FOSSIL HYSTRICOIDEA FROM THE MAKAPA VALLEY, TRA SV AAL By MARJORIE GREENWOOD (Jinja, Uganda) ABSTRACT In this paper are described three Hystricomorphs from the Australopithecus prometheus layer of the Makapan Valley Limeworks Quarry, Transvaal. Besides Hystrix cf. H. a.fricae .. australis, a ne·w genus and species Xenohystrix crassidens and a new species Hystrix major are recognized. INTRODUCTION Although remains of small rodents fiom tire Limeworks Quarry of the Makapan Valley, Potgietersrus District, are abundant in certain breccias, they are rarely found in association \vith the larger mammalian fossils. These "rodent breccias ' are apparently formed by the accumulation of fossilized owl pellets. He 1~ ever, large Hystricoidea, which could not have been the prey of owls occur together vlith the remains of Primates, Subungulates, Ungulates and Carnivora. The larger mammalian fauna occurs in three calcified and cemented layers. At thE time of writing, the remains of Hystricoidea were confined to the Australo­ pithecus prometheus layer, "\vhich is both the lowest and the richest zone of the three. Maxillary and mandibular fragments with teeth still in position as well a ~ isolated teeth are available for study. Some fragments were fossilized in their present imperfect state, \vhilst others \\rere broken subsequently during quarrying operations. Amongst the large collection of Ungulate bones from the quarry are several bearing evidence of rodent gnawings. These incisor marks, which antedate fossilization, are of different sizes. Obviously, therefore, both large and small rodents frequented the caves as ' ell as being the prey of other in ­ habitants. It is apparent from the remain preserved that at least three members of the Hy tricomorph group are represented in these Plio-Pleistocene limestone deposits . CLASSIFICATION Order RODE TIA Sub-order HiTSTRICOIDEA Family HYSTRICJDAE Sub-family H ystricinae Hystrix cf. H. africae-australis Peters 1852) Porcupine material in the collection comprised a fragment of a right lower javv ,vith an incomplete M2 and an unworn M3 (No. M.lOOO B.P.I.) and a small portion of an upper incisor ( o. M.lOOl B.P.I.). 77 Owing to the fact that fossil material is SG- meagre, it is not possible t&··--decide \vith certainty whether or not the Makapan spec1mens differ specif1cally from H ystrix africae-australis. The measurements of the teeth preserved lie within the size range of H. africae­ australis. The enamel pattern of these specimens is closely similar to that in teeth sho,ving a comparable degree of wear in the living porcupine. Since this pa.tttern is known to be a very variable feature it cannot be considered of any great taxonomic importance. Broom (1946) remarked on the occurrence at Sterkfontein of a porcupine which differs little from the living species. Provided that there is sufficient rock or bush cover the South African Porcupine inhabits country varying from arid rnountain areas to thick for est; and the species is present in the Makapan valley today. No information about environmental conditions in any earlier period can be deduced from an animal with such a wide Jange of tolerance as Hystrix africae-australis. Hystrix major sp. nov. Specific diagnosis. An extinct Hystrix with cheek teeth appro::--imately one third larger than Hystrix a fricae-australis. 11 olo-ty pe : A left mandibular fragment with a damaged incisor, a perfect P 4 and portion of the l\1 1 in full wear ( o. M.l002, B.P.I. See figures 27 A and B . Locality: Limeworks quarry, Makapan Valley. Horizon: Fig. 27 .- Hystrix major, holotype (No. M.roo2). Left mandibular fragment with It, P 4 and M1 in situ. A, Labial view and B, Occlusal view. Natural size. Australo pithecus prometheus zone of the Limeworks quarry. Discussion of specific characters: The type specimen, together with a mandibular fragment bearing a well-worn M3 and a broken M2 , are the only remains of this large porcupine so far recovered from the deposit. 