with expanded ends and a slight waist. The cleithrum is a weak splintlike bone. The interclavicle is a moderately light bone, moderately expanded anteriorly with a long slender to fairly slender stem. A shallow groove receives the lower end of the clavicle. The humerus is fairly light to moderately massive; it varies in size from small to large (length 276-588 mm.), but is never short and squat, and some- times long and fairly slender; there is never an ectepicondylar foramen; the capitellum is weak to moderately strong and usually poorly modelled and it never extends proximally along the ventral face but remains situated 206 ANNALS OF THE SOUTH AFRICAN MUSEUM much distally; the processus medialis always lies far proximally, nearly in the same plane as the caput; the processus lateralis always les far proximally; the proximal expansion is moderately great to great; the epi- condyles are moderately to greatly expanded; the shaft is short to long, narrow to very broad; the ventral opening of the entepicondylar foramen is always slitlike. The ulna is small and light to moderate and fairly strong, or is short, massive and squat. The radius is long and slender or fairly short with strongly expanded ends. In the manus the carpal formula is 4, I-2, 5, and the phalangeal formula, 18.2, 2). 2818.2 The pelvis has a fairly long pubo-ischiadic plate; the iliac blade is low to only moderately high; the ilio-fibularis ridge has a fairly sharp edge and is only moderately strong and usually lies nearly horizontally; the anterior iliac process is only slightly to moderately everted; the posterior iliac process is fairly long and low; the outer iliac face is only slightly concave in antero- posterior direction. | The femur varies from fairly small to fairly large (length 276-468 mm.), light, moderate to fairly robust, sometimes short and squat; the internal trochanter undeveloped or forming and elongated tubercle situated well away from the preaxial border; the width over the external trochanter is small (95 mm.) to great (256 mm.), and without a notch. The tibia is massive and short (232 mm. in length) or slender or moderate (length 180-270 mm.). The fibula is light and slender (length 222-288 mm.). In the pes the tarsal formula is 2, 1, 4, and the phalangeal formula 2, 3, 3, 3) 3: Tapinocephahdae Family Characters of the Girdles and Limbs. The pectoral girdle is only moderately massive; the scapula is low, with its posterior edge fairly straight. The humerus is fairly massive but fairly long. The ulna is moderate and fairly long or it is massive and squat. The iliac blade is low with both the anterior and posterior processes long and low; the ilio-fibularis ridge is strong and prominent. The femur is either fairly massive and fairly long or massive and squat. The above statement on the characters of the girdles and limbs of the Tapinocephalidae is based on very inadequate material, as will become evident below. GIRDLES AND LIMBS OF DEINOCEPHALIA 207 Genus Tapinocephalus Owen (Figs. 1 a-b and 2) In the collection of the South African Museum there is only one specimen in which there is an identifiable skull associated with some parts of the postcranial skeleton. This skull (S.A.M. 2344) is associated with two humeri and a scapula (S.A.M. 3355). Scapulae. a. Tapinocephalus atherstonei, S.A.M. 3355. Lateral view. (x 3.) b. Tapinocephalus atherstonei, S.A.M. 3355. Posterior view. (x %.) c. Keratocephalus moloch, S.A.M. 11937. Lateral view. (x %.) Note: All the figures in this paper are orthoprojections and not perspective drawings. In the girdles the figures labelled ‘‘lateral view’’ are of the bones in natural position projected on to the median plane, “‘anterior’’ and ‘‘posterior’’ are on to the one ‘ plane at right angles to the median plane and ‘‘dorsal’’ and “‘ventral’’ on to the other plane at right angles to the median plane. The scapula (Fig. I a-b) is relatively low (530 mm.), with a broad blade (as reconstructed the width in its upper part is 270 mm.); the tricipital ANNALS OF THE SOUTH AFRICAN MUSEUM 208 ‘om} 4SIIT ou} 0} so[sue 4Yy81I ye ourfd Jay}Oo ay} uO pue oaoqe oy} 0} sapsue yysi1 ye oueld Terxe oy} 0} uO st ,,JoroyUR,, ‘saApuoorda oy} Jo soKzIns [esiop 94} Jo surfd sy} 0} uO _ (eurxoid a, suotyefoid ore ,[esiop,, pue ,[eI}UaA,, ToUINY 94} Ul :9}0N ‘MarIA [eIZUZA *LoOS ‘NV'S “9 ‘MoIA TeulrxoIg ‘“SSEE “FWeWwsS ‘Pp ‘mata olayUy "SSEE CIWS “9 ‘mora [esioq “SSEE ‘Wy'S ‘q ‘Mora [erzUOA “SSEE “pry's ‘ke (2X) “tauopzssayjzn snjyoydasourdyy jo trounzy tA Se amr ee aonste ade, GIRDLES AND LIMBS OF DEINOCEPHALIA 209 ridge forms a strong but low mound; the internal opening of the foramen supracoracoideus does not open into the subscapular groove, which is very shallow indeed; the glenoidal facet is small and faces postero-ventrally and but very little externally; the upper part of the blade is fairly straight with little curvature to fit round the thorax. The humerus (Fig. 2). Associated with the skull from Uitkyk (S.A.M. 2344), described by Haughton and correctly referred to Tapinocephalus, there is a fairly complete left humerus and a weathered proximal half of the right humerus (S.A.M. 3355). This association enables us to clear up the confusion which exists in regard to the Tapinocephalus humerus. The story of this confusion is as follows: In the first instance Owen (23) mistakenly referred the humerus B.M. 43525p from Varsfontein, Prince Albert, to Pareiasaurus. Lydekker (22), in correcting this obviously mistaken identi- fication, himself erroneously referred it to Tapinocephalus. Broom (12), accepting Lydekker’s identification, subsequently referred a specimen which he sold to the American Museum (A.M.N.H. 5611) to Tapinocephalus. Later Broom (13) figured a humerus (collection not stated) and states that this represents the true humerus of Tapinocephalus and at the same time refers B.M. 43525 to Titanosuchus. To make the confusion more confounded Broom then contradicts Seeley’s (28) identification of B.M. 49369 as the humerus of Titanosuchus and refers this humerus to Tapinocephalus. The Uitkyk humerus (S.A.M. 3355) associated with a Tapinocephalus skull, has no ectepicondylar foramen. In this feature it agrees with the known humeri of all the other members of the infra-order Tapinocephalia. Whereas all the humeri known to be associated with skulls classified as belonging to the infra-order Titanosuchia do have a well-developed ect- epicondylar foramen. Thus quite apart from other features still to be considered below, the humeri A.M.N.H. 5611, B.M. 49369 and Broom 1928 (no collection or number given) must all belong to some genus of the Titanosuchia. B.M. 49369 should be left where Seeley put it, viz. in the genus Jitanosuchus and A.M.N.H. 5611 in Jonkeria and Broom’s 1928 humerus in Scapanodon and B.M. 43525 is the humerus of Phocosaurus. The humerus of Tapfinocephalus may be described as follows: very large (length 520-590 mm.); not very massive; proximal expansion great (width 276 mm.); distal expansion moderate (width 246-282 mm.); shaft fairly long and moderately thick (diams. 105 xX 90 mm.); the delto-pectoral crest long, strong, terminating well proximal of the ventral opening of the entepicondylar foramen; caput oval in outline and lying in the same plane as the processus medialis and the processus lateralis; the processus medialis thus lies far proximally; the capitellum (radial condyle) is not well modelled, is weak, and lies much distally with its ventral part lying well distal of the plane in which the entepicondylar foramen opens; twist on shaft fairly 210 ANNALS OF THE SOUTH AFRICAN MUSEUM great (35°—~40°); the lateral median line is weak and the antero-dorsal ventral line is fairly weak; the entepicondyle is not greatly expanded and the foramen enters ventro-postaxially and leaves in a narrow ventral slit; the ectepicondyle is moderately expanded to form a thin unperforated. curved plate with a sharp edge. No other bones of the girdles or limbs in Tapinocephalus are known with any certainty, but below it will be seen that certain elements could probably be those of Tapinocephalus. Tapinocephalus atherstoner Owen The specific description is as for the genus. Referred specimens in the S.A.M. collection: S.A.M. 3355. An imperfect left.scapula (Fig: 1 \.a-b)jaay game. leit humerus (Fig. 2 a-d) and the weathered proximal end of the right humerus, associated with a good skull. Uitkyk, Beaufort West. Low Tapinocephalus zone. Coll. Haughton. | S.A.M. 5007. An isolated good left humerus (Fig. 2e). Wilgerfontein, Prince Albert. Low Tapinocephalus zone. Coll. Haughton. Genus Phocosaurus Seeley This description is based mainly on the Varsfontein material in the British Museum (B.M. 43525), incorporating the accounts of Seeley (27), Lydekker (22) and Watson (20). The pectoral girdle is large and massive; the scapula probably fairly low (lower than reconstructed by Watson, Fig. 11, which is given x 4, whereas it is in fact x 4); the scapular blade is fairly broad (240 mm.); the tricipital ridge is low; the glenoidal facet faces well externally. The precoracoid is inadequately known; ‘‘foramen opens into a distinct pit on the visceral surface’ (Watson). The coracoid is massive with the facet facing much externally. The humerus (Fig. 3) is large and massive (length 492 mm.); proximal (276 mm.) and distal (306 mm.) expansions large; shaft short and wide (diams. 168 X 96 mm.); the delto-pectoral crest very long and nearly reaching the plane of the ventral opening of the entepicondylar foramen; the caput is widely oval and lies in the same plane as the processus medialis and the processus lateralis, which also lies far proximally; the radial condyle is strong, fairly thick and well modelled, but situated much distally and not extending much proximally along the ventral face and thus well distal of the plane in which the entepicondylar foramen opens; the twist on the 2i1 GIRDLES AND LIMBS OF DEINOCEPHALIA "MOIA [CUITXOIg D “MOTA joweyuy "9 tee 4, “7, 4, *MOTA at 4,044 5 lesiog “q ‘Me IA [eI}UOA “Be ‘SZSEP . Wa (%%* ) UOLYISIBAUL SNANDSOIOYY JO sniduIn fy 2—Annals 212 ANNALS OF THE SOUTH AFRICAN MUSEUM shaft moderate (25°); the L.M.L. is fairly distinct and the A.D.V.L. is not prominent; the entepicondyle is strongly developed as a thick plate of bone and the ventral opening of the foramen is slitlike and situated well away from the edge of the bone; the ectepicondyle is developed as a greatly flaring thin sheet of curved bone The ulna (see Seeley’s Pl. 22) is short (320 mm.); the dorsal lip to the sigmoid face is fairly strong; the styloid ridge is prominent; width over coronoid process is moderate (204 mm.). The pelvis (see Seeley, Pl. 21, Lydekker, Fig. 17 and Watson, Fig. a The pubo-ischiadic plate is short (as reconstructed + 80% of the total height of the pelvis); the ilium appears to have been fairly low in its supra- acetabular part and fairly long antero-posteriorly; the ilio-fibularis ridge on the posterior iliac process lies fairly horizontally and appears to have been thickly rounded; the antero-ventral edge of the pubis is strongly everted with the tuberculum pubis confluent with the thickened antero-ventral edge curving in towards the median line. The femur (see Owen (24), Pl. 17, Fig. 8) is only represented by a proximal half. The proximal expansion is fairly broad (225 mm.); the preaxial face is fairly deeply concave with the caput directed well pre- axially; caput fairly massive; external trochanter not distinctly separated from the proximal face; the internal trochanter lies well in and is developed as a prominent oval tubercle; the shaft is fairly slender (diams. 112 xX 72, mm.). Phocosaurus megischion Seeley The specific description is as for the genus. Type: B.M. 43525. Incomplete coraco-scapulae, humeri, mght ulna, incomplete pelvis and the proximal half of the mght femur. Vars- fontein, Prince Albert. Middle Tapinocephalus zone. Coll. Atherstone. Referred specimens in the S.A.M. collection: S.A.M. 11300. (Figs. 4 and 5.) A right humerus, showing a pathological lesion and an incomplete ilium. Deesweesfontein, Laingsburg. Low Tapinocephalus zone. Coll. Boonstra. S.A.M. 11988. Proximal half of an isolated left femur. Boeteka. Beaufort West. High Tapinocephalus zone. Coll. Boonstra. Genus Keratocephalus v. Huene Although there are four skulls of Keratocephalus in the collection of the South African Museum there are only a few bones of the girdles and limbs, 213 GIRDLES AND LIMBS OF DEINOCEPHALIA “MaIA [esIog) “qd SUIMOL [s pu « 3) [eystd) "OOfII ‘MOIA JoIN}Uy °9 uoIse, Teorsoroyzed (2X) "MOTA [eI]JUIA “ke WYVS ( “UOLYyISId at v UWIOIf SBUTZ[NSeI UWoTsor SNHANDSOIOY JO SNIIUIN]{ wre eh F! eee Ae: 214 ANNALS OF THE SOUTH AFRICAN MUSEUM but, together with von Huene’s account of some bones in Tiibingen, the following incomplete description can be given. Of the pectoral girdle only a piece of a scapula (Fig. 1c) is known and this is very similar to the corresponding part of the scapula of Tapinocephalus. In the collection there are two imperfect ulnae (Fig. 6a). The ulna is massive and squat (length + 350 mm.); the styloid ridge is prominent; broad over the shaft and very broad over the coronoid process. The only known radius Fig. 5. (Fig. 6 b-d) is massive and : squat (length 240 mm.); the ventral face is concave with a strong longitudinal ridge; the flange (for the insertion of the biceps) on. the proximo- postaxial corner is strong; the proximal facet is convex. With no pelvis in this col- lection I am _ extracting the characteristic features from von | Huene’s (21) account. The supra-acetabular part of the ilium is low and very long (the height is only 52% of its antero-posterior length); the anterior process of the iliac blade is very long and low and Ilium of Phocosaurus megischion. (x %.) is fairly strongly everted; the S-A.M. 11300. Lateral view. low posterior process is fairly long, but much shorter than the anterior process, with the ilio-fibularis ridge sharp and prominent and lying horizontally; the dorsal iliac edge is not folded over laterally; on the inner face of the anterior process there is a large face to receive a rib lying anterior to the main sacral mb. The femur (see v. Huene’s Figs.) is short (400 mm.) and broad; the width over the external trochanter is 256 mm.; the preaxial face is deeply concave with the caput much preaxially directed; the external trochanter is not demarcated from the proximal face, which extends far outwards from the caput; the internal trochanter developed as a prominent tubercle; the shaft is short and wide (diams. 132 x 84 mm.); the distal facets are terminal. With only one crushed tibia in our collection (Fig. 6 e-f) and the one figured by von Huene, the Keratocephalus tibia appears to be massive GIRDLES AND LIMBS OF DEINOCEPHALIA 215 but short (length 232-276 mm.) with a broad shaft and a massive cnemial eminence. Keratocephalus moloch. (x %.) a. Ulna of S.A.M. 11937. Dorsal view. b. Radius of S.A.M. 11937. Dorsal view. c. Radius of S.A.M. 11937. Ventral view. d. Radius of S.A.M. 11937 Posterior view. e. Tibia of S.A.M. 8946. Dorsal view. f. Tibia of S.A.M. 8946. Posterior view. Keratocephalus moloch von Huene The specific description is as for the genus. Type: Tiibingen. An ilium, femur and tibia, associated with parts of the skull. Abrahamskraal, Prince Albert. Low Tapinocephalus zone. Coll. v. Huene. Referred specimens in the S.A.M. collection: S.A.M. 8946. An imperfect left ulna and a crushed left tibia (Fig. 6 e-f), associated with a skull. Mynhardtskraal, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. 216 ANNALS OF THE SOUTH AFRICAN MUSEUM S.A.M. 11937. An imperfect right scapula (Fig. 1c), an imperfect left ulna (Fig. 6a) and a good nght radius (Fig. 6 b-d), associated with a skull. Buffelsvlei, Beaufort West. Low Tapincephalus zone. Coll. Boonstra and Marais. Genus Pelosuchus Broom A part of the scapula preserved indicates a fairly close similarity to the scapula of Keratocephalus, as does also the incomplete weathered coracoid. The femur (Fig. 7) is fairly j short (420 mm.) and broad; the Fig. 7. width over the external tro- chanter is 215 mm.; the preaxial face is deeply concave with the caput directed much preaxially; the external trochanter does not appear to be demarcated by a notch from the proximal face; the internal trochanter lies near the middle of the bone far away from the preaxial border and is a prominent and strong tubercle; the shaft is fairly strong and broad (diams. 132 x 80 mm.); the distal facets are terminal. A distorted tibia appears to be stout and short with a strong cnemial eminence. Pelosuchus priscus Broom The specific description is as Femur of Pelosuchus priscus. (x #.) S.A.M. 918. a. Ventral view. b. Anterior for the genus. view. Note: In the femur ‘‘dorsal’’ and Type: S.A.M. gre. “Part or “‘ventral’’ are projections on the plane . in which the ventral faces of the condyles = 4 scapula, coracoid, : lie and “‘anterior’’ on to the axial plane fairly good femur (Fig. 7) at right angles to the above. and distorted tibia, associated with a weathered dentary. Bokfontein, Prince Albert. Middle? Tapinocephalus zone. Coll. Cairncross. Generically Undetermined Specimens: S.A.M. 2753. A middle portion of a humerus (Fig. 12a) and the distal end of a femur. Viviers Siding, Beaufort West. Mid Tapino- cephalus zone. Coll. Haughton and Whaits. S.A.M. 9097. An isolated femur (Fig. 8). This is probably a femur of Tapinocephalus. Merweville Commonage. Low Tapinocephalus zone. Coll. Boonstra. GIRDLES AND LIMBS OF DEINOCEPHALIA 217 Above: Femur, probably of Tapinocephalus. (x &-) S.A.M. 9097. a. Dorsal view. b. Ventral view. c. Anterior view. Left: Interclavicle of ? Tapinocephalus. (xX &-) S.A.M. 9153. a. Ventral view. b. Lateral view. 218 ANNALS OF THE SOUTH AFRICAN MUSEUM S.A.M. 9153. An interclavicula (Fig. 9) and a distorted femur (Fig. 10). The interclavicle has a long slender stem and the femur a ridgelike internal trochanter. These may represent a genus lying between Tapinocephalus and Struthiocephalus. Jacobskraal, Prince Albert. Low Tapinocephalus zone. Coll. Boonstra. Fig. 10, ° Cigtete teks, aneiee eeoeet 6 . “e ° % eens 7.5.4 8 < @e os e *. * 4, . ae s ee ru oe oe Femur of ? Tapinocephalus. (x #@.) S.A.M. 9153. (Slightly distorted.) a. Dorsal view. b. Ventral view. c. Anterior view. S.A.M. 9164. The middle portion of a humerus. Wakkerstroom, Prince Albert. Low Tapinocephalus zone. Coll. Boonstra. S.A.M. 11303. An isolated imperfect femur. Buffelsvlei, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. S.A.M. 11306. An isolated good femur (Fig. 11), which agrees well with the proximal end of the femur (B.M. 43525) which I am including in the type material of Phocosaurus. The femur is quite robust and fairly long (498 mm.); the width over the external trochanter is moderate (234 mm.); the preaxial face is concave, with the caput directed well preaxially; the caput is massive (164 x II4 mm.); the shaft is fairly long and narrow (diams. 