78 The well-worn surfaces of the teeth of both specimens have the re-entrant folds forming "islands" of enamel completely separated from the surrounding enamel border. At an equivalent stage of wear the pattern formed by these "islands" is similar to that of the living South African species. As in H. cristata Linnaeus and H. africae-australis, the mandibular teeth are semi-rooted. However, in the ne'v species the roots of the M3 of specimen No. M.l003 B.P.I. are proportionately more robust and divergent than those of H. africae-australis M3 , but compare well with those of H. cristata. The apparent brachyodonty is obviously due to wear, but it should be noted that the lower extremity of the external enamel fold almost reaches the junction of the root and crown, unlike that of the hypsodont teeth of the living South African porcupine. As far as can be determined, the conformation of the lower jaw is identical with that of the living local porcupine, except that the horizontal distance bet,veen the anterior aspect of the premolar base and the inner margin of the symphysial surface is shorter, in spite of its larger size. The first fossil porcupine from Africa, Hy~rix a-stasobae (from Abu Hugar on the Blue Nile) was described by Bate 1951, "vho remarked on the shortness of the ~nout as one of the senual primitive characters exhibited in the single specimen available. Unfortunately it is not possible to compare H. major with H. astasobae, since the latter is only known from a cranial fragment. In size the P 4 agrees closely with H. primigenia Wagner, from Pikermi, Greece, but it is manifestly more elongated antero-posteriorly and the curve of the diastemic surface is not as shallow. It exceeds in length and breadth an unnamed ~pecimen briefly described by Colbert, fron1 the Pleistoc~ne of Mogok, Burma. On the other hand the advanced development of the roots is · similar to the condition found in the Burmese specimen and to H. silvalensis Lydekker, from the Middle Siwaliks of India. Two Hystrix fossils, an incisor fragment (B.M.16594 and an isolated M2 (B.M. 16595) from Olduvai Gorge, Kenya, are apparently from an individual of comparable size. The Olduvai molar is some,vhat more hypsodont than that of Hystrix major. For c-omparison, the dimensions of some correspoding premolars are given belo·w H. africae-australis - mean 5 specimens Longest diameter Anterior-posterior 9.20 H. africae-australis- maximum in 5 specimens ..... . 10.0 H. cristata - N. Africa H. cristata - W. Uganda Hystrix (sp. indet.) Burma A.N.S.P. 14646, after Colbert H. primigenia - drawing in Abel H. maJor - sp. nov 10.5 10.5 10.5 12.0 13.0 1n millimeters Transverse 7.35 8.0 7.5 8.0 8.5 11.0 9.5 The longest antero-posterior dimension of the specimen differs significantly (P.01) from the n1ean of the dimensions in a sample of five specimens of Hystrix africae-australis. 79 .. co 0 A - le M .- D -t err a.- ~ . . _ , - ·· - te nt .- - fc m .- .: ~ .. ..... _ -- _: _· ~· .. .. ::. F ig . 28 .- X en oh ys tr ix c ra ss id en s. A , S y n ty p e A , N o . M .1 00 4, ri g h t m ax il la ry f ra g m en t sh o w in g M l w it h r oo ts ~ x p o ~c d an d M ~ , la bi al v ie w . B , S y n ty p e B , N o . M .IO O S, l ef t m ax il la ry f ra g m en t, P t., M 1 an d M 2 in si tu , oc cl us al vi ew . C , S y n ty p e C , N o . M .1 00 6, p o rt io n o f le ft p re m ax il la an d m ax il 1a w it h 1 1 in si tu , la bi al vi ew . D , S y n ty p e D . N o . M .1 00 7, lt ft m an d ib u la r fr ag m en t, P ,, M t an d M 3 in si tu , li n g u al as pe ct . E , M an d :b u la r fr ag m en t No ,_. M .I o o 8 , sh o w in g P .t an d al ve ol us o f M J , d o rs o .. J at er al vi ew . A ll sp ec in 1c ns na l: ur a] si ze . XENOHYSTRIX gen. nov. Generic diagnosis : A large extinct Hystricoid vvith rooted upper molar · transverse and anterior posterior diameter of the upper teeth subequal, with occasionally the transverse diameter greater; diastemic portion of the premaxilla and the maxilla strongly arched, and mandibular diastema with corresponding degree of curvature. Type species: Xenohystrix crassidens sp. nov. Xenohystrix crassidens sp. nov. Diagnosis: Teeth of large size: overall length of P4-M3 series 40 mm. or more the mean length of the