120 x 86 mm.); the tibial facets are GIRDLES AND LIMBS OF DEINOCEPHALIA 219 directed much distally and the postaxial epicondyle lies further distally than the preaxial epicondyle; the femoro-tibialis ridge is fairly strong and the area for the insertion of the ilio-femoralis is narrow. Boesmansrivier, Beaufort West. Mid Tapinocephalus zone. Coll. Boonstra. | Femur of ? Phocosaurus. (x 3.) S.A.M. 11306. a. Dorsal view. b. Ventral view. c. Anterior view. S.A.M. 11702. An isolated distal end of a humerus (Fig. 12 b-d) which agrees fairly well with that of Tapinocephalus. Elandsberg, Suther- land. Low? Tapinocephalus zone. Coll. Boonstra. S.A.M. 11993. An isolated good left humerus (Fig. 13) with a long delto-pectoral crest and a well-modelled radial condyle as in Phoco- saurus, but otherwise much like the humerus of Tapinocephalus. Locality and collector unknown. S.A.M. 11997. An incomplete ilium (Fig. 14), distal end of a femur and the distal end of a radius, associated with a fairly good but as yet unprepared skull. ANNALS OF THE SOUTH AFRICAN MUSEUM 220 “MOIA JOUEIUY “ZOLII “W'V'S “P ‘MIA [esIOG *zoLiII WW'S °9 "MOIA [eIZUIA ‘cOdIT ‘W'Y'S “Q “mata Te}UeA “€Sz “Wry's “e (- X) ‘snpoyderourdvy ¢ Jo iewnyzy ‘tT “Big GIRDLES AND LIMBS OF DEINOCEPHALIA 221 The ilium has most of its blade missing, but on the lower edge of the posterior iliac process there is a strong ilio-fibularis ridge lying horizontally; the acetabulum is very large but shallow. In its unprepared state the skull cannot be definitely identified; it is certainly not that of Keratocephalus and in shape differs from that of Tapinocephalus and may thus very possibly be near Phocosaurus. Locality and collector unknown. Fig. 15. Humerus of ? Tapinocephalus. (x é-) S.A.M. 11993. Ventral Ilium of ? Phocosaurus. (x %.) view. S.A.M. 11997. Lateral view. Struthtocephalidae Family Characters of the Girdles and Limbs. With much more material at our disposal than in the case of the Tapinocephalidae a much fuller account can be given of the girdles and limbs of the Struthiocephalidae. The pectoral girdle is small to medium sized and light to only fairly massive. 222 ANNALS OF THE SOUTH AFRICAN MUSEUM The scapula is small and low (310 mm.) to fairly large and high (480-500 mm.) with the upper part of the blade expanded (120-230 mm.); its posterior border is fairly concave; the tricipital ridge is not very strongly developed; the subscapular groove is shallow and confluent with a groove lying in the visceral face of the precoracoid, dorsally of the inner opening of the supracoracoid canal; the glenoidal facet of the scapula is of medium size, concave and with a fairly well-moulded and raised external rim and it faces postero-ventrally but not externally. The coracoidal plate is long antero-posteriorly. The precoracoid is fairly large and consists of a fairly thin plate of bone, but is thickened at the apex, which lies dorso-posteriorly of the outer opening of the supracoracoid foramen; its outer face is only very slightly convex; the supracoracoid canal is not directed very obliquely and its inner opening still lies in the precoracoid near the precoracoid-scapular suture, but has a groove dorsally confluent with the subscapular groove. The precoracoid is excluded from the glenoid and on the apex it carries a ridge which limits the anterior movement of the humerus. The coracoid is a fairly strong element with a fairly large glenoidal facet which faces dorso-posteriorly and also externally. No cleithrum is preserved but in all probability was a splintlike bone as in Moschops. A pair of clavicles is preserved in Struthiocephalus whaitsi, but as is evident from Fig. 17 they have suffered from distortion, so that the right one appears to be a much squatter bone than the left one. The left clavicle, which I believe shows the natural form more truly, is a fairly light bone with an expanded lower end, which is applied to the outer face of the upturned interclavicular antero-lateral corner; the upper end has its anterior edge thickened and posteriorly there is a thin flange, which is applied to the outer face of the scapula; internally a ridge limits the posterior movement of the clavicle over the scapula. The interclavicle is a strong bone with a fairly thick and long stem, which is, however, only moderately wide; the anterior spatulate end is considerably expanded and curves strongly upwards; the moderately hollowed surface for the reception of the clavicle is dorso-posteriorly bounded by the thickened raised edge of the bone; the upper surface of the stem has no median ridge for the lower edge of the coracoidal plate to abut against. The humerus of this family varies considerably; in some cases it is fairly short and broad and in others long and fairly slender; in length it varies from small to fairly large (276?-475 mm.); the proximal expansion is moderate to large (156-270 mm.); the distal expansion is moderate to very large (174-288 mm.); the shaft is short to long and narrow to very broad GIRDLES AND LIMBS OF DEINOCEPHALIA 223 (width 84-125 mm.); the delto-pectoral crest is short to fairly long and it terminates, in all cases, well proximal to the plane in which the ventral opening of the entepicondylar foramen lies; the caput is narrowly to moderately narrowly oval; both the processus medialis and the processus lateralis lie far proximally, more or less in the same plane as the caput; the radial condyle is weak, not well modelled, and does not extend much proximally along the ventral surface and thus always lies well distal of the plane in which the entepicondylar foramen lies; the twist on the shaft is small to moderate (8°—-20°) and the one case in which it is 50°, this is undoubtedly due to postmortem distortion; the L.M.L. is weak to fairly strong; the A.D.V.L. is moderately to well developed; the entepicondyle is weak to moderate and the foramen slitlike; the ectepicondyle is little. to greatly expanded. The ulna is very inadequately known and is apparently fairly light and relatively long (258-366? mm.). The radius is only known from a couple of ends. The pelvis, hitherto unknown except for Haughton’s (20) photograph of the mounted skeleton of Struthiocephalus and Broom’s (15) restoration sketch based on it, is represented in the collection by three fairly well preserved specimens and a fourth with two iliae. The three bones of the pelvis are weakly united and the symphysis between the two halves is weak. The pubo-ischiadic plate is relatively fairly long (the antero-posterior length is 9g0-102% of the height ot the pelvis); the supra-acetabular part of the ilium is low (45-65% of its antero-posterior length) and long antero- posteriorly; the anterior process of the ilium is long and low and only slightly everted; the posterior process of the ilium is not much shorter than the anterior process, but is much lower, with its posterior edge lying nearly horizontally and partially everted to form a weak ridge (for the insertion of the m. ilio-fibularis), which also lies nearly horizontally; the dorsal iliac edge is folded over laterally to overhang and limit the gluteal area; on the: inner face of the anterior iliac process there is a strong attachment of a mb anterior to the main sacral rib. In the pubis the upper part of the anterior edge is strongly everted to form an elongated tuberculum pubis, which ‘is: ventrally demarcated from the slightly thickened antero-ventral edge by a’ distinct step; in the median line the middle part of the pubis meets its fellow in a very weak pubic symphysis; the middle part of the ventral’ edge of the ischium is thickened to form a large sutural face for a fairly strong ischial symphysis; between the pubic and ischial symphyses there is a large quadrangular fenestra, in life filled with cartilage. The femur of the Struthiocephalidae shows considerable variation: small: (length 276 mm.), light and slender or medium sized (length 408-440 mm.},: 224 ANNALS OF THE SOUTH AFRICAN MUSEUM fairly robust and broad or fairly long (468 mm.), fairly light and fairly slender; the width over the external trochanter is small (95 mm.) to great (210 mm.); the preaxial face is moderately to strongly concave with the caput only slightly or moderately preaxially directed; the external trochanter is not separated from the proximal face by any notch; the internal trochanter is undeveloped or forms a strong elongated tubercle, situated well away from the preaxial border; the shaft is fairly long to short and fairly slender to broad (55-125 mm.); the tibial facets are much distally directed; the m. femoro-tibialis originates on a fairly strong ridge; the ilio-femoralis area of insertion is narrow or broad. The tibia is small (length 180 mm.) and slender to medium sized (length 240-270 mm.) and fairly slender with a well-developed cnemial ridge or eminence, and cnemial groove. The fibula is light, fairly short to long and slender (228-288 mm.). Genus Struthiocephalus Haughton The pectoral girdle is of medium size and only fairly massive (total height as projected on to the median plane 696 mm.). The scapula is fairly massive and high (480 mm.); its posterior border is deeply concave. As the family description is based mainly on specimens of this genus refer back for additional characters of the pectoral girdle. The humerus is preserved in two specimens with skulls associated, but in one, in addition to being juvenile, the humerus has undergone considerable postmortem distortion and in the other proximo-distal compression has shortened the bone with concomitant widening. The humerus is fairly large (length 393-430 mm.) and fairly massive; the proximal expansion is large (270 mm.) and the distal expansion very large (288 mm.); the shaft is fairly long to short, moderately thick (diams. 102 X 60 mm.) to robust (diams. 126 x 60 mm.); the delto-pectoral crest is of medium length, terminating fairly far proximal of the ventral opening of the entepicondylar foramen; the caput is narrowly oval and lies slightly proximal to the processus medialis; the twist on the shaft is moderate (20°; in the deformed S.A.M. 11493 it is 50°); the L.M.L. is fairly strong; the A.D.V.L. is well developed; the entepicondyle is fairly strong with the foramen entering dorso-postaxially and leaving as a ventral slit; the ect- epicondyle is greatly expanded as a thin sheet of bone. The ulna and radius are inadequately known. The pelvis is as described for the family; it may be noted here that in both specimens known, but in particular in the juvenile one, the ilium has not grown out to meet the other two bones in the central part of the GIRDLES AND LIMBS OF DEINOCEPHALIA 225 acetabulum, whereas the three bones meet in the pelvis of Struthtocephatellus which on account of its small size might have been considered a not fully grown Siruthiocephalus. Fig. 15. A.M. 3012. Lateral view. ( b. Coraco-scapula of Struthiocephalus whaitsi. S.A.M. 11493. Lateral view. ( a. Pectoral girdle of Struthiocephalus whaitsi. S. The femur is of medium size (length 440 mm.) and fairly broad and robust; the width over the external trochanter is 204 mm.; the preaxial face is moderately concave and the caput only slightly preaxially directed; the internal trochanter forms a fairly strong ridgelike tubercle; the shaft = ~weeecoeo ®* ~» 226 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 16. Interclaviclae of Struthiocephalus whaitsi. (yx 4.) a. S.A.M. 3012. Ventral view. b. S.A.M. 3012. Lateral view. c. S.A.M. 11579. Ventral view. d. S.A.M. 11579. Lateral view. GIRDLES AND LIMBS OF DEINOCEPHALIA 227 short and broad (diams. 125 xX 50 mm.); as preserved the postaxial tibial facet lies much distally; the ilio-femoralis area of insertion is broad. The tibia appears to be fairly short (240 mm.). The fibula is fairly short (230 mm.) but fairly robust with a deeply concave preaxial border. Struthiocephalus whaitsi Haughton The specific description is as for the genus. Referred specimens in the S.A.M. collection: S.A.M. 3012. A nearly complete and fairly good pectoral girdle (Figs. 15a, 16 a-b and 17 a-d), a good humerus (Fig. 18), a fairly good pelvis (Fig. 19), two somewhat crushed femora (Fig. 20), a fair tibia (Fig. 21) and fibula (Fig. 22), associated with a fairly good but crushed skull. Abrahamskraal, Prince Albert. Low Tapino- cephalus zone. Coll. Haughton. a b Cc d e Claviculae. (x 4.) a. Struthiocephalus whaitsi. S.A.M. 3012. Lateral view of right clavicle. b. Struthiocephalus whaitsi. S.A.M. 3012. Inner view of right clavicle. c. Struthiocephalus whaitsi S.A.M. 3012. Lateral view of left clavicle. d. Struthiocephalus whaitst. S.A.M. 3012. Inner view of left clavicle. e. Moschops capensis. S.A.M. 11972. Inner view of left clavicle. ANNALS OF THE SOUTH AFRICAN MUSEUM 228 *MOTA "MOIA [CUITXOIg ‘Pp TerqusA °C (RX) ‘z10€ ‘MOIA JOMOJUy ‘9 ‘MITA [eSIOg ‘q ‘WV'S ‘isquvym = snjoydar014jnM4S jo snsoUIny;y GIRDLES AND LIMBS OF DEINOCEPHALIA 229 S.A.M. 11493. A fair coracoidal plate (Fig. 15b), a distorted humerus (Fig. 23) and a good pelvis (Figs. 24 and 25), associated with a fairly good skull. Juvenile. Mynhardtskraal, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. Fig. 19. Struthiocephalus whaitsi. S.A.M. 3012. (x %.) a. Pelvis in lateral view. b. Ischium in inner view. c. Ilium in dorsal view. S.A.M. 11572. A fair interclavicle, most of both coracoidal plates and the proximal ends of both scapulae (Fig. 26). Klein-Koedoeskop, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. S.A.M. 11579. A good proximal two-thirds of a humerus, a good precoracoid and a good interclavicle (Fig. 16 c-d). Buffelsvlei, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. 230 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 20. 5 STS) Ly Ohpiee, i we ‘ te e @e, o oe i O Sen. PO LO AES Dra dae eo sate e Oe rth Kone Q eta Femora of Struthiocephalus whaitsi. S.A.M. 3012. (x @.) a. Left femur in dorsal view. b. Right femur in ventral view. c. Left femur in anterior view. d. Right femur in anterior view. a b Cc Fibula of Struthiocephalus wha ae : we S.A.M. 3012. (x %.) a. Dag Tibia of Struthiocephalus whaitsi. S.A.M. 3072.. (x 4:) view. b. Ventral view. c. Anteri a. Dorsal view. b. Ventral view. c. Posterior view. view. 4 231 GIRDLES AND LIMBS OF DEINOCEPHALIA TeyuIA *e “MOIA [PUITXOIG ‘Pp (¥ Xx) “C6PrI ‘W "MOIA JOWOJUW “9 “MOIA [RSIOG] "Q “MOIA Vs “IsqDym Snioydar01yinséIG atueAnl we JO sniswnyyT 232 ANNALS OF THE SOUTH AFRICAN MUSEUM Struthiocephalus whaitsi. Juvenile. S.A.M. 11493. (x 4.) a. Ilium in dorsal view. b. Pelvis in lateral view. c. Pelvis in anterior view. GIRDLES AND LIMBS OF DEINOCEPHALIA 233 S.A.M. 11939. A weathered humerus, ulna (Fig. 27 a-b) and femur (Fig. 27 d-f), associated with a jaw fragment. Dikbome, Laingsburg. Low Tapinocephalus zone. Coll. Boonstra. weer: .) SE ai we ease Crs Pelvis of Struthiocephalus whaitsi. Juvenile. S.A.M. 11493. (x %.) Ventral view. Specifically undetermined specimen: The following specimen approaches sufficiently close to warrant pro- visional inclusion in the genus, but not in the species — whattsz. S.A.M. 11941. se ee, CBee SLOT wom F501 Interclavicle of Struthiocephalus sp. S.A.M. 11941. (x 4.) a. Ventral view. b. Lateral view. Genus Struthiocephaloides Boonstra The pectoral girdle is only known from a single scapula; the scapula is fairly large and high (500 mm.); the upper part of the blade is expanded (210 mm.) and the posterior face is not deeply concave. No other part of the girdles and limbs is known. Struthiocephaloides cavifrons Boonstra Specific description as for the genus. Type: S.A.M. 5607. A left scapula (Fig. 29 a-b), associated with a good skull. Lammerkraal, Prince Albert. High Tapinocephalus zone. Coll. Haughton. GIRDLES AND LIMBS OF DEINOCEPHALIA 237 Fig. 29. = eco @ Scapulae. (x %.) a. Struthiocephaloides cavifrons. S.A.M. 5607. Lateral view. b. Struthiocephaloides cavifrons. S.A.M. 5607. Posterior view. c. Struthio- cephalellus parvus. S.A.M. 5006. Lateral view. Genus Struthiocephalellus Boonstra The pectoral girdle is only known from an imperfect scapula and frag- ments of the coracoidal plate; the scapula is small (height 310? mm.) and the width of the blade is 210? mm.; the posterior face of the scapula is only slightly concave. The humerus is small (length 276? mm.) and is lightly built; the proximal expansion is fairly great (156 mm.); the distal expansion is relatively great (174 mm.); the shaft is fairly long and fairly wide (diams. 66 x 48 mm.); the delto-pectoral crest is only moderately long and is fairly weak and terminates well proximal of the plane in which the ventral opening of the entepicondylar foramen lies; the relations of caput, processus medialis and lateralis are unknown; the radial condyle is weak (thickness 48 mm.), distally situated and curving round on to the dorsal surface; the twist on the 238 ANNALS OF THE SOUTH AFRICAN MUSEUM shaft is moderate (20° ?); the L.M.L. is weak and the A.D.V.L. fairly strong; the entepicondyle is expanded as a thin sheet of bone and the foramen enters ventro-postaxially and leaves as a ventral slit well away from the edge of the bone; the ectepicondyle is expanded as a thin curved sheet of bone. me The ulna has only the proximal part preserved; it is lightly built; proximal to the coronoid process a sharp longitudinal ridge separates a groove from the sigmoid surface; the ventral surface is deeply excavated longitudinally. The pelvis has a relatively long pubo-ischiadic plate (97% of the height of the pelvis); the supra-acetabular part of the ilium is low (126 mm.) and long (200? mm. as restored) and the height is thus 63% of the length; although neither the anterior nor posterior iliac processes are fully preserved both appear to have been long and fairly low. The pubis is anteriorly slightly everted with a prominent tuberculum pubis, clearly demarcated from the medially directed antero-ventral edge of the pubis; the pubic symphysis is fairly long but is weak. The ischium is long with a long but weak symphysis; the postero-dorsal edge of the ischium is strongly thickened. The femur is small, slender and fairly lightly built (length 276 mm.); the width over the external trochanter is small (95 mm.); the preaxial face is only slightly concave and the caput, which is relatively weak (55X25 mm.), is directed much dorsally as well as preaxially — there is thus a twist on the shaft; the external trochanter is indistinctly developed and the internal trochanter not developed at all; the shaft is long and fairly narrow and thin (diams. 55 xX 48 mm.); the tibial facets face ventro-distally with the postaxial one lying further distally; the femoro-tibialis mdge is moderately strong and the ilio-femoralis area of insertion very narrow. The tibia has its proximal face divided into two distinct concave facets by a sharp and strong dorso-ventral ridge; a prominent cnemial protuberance is present, continued distally as a sharp cnemial ridge, with a deep groove lying postaxially of this ridge; all these features are well and sharply modelled. Struthocephalellus parvus Boonstra Type: S.A.M. 5006. A partial scapula (Fig. 29c), a fairly good humerus (Fig. 30), the proximal end of an ulna (Fig. 27c), a fairly good pelvis (Fig. 31), a good femur (Fig. 32) and the proximal end of a tibia (Fig. 33), associated with a fair skull. Abrahamskraal, Prince Albert. Low Tapinocephalus zone. Coll. Haughton. GIRDLES AND LIMBS OF DEINOCEPHALIA 239 Generically undetermined specimens: S.A.M. 8947. This specimen, which consists of two fairly good femora (Fig. 34 a-c), a good tibia (Fig. 34 d-f) and a good fibula (Fig. 34 g-i), appears to be of a Struthiocephalid, larger and of somewhat heavier build than Struthiocephalus whaits:. The tibia is note- worthy for its very strong cnemial protuberance. Mynhardtskraal, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. Fig. 30. Cc Humerus of Struthiocephalellus parvus. S.A.M. 5006. (x 3.) a. Ventral view. b. Dorsal view. c. Anterior view. Fig. 3], The following four specimens are of Struthiocephalids which are much longer in the limb than Struthiocephalus. S.A.M. 3614. A_ good isolated humerus (Fig. 35 b-e). This humerus is relatively long and slender (length 480 mm.); both the proximal (width 192 mm.) and the distal (width 198 mm.) expan- sion is small; the shaft is fairly long and slender (diams. 95 x 78 mm.); the twist on the shaft is small (8°); the capitellum is very weakly modelled and situated far é distally; both epicondyles are weak (x 4) and the entry of the entepicon- parvus. S.A.M. 5006. Lateral view. 240 ANNALS OF THE SOUTH AFRICAN MUSEUM dylar foramen is clearly visible in dorsal view. Abrahamskraal, Prince Albert. Low Tapinocephalus zone. Coll. Haughton. Fig. 32. Right: Femur of Struthio- cephalellus parvus. S.A.M. 5006. (x ¢-) a. Dorsal view. b. Ventral view. c. Anterior view. Left: Tibia of Stvuthio- céphalellus parvus. S.A.M. 5006. (Sees) Ca orsal view. b. ~Ventral view. c. Posterior view. S.A.M. 5009. A fair humerus with the proximal end not in contact with the shaft (Fig. 35a). Wolwefontein, Prince Albert. Low Tapino- cephalus zone. Coll. Haughton. Although the delto-pectoral crest is shorter than in S.A.M. 3614 and the processus medialis situated more nearly in the same plane as the caput these two specimens probably are of the same species. S.A.M. 4349. Two fairly good iliae (Fig. 36 a-b), a good femur (Fig. 36 c-e), the proximal halves of both ulnae (Fig. 36f) and of both tibiae and a fibula (Fig. 36g). GIRDLES AND LIMBS OF DEINOCEPHALIA 241 | Fig. 34. | 2? Struthiocephalus. S.A.M. 8947. (> }.) a. Femur in dorsal view. b. Femur in ventral view. c. Femur in anterior view. d. Tibia in \ dorsal view. e. Tibia in ventral view. f. Tibia in posterior view. | g. Fibula in dorsal view. h. Fibula in ventral view. i. Fibula in anterior view. ANNALS OF THE SOUTH AFRICAN MUSEUM 242 ‘MOIA [CUWITXOIg ‘PIQE “IN'V'S ‘(9 ‘“MOIA ‘MOIA [esiog ‘FIQE ‘WW'S ‘9 ‘“MOIA [eI}UIA 600 “Wyss “@ (¥ X) ‘oeprpeydasoryynsys 94} jousjuy “bIg€ “W'V'S ‘P ‘PIOE CIN'W'S ‘GQ ‘MOIA [eI}JUOA jo snues powreuun ue jo touImNyy GIRDLES AND LIMBS OF DEINOCEPHALIA 243 Fig. 36. ees lal i An unnamed Struthiocephalid. S.A.M. 4349. (xX 4.) a. Ilium in dorsal view. ' b. Ilium in lateral view. c. Femur in dorsal view. d. Femur in ventral view. Le ° . ° . : . e . e. Femur in anterior view. f. Ulna in dorsal view. g. Fibula in ventral view. 3—Annals 244 ANNALS OF THE SOUTH AFRICAN MUSEUM 46 2° Sou scare An unnamed Struthiocephalid. S.A.M. 9008 Ose 3.) “ave Femur in dorsal view. b. Femur in ventral view. c. Femur in anterior view. d. Tibia in dorsal view. e. Tibia in ventral view. f. Tibia in posterior view. g. Fibula in ventral view. GIRDLES AND LIMBS OF DEINOCEPHALIA 245 The ilium is low (supra-acetabular height 210 mm.), with both the anterior and the posterior process long and low (antero-posterior length 335? mm.); with the height thus 62% of the length; the anterior process is slightly everted and on its inner face receives a rib lying anterior to the main sacral rib; the origin of the m. ilio-fibularis hes on the everted horizontal ridge on the posterior iliac process; the dorsal edge of the ilium is folded over the upper limit of the gluteal area. It is thus apparent that this ilium is very similar to that of Struthiocephalus. Scapulae of two indetermined Struthiocephalids. (x %.) a. S.A.M. 10197. Lateral Whew... De) SoA M. 101974 . Posterior, view.- c..'S.A.M. 211700. Lateral ..view. d: S.A.) 11700. ~ Posterior "view. The femur is long (468 mm.) and slender; the width over the external trochanter apparently small (Ig0? mm.); the preaxial face is quite strongly concave and the caput directed well preaxially; the internal trochanter is 246 ANNALS OF THE SOUTH AFRICAN MUSEUM situated fairly near the preaxial border; the shaft is long and narrow (diams. 102 X 66 mm.); the tibial facets are directed well distally with the postaxial one situated furthest distally. The ulna is apparently long with a strong longitudinal ridge and a deep excavation on its dorsal face. The tibia and fibula are both long and slender bones. Leeurivier, Beaufort West. Low Tapinocephalus zone. Coll. Haughton. S.A.M. 9008. A good femur (Fig. 37 a-c) and tibia (Fig. 37 d-f) and the proximal end of a fibula (Fig. 37g). Klein-Koedoeskop, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. Fig. 39. Femur of uncertain affinity. S.A.M. 11880. (x 2.) a. Dorsal view. b. Ventral view. c. Anterior view. This specimen is co-specific with S.A.M. 4349. These two specimens thus represent, together with S.A.M. 3614 and S.A.M. 5009, a Struthio- cephalid with an ilium very similar to that of Struthiocephalus but with much longer pro- and epipodials. The following two specimens both come from the Nieueveld, where the fauna of the Tapinocephalus zone is very poorly known. S.A.M. 1to197. An isolated well-preserved scapula and coracoid (Fig. 38 a-b). Grootfontein, Fraserburg. Low? Tapinocephalus zone. Coll. Boonstra. GIRDLES AND LIMBS OF DEINOCEPHALIA 247 S.A.M. 11700. < ai) a. Dorsal view. b. Ventral view. c. Anterior view. Left: Scapula of a Moschopid. Gen. Indet. S.A.M. 9003. (x 4.) a. Lateral view. b. Posterior view. 262 ANNALS OF THE SOUTH AFRICAN MUSEUM This is a medium sized (length 414 mm.) fairly robust bone (width over the external trochanter 180 mm.); the shaft is fairly long and moderately broad (diams. 102 X 72 mm.); both condyles lie in practically the same plane; the ilio-tibialis ridge is strong. This femur appears to lie about midway between that of Moschops and Pnigalion. Letjiesbos, Beaufort West. Mid Tapinocephalus zone. Coll. Maddison. Fig. 54. Above: Humerus of ? Mos- chops., S.A.M. 0157.7 (X-3-) a. Ventral view. b. Dorsal view. c. Anterior view. Left: Radius of ? Moschops. S.A.M. | 915750" ( p@teaen Dorsal view. b. Ventral view. c. Posterior view. GIRDLES AND LIMBS OF DEINOCEPHALIA 263 S.A.M. 9003. A good isolated scapula (Fig. 53). This scapula is high (540 mm.) with a fairly narrow blade (width 156 mm.) and with its posterior border fairly straight and the glenoid not facing externally. Klein-Koedoeskop, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. S.A.M. g101. The distal end of a fibula. Rietfontein, Prince Albert. Low Tapinocephalus zone. Coll. Boonstra. S.A.M. 9124A. A good isolated ulna (Fig. 40 a, a’). Very similar to the ulna of Moschops capensis. Voélfontein, Prince Albert. Low Tapmocephalus zone. Coll. Boonstra. 3 S.A.M. 9157. A good humerus (Fig. 54), radius (Fig. 55), ulna (Fig. 40 b, b’) and part of the carpus. I (4) have recently described the epipodial and the carpus. The humerus is of medium size (length 384 mm.); the proximal expansion is fairly large (210 mm.) and the distal expansion is moderate (212 mm.); the shaft is fairly wide and short (diams. 91 x 66 mm.); although the ulna is very similar to that of Moschops the radius is a much more slender bone. This specimen thus lies near Moschobs. Wolwefontein, Prince Albert. Low Tapinocephalus zone. Coll. Boonstra. S.A.M. 11581. An isolated flattened femur (Fig. 56). This small femur (length 372 mm.) is most probably of a juvenile Moschops. Buffels- vlei, Beaufort West. Low Tafinocephalus zone. Coll. Boonstra. S.A.M. 11987. An isolated weathered distal two-thirds of a humerus. The groove on the under surface of the ectepicondyle for the passage of the radial nerve is well developed. Buffelsvlei, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. S.A.M. 11995. An isolated radius (Fig. 57) with a well-developed proximo-postaxial flange. Stouter than S.A.M. 9157. Sutherland District. Low? Tapinocephalus zone. Coll. Boonstra. S.A.M. 3308. An isolated good fibula (Fig. 58). With a length of 252 mm. and its slender build it comes very near that of Moschobps. Uitkyk, Beaufort West. Low Tapinocephalus zone. Coll. Haughton. 264 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 57. Above: Radius of ? Moschops. S.A.M. 11995. (x ¢.) a. Dorsal view. b. Ventral view. c. Posterior view. Left: Femur of ? Moschops. S.A.M. 19581. (x%.) Ventral view. ao Job Cc Fibula of ? Moschops. S.A.M. 3308. (x %.) a. Dorsal view. b. Ventral view. c. Anterior view. Titanosuchia Infra-ordinal Characters of the Girdles and Limbs. The pectoral girdle is large and massive; the scapula is fairly to very high (530-690 mm.) with the upper part of its blade fairly to very broad GIRDLES AND LIMBS OF DEINOCEPHALIA 265 (220-426 mm.); the scapular head of the triceps originates from a fairly to very prominent ridge; the internal opening of the foramen suprecoracoideus opens into the deep subscapular groove, i.e. it crosses the suture between the precoracoid and the scapula; the glenoidal facet of the scapula is large, facing ventro-posteriorly and not, or but little, externally. The coracoidal plate is of great antero-posterior extent. The precoracoid is large and strong, with its anterior border thickened, and its greatly thickened dorso-posterior apex forms at most the anterior depressed rim of the glenoid; the foramen supracoracoideus penetrates the bone very obliquely, so that internally it forms a groove in the sutural face crossing to open into the subscapular groove; the outer precoracoidal face is only moderately convex. The coracoid is massive with its large glenoidal facet facing dorso-posteriorly and also much externally. These three bones have greatly widened sutural faces near the glenoid, but distally from this corner the bones become abruptly thinner along the sutural lines. The clavicula is a medium-sized to large bone; flat with expanded ends and a constricted waist. The cleithrum is a strong element with its upper end expanded and projecting beyond the upper scapular border. The interclavicle is massive to very massive with a greatly widened stem; broadly spatulate anteriorly, and here curving sharply upwards. In some specimens a ridge along the median line on the dorsal surface of the stem limits the movement of the coracoidal plates. A deep groove receives the posterior edge of the lower end of the clavicle. The humerus is always massive, but varies in size from fairly small, short and squat to very large and long (312-575 mm.); an ectepicondylar foramen is always present; the radial condyle is always strong, thick and well modelled and extends far proximally along the ventral face, in some cases nearly reaching the base of the delto-pectoral crest; the processus medialis lies either nearly in the same plane as the caput or in a more distal plane, i.e. it is proximally or not proximally situated; the processus lateralis is always situated far proximally; the proximal expansion is relatively moderately or very great; the epicondyles are greatly expanded; the shaft is either fairly long or very short, fairly slender or very broad; the ventral opening of the entepicondylar foramen is slitlike or broadly oval; except in one case, the delto-pectoral crest is long; the ectepicondylar foramen is always small and circular. The ulna is large and massive (length 320-402 mm.). The radius is large and massive (length 282-318 mm.). The manus is unknown. 266 ANNALS OF THE SOUTH AFRICAN MUSEUM The pelvis has a short pubo-ischiadic plate; the iliac blade is high; the ilio-fibularis ridge is rounded and strong and usually lies vertically; the supra-acetabular buttress is strong with a well-developed notch; the aceta- bulum faces outwards; the anterior iliac process is moderately to strongly everted; the posterior iliac process is fairly short and high; the iliac blade is concave in antero-posterior direction. The femur is large and massive (length 498-605 mm.) with the internal trochanter developed as a strong rounded tubercle, situated well away from the preaxial border; the width over the external trochanter, which sometimes has a notch separating it from the proximal face, is great (234-306 mm.). The tibia is large and massive with strongly expanded ends (length 255-355 mm.). The fibula is large and stout with a deeply concave anterior face (length 235-345 mm.). The pes is incompletely known. Titanosuchidae Family Characters of the Girdles and Limbs. The Titanosuchid pectoral girdle is only adequately known in one genus. The pectoral girdle is large and massive; the height is 860-890 mm., and the length of the coracoidal plate is 560-580 mm., which is thus 62-67% of the height. The scapula is high (680 mm.) and the upper end of the blade is broad to very broad (372-426 mm.); the scapular head of the triceps is attached to a sharp ridge or a prominent mound. The precoracoid is large and massive (340 X 342 mm.). The coracoid is of medium size but massive (250 X 252 mm.). The clavicle is a large bone (length 505 mm.); it is medio-laterally flattened, with expanded dorsal and ventral ends, but the waist not greatly constricted; the ventral spatulate end curves inwards to fit over the outer face of the upturned antero-lateral corner of the interclavicle; the dorsal end has its anterior end greatly thickened and is produced dorsally as a short, strong process, which presumably fits into a groove on the lower end of the cleithrum, which is however not preserved. The interclavicle is a large and massive bone; the length is 570 mm.; width over anterior expansion 455-545 mm.; width over posterior end of stem 215-310 mm. and over the waist of the stem 155-180 mm.; the lateral horns have a thickened postero-lateral edge, anterior to which there is a fairly deep groove to house the ventro-posterior edge of the spatulate end of the clavicle. GIRDLES AND LIMBS OF DEINOCEPHALIA 267 The humerus is long and large (length 480-575 mm.); the radial condyle, although strong and extending far along the ventral face, does not reach the plane of the entepicondylar foramen, and lies well distal of the base of the delto-pectoral crest; the processus medialis always lies far proximally — nearly in the same plane as the caput; the proximal expansion is moderately great (240?-310 mm.); the shaft is fairly long and relatively not very broad; the ventral opening of the entepicondylar foramen slitlike or oval. The ulna has a short massive shaft and a long sigmoid face with a rounded lip. The radius is a long robust bone. In the massive pelvis the upper edge of the ilium does not slope strongly downwards in posterior direction; the anterior process is moderately everted; the ilio-fibularis ridge is massive; the ilium forms much the greater part of the acetabulum. The femur is very large (length 564-605 mm.); the width over the external trochanter is great (275-306 mm.); the preaxial face moderately to deeply concave. The tibia and fibula large and strong. Genus Titanosuchus Owen The pectoral girdle poorly known; the coracoid is massive with the glenoidal facet facing much externally. The humerus is very large (length 530 mm.); the shaft is short and wide (130 mm.); the width over the proximal expansion is 308 mm.; the delto- pectoral crest is fairly short and its extremity is knoblike; the processus medialis is situated well proximally; the capitellum is strong and extends far proximally along the ventral face of the bone; the opening of the entepicondylar foramen ventrally is slitlike; the ectepicondyle is developed as a fairly thin sheet of bone and the foramen is situated well away from the edge of the bone. The epipodial bones and the manus are unknown. The pelvic girdle is unknown. The femur is very long (605 mm.); the width over the external trochanter is 275 mm.; the shaft is fairly narrow (135 mm.); the preaxial face is not very deeply concave, but the caput is preaxially directed. The fibula is large and massive (295 mm.). The tibia and pes are not known. 268 ANNALS OF THE SOUTH AFRICAN MUSEUM Titanosuchus ferox Owen The specific diagnosis is as for the genus (see Seeley’s (28) plates). B.M. 49367, coracoid; 49369, humerus; 49368, femur; and 49367b, fibula; all said to be associated with the type cranial material. Koedoeskop, Beaufort West. Mid Tapinocephalus zone. Coll. T. Bain. Referred specimens in the S.A.M. collection: S.A.M. 739. A fairly good isolated femur, with a length of 570 mm. and shaft diameters of 157 XxX 65 mm. _ Beaufort West District. Tapinocephalus zone. Coll. Oakley. S.A.M. 11491. A good isolated right femur (Fig. 59) with a length of 540 mm.; 294 mm. over the external trochanter, which is separated from the proximal face by a distinct notch; the shaft diameters are 144 X 84mm. Mynhardtskraal, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. Fig. 59. Femur of Titanosuchus ferox. S.A.M. 11491. (x %.) a. Dorsal view. b. Ventral view. c. Anterior view. GIRDLES AND LIMBS OF DEINOCEPHALIA 269 Genus Scapanodon Broom All that is known of the pectoral girdle is a single incomplete scapula which shows that the scapular head of the triceps originated from a sharp ridge on the posterior face of the scapula just above the glenoid. The humerus is fairly long to long (480?-522 mm.); the proximal expansion is not very great (240-264? mm.); the distal expansion fairly great (282-290? mm.); the shaft is fairly short to long and not very broad (diams. 114-118 X I10-12I mm.); the delto-pectoral crest is long with its extremity extending to fairly near the entepicondylar foramen or terminating well proximally of this plane; the caput is oval and fairly massive and lies in nearly the same plane as the processus medialis, which is thus well proximally situated; the twist on the shaft is great (40°); the L.M.L. is strong, forming a pronounced bulge on the dorsal surface of the shaft; the A.D.V.L. is strong and sharp; the entepicondyle is developed as a moderately thick, but not greatly flaring flange of bone, with the foramen ventrally showing a slitlike opening not very far removed from the edge of the bone; the ectepicondyle is a greatly flaring thin sheet of bone pierced vertically by the foramen lying well removed from the edge of the bone. The epipodial bones of the forelimb and the manus are not known. The pelvic girdle is massive and high; the pubo-ischiadic plate is short (80% of the height of the pelvis as restored); the supra-acetabular part of the ilium is high and relatively short (the height is 74% of the antero- posterior length); the anterior iliac process, although not fully preserved, appears to have been fairly short, but fairly high, and it is only moderately everted; the posterior iliac process is short and has its posterior edge directed much upwards and this edge is folded over to form a massive ilio-fibularis ridge, which forms a prominent feature on the outer iliac face; anterior to this ridge the iliac blade is deeply concave where the gluteal muscle is attached; the inner face of the anterior process is irregularly pitted for the reception of a rib lying anterior to the main sacral rib; the iliac portion of the acetabulum is very great, forming over two thirds of the acetabulum. The antero-ventral edge of the pubis is strongly everted with the tuber- culum pubis confluent with the strongly thickened outwardly directed antero- ventral edge, which thins as it proceeds towards the median line. The pubic symphysis must have been short and weak. The pubic foramen is a large oval opening. Only the upper half of the ischium is preserved; it forms only a small part of the acetabulum; its postero-dorsal edge is greatly thickened and it would appear that the ischial symphysis was weak. The femur is massive and long (564 mm.); its proximal expansion is great, with the width over the external trochanter, which has a notch 270 ANNALS OF THE SOUTH AFRICAN MUSEUM separating it from the proximal face, 306 mm.; the preaxial face is fairly strongly concave with the caput moderately thick (134 mm.) and directed moderately preaxially; the shaft is fairly long but very broad (162 mm.); the preaxial condyle is much weaker than the postaxial condyle, which also lies further distally and is very massive (174 mm. thick); the femoro-tibialis ridge is strong, but not strongly bulging on the dorsal surface of the shaft. No tibia, fibula or pes is known. Scapanodon duplessis: Broom Of the girdle and limb-bones only the humerus is known. The humerus is long (520-522 mm.); the delto-pectoral crest with a swollen end terminates well proximally of the entepicondylar foramen. S.A.M. 772 and 773. Two fairly good left humeri (Fig. 60 b and c) said to be associated with the poor type jaws. Seekoeigat, Prince Albert. High Tapinocephalus zone. Coll. du Plessis. Fig. 60. Aaa es ea w A Sw Humeri in ventral view. (x 3.) a. Scapanodon septemfontis. S.A.M. 5001. b. Scapanodon duplessisi. S.A .M. 772. c. Scapanodon duplessisi. S.A.M. 773. —e—B_ GIRDLES AND LIMBS OF DEINOCEPHALIA 271 Scapanodon septemfontis Sp. Nov. I am making the specimen (S.A.M. 5001), which Broom (13) erroneously described as being referable to Tapinocephalus atherstone1, the type of a new species of Scapanodon. Nothing is known of the pectoral girdle. The humerus is large and fairly long (480? mm.); the delto-pectoral crest terminates far distally, but still well away from the ventral opening of the entepicondylar foramen. Fig. 61. OS PES SS . Cis = . 3 ee ee. Oe tes ai woe . - Sac J ¢ (FeO r ‘ UG ‘ OC eo: —t) *o.° 5 e o Ceo mse’ vy eee Pelvis of Scapanodon septemfontis. S.A.M. 5001. (x ¢@.) Lateral view. 272 ANNALS OF THE SOUTH AFRICAN MUSEUM The rest of the forelimb is unknown. The pelvis and femur are as described in the generic diagnosis. Type: S.A.M. 5001. An incomplete humerus (Fig. 60a), a nearly complete pelvis (Fig. 61) and a good femur (Fig. 62). Sewefontein, Prince Albert. Low Tapinocephalus zone. Coll. Haughton. Referred specimens in the S.A.M. collection: S.A.M. 1203. A pubis. Letjiesbos, Beaufort West. Mid Tapinocephalus zone. Coll. Maddison. S.A.M. 11578. Part of a scapula and the major part of a humerus. Aasvoélbos, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. Femur of Scapanodon septemfontis. S.A.M. 5001. (x %.) a. Dorsal view. b. Ventral view. c. Anterior view. GIRDLES AND LIMBS OF DEINOCEPHALIA 273 Se Dn a te . “0 414,°" 7 “ere Precoracoid of Parascapanodon avifontis. S.A.M. 9127. (x #%.) a. Outer view. b. Outline of sutural face for the scapula. c. Outline of sutural face for the coracoid. d. Inner view. e. Lateral view (as projected on to the median plane). 274 ANNALS OF THE SOUTH AFRICAN MUSEUM Genus Parascapanodon Gen. Nov. The generic characters of the pectoral girdle are as described for the family. The humerus is very large and massive; the length is 575 mm. and the ends greatly expanded (proximal 310? mm., distal 312 mm.); the shaft is fairly long but very robust (diams. 144 x 142 mm.); the delto-pectoral crest is long, but terminates well proximal of the plane in which the ent- epicondylar foramen lies; the caput is very massive, but short; the processus medialis lies just a little distally of the plane in which the caput lies; the capitellum is very strong and massive and extends far along the ventral face, but does not reach the plane of the entepicondylar foramen; the ““twist’” on the shaft is large (40°); the L.M.L. is strong, with a massive swelling on the dorsal surface of the shaft; both epicondyles are strongly developed; the ventral opening of the entepicondylar foramen is large and broadly oval, and the ectepicondylar foramen is small and situated well away from the edge of the bone. Right clavicle of Parascapanodon avifontis. S.A.M. 9127. (x %.) a. Outer view. b. Inner view. c. Anterior view. "MOTA JOMOJUY “9 “MOTA : eusA *q “MaIA [esIog “e (‘fF X) L216 “W'S ‘syuofian uopoundvaspang JO Inuls 7 275 GIRDLES AND LIMBS OF DEINOCEPHALIA 4—Annals 276 ANNALS OF THE SOUTH AFRICAN MUSEUM The ulna is a large and massive bone (length 372 mm., width over the coronoid process is 200 mm.); the sigmoid face is long, with its ventral part broadly rounded; the coronoid process is situated far distally and the shaft is massive, broad and short. The radius is a long robust bone (length 294 mm.) with a strong proximo- postaxial flange. No pelvis is known. The femur is very long and massive (length 595 mm.); very broad over the external trochanter (300 mm.); the preaxial face is deeply concave, with the caput much preaxially directed and massive (diams. 215 x 167 mm.); the external trochanter is indistinctly separated by a notch from the proximal face; the shaft is fairly long and broad (breadth 150 mm.); wide over the massive distal facets; the area of origin of the femoro-tibialis forms a strong bulging ridge. The tibia is large and massive (length 330-355 mm.); the cnemial eminence very massive, continued distally as a strong ridge, with a deep groove lying postaxially. The fibula is large and stout (length 330-345 mm.). Parascapanodon avtfontis Sp. Nov. The specific characters are as for the genus. Fig. 66. Left fibula of Parascapanodon avifontis. S.A.M. 9127. (x ¢.) a. Dorsal view. b. Ventral view. c. Anterior view. GIRDLES AND LIMBS OF DEINOCEPHALIA 277 ee Pm es Se “~ 5 “ae Us te be eh ee le Scapulo-coracoid of Parascapanodon avifontis. S.A.M. 9106. (x $.) Lateral view. 278 ANNALS OF THE SOUTH AFRICAN MUSEUM Type: S.A.M. 9127. A very good precoracoid (Fig. 63), a good clavicle (Fig. 64), a good femur (Fig. 65), a well-preserved fibula (Fig. 66), associated with parts of a large skull. Voélfontein, Prince Albert. Low Tapinocephalus zone. Coll. Boonstra. Paratypes: Although lacking cranial parts, and direct comparison with the type skull parts thus impossible, I have used the following specimens in drawing up the generic description of Parascapanodon: S.A.M. 9106. Parts of the scapula, coracoid and precoracoid (Pig. - 67). Veldmansrivier, Prince Albert. Low T apinocephalus zone. Coll. Boonstra. S.A.M. 11488. Parts of the precoracoid, coracoid and _interclavicle (Fig. 68) and a tibia (Fig. 69 a-c) and fibula (Fig. 69 d-f). Voélfontein, Prince Albert. Low 7 apinocephalus zone. Coll. Boonstra. Fig. 68. Interclavicle, precoracoid and coracoid of Parascapanodon avifontis in ventral view. S.A.M. 11488. (x 4.) GIRDLES AND LIMBS OF DEINOCEPHALIA 279 Fig. 69. tT ee _ Parascapanodon avifontis. S.A.M. 11488. (x §.) a. Right tibia in dorsal view. b. Right tibia in ventral view. c. Right tibia in posterior view. d. Right fibula in dorsal view. e. Right fibula in ventral view. f. Right fibula in anterior view. These two specimens are included in this species on the ground of the great similarity of their precoracoids to that of the type. 280 ANNALS OF THE SOUTH AFRICAN MUSEUM On the similarity of its fibula to that of the type I have also included: S.A.M. 9163. A good fibula (Fig. 70 a-c) and a fair ulna (Fig. 70 d-e). Wakkerstroom, Prince Albert. Low T apinocephalus zone. Coll. Boonstra. Fig. 70. . Loc cosmos ‘ oot bras a Pe oye F 8 ee a s, tee 26s, Oleh = Parascapanodon avifontis. S.A.M. 9163. (x #.) a. Left fibula in dorsal view. b. Left fibula in ventral view. c. Left fibula in anterior view. d. Left ulna in dorsal view. e. Left ulna in anterior view. GIRDLES AND LIMBS OF DEINOCEPHALIA 281 On the similarity of its tibia to that of S.A.M. 11488 I have also included : S.A.M. 11299. A good tibia (Fig. 71 a-c) and a good radius (Fig. 71 d-f). Boesmansrivier, Beaufort West. Mid Tapinocephalus zone. Coll. Boonstra. Parascapanodon avifontis. S.A.M. 11299. (xX @-) a. Right tibia in dorsal view. b. Right tibia in ventral view. c. Right tibia in posterior view. d. Right radius in dorsal view. e. Right radius in ventral view. f. Right radius in posterior view. ANNALS OF THE SOUTH AFRICAN MUSEUM 282 ‘MOTA [PUITXOIG *9 (%#X) ‘rggrr ‘wy's ‘MOIA TesIOg ‘q ‘syuofian uopoundvosvavg JO snzreuInyzT ‘ BPE) oy ay Vide he GIRDLES AND LIMBS OF DEINOCEPHALIA Scapulo-coracoid of Parascapanodon. S.A.M. goto. Lateral view. 283 284 ANNALS OF THE SOUTH AFRICAN MUSEUM I am including the following isolated humerus as a paratype: S.A.M. 11881. WH o q q -e | Outer view of right d. Inner view of right Jonkeria haughtoni. a. Outer cleithrum. cleithrum. c. clavicula. (xX é-) clavicula. 291 "MOIA [VUNXOIG “Pp ‘MOTA IOLIoyUy “9 “MOTA esioqd ‘q ‘“MOIA [eIJUIA “e (GIG) =EKeve IKE Ves “uojysnoy viuayuof JO SNIsUIN}{ GIRDLES AND LIMBS OF DEINOCEPHALIA 292 ANNALS OF THE SOUTH AFRICAN MUSEUM LAPS Er +f ~ "eae tee: Ulnae of Jonkeria haughtoni. (x %.) a and a’. Dorsal and anterior views of S.A.M. 4343. b and b’. Dorsal and anterior views of S.A.M. 11464. c and c’. Dorsal and anterior views of S.A.M. go02. GIRDLES AND LIMBS OF DEINOCEPHALIA 293 eu, os whats © < see F . * Above: Jonkeria haughtom. S.A.M. 4343. (x#-) a. lium in dorsal view. b. Ilium in posterior view. c. Ilium in lateral view. d. Ischium in lateral view. Left: Right fibula of Jonkena haughion. S.A.M. 4343. (x #4 a. Dorsal view. b. Ventral view. c. Anterior view. 2904 ANNALS OF THE SOUTH AFRICAN MUSEUM Referred specimens in the S.A.M. collection: S.A.M. 4342. An isolated scapula (Fig. 81). lLeeurivier, Beaufort West. Low Tapinocephalus zone. Coll. Haughton. ( f ‘ i) c ( ae ( ~ i] ~ = =~ ese -—- - ee Scapula of Jonkeria haughtoni. S.A.M. 4342. (x #@.) a. Lateral view. b. Posterior view. GIRDLES AND LIMBS OF DEINOCEPHALIA 295 S.A.M. gooz. An isolated scapula (Fig. 82) and a distorted ulna (Fig. 78 c, c’). In outline the posterior border of the scapula has a distinctive step also shown in S.A.M. 4342, and probably also in the type where this edge is not preserved. Klein-Koedoeskop, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. Fig. 82. ye ott oe : b Ue para le» ee ete eit’. Sari ar recat’ yas ee me eed, e*. ore eye 8 8 8 y- . = - Men e Coe Scapula of Jonkeria haughtoni. S.A.M. 9002. (x #@.) a. Lateral view. b. Posterior view. 206 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 83. Left: Scapula of Jonkeria haughtoni. S.A.M. 19207. (x ¢.) Lateral view. Below : Right radius of /Jonkena haughtoni. S.A.M. 9145. (x @.) a. Dorsal view. b. Ventral view. c. Posterior view. GIRDLES AND LIMBS OF DEINOCEPHALIA 297 S.A.M. 11297. An isolated scapula (Fig. 83). Boesmansrivier, Beaufort West. Mid Tapinocephalus zone. Coll. Boonstra. S.A.M. goog. An isolated precoracoid. This medium-sized bone, composed of a thin sheet except at the apex, where the sutural faces for the coracoid and scapula are fairly strong, is very similar to that of J. truculenta but larger, and is probably that of J. haughtoni. Klein-Koedoeskop, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. S.A.M. 9145. A good radius (Fig. 84) and an incomplete pubis probably belong to this species. Seekoeivlei, Beaufort West. Low Tapino- cephalus zone. Coll. Boonstra. S.A.M. 9147. An isolated ulna with a damaged proximal end agrees sufficiently well with that of the type to be included in this species. Seekoeivlei, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. S.A.M. 11464. A very well preserved ulna (Fig. 78 b, b’); although somewhat shorter than that of the type it can safely be included in this species. Koedoeskop, Beaufort West. Mid Tapinocephalus zone. Coll. Boonstra and Avenant. Jonkenia ingens (Broom) In the specimen described by Broom (14) as Jonkeria pugnax but by me (1) considered a synonym of Jonwkena ingens Broom mentions the scapulae but without figuring or describing them; and of the humerus he states that it resembles that of Jenkeria crassus very closely. In this collection there are four humeri, which in point of size can best be tentatively referred to this large species of Jonkeria, rather than to any other of the Titanosuchids. The humerus in these specimens is large, massive, but not very long (length 504-510 mm.); both ends are greatly expanded (proximal 330-336 mm., and distal 3122-324 mm.); the shaft is short and broad (diams. 130-156 x Ir0 mm.); the delto-pectoral crest is long and powerful, but terminates well proximal of the plane in which the entepicondylar foramen lies; the caput is large and widely oval; the processus medialis lies very far distally of the plane in which the caput lies; the capitellum is strong, thick and very well modelled and it extends far proximally along the ventral face, but does not reach the plane of the entepicondylar foramen; the L.M.L. is strong; the A.D.V.L. is not very sharp; the entepicondyle is strongly developed as a thick flange of bone, with the ventral opening of the entepicondylar foramen fairly large and oval, lying fairly near the edge of the bone well removed from the base of the delto-pectoral crest; the ectepicondyle is a flaring sheet 298 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 85. ast . ee ‘ oe Dna ex co O10 COPEL Cis Sy Cc. Humeri of Jonkeria ?ingens. CX es) jal.) SHACME iazia3t Ventral view. b. S.A.M. 738. Ventral view. c. S.A.M. 738. Dorsal view. d. S.A.M. 738. Proximal view. GIRDLES AND LIMBS OF DEINOCEPHALIA 299 of bone pierced vertically by the small foramen lying well away from the edge of the bone. Referred specimens: A.M.N.H. 5608. Much of two scapulae and a humerus, associated with a skull. Kookfontein, Prince Albert. Mid Tapinocephalus zone. Coll. van Wyk. S.A.M. 738. A good isolated humerus (Fig. 85 b-d). Gamka River. Low? Tapinocephalus zone. Coll. Cloete. S.A.M. 3433. A good isolated humerus (Fig. 85a). Janwillemsfontein, Prince Albert. Low Tapinocephalus zone. Coll. Haughton. S.A.M. g006. An isolated proximal half of a humerus. Klein- Koedoeskop, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. S.A.M. 11994. An incomplete isolated humerus with a good distal end. Welgemoed, Prince Albert. Low Tapinocephalus zone. Coll. Haughton. S.A.M. 9348. An incomplete and distorted humerus and an ilium lacking both the anterior and posterior processes (Fig. 86). The ilium is larger and more strongly built than those of the other species of Jonkeria and may provisionally be considered to represent that of Jonkeria ingens. Mierfontein, Beaufort West. Mid Tapinocephalus zone. Coll. Boonstra. Fig. 86. ee ae, aS oe Ilium of Jonkeria ?ingens. S.A.M. 9348. (x %.) Lateral view. ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 87. 300 Re c . Ca eer Yt rh 2. feuet ve “Neg see (x 8.) a. Pelvis in lateral view. Jonkeria koupensis. S.A.M. 9004. : b. Ilium in dorsal view. c. Pelvis in anterior view. GIRDLES AND LIMBS OF DEINOCEPHALIA 305 Jonkeria koupensis Sp. Nov. This new species appears to be very closely related to J. haughtoni, but the differences in the pelvis, as will be evident from the accompanying characterisation, warrant treatment as a distinct form. In the pelvis the pubo-ischiadic plate is probably short (83% of the height as reconstructed); the supra-acetabular part of the ilium is high (264 mm.) and the antero-posterior length of the iliac blade is very short (282 mm.), so that the height is 93% of the length; the anterior iliac process is fairly short, but appears shorter than it really is because of its strong eversion antero-laterally, and is fairly high; the posterior process is short and fairly low, with its postero-ventral edge moderately strongly folded over to form a fairly strong ilio-fibularis ridge on the outer face; this ridge is directed obliquely upwards, with its upper end not strongly bulbously thickened; a slight groove on the inner face of the everted anterior iliac process indicates the attachment of a rib anterior to the main sacral rib; antero-posteriorly the outer face of the iliac blade is deeply concave. The antero-ventral edge of the pubis is strongly everted, with the tuberculum pubis confluent with the thickened part of the antero-ventral edge, which stretches to the median line where the pubes meet, but do not form a real symphysis. The ischium is not preserved but probably is as reconstructed. The ulna has the distal two-thirds preserved; it is a massive bone with the dorsal lip of the sigmoid face developed into a massive swelling in its preaxial part. Fig 88. Type: S.A.M. 9004. A good pelvis, ; lacking only the ischia (Fig. 87) and the distal two-thirds of the ulna. Klein-Koedoeskop, Beaufort West. Low Tapino- cephalus zone. Coll. Boonstra. Referred specimen: S.A.M. 11983. An isolated ilium (Fig. 88). | Abrahamskraal, Prince Albert. Low Tapino- cephalus zone. Coll. Haughton. Illium of Jonkeria koupensis. S.A.M. 11983. (x %.) Lateral view. ANNALS OF THE SOUTH AFRICAN MUSEUM 302 "MaIA [esIogd “q ‘MOTA [PUIITXOIG "Pp ‘MOIA IOLIOJUY ‘9 “MOIA [eI]UDA “Be (°2 X) ‘6b16 “WV'S ‘pasvg visayuof Jo snxsWINn;T GIRDLES AND LIMBS OF DEINOCEPHALIA 393 Jonkena parva Sp. Nov. Although only a single isolated humerus is known, this bone is so distinctive that I do not hesitate to make it the type of a new species of Jonkena. This humerus is the smallest Jonkeria humerus as yet known; its length is only 312 mm., but it is a very massive element with both the proximal (222 mm.) and the distal (252 mm.) ends very greatly expanded; the shaft is very short and the bone is greatly constricted in the waist; the diameters of the shaft are 84 x 78 mm.; the delto-pectoral crest is fairly short and it terminates very far proximal of the plane in which the entepicondylar foramen lies; it has a very massive ventral edge and it terminates as a very thick knob. The caput is weak and its face is straplike, but it forms the most proximal part of the bone; the processus lateralis lies more distal than the caput; the processus medialis, as in all the Jonkerias, lies well distally of the caput; the capitellum is very massive indeed and it extends very far proximally along the ventral face, with its proximal border lying in a plane proximal to that in which the entepicondylar foramen lies; posteriorly of the well-modelled capitellum there is a deep groove in which the coronoid process moved when the ulna was flexed; the twist on the shaft is great (30°); the L.M.L. is distinct, with a large moundlike muscle scar on the dorsal surface of the shaft; the A.D.V.L. is very strong and forms a prominent ridge; the entepicondyle is strongly developed to form a greatly outflaring thick sheet of bone; the ventral opening of the entepicondylar foramen is oval and it lies well postaxially, near the edge of the bone; the ectepicondyle is developed as a greatly flaring thin sheet of bone penetrated in its thinner part, near the edge, by a small round ectepicondylar foramen. Type: S.A.M. 9149. An isolated right humerus very well preserved (Fig. 89). Saairivier, Prince Albert. Low Tafinocephalus zone. Coll. Boonstra. Jonkeria rossouwi Sp. Nov. The specimen on which this new species is founded was found by Broom (14) to be different from the then known species of Jonkeria, but Broom, with a restraint unusual for him, did not name it. I am naming it Jonkeria rossouw?t for Mr. P. J. Rossouw of the Geological Survey of the Union in recognition of the importance of his stratigraphical work on the Tapino- cephalus zone. In this species the pectoral girdle is fairly large and fairly massive. The scapula is fairly low (height 552 mm.) and the upper part of the blade is greatly expanded (width 324 mm.); the tricipital bulge is very prominent; the supra-glenoidal edge forms a strong raised rim; the internal opening of 304 ANNALS OF THE SOUTH AFRICAN MUSEUM the supracoracoid foramen opens into the deep subscapular groove; the glenoidal facet of the scapula faces ventro-posteriorly but not externally. The precoracoid is long but low (somewhat affected by crushing); the foramen pierces the bone very obliquely. The coracoid is small but massive, with a large glenoidal facet facing well externally. The interclavicle is massive but short (480? mm.) with the stem wide posteriorly and with a narrowed waist anteriorly; the anterior spatulate end curves upwards very sharply and has a deep groove on its outer antero- Fig. 90. Jonkeria vossouwi. S.A.M. 5014. (> %.) a. Scapulo-coracoid in lateral view b. Scapula in posterior view. GIRDLES AND LIMBS OF DEINOCEPHALIA 305 lateral face for the reception of the postero-ventral edge of the clavicle; on the dorsal surface of the stem there is a strong medial ridge against which the precoracoids abut. The clavicles and cleithrum are not known. The humerus is fairly short (378 mm. in length), but massive with greatly expanded proximal (330 mm.) and distal (276? mm.) ends; the shaft is very short, thick and broad (132 x 84 mm.); the delto-pectoral crest is very long and nearly reaches the plane in which the entepicondylar foramen lies; the caput is broadly oval; the processus lateralis lies well proximally, in the same plane as the caput, whereas the processus medialis lies well distally; the capitellum is fairly massive and extends well along the ventral face and nearly reaches the plane of the entepicondylar foramen; the ‘‘twist’’ on the shaft is fairly small (15°); the L.M.L. is fairly strong with muscle scars on the dorsal surface of the shaft; the A.D.V.L. is well developed; the entepicondyle is strong with the ventral opening of the foramen large and nearly round; the ectepicondyle forms a thick flange pierced by the small foramen situated well away from the edge of the bone. (The dorsal surface of the ectepicondyle shows a pathological lesion in the form of an irregular deep excavation and this is accompanied by a concomitant outgrowth of the olecranon of the ulna. In another humerus, that of Phocosaurus S.A.M. 11300, the disto-ventral face of the capitellum is deeply eroded with subsequent healing and the formation of a new but concave instead of a convex articulatory facet. (Osteitis fibrosa? Osteomalachia ? )) The ulna has suffered pathological deformation so that the olecranon is proximally prolonged as a thick outgrowth. The normal ulna would appear to have had a more slender shaft and a weaker coronoid process than the other known species of Jonkenia. The radius is only slightly pathologically deformed. Its length is 312 mm. and the flange on the proximo-postaxial corner is weak. The manus is not known. In the pelvis the supra-acetabular part of the ilium is high (288 mm.) and relatively short (336 mm.) so that the height is 87% of the length; the anterior iliac process is relatively short, but fairly high and strongly everted; the posterior process is short and fairly low, with its postero-ventral edge folded over strongly to form a strong vertical ridge, which is dorsally strongly bulbous, and it projects strongly laterally; on the inner face of the anterior iliac process no distinct facet is preserved for the attachment of a rib lying anterior to the main sacral rib. No pubis or ischium is preserved. The femur is fairly long (504 mm.); fairly broad over the external trochanter (264 mm.), which is not separated by a notch from the proximal 306 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 91. oa: =a: e] ry = <~ ~ . “oe ae. aan we ° aly wit) ¢ om! ee Interclaviculae. (x %.) a. Jonkeria rossouwi. S.A.M. 5014. Ventral view. b. Jonkeria rossouwi. S.A.M. 5014. Lateral view. c. Jonkeria sp. indet. S.A.M. 9124. Ventral view. d. Jonkeria sp. indet. S.A.M. 9124. Lateral view. wr, GIRDLES AND LIMBS OF DEINOCEPHALIA jesioq “q "MOIA [VWIXOIG “p “MOIA [eIjUeA “eB (°2 X) ‘MOIA IOWSJUY “9 “AMOIA ‘troS “JuTW'S ‘“tanossos pisayuof JO SnisUIN}{ 5—Annals 308 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 93: Left radius of Jonkeria rossouwi. S.A.M. 5014. (x 2.) a. Dorsal view. b. Ventral view. c. Posterior view. Right ilium of Jonkeria rossouwi. S.A.M. 5014. (x #.) a. Dorsal view. b. Posterior view. c. Lateral view. d. Anterior view. GIRDLES AND LIMBS OF DEINOCEPHALIA 309 face; the caput is fairly thick (114 mm.) and is directed much preaxially; the shaft is broad but flat (diams. 156 x 84 mm.); the femoro-tibialis ridge is fairly strong; the distal facets of the femur are small and directed much distally, especially the ectepicondyle which lies far distally. (These features of the distal end of the femur, differing as they do from the other Jonkerias, appear to be due to some measure of pathological deformation.) The tibia is fairly robust (length 300 mm.); the proximal face is inclined much postaxially to correspond with the distally situated postaxial facet of the femur. The fibula is fairly slender and long (330 mm.). The pes is not known. Type: S.A.M. 5014. The left scapula and incomplete precoracoid and coracoid (Fig. 90), an imperfect interclavicle (Fig. 91 a, b), a diseased left humerus (Fig. 92), a deformed left ulna and a left radius (Fig. 93), a right ilium (Fig. 94), a left femur (Fig. 95), a left tibia (Fig. 96) and a left fibula (Fig. 97). Abrahamskraal, Prince Albert. Low Tapinocephalus zone. Coll. van der Byl. Fig. 95. Left femur of Jonkeria rvossouwi. S.A.M. 5014. (X é-) b. Ventral view. c. Anterior view. | 6—Annals 310 ANNALS OF THE SOUTH AFRICAN MUSEUM Referred specimens: S.A.M. 11979. An incomplete interclavicle (Fig. 98 a, b) and the proximal ends of an ulna and fibula. Kruisvlei, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. S.A.M. 11982. A fairly good interclavicle (Fig. 98 c, d). Abrahamskraal, Prince Albert. Low Tapinocephalus zone. Coll. Haughton. Fig. 96. Left tibia of Jonkeria rossouwi. S.A.M. 5014. (xX view. b. Ventral view. c. Posterior view OI Re eee] je) o (2) eS (op) job) — Left fibula of Jonkeria rossouwi. S.A.M. 5014. (x @.) a. Dorsal view. b. Ventral view. c. Anterior view. art GIRDLES AND LIMBS OF DEINOCEPHALIA ‘SG 7. Ventral Ventral view. c. S.A.M. 11982. (SC at) ao Sra rT arg7 0: 11982 Lateral view. 11979. Lateral view. d. S.A.M. Interclaviculae of Jonkeria rossouwt. bz SAM. view. 312 ANNALS OF THE SOUTH AFRICAN MUSEUM Jonkena Sp. Indet. S.A.M. 9124. A good interclavicle (Fig. 91, c, d). Voélfontein, Prince Albert. Low Tapinocephalus zone. Coll. Boonstra. This interclavicle is larger and stronger than those of the oe species of Jonkeria and may thus possibly be J. ingens. S.A.M. 11984. An imperfect isolated ilium (Fig. 99) of an unknown locality. It would appear to lie fairly close to the ilium of J. haughtont. Right ilium of Jonkeria sp. indet. S.A.M. 11984. (x %.) Lateral view. S.A.M. 11886. A nearly complete interclavicle (Fig. Ioo c, d). Vindraersfontein, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. This small bone with a narrow waist and the strongly expanded posterior end of the stem is apparently of a small species of Jonkeria — possibly of J. parva. Titanosuchia Genera Indet. 5.A.M. 11571. A large isolated interclavicle with a strongly upturned anterior end and a strong median keel on the dorsal face of the stem (Fig. 100 a, b). It has the long stem of the Tapinocephalians but the width is that usually seen in the Titanosuchians. Probably a large Titanosuchid. Klein-Koedoeskop, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. GIRDLES AND LIMBS OF DEINOCEPHALIA S53 : Re ice ete: Sites eH, SoS eee RIOR ROC = ‘ at Mee: ae aS . a) ° Di Oe! > . « é OS eed “eae a Sci ° e ‘8; S- eee © qs . 4 alte A a . a . i ike cot a ‘ i ‘is v . -« . t . . . f= ? sy fe eee ‘ ee helt ree Che <3 = 3 = . ¢ Cy ena o ¥ a” Sgt . . . ° © . ~ Fi ' L = ‘ . ea u £4 S > wo 4% Interclaviculae. (x %.) a. Titanosuchian gen. indet. S.A.M. 11571. Dorsal view. b. cf. parva. Titanosuchian gen. indet. S.A.M. 11571. Lateral view. c. Jonkeria S.A.M. 11886 Ventral view. d. Jonkeria cf. parva. S.A.M. 11886. Lateral view. 314 ANNALS OF THE SOUTH AFRICAN MUSEUM S.A.M. 11986. An isolated ilium (Fig. roz). Letjiesbos, Beaufort West. Mid Tapinocephalus zone. Coll. Boonstra. This ilium has on the outer face of the posterior iliac process neither the horizontally placed ilio-fibularis ridge, usually seen in the Tapinocephalia, nor the obliquely placed ridge, usual in the Titanosuchia. The area of origin of the ilio-fibularis is rather a thickening of the postero-ventral edge without any eversion or folding over. In outline this illum agrees more with those of the Titanosuchia. Ilium of a ? Titanosuchian. S.A.M. 11986. (x €.) \ a. Lateral view. b. Dorsal view. Anteosauria Infra-ordinal Characters of the Girdles and Limbs. The pectoral girdle is light; the scapular head of the triceps originates from a prominent tubercle; the outer face of the precoracoid is strongly convex and the glenoidal facet on the coracoid is bipartite. GIRDLES AND LIMBS OF DEINOCEPHALIA B15 The humerus is fairly light; without an ectepicondylar foramen; the radial condyle is well modelled, but situated well distally; both the processus lateralis and medialis lie far proximally; the delto-pectoral crest is very short and is directed much postaxially. In the manus the carpal formula is 3, I, 5, and the phalangeal formula 3) 3, 4, 42, 2. The iliac blade is low and the acetabulum faces much ventrally; the ilio-fibularis ridge lies horizontally; no supra-acetabular buttress is developed. The femur is light, long and slender with the internal trochanter developed as a ridge; the width over the external trochanter is very small. In the pes the tarsal formula is 2, 1, 5 and the phalangeal formula ora 4!» 2. Anteosauridae Family Characters of the Girdles and Limbs. The Anteosaurid pectoral girdle is only imperfectly preserved in two specimens; but, even so, it is evident that the Anteosaurid girdle differs greatly from that of all the other Deinocephalia. The girdle is of moderate size and is lightly built. No complete scapula is known but it would appear that it was of moderate width and height (455-475 mm. as reconstructed); instead of a tricipital mound or ridge, as in all the other Deinocephalia, there is developed (for the origin of the scapular head of the triceps) a prominent tubercle on the posterior edge of the scapula 25-35 mm. above the glenoidal edge; the internal opening of the foramen supracoracoideus opens into the fairly deep subscapular groove; the glenoidal facet of the scapula is large and faces ventro-posteriorly, but also much externally, and the outer and posterior margins are sharp, prominent and well moulded. The coracoidal plate is fairly long antero-posteriorly. The precoracoid is relatively large; its dorso-posterior apex, just posterior to the outer opening of the foramen supracoracoideus, forms a fairly long margin to the glenoid; here the outer face of the precoracoid forms a decided depression or recess to house the processus lateralis of the humerus, when this bone is in its forward position; the foramen supracoracoideus penetrates the bone obliquely so that it internally forms a groove in the sutural face and thus opens into the subscapular groove; the outer precoracoidal face is strongly convex, so that its lower edge is directed much medially. The coracoid is small and light, but greatly thickened where it houses the lower half of the glenoid; this facet is bipartite — the inner and posterior moiety receives the head of the humerus in its posteriorly disposed position, and the outer and anterior moiety receives the humerus when it is disposed 316 ANNALS OF THE SOUTH AFRICAN MUSEUM anteriorly; the inner part of the facet is directed dorso-posteriorly and slightly externally, whereas the outer part is directed much externally. Nothing is known of the cleithrum, clavicula and interclavicula. __ The humerus, known only in three specimens, is fairly lightly built and fairly small to medium (length 300?-375 mm.); the proximal expansion is fairly small (width 162-168 mm.); the distal expansion is somewhat greater (width 200-205 mm.); the shaft is fairly long and slender (diams. 76-84 xX 48-65 mm.); the delto-pectoral crest is very short and terminates a consider- able distance proximal to the ventral opening of the entepicondylar foramen, and is directed much postaxially; the caput is narrowly oval; the processus medialis lies in the same plane as the caput, whereas the processus lateralis lies a little further proximally; the radial condyle is situated distally, but extends a little along the ventral face and also curves on to the dorsal face; the ‘“‘twist’’ on the shaft is large (40°); the L.M.L. is fairly definite and the A.D.V.L. is sharp and prominent; the entepicondyle is fairly greatly expanded, with the foramen entering dorso-postaxially and leaving as a ventral slit; the ectepicondyle is well expanded as a thin plate of bone housing on its ventral face a groove for the passage of the radial vessels. The only radius known is a light, slender featureless bone and the only known ulna (length 260 mm.) is crushed flat. The manus in the only specimen known has the carpal formula 3, I, 5 and the phalangeal formula 3, 3, 4, 4?, 2. The pelvis is only known from one imperfect ium. This, however, shows that the Anteosaurids had an ilium of a type radically different from any known in the South African Deinocephalia. It is without the strong supra-acetabular buttress so typical of the Deinocephalia; instead, the acetabulum is dorsally bounded by a continuous curved ridgelike margin without a supra-acetabular notch; the thickly rounded posterior edge of the fairly short posterior process, from which the ilio-fibularis originates, lies horizontally as in the Tapinocephalia; the iliac blade was presumably short and low. In Fig. ro4 an orthoprojection, taken by pantograph, vertically on to the iliac part of the acetabulum (as is the case in all the figures showing the lateral aspect of the pelvis, i.e. a projection on to the median plane) is shown. But I believe that in the naturally articulated skeleton the iliac portion of the acetabulum, in this case, would be directed much more ventrally so that the disposition of the femur in the Anteosaurids would be radically different from the condition in all other Deinocephalia. The femur of the Anteosaurids looks very different from that of the other Deinocephalians; although not very long (210-420 mm.) it appears very long because of its general slenderness and long shaft; the width over GIRDLES AND LIMBS OF DEINOCEPHALIA a7 the external trochanter is small and varies from 47 to 120 mm.; the shaft is narrow and has diameters ranging from 28 x 18 to 75 X 45 mm.; the external trochanter is situated far proximally and nearly in the same plane as the caput; the caput is short pre-postaxially and flows directly into the external trochanter; with the preaxial border deeply concave and the postaxial border convex the femur is a curved bone with the caput well preaxially and somewhat dorsally directed; the internal trochanter forms a low ridgelike tubercle situated well away from the preaxial face, with a resulting reduced inter-trochanteric fossa; the femoral condyles face much distally with the external one further distally than the internal one; the popliteal fossa is elongated; the epicondyles, in one specimen, form a thin sheetlike expansion, both pre- and postaxially; the femoro-tibialis ridge is weak and the area for the insertion of the ilio-femoralis is narrow; there is a strong ridge for the insertion of the pubo-ischio-femoralis internus. The tibia is unknown. The fibula is a lightly built long and slender bone with its proximal end strongly expanded and with the facets for the femur nearly terminal. Whereas the proximal end is flattened, the distal facet is broadly oval in outline; the preaxial face is fairly straight, whereas in the other Deinocephalia this is deeply concave. The pes has the tarsal formula 2, I, 5 and the phalangeal formula uneeriaim but probably 2?, 3°, 3°, 42, 2. Anteosauridae The Family Characters are as for the infra-order. Genus Anteosaurus Watson In this genus the girdles and limbs are very inadequately known. The generic characters that can be determined from an incomplete scapula, an incomplete and distorted humerus, three coracoids, an incomplete ilium, two good and two incomplete femora are as follows: Tricipital tubercle elongated, 35 mm. above the edge of the glenoid; humerus fairly small (length 300° mm.); the shaft is slender (diams. 60 x 48 mm.); the ilium is as described for the infra-order; the femur is slender (length 402?-420 mm.); width across the external trochanter 108-120 mm.; the epicondyles with moderate or pronounced sheetlike expansions. Anteosaurus acutirostris (Boonstra). Femur 420 mm. in length, with flaring epicondyles. S.A.M. 11977A. Two good femora (Fig. 102 a-c), associated with a good skull (S.A.M. 9329). Kruisvlei, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. 318 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 102. Femora. (x %.-) a. Anteosaurus acutirostris. S.A.M. 11977A. Dorsal view. b. Anteosaurus acutivositns. S.A.M. 11977A. Ventral view. c. Anteosaurus acutivostris. S.A.M. 11977A. Anterior view. d. Anteosaurus sp. SaA.Miin 2753 A. Ventral view. e. Paranteosaurus primus. S.A.M. 11485. Ventral view. f. Anteo- saurus sp. S.A.M. 5614. Ventral view. GIRDLES AND LIMBS OF DEINOCEPHALIA 319 ‘Anteosaurus lotz: (Broili and Schroder) Alte Akademie. No. ?. Parts of peaalinats pubis and femur, associated with some skull pieces. Brakleegte, Beaufort West. High? Tapino- cephalus zone. Coll. Schroder. Little can be said about these fragments, but if the pubis is correctly determined and is as figured by Broili and Schroder, then this element is radically different from the pubes of other Deinocephalia. Anteosaurus spp. _§.A.M. 2752. A partial distorted humerus and a coracoid, associated with a skull. Viviers Siding, Beaufort West. Mid Tapinocephalus zone. Coll. Haughton and Whaits. Fig. 103. Scapulo-coracoids. (xx %.) a. Anteosaurus sp. S.A.M. 5614. Lateral view. b. Eccasaurus priscus. S.A.M. 11597. Lateral view. 320 ANNALS OF THE SOUTH AFRICAN MUSEUM S.A.M. 2753A. Proximal and distal ends of a femur (Fig. 102d), associated with a skull fragment. Vjiviers Siding, Beaufort West. Mid Tapinocephalus zone. Coll. 2 Haughton and Whaits. Fig. 104. S.A.M. 5614. Part of a scapula (Fig. 103a), an incomplete ilium (Fig. 104) wn ores and the proximal end of a femur (Fig. ait, i iba ecg 1o2f), associated with two dentaries. ates eh alta BSc S30 a ri : bitte “G te seis Abrahamskraal, Prince Albert. Low Tapinocephalus zone. Coll. Haughton. S.A.M. 7396. An isolated coracoid. Abra- hamskraal, Prince Albert. Low Tapinocephalus zone. Coll. van der Byl. S.A.M. 11887. An _ isolated coracoid. Vindraersfontein, Beaufort West. Low Right ilium of Anteosaurus sp. Tapinocephalus zone. Coll. Boonstra. SN Genus Paranteosaurus Boonstra Paranteosaurus primus Boonstra S.A.M. 11485. The proximal end of a femur (Fig. 102e), associated with a skull. Mynhardtskraal, Beaufort West. Low Tapinocephalus zone. Coll. Boonstra. This fragment shows no features by which it can be distinguished from the femur of Anteosaurus. Genus Micranteosaurus Boonstra The femur is slender and curved (length 210 mm.); the width over the external trochanter is 47 mm.; the shaft is long and narrow (diams. 28 x 18 mm.); a ‘‘twist’’ on the shaft causes the caput to be directed much dorsally; the ectepicondyle (and probably also the entepicondyle) is without any sheet- like expansion. The fibula is very long (138 mm.) and slender. The pes has the tarsal formula 2, I, 5 and the phalangeal formula 22,38) Heats 2 The radius (105 mm.) is much shorter than the fibula. The manus has the carpal formula 3, I, 5 and the phalangeal formula 3, 3) 4, 4?, 2. GIRDLES AND LIMBS OF DEINOCEPHALIA 320 Micranteosaurus parvus Boonstra As I (4) have only recently described this form, the above generic diagnosis of the limbs is extracted from that account, which consult for the figures. Type: S.A.M. 4323. A coracoid, radius, manus, femur, fibula and pes, associated with a snout. Commonage of Merweville, Beaufort West. Low Tapinocephalus zone. Coll. Haughton. Genus Eccasaurus Broom Notwithstanding its very distinctive humerus (which constitutes the type specimen) Broom (9) included this genus in the Tapinocephalia and later even referred a typical Tapinocephalian tooth to this genus. Although Broom in his original description only mentions the humerus there is in the South African Museum collection, associated with the type humerus, some skull pieces, two imperfect and weathered femora, a flattened radius?, some vertebrae, the distal end of a fibula, some other fragments and a long slender simple pointed tooth, which is probably an incisor. The generic characterisation below is based on the type supplemented by features determined from a second specimen (S.A.M. 11597). In the scapula the tricipital tubercle is oval in outline and is situated 25 mm. above the edge of the glenoid; the other features of the scapulo- coracoid are as described above for the family. The humerus is 348-375 mm. long and the width of the shaft is 76-84 mm.; other features as for the family. The only known ulna is crushed dorso-ventrally; the length is 260 mm. The pelvis is unknown. The femur is 355 mm. in length and fairly slender; the width over the external trochanter (120 mm.) is larger than in Anteosaurus; the epicondyles are not laterally expanded as thin flanges. There is thus no doubt that Eccasaurus with its long pointed incisor, its humerus with the delto-pectoral crest very short, its slender femur, where the external trochanter is situated so far proximally and so little laterally, and in the referred specimen (referred because of the great similarity in the humeri) the distinctive tricipital tubercle on the scapula, cannot possibly be a Tapinocephalian but is in fact an Anteosaurian. The tooth, which Broom has referred to Eccasaurus, does not belong to this genus but is typically Tapinocephalian. Eccasaurus priscus Broom The specific characters are as for the genus. 322 ANNALS OF THE SOUTH AFRICAN MUSEUM Type: S.A.M. 915. -05 breadth T°2 7:56 | -41 -88 | >-05 © length 15-3 15-2 | 17-16 | 1-14 | 1-86 | >-05 breadth W262) ie U9 yA 3235 -84 | 1-05 | >-05 P, length 14-8 | 15-1 15:55 -50 | 1-50 | >-05 breadth .. ne 10-0 | 10-5 | 11-03 -50 | 2:03 | <-05>-04 length : breadth ratio 1-48 1-44) 1-41 06 | 1-16 | >-05 P, length 19-6 21-17 ‘40 | 3:70 | <-0004* breadth .. ay 12-7 |.12°7 114-40 -60 | 2°83 | =-005* length : breadth ratio 1-54 1-47 | -06 | 1-20 | >:05 P, length D322 4 | 23e lyk lean 52 -56 88 | >-05 breadth .. te: 12-4 | 12-4 | 13-80; -51 | 2:75 | =<-01>-005* length : breadth ratio 1-87 1-86 1-71 -06 | 2:67 | <-01>-005* M, length .. dis et ey 224 | 2204. 2a D6 -89 | 2-43 | <:02>-012* breadth .. i, We See IAS IEA «| 12761 -43 | 3-05 | <-:003>-001* | E.C. 4 | E.C. 6 | | | | P? length .. oh BN Dp PZB AL GIO PAIN 9.220105) °63 | 1:08 | >-05 breadth .. he Ps «. | 15°9 | 15:9 | 15-98 -70 -11 | >-05 length : breadth ratio .. a 1-45 1-40 1-43 -05 ‘06 | >-05 P'dength .. sia si Le ors 35-31 | 1-08 -45 | >-05 breadth .. oy, ao Cea ae) ite | 21-48 68 -56 | >-05 M' length .. lid ie ie 6-0 5-62 -38 | 1:00 | >-05 => -05 breadth .. 2 hs .. | 14-0 13-07 enh | LeO7, ee ‘ i A (11) Crocuta spelaea (Goldf.). Specimen E.C.1: posterior half of skull, broken off at about the level of the post-orbital constriction. The zygomata are missing and the auditory bullae broken, but otherwise the specimen is almost perfect (Pls. XXVII- XXX). 338 ANNALS OF THE SOUTH AFRICAN MUSEUM Specimen E.C.g: mandibular fragment with much damaged P., Ps, P. and the alveoli of M,. Specimen E.C.8: mandibular fragment with the anterior two thirds of M, (Pls. XXX, XXXI). | Specimen E.C.5: maxillary fragment with P* and damaged P* (Pls. XXX, XXX). The skull is large and heavily built, the occiput high and narrow, and very distinctly ‘‘shouldered’’ (see Pl. XXVIII). The lambdoid and sagittal crests are well developed, and the latter is very high posteriorly. The con- formation of this region is very similar to that of a skull of Crocuta spelaea from Sundwig in the British Museum of Natural History (No. 28558) and an occipital fragment (M. 4570) from Torbryan Caves also shows similar development of the crests. Goldfuss (1821) notes as a characteristic of Crocuta spelaea the large development of the post-glenoid process. This is not at all an easy character to assess or to measure, particularly since in fossil material the tip of the process is commonly slightly damaged. From an examination of the C. spelaea material in the British Museum of Natural History it appears that, apart from its size, the post-glenoid process of C. spelaea differs from that of C. crocuta (Erxl.) in its orientation. In C. spelaea the process lies almost in the transverse plane of the skull, whereas in C. crocuta it slopes distinctly forward from its outer to its inner end. In specimen E.C. 1 the process is. orientated as in C. spelaea (Pl. XXIX). | Unfortunately both the carnassials are damaged posteriorly, but the presence of posterior roots makes it possible to restore the missing portions. with a fair degree of accuracy, and at least to determine a minimum length. In specimen E.C. 5 the palate is somewhat damaged at the posterior end of P*, and it is therefore impossible to be quite certain that M’ was absent, but clearly if present at all it can only have been very small. The upper carnassial possesses the long metacone and relatively short parastyle characteristic of advanced Crocutas, but the protocone does not slope forward as much as is usual in C. spelaea. The lower carnassial, apart from its large size, 1s remarkable only for the structure of the anterior cingulum. This forms a sharp shelf-like excrescence round the antero-external margin of the tooth extending for just over 2 mm. and then ceasing abruptly at either end. Since the posterior portion of the tooth is missing the characters of the talonid are unknown. The premolars do not provide any distinctive characters but closely resemble those of Crocuta crocuta, except for their larger size. The posterior part of P. is missing, but the posterior root is very broad: P. in C. spelaea is commonly very wide at the posterior end. FOSSIL CARNIVORA FROM HOPEFIELD 339 The measurements of the specimens are given below, compared, in the same way as before, with the corresponding figures for C. spelaea (table 2). The only points of difference are that the premolars are a trifle narrower in the Hopefield than in the European specimens. In this character the Hope- field specimens resemble the capensis subspecies of C. spelaea described by Broom (1939) (see also Ewer 1954) from Kromdraai. C. spelaea capensis, however, differs from typical C. spelaea also in the large and almost square M’ and in the very long P’, neither of which characters is shown by the Hopefield specimens. In addition the protocone of P* in C. spelaea capensis is large and slopes forwards very considerably. The general narrowness of the premolars is a primitive character, and cannot, by itself, be taken to indicate any particular close relationship of the Hopefield specimens to C. spelaea capensis. The former are best regarded as closely resembling the typical European C. sfelaea, differing only in the slightly more primitive character of the premolars. This difference does not seem of sufficient importance to & TABLE 2. Measurements of Hopefield specimens of Crocuta spelaea. The dimensions of the teeth are compared with those of a sample of C. spelaea from various European localities: the figure in brackets after the mean for the latter gives the number of specimens on which the mean is based. Other symbols as before. Specimen E.C. 1. Maximum vertical height of sagittal crest above upper edge of foramen magnum 81:6 mm. Maximum vertical height of external occipital protuberance above foramen magnum 55:0 mm. Maximum width across occipital condyles 47-0 mm. Maximum width of skull (intersquamosal) 95-0 mm. Maximum width across mandibular fossae ca. 124-0 mm. | European sample 4 | Mean S.D. | oO P Specimen E.C. 9 Pyiength .. Ne ie Sg Meee 7/3) 16-9 (31) 85 TPA SPOS Peneneth |. a BY. ee | 22D 227) (35) -78 -64 | >-05 breadth .. 8 ih abt 14-6 | 16:5 -93 | 2:04 | =-04 length : breadth ratio .. Weel 2 1-38 -054 | 2-59 | =-01* Pare er rs A oaes 193.5) (32). 1131) 89, 1s s8205 breadth .. ie Me: MY 13-5 14-9 87 1-61 | >-05 Jength : breadth ratio .. an 1-8] 1-58 -088 | 2-61 | =-O1* Specimen E.C. 8 | M, length .. a i re Caz 30°'S.) id 24i (26) +2 1,233 1:43 | >-05 breadth .. ae ie Ayal 12-8 Sp) 67/5) 1-23 | >-05 Specimen E.C. 5 | PoOMenethy ..),. me i! bien 24:3 | 24-1 (19) | 1-34 “15, |=="05 ‘breadth .. ie Ne chee 16:6 | 17-9 a7 1-11 | >-05 length : breadth ratio .. 5 1-46 1s -§2 | 2:11 | =<:04—:03 Pedenpty ii)5 . ‘. hs vol ca 40 40:9 (27) “92 -98 | >-05 225) L222 es L238) e205 breadth .. a ie ABA ce OA 340 ANNALS OF THE SOUTH AFRICAN MUSEUM warrant the erection of a new subspecies for the Hopefield specimens, but should be borne in mind in considering the probable age of the deposit. 3. Family Felidae (°?) Leptatlurus serval (Schreber). Specimens E.C.15, E.C.17: mandibular rami, incomplete anteriorly and posteriorly, bearing P; Ps and M,. Specimen E.C.16: portion of mandibular ramus with P. and Mh. The specimens all belong to a small felid, and the teeth do not show any | peculiar characters. A set of skulls is not available to us for detailed comparison, but, judging from the measurements given by Roberts (1951), the specimens are a trifle too large to belong to the Cape wild cat, Fels cafra Desmarest, and fall within the range of Leptailurus serval (Schreber), to which species they are tentatively referred. TABLE 3. Measurements of fossil specimens of Leptailurus serval. Specimen No. | E.C. 15 | E.C. 16 | E.C.17 Depth of mandible between P, and M,.. ie Ag Ne 13°5 1562), eae +S P, length .. vy oy uth i Wy uk re 8°7 7°5 breadth .. uM et iG i tei) an 4°8 3°9 P, length .. Me ws ae i, A 4) in 10:5 10-7 9-7 breadth .. a Hs Mi aD Li Hs gi 5-8 5-1 5-1 M, length .. Mi eit she ir i, a AN 12-3 11°4 11-5 breadth. . bs uh ae a i ua 6:2 4-9 5°5 4. Family Canidae (i) Lycaon pictus (Temm.) n. subsp. magnus. DIAGNOSIS: A subspecies of Lycaon differing from the extant form in the greater length of the mandible, larger lower canines and incisors and longer but relatively narrower lower premolars. Specimen E.C.13: portion of left mandibular ramus, incomplete below and bearing P, to P, and the anterior root of M,. (Pls. XXX, XXXI.) Specimen E.C.121: right mandibular fragment bearing I., I, and C. Specimen E.C.30: isolated damaged right M,. All three specimens show moderate wear and are in a very similar state of preservation: it is probable that they belong to a single individual. Apart FOSSIL CARNIVORA FROM HOPEFIELD 341 from their distinctly greater length the teeth differ in no way from those of the living Lycaon pictus (Temminck). The measurements are given below, compared as before with those of the extant species. It will be seen that the fossil jaw is considerably longer than that of the extant species and that the teeth differ significantly from those of the latter in the following points: the premolars are considerably longer, but not much broader; the canine and I. are significantly larger in both dimensions while the significance of the slightly greater breadth of I. is less certain. It is considered that these differences are sufficiently clear cut to warrant placing the fossil specimens in a distant subspecies. Wells & Cooke (1942) record from Vlakkraal a damaged M, and a worn I,; the latter is said to be “‘a little larger than any specimen of this genus actually axamined.’’ It seems not improbable that this belongs to the same subspecies as the Hopefield material. TABLE 4. Dimensions of teeth of fossil Lycaon compared with those of a sample of 1 extant Lycaon pictus. Symbols as before. Extant | Specimen No. L. pictus | F )EC. 12 EC13/ Mean | SD. | o P Length from back of P, to front of P, .. ws ee i 52-8), |40-7 1:79 | 6-76 | <-0004* I, length 555) 4-48 “57 04 | >-05 breadth 5-6 4-71 -39 | 2-29 | <-:03>-02 _ I, length re mt te 7:6 6:03 -47 | 3-34 | <:-001> -0004* breadth ct sf ia 6°5 5-66 -30 | 2-80 | =-005* C length a a .. | 13-0 10-61 ‘72, | 3-32 | <:001> -0004* breadth a! a ib 9-1 7-80 53 | 2:45 | <:02>:012* P, length 8-1 6:45 -50 | 3:30 | <-001> -0004* breadth if ae 5199) 4:71 350 WV Ned ipl 205 length : breadth ratio 1-560) 373 *119 | 1-57 | >-05 P, length 12-4 10-05 -62 3:79 | <-0004* breadth nas ap Bop) 5:35 “25 60 | >-05 length : breadth ratio 225 1-879 -076 | 4-88 | <-0004* P, length 14-3 11-95 54 | 4-37 | <-0004* breadth (01) |. 6-3 | 6-08 | -39 56 | >-05 length : breadth ratio 224 1-970 | -130 | 2:30 ; <-03>-02 | P, length 16:5 13:75 | -79 3°48 | <-0004* breadth om My 7:3 7-11 | 47 40 | >-05 length : breadth ratio 2:26 | 1-938 : 100 | 3:22 | <-003>-001* M, breadth... ye is 10-4 992 342 ANNALS OF THE SOUTH AFRICAN MUSEUM (ii) Canis mesomelas Schreber. Specimens E.C.18-20, 22-27 and 29; various incomplete mandibular fragments. : Specimen E.C.31: isolated Mi. Specimen E.C.32: isolated Py. Specimen E.C.21: maxillary fragment bearing M’ and M’. In another paper (Ewer, in the press) only a single character was found in which there was no overlap between the two living species of jackal, Cams mesomelas Schreber and C. adustus Sundevall, and which therefore allowed of a certain assignment of a single specimen to one or the other species. This character is the length of the carnassial tooth relative to that of the succeeding molar. This is referred to as the ‘‘carnassial: molar ratio’’ and either upper or lower ratios serve to separate the two species. In the case of the material under discussion at present four specimens, E.C.18, 22, 25 and 26, include both M, and M: and therefore allow of the calculation of the lower carnassial: molar ratio. The values found are 2.03, 1.98, 2.10 and 2.30 respectively. For the samples of 14 C. mesomelas and 10 C. adustus used in the previous study the values range from 1.99 to 2.57 for the former and 1.60 to 1.81 for the latter. The Hopefield specimens clearly belong to mesomelas, and in the table of measurements the mean and range for the sample of extant mesomelas are given for comparison (table 5). It will be seen that although the means for the Hopefield specimens lie within the normal range for mesomelas a number of individual measurements lie beyond the upper size limit found for the living species. A fossil subspecies of C. mesomelas has been described (Ewer, in the press) from the Transvaal deposits, differing from the living form principally in the fact that relative to the length of the carnassial M, is less reduced and the premolars are longer. It is therefore of some interest to compare the lengths of M. and of the premolars in the Hopefield specimens with the corresponding values for the Transvaal fossils. Figure 1 shows the mean values for the relevant dimensions, with those of the extant form for comparison. It will be seen that in each case the Hopefield value is inter- mediate between those for the Transvaal subspecies and for the living form. There seems to be little doubt that we are here dealing with a phylogenetic series in which a reduction of the molars and a shortening of the premolars is taking place; the Hopefield specimens representing a form ancestral to the living black-backed jackal and the Transvaal form in turn ancestral to the Hopefield. The functional significance of the changes is not at once apparent, and it may be that they would be better expressed by saying that a general reduction in overall size is taking place, but without any reduction in size of the carnassials, which therefore show an increase in relative size. Without FOSSIL CARNIVORA FROM HOPEFIELD 345. tig. 1 P, AXA ! GI fransvaal subspecies. x Hopefield specimens. A Living GC. mesomelas. Fe Aye! aS a 2 P. R= "3 AX———_f] & @ ; e re) PR sa 4: A—xX—) M, A a 4. Sar “ta 3 10 7 12 Tooth length (mm) VALUES. Length(mm.) Transvaal Hopefied. Living. Pen 4.5 4.2 4.0 P, 9.0 8.3 8.0 Sela 9.8 8.9 8.8 Pp, Th. 10.8 10.3 M, 9.6 9-4 8.5 whole skulls and skeletal material available it is not possible to decide whether this latter formulation is the more correct, It may be noted in passing that the Hopetield Lycaon also differs from the living species in possessing a longer premolar row. | : A second fossil subspecies of C. mesomelas is known from African deposits. This is Canis mesomelas latirostris (Pohle) from Olduvai and Serengeti. This subspecies is distinguished from the living form by its very broad snout and shorter and less inflated auditory bulla. In the absence of a complete skull no adequate comparison of our material with this subspecies is possible, but Pohle (1928) gives measurements for the lengths of the teeth of his three specimens. The lower premolars resemble those of the Hopefield specimens in being rather longer than is usual in living C. mesomelas, but M. (present in Annals—2 | : | | | co Lor) = an La) = SS | ES | | ANN en, | Am Rae | nN | ER pNeReee | nnn | ene nes | eeatnnensineememet | erie rete | EGeT eenncem eS | atmpceminnenel Secon | -05 Palate length ; ae $e Ris 50-4 60-65. | 4-389 | 2:34 | ==-02* Posterior palate width? dT a Me 16-6 19-45 1-354 | 2-10) —-04=>-03 Postorbital width é “a LY. 28'-2 34-47 1-533 | 4:09 | <-0004* Maximum cranial width Ne een ere (CIS aN 65:81 3-540 712 i = 0S Width of nasal aperture. ¥ Tee aor £3709 1-579 | 1°48: |) "05 Length of P4 & tt 2048 12:67 1-133 |: 1-65) 3-05 Width of palate at level of P! NU oS Getta 45-33 3-114 | 1-84 | >-05 Palate length as % of basilar lengih .. 44-60 49-226 |" 1-990") 2432 | 202" Posterior palate width as % of basilar length .. 14-69 15-780 -968 | £13 |S =05 Postorbital width as H4 of basilar length 24-96 28:026 | 1-704 | 1-80 | =>-05 1 This is the width acros the posterior end of the hard plate. III. DzIscussIon. The faunal remains described above may be considered in terms of the light they may throw on two problems: the date at which the Hopefield deposit was laid down, and the ecological nature of the locality at that time. As regards the latter point the carnivore fauna adds little to our knowledge. All the species described are very similar to species which have existed within the area during historic times. One curious point is that although hyaenas and jackals are present, no remains of large carnivores, such as leopard and lion, have to date been discovered. Artiodactyl and horse remains are abundant, and it seems very improbable that in fact no large carnivore predators were present. It seems likely that further work ° may bring lion or leopard remains to light, and their absence cannot at this stage be taken as established. As regards the date of the deposit it is quite clear on the one hand that it is much more recent than the Transvaal deposits. The species represented are all closely related to living forms, and no trace has been found of the archaic hyaenids, Lycyaena and Leecyaena, and the sabre-tooths which exist in the latter deposits. At the same time the fact that most of the specimens show slight differences from their living counterparts indicates that the deposit cannot be of very recent origin, but is likely to be at least as old as the upper Pleistocene. The Vlakkraal deposit in which was found the large Lycaon incisor previously mentioned is estimated to belong to the upper Pleistocene (Wells & Cooke 1942). Chemical analyses, carried out through the courtesy ° of Dr. K. Oakley of the British Museum, on a fragment of Hyaena brunnea mandible indicate that the fluorine content does not differ from that of the FOSSIL CARNIVORA FROM HOPEFIELD 347 human skull and of Mesochoerus lategani. The carnivore fossils per se do not provide any grounds on which it would be possible to decide whether the deposit belongs to the upper Pleistocene, as has been suggested (Singer & Keen, 1955), or is as old as the top of the middle Pleistocene. SUMMARY All the carnivore remains so far recovered from the Hopefield site are described. These include a new subspecies of Lycaon, Lycaon pictus magnus, a Crocuta spelaea and a jackal showing characters intermediate between the fossil subspecies from the Transvaal cave deposits and the living form, together with a Hyaena brunnea and a Mellivora capensis, each differing very slightly from the corresponding living form. In addition there are fragmentary remains of a viverrid resembling Herpestes ichneumon and a felid, probably Leptailurus serval. It is concluded that the carnivore fauna is consistent with the upper Pleistocene dating which has previously been suggested for the deposit. ACKNOWLEDGEMENTS We are indebted to Dr. Hewitt of the Albany Museum, to Dr. K. H. Barnard of the South African Museum and to Mr. Skead of the Kaffrarian Museum for their allowing us every facility for studying skulls in their collections and for the loan of material. One of us (R.F.E.) is also indebted to the authorities of the British Museum. (Natural History) for facilities for examining and measuring material in their collections, and to the South African Council for Scientific and Industrial Research for a research grant. Mr. G. McManus, Department of Surgery, University of Cape Town, kindly photographed the specimens. : Part of a research grant from the Dr. C. L. Herman Research Fund of the University of Cape Town allocated to Drs. Singer and Keen was utilized for studying the carnivora. One of us (R.S.) is indebted to the Wenner-Gren Foundation for Anthropological Research, New York, whose motor vehicle, donated to the University of Cape Town, is being extensively used for collection of material from the site. Publication of this paper was assisted by a grant-in-aid made by the Council of the University of Cape Town. REFERENCES Broom, R. 1939. A preliminary account of the Pleistocene carnivores of the Trans- vaal caves. Ann. Transv. Mus. r9, 331-338. Drennan, M. R. 1954. Saldanha Man and his Associations. Amer. Anthropologist 56, no. 5, 879-884. DRENNAN, M. R. and Sincer, R. 1955. A Mandibular Fragment, probably of the Saldanha Skull. Nature 175, 364. Ewer, R. F. 1954. The fossil carnivores of the Transvaal caves. The Hyaenidae of Kromdraai. Proc. Zool. Soc. Lond. 124, 565-585. Ewer, R. F. 1955. The fossil carnivores of the Transvaal caves. The Hyaenidae, other than Lycyaena, of Swartkrans and Sterkfontein. Ibid. 124, 815-837. Ewer, R. F. (in press). The fossil carnivores of the Transvaal caves. Canidae. Ibid. Gotpruss, G. A. 1821. Osteologische Beitrage zur Kenntniss verschiedener Satige- thiere der Vorwelt. Uber die Héhlen-Hyane (Hydne spelaea). Nova. Acta Akad. Caes. Leop. 456-462. Poute, H. 1928. Die Raubtiere von Oldoway. Wiss. Ergeb. Oldoway-Expedit 1913. N.f. Hfr. 3. 45-54. Roperts, A. 1951. The Mammals of South Africa. Published by the Trustees of the ‘‘Mammals of South Africa’’ Book Fund. SINGER, R. 1954. The Saldanha Skull from Hopefield, South Africa. Amer. J. Phys. Anthrop. n.s. 12, no. 3, 345-362. SmncER, R. and Keen, E. N. 1955. Fossil Suiformes from Hopefield. Ann. S. Afr. Mus. 42, 169-179. WELts, L. H. and Cooke, H. B. S. 1942. The associated fauna and culture of the Vlakkraal thermal springs, O.F.S. Trans. Roy. Soc. S. Afr. 29, 203-233. ‘ Ni oes a iy } iy A 2.1 Ai | : La aya aaig i r Bt Mas Nth { } ASTI ‘ i x WP een } a] a * ih ‘ ¥ ; % 8 q i 4H ANCE V AN, Prate XXVII Name oe arr. Mus. Vol. XLII Pirate XXVIII. Crocuta spelaea Goldf, Side view of specimen E.C.1, ANG, S\, Aver, WIE, Woll, SGU PLATE XXVIII. Crocuta spelaea Goldf. PEATE SOGviial Posterior view of specimen E.C.r. So PLATE XXIX Nii SmeAdr. Mus), Volk XETI scale cm 1 2 3 Prare XXIX. Crocuta spelaea Goldf, Ventral view of specimen E.C.1. ANG, S), Boe, Whos, Woll, SLICE PLATE XXX. Lingual view of: Top: Lycaon pictus magnus n.subsp. Specimen Centre: Crocuta spelaea. Specimen E.C. 5. Bottom: Crocuta spelaea. Specimen E.C.8. Nimes Air. Mus Vol. XII PLaTE XXXI cee PATE Oe Labial views of the specimens illustrated on Plate XXX. wc AWW, S, Ase, Wins, Woll, SILI PrATE DOxOxXi a Mt 1 — PLATE XXXII. Mellivora capensis (Schreber). IS (Caikal. Dorsal and palatal views of specimen i 18. Further Fossil Suidae from Hopefield. By E. N. KEEN and R. SINGER, Anatomy Department, University of Cape Town. (With Plates XXXIII-XXXV.) INTRODUCTION In a previous publication (Singer and Keen, 1955) the available suid material from the fossil site on the farm ‘‘Elandsfontein’’ was ascribed to a single new species Mesochoerus lategant. Subsequently, on a number of field trips, one of us (R.S.) and a medical student, A. J. van Niekerk, recovered not only further specimens of the above genus, but also a fragment of a mandible containing deciduous third and fourth molars and a permanent unworn first molar (specimen S. 22) of this genus, as well as a second and third molar (probably upper) of a different genus, namely Tapinochoerus. It is the purpose of this paper to describe the new material in the Anatomy Department : — A. The additional Mesochoerus specimens so as to increase the range of variation in Mesochoerus lategani and to evaluate the status of this species better, and to describe a specimen (S.28) of Mesochoerus paiceae. B. The milk dentition in the fragment of mandible numbered $.22. C. The Tapinochoerus specimen (S.26) and to establish its specific rank. DESCRIPTION AND DISCUSSION A. Genus MESOCHOERUS Shaw and Cooke DESCRIPTION 1. Mesochoerus lategant Singer and Keen. The general description of these four specimens (S.23, 24, 25, 27) is the same as in our previously published material (1955), and it is only necessary to note the specimens and their dimensions, and to outline any additional distinctive features. - ae ame et we FURTHER FOSSIL SUIDAE FROM HOPEFIELD So Specimen S.23: A left upper third molar in which the posterior part of the talon consists of only one pillar behind the third lateral pair. This single pillar is vertically grooved as if in partial separation. The anterior median pillar and its outriders project beyond the root margin giving the tooth a pronounced rounded bulge anteriorly, well seen in side view (Pl. XXXIII A). On reviewing our specimens, this feature is seen in all the third molars particularly when in early wear. It is also observed in Metridiochoerus as well as in Mesochoerus olduvaiensis and paiceae, while in Sus limnetes from Omo (now classified as Kowropotamus = Potamochoerus by Hopwood and Hollyfield, 1954) the anterior median pillar and its outriders form a separate plate. In Notochoerus and Phacochoerus species the bulge is absent. Specimen §.24: A left upper third molar in which the talon behind the third lateral pair consists of 5 small pillars. The buccal pillar of the second lateral pair also has an accessory nodule anterior to and confluent with it. Specimens S.23 and S.24 are in approximately the same stage of late wear, yet in S.23 the posterior pillars are fused into one massive posterior median pular. This variation in number of pillars in the talon has already been commented on in our previous publication. --Specimen $.25: A right upper third molar whose front portion is missing — as it is broken through the anterior pair of lateral pillars. The lingual surface of the 2nd and 3rd lateral pillars shows two deep vertical grooves, giving an exaggerated appearance not yet encountered in this species and suggestive of incomplete fusion of the cones which form the complex occlusal pattern of the teeth (Pl. XXXIII B). Viewed from the occlusal aspect the tooth presents an unusual concave curvature on the labial side of the posterior part of the tooth (Pl. XXXIII C) which is produced by a bulge at the crown-root junction of the 2nd lateral pillar. The roots are missing. Specimen $.27: An almost complete right upper third molar. The anterior pair of roots are broken off, and a small piece of the central portion of the first lateral pillar on the lingual side is missing. The part of the talon behind the 3rd lateral pair in this massive tooth also consists of 5 pillars. The anterior median pillar and its outriders are so large as to almost form a separate plate, while the 2nd lateral pillar on the lingual side has an accessory cone in front of it (compare S.24) while the pillar on the buccal side has a small nodule projecting up in front of it at the cingulum. The buccal pillar of the 3rd lateral pair has 3 small accessory nodules on its outer side. The outer root of the 2nd pair of lateral pillars, though broken near its tip, is projecting back acutely to overlap and cover the plate-like roots of the 3rd pillar with which it is partially fused just below the crown-root junction (Pl. XX XIII D). 352 ANNALS OF THE SOUTH AFRICAN MUSEUM Maximum Maximum Occlusal Height of unworn Specimen. Length. Breadth. Breadth. pillars above root. Se 7s Ot Re MM RD TORY De URN o's LORD wh NT OEE 25-4 212 — SOSA Ty atin aces WVelee te Mie ete Neen oa yy CO) 25-9 21.4 26.0 (posterior) SS HTS EES AC CE PE: — C27 C.25 30.0 (posterior) SA Ant ee) Gent as MTN Cah oe, MI oS Bs 29.7 24.2 28.0 (posterior) Range of previous specimens (7) 63-70 24-28 20-23 22-35 TaBLE I: Dimensions of the crowns of the upper third molars (mm.). The length of S.23 and S.24 might appear to exclude them from the range of variation revealed by the previous series. However, in the previously described series of lower third molars, of which 6 could be measured, the smallest was 67 mm. long, while the other 5 ranged from 73 to 77 mm. (mean 75 mm.), but there seemed to be no other reasons for excluding the short tooth from the series. In the same way, the shortness of S.23 and S. 24 will not exclude these two upper molars, identical in all other respects, from the species. The range of variation in length is now increased to 59-70 mm. Bearing in mind the following description of a different species of Mesochoerus from our site, and also that no upper teeth of Mesochoerus paticeae have previously been described for comparison, there is a possibility that the smaller upper teeth which we have included in our range for Mesochoerus lategani (e.g., S.24) may yet prove to be the uppers of Mesochoerus paiceae. From table 1 it is also seen that $.25 and S. 27 are at the upper end of the range of variation for maximum breadth, and actually $.27 outstrips the other specimens and increases the upper end of the range to 29.7 mm. 2. Mesochoerus paiceae Broom. Specimen S.28: A mandibular fragment containing a right third molar tooth in early wear. There are only 4 pairs of lateral pillars and these are more widely separted from each other than in Mesochoerus lategani specimens. The pillars taper to a pointed occlusal end, their bases being wide and partially fused at the cingulum. The 2nd lateral pillar on the labial side has an accessory cone anterior to it (Pl. XXXIII E). The anterior median pillar is pushed somewhat to the lingual side by a large outrider which is continuous with 6 low nodules fused together to form a plate anterior to the anterior median pillar. Dimensions of S.28 (mm.):— Maximum length 9) 0 OS ty A Maximum’ breadth... 0° S0 oe" Ve Re eke Occlusal breadth EUV AMER ec ha 179 Height of unworn 3rd pillar above root .. .. .. .. 31.5 FURTHER FOSSIL SUIDAE FROM HOPEFIELD 353 The individual dimensions of this tooth fall within the previously reported range of Mesochoerus lategani, with the exception of the maximum length which is 2 mm. short of the lower end of the range. In itself this would have been of minor importance. Hower, there are only 4 pairs of well- separated lateral pillars, as compared with 5 pairs in Jategani, and behind the fourth pair is a single undivided median pillar. In addition, despite its shortness, the breadth is at the upper limit of our previous range. Taking these 3 points into consideration, it cannot be distinguished from the 2 specimens (Broom’s type and Shaw and Cooke’s neotype) on which Mesochoerus patceae is based. DISCUSSION Since our previous publication on the Hopefield Suidae, Hopwood and Hollyfield’s monograph (1954) has been received. In this they classify Mesochoerus with Hylochoerus, hereby following Arambourg (1947), although the latter was not quite certain of this attribution. We have weighed up this opinion in the light of the reasons which led Shaw and Cooke (1941) to decide that their genus should be separated from Hylochoerus. Briefly, these were that the third molars were elongated and not so brachyodont as in Hylochoerus, and that the premolars were not reduced as in the modern animal. These points remain, in our opinion, ample justification for maintaining the genus Mesochoerus which now comprises three species, namely, patceae, olduvaiensts and lategani. Leakey’s Mesochoerus heselont has been attributed by Arambourg (1947) to Omochoerus. However, Leakey (personal commu- nication, 1955) still maintains that Omochoerus should be included under Mesochoerus. Consequently the position of heselont is held in abeyance. On a phylogenetic sequence, Mesochoerus would seem a logical intermediate stage between Hylochoerus and Metridiochoerus. This problem may finally be resolved by further discoveries, particularly of tusks and other skeletal remains. | The specimen S.28, as described above, cannot be grouped with the previously described M. specimens of Mesochoerus lategani. Its dimensions and general appearance are such that it must be identified as Mesochoerus paiceae. In our previous series S.21 was included on the assumption that a 5th pair of lateral pillars was broken away posteriorly. Re-examination of this specimen, in the light of the fresh discovery, makes this seem less likely, and S.2t may represent a second example of a tooth which must be referred to the genotype patceae. 354 ANNALS OF THE SOUTH AFRICAN MUSEUM B. MESOCHOERUS. Milk Dentition Specimen $.22: A left mandibular fragment bearing an unworn M:, Dy and a root fragment of D, (Pl. XXXIV). | Dimensions of the fragment :— Potal engsth (2 Pay Os” ED: ae Maxamiunt breadth’), 2") 2°05), REI Qo) Oe ht Height “or mandible at IMG) 08 ey er eae The first molar is unworn and is composed of 2 pairs of lateral pillars with a double median pillar between and separating them, small anterior outriders, and a large complex posterior median pillar. The posterior lobe of the tooth is broader than the anterior, and its 2 pillars are more separated from each other. The enamel is coarsely rugose and its irregularities form a pattern which is best seen on the labial side of the posterior lobe. The anterior and posterior lateral pillars fuse with each other about 4 mm. above the cingulum, and the posterior median pillar fuses with the posterior lateral pillar on both sides. The cingulum is only slightly developed. There are 4 roots which cannot be directly inspected as they are embedded in the mandible, but they were examined on a skiagram. The posterior roots are larger than the anterior. -Dimensions of M, (mm.):— Maximum length Rian wate uniac mre Ol ieee. 23 Maximum breadth (posteriorly) )0s" 2...) toe 13 Height "ortrown (amterionhyy: Ve yey vee 16 Height" of ‘crown’ (postenorly)) "2 Ve Root length er ee So ee rr Behind the tooth is a broad incomplete cavity which presumably contained the developing M.. The breadth of the mandible diminishes noticeably behind M., and there are no signs of root canals from the cavity for M:. © ; The last (fourth) deciduous molar is a trilobed tooth with 3 roots on the labial side and 2 on the lingual side. It is in moderate wear. The 3 pairs of lateral pillars increase in size and decrease in wear from front to back. The 2nd lingual pillar is broken. The general appearance of the tooth is seen in Plate XXXIV. Examination of the fragment and of the skiagram reveals that the roots of this tooth straddle a cavity which one would have expected to contain the developing tooth germ of P.. This cavity commu- nicates widely with the mandibular canal and it seems logical to assume escape of the tooth germ through this opening. A skiagram of the mandible of a modern domestic pig at about the same stage of development shows an unerupted tooth resting in a cavity at exactly the same position in relation to D, as in the fossil specimen. FURTHER FOSSIL SUIDAE FROM HOPEFIELD 355 Dimensions of D, (mm.) :— Mermum tenethe 2.) OF... Mie TAS ae 24 Maximum breadth at bier th (hdsterioniyy) ce) ee Te Mersin lensth: of roots i(irem! Xray) NP Ae err 8 The third deciduous molar is only represented by a root fragment resting in one of the 2 alveolar root canals in front of D,. The bulk of the suid remains collected from Elandsfontein have proved to belong to the genus Mesochoerus. The dimensions of M. of $.22 compare very favourably with those of the 2 worn M, teeth of Mesochoerus lategan previously described, especially if the measurements are taken close to the cingulum. The unworn height of the crown (16 mm.) compares reasonably with the unworn height of M; (3rd pillar), which averaged 29 mm. in 4 specimens in which it could be accurately measured. The ratio of the unworn height of M,/M; is thus 1:1.8. There seems good reason to conclude that this specimen is derived from a young Mesochoerus. C. Genus TAPINCCHOERUS van Hoepen and van Hoepen Tapinochoerus meadowsi Broom DESCRIPTION Specimen S.26: A third and a second molar, found lying in apposition, were subsequently joined together with plaster-of-paris. The second molar is partially embedded in bone. The third molar resembles the tooth on which Broom (1928) originally based the species Notochoerus meadows? ( = Tapinochoerus meadowsi, Cooke, 1949). One of us (R.S.) examined specimens of Tapinochoerus meadowsi from the Transvaal Museum ee Ske) 286) bE 1), now, being ‘studied by Dr" KR. FF. Ewer, Rhodes University, Grahamstown. The former two specimens are lower M. probably from the same jaw, and B.F.1 is a part of a skull, briefly described by Broom, 1948, containing unerupted M° teeth and partially worn M’ teeth. As a result of careful comparison, S.26 is considered to consist of left upper molars. Left upper third molar: The general structure of the tooth is phacochcoeroid consisting of two rows of lateral pillars separated by intermediate pillars. The anterior pair of lateral pillars is in full wear and constitutes the first lobe. The pair behind this is just coming into wear (Pl. XXXV). _ Posterior to. this the unerupted portion of the tooth falls away sharply at an angle of | about 45°. This part of the tooth consists of a large number of pillars (probably constituting 3 lobes) on each side with an intermediate pillar between . 356 ANNALS OF THE SOUTH AFRICAN MUSEUM each pair. These lobes are less separated than the type specimen, but exhibit a similar appearance to S.K.? 387. The extreme posterior end of the tooth is broken off. The anterior lobe is broader than the second, and is rounded anteriorly in the occlusal view, with a slight constriction separating it from the 2nd lobe. The anterior extremity of the lobe consists of a number of circular and irregularly shaped nodules which cause an anterior bulging obvious in the lateral view (Pl. XXXV). The worn surface of the lateral pillars of this lobe are H-shaped with the inner pair of lobules equal in size and parallel to the outer pair. The bases of all the pillars are flattened from side to side, and are open; no roots are apparent. This is a typical phaco- choeroid appearance. The cement covering the tooth is scanty. Three third molar teeth from the Olduvai Gorge (Coryndon Museum, Old. B.K.II, Ex. 1953, nos. 109, 160, 448), studied by R. S., are almost identical in general appearance, with the posterior unworn part of the tooth falling away sharply at an angle of about 45° from the anterior plane of wear. The degree of separation of the pillars varies in the 3 specimens, being more marked in 109 and 448, but the same as S.26 in 160. However, in the latter the outer enamel is thicker and the tooth as a whole is broader. Maximum Maximum Occlusal Height of worn Maximum Specimen. Length. Breadth. Breadth. anterior pillar. Height. S20 NE eee eae Ran eae et 19.2 63.8 70.0 (2nd lobe) Type specimen .. .. 76 19 16 55 64 poo Mine | oly Lin Mn od 19.1 15.7 53.1 68.3 (3rd lobe) Sue ABS ay wie eek Sse IQ.1 15.9 53-0 c.67 (3rd lobe) Old ryraes yt f) Ye Wee Webs ep y ay “4 PEN Meh w aed ; ‘ Mi PENSE HEY a Mii i B HP AS 5 : Hy H Hed Hie ibe A el eh a san A ‘ . SH hte heb fly Hay ‘ Le Lee ee ! 6 , ¥ sth ic i lee " pair bes we? ’ onus e) Ai . » ob bi hewmen ¥ » y b Pa beri YE: q ee By tt + oi oie ees ate eee Ce Se a yur F\ eer.’ } h mitt ty Heh Bit a eae at hala o> PIR DL bop hy SPR EAE hy b heaves UR a ta AG he neh die bib ah & 4 ’ Um DA ey 5 stb! ' ’ i sofa Re wt \ \ ’ Ls Mite. who Hei } » WERE ; b> vee oY b ety eee w flu L } A ytd , Best yet , ty ' 4 i 4 rt 5 ! nite ie at sity ‘ yeh ip t we oye nepess Beh hs ANY whe ony dR, si ae i a ‘ RUM: 6 LS Web & wee t > pe awa WEE ! Buen a Sey , he VR pos irk pe citaey kL wept Ae tem , > yy RARER Fe ¥ Web testis etek Meena tE Dd eh Pe aie y Br Hie ‘ Hew Keak We eat mead b ; : yee i) i Wh oe ent ee) i" h t eb 4 f Bit Y t “teh Ret ABER PAN felbeglS Sy 1 " + nie t6 al WLS low i : 3 fh He Wit Uae Prey pes ive Roh i" ‘ ts bitye eR Wey s ‘ int RUA ee Os th uP NA Wage Wi (it OA! AN Me Ne a } Bry tan PERRY i » heb es: ” vii ‘ ne a 1 y dus a A 1H ey Peya Mae eae 4 hey eae ih Po ih eS Li a rea . i aiieiee OA EOE ast ip ta Y Lie ite net t i i PL Mote: i a De AA et Waa OB EUS a REC Da Wiaeaseiity oy eit Lat t f HRM NED DB, Nie he ER URL bee bm hho jon the " Ma preemie h' LoTR) (Se Te ee phy VR dey hee ot OF) ys . m eee FEA fy eye wi ‘aa i Pee | Wa eR RW AB tet e i) Sha eb w eee teen Mat feo Ae BM eibehy (Bete? + hh Seed) H hey yee hh (ek n y EOUE Ds Uey f HEATH AE ited oh hey th & & ; } } fi PAO R Pee he pen boy \ bop Bb ly ef iit i ‘ a f y ‘ i t COSY hate ! Taga cat OR jh yt WEB WLS Me Be Re % tf ib ho Whe 4 {i ied eat hh YS hive yi + t i Aa) $ u iv 3 hee y 409 ) : : i KH bY » f Prin \ t i Rt ih pin ho 6 1 } bee} ‘ i GOOG A Ae i are ee wis " Oe SOPRA ca RM te bine ' Aa Get UE tO MO Gel OS AMADIS i 1 oe et Le 7 Hye bi heb yeh b Be WV Aaah Weg G hi ; Ft tthe ‘ f f } bi} , i) Serratia " n oa (eo MEEKS ‘ WWW RP nee Y ) Heel y ese MeL RS aA ag PO SA OA eB yh Wee EB ah Benet yeh ver it bh $ : Th at He, eo t r he ue ty PEGE al LAL! : (gage be Hie nehed v% Pepe bib W pel + + PRES p v . URS tate t , i ' i 4 J " ‘ . ‘ F iH J : , ot j fag ae eck ree i, i RE De ha vf ) ah ihe Hh Pee on as f why ‘ 3 ang i, Ur Ay i nape ive Le Geb EEE ihe Oe) EL ee aa Nesters y H(A be Ve ORE bly HR PP Bld BPG ah Pe Gah LR SO hea ha 29 E hob Hit ak in bh een Nar + hen te ay ” b WTP eR bt } vb 5 Hi Wie Woh ech \ Vien t ‘ Eo ri Hye , SHEER LO bb bebe fe bee ay eek alge he HUE VID AIT Ad Phebe bh Re A bl Ob ASB HM yo) LF hide bobed AN he ay theron a ‘ t wv WW BM y! ahi " ay, i if pee wal PL yeeneh i i ts." A ved) We Be pig dy) ty UU Op Le eb d peoped bhAcih ib (bg . Le aU a ey {ok cb be tee 6 Be pe OR a ~ re a) Ret Aa GW Peet Naty ry ATA La PTO hh bt APL ee . beh, waeky WR aE yet Bh bbe bk 2 iit Der aaah Site Hew he etd PE UME AUS A \ : wea ae My he kb be ou \ » \ \ ete dee Ley ei teal AOU Gboh Bari bE elt EN De oe WO PUVA 1 70 is ca aL ‘ Lard uae | ve oft a Lhe | E Boe ie - : ee PUMCMU DEUS TOU SOULE Cok UA SR tO SO TA BE OLA ROR op nk io Ae PRA EO WD Oe RA ee We Lot UR WAL LRT OA Rr a OPC a WhO & CUO ER TOP Oe AG ep PA em he Re Se te TU Bek or Bb pele it Bonet ne Wt NE oe el CA LL Oe COO I ; hi athe} Op RE bh bee hb. eg PEW HEE DE Geb she eek NN) BW) Wie BA) hebe ts 4 Py Leb t pra heb rnin eG jek ® wan Like i Wa eR hte ne} ‘h Fi WHALh: ANY ay WN a Aan eo 1 eo wt OR eh Gin We LE eee 1 igen be WV Medd be HED Weil Deke eee ae hie + diet Ws i Ao Web Ap GSU Woe 3 " Woh ie be Cr mL | Pah bee ‘i et i » ‘ PRL ern | bet el ee ss) ia Sia ae SNR ase +i) ay a "vidi eA A HOU ae ’ qi 4 een rar’ y ae J +h na wi 4, slp r } Nac 4s BLE ly be Poe, he : ‘ ' < Ae fe ‘ oh wie mirigy GAARA A heb bit rs Me Whew aye I . ar 1 ee On aie ei ietstiene ! » 1 We Thi hee a wi PE ty we A Way \ { Woitek 4 ¥ i j pare HAD a ei be ke ie Pi WRAY BR Re ype ee) DC Siar) Ate et to wi A Bae he ey toi we PHBL ELH okie @ bebe ger tt Wi) 4 ‘ ow Ye, ti A » ar he : a mn 4 ‘eke \ ‘ {AS OUD bay hh UBB pe Bee bb oe aN at) ; he MACAU : | 0h en Me WHE: : NN OW YAN Wh Gil beh A rai Ane es i f i EY bh bbe beh ee? Uy iy i yh ee f U d ‘iy ; ’ We Al oy wy Li tebe nS ’ i 5 wee) | @hteh be heed Wine ht be