P4-M3 as determined from a ample of eleven Hystrix africae-australis is 33.8 mm.). Syn-type A: A right maxillary fragment with Ml and M2 in position. The root of the Ml have been exposed ( o. M.l004, B.P.I. ee figure 28a . Syn-type B: A left maxillary fragment with P4 Ml and M2 in full wear ( o. M.lOOS, B.P.I., see figure 28b). Syn-type C: A left diastemic curve of maxilla and premaxilla 'vith an almost complete i11cisor in position (No. M.l006, B.P.I., see figure 28c). Syn-type D: A left mandibular fragment with the intra-alveolar portion of 11 remaining. P 4 in early wear, M1, M2 and a fragmentary M3 in full ,wear ( o. M.l007~ B.P.I., see figure 28d . 14dditional material : Other specimens from the arne deposit include a mandibular fragment shovving the diasteme and a slightly \vorn P 4 , and the alveolus of M 1 ( o. 1008, B.P.I., see figure 28e), and a few isolated cheek teeth and broken incisors. One almost complete left upper incisor shovvs indentical curvature 'vith that of Syn-type C. Locality: Limeworks quarry, Makapan Valley. Horizon: Australopithecus pron2etheus zone of the Lime"Torks quarry. 81 T A B L E V I M A X IL L A R Y T E E T H p I p4 I M l I M z M a L. B. L . B . L . B. L . B. L. B. - - - - - - · - - - H ys tr ix a fr ic ae -a us tr al is M in . .. . . 6· 3 6· 0 8· 2 7· 3 7· 3 7· 0 6· 5 7· 0 8· 0 6· 5 M ax . .. . . 8· 5 7· 4 1 I · 0 10 ·5 9· 5 9· 0 9 ·0 8· 5 9· 0 7 · 5 I S ou th er n A fr ic a M ea n .. . . 7· 4 6· 5 9· 6 8· 5 8· 2 8· 2 8· 5 7· 9 8· 2 7· 5 N . . .. . . 7 1 II 10 10 H . cr is ta ta .. . . . . . . . . 7· 3 7 ·I 10 ·5 9· 0 8· 0 8· 0 9 ·0 8· 5 8· 5 7· 0 N . A fr ic a H . cr is ta ta . . . . . . . . .. 8· 0 7· 0 9· 5 9· 0 8· 0 9· 0 8· 5 9· 0 8· 5 7· 0 W . U ga nd a t H . pr im ig en ia 1 ) • . . . .. . . . . - - 11 .0 11 · 0 9· 75 11 · 0 9· 5 10 ·7 5 9· 5 9· 75 P ik en ni , G re ec e t H . pr in 1i ge ni a 2 ) • • . . . . .. . . 8· 0 - 9· 5 - 10 ·0 - 9· 0 - 8· 0 t H . si va le ns is 3 ) . . . . . . . . .. - - - - - - - - - . - S iw al ik R an ge , ln d ia t H . si va le ns is 4 ) . . . . .. . . . . - - - - - - - - - - M og ok , B ur m a t H . la gr el ii 6 ) . . . . . . . . .. - - 6· 0 5· 5 5· 3 5· 4 I 5· 3 5 ·2 I 4· 8 4· 3 H on an , C hi na co tv tH . sp . B .M . 16 59 3 9· 3 8· 0 - - - - - I - O ld uv ai , K en ya . . .. . . . . I 9. "5 I - tH . sp . B .M . 16 59 4 . . .. . . . . - - - - - - ll . 0 - - O Jd uv ai , K en ya tH . sp .6 ) • • . . . . .. . . . . - - - - - - - - - - M og ok , B ur m a t H . cf . H . aj i-i ca e- au st ra li s (N o. M . 10 00 , B .P .I. ) - - - - - - - - - - M ak ap an , T ra ns va al " " (N o. M . ·10 01 , B .P .I .) 8· 0 7· 5 - - - - - - - - " " tH . m aj or (N o. M . 10 02 , B .P .I .) I I I " " " (N o. M . 10 03 , B .P .I .) - - - - - - - - - - " " t X en oh ys tr is c ra ss id en s (N o. M . 10 04 , B .P .I .) - - - - 12 ·5 12 ·0 13 ·0 12 ·0 " , , ' " (N o. M . 10 05 , B .P .I .) - - 12 ·0 12 ·0 10 ·5 12 ·0 11 . 5 12 ·0 - - " " " " (N o. M . 10 06 , B .P .I .) 11 . 5 10 ·5 - - - - - - - - " " " " (N o. M . 10 07 , B .P .I .) - - - - - - - - - - " " " " (N o. M . 10 08 , H .P .I .) - - - - - - - - - - " " " " (N o. M . 10 10 , B .P .I .) - - - - - - - - - - " " " " (N o. M . 10 09 , B .P .I .) 12 ·0 11 · 0 - - - - - - - - " " - - - - M ea su re m en ts i n m il li m et er s. 1 ) M ea su re m en ts m ad e on f ig ur e in A be l, fr om G au dr y. 2 ) M ea su re m en ts t na de o n fi gu re b y G au dr y. 3 ) M ea su re m en ts m ad e fr om L yd ek ke r (s pe c. G .S .I ., N o. D . 96 ). 4 ) M ea su re m en ts m ad e fr om C ol be rt ( sp ec . A tn er . M us . N o. 1 99 09 ). 5 ) M ea su re m en ts m ad e fr ot n L on nb er g. 6 ) M ea su re m en ts n 1a de f ro n: C ol be rt ( sp ec . A n1 er . M us . N o. 1 46 46 ). co w - - - - - - - ·· - - - - - - -- -- M A N D IB U L A R T E E T H Jl L . B . H ys tr ix a fr ic ae -a us tr al is M in . . . .. 6· 2 M ax . .. . . 8· 0 M ea n . . .. 6· 8 N . . . . .. 8 H . cr is ta ta . . . . . . .. . . 6· 5 H . cr is ta ta . . . . . . . . .. 7· 5 t H . pr im ig en ia 1 ) • • . . . . .. . . 8· 5 t H . pr im ig en ia 2 . . . . .. . . . . - tH . si va le ns is 3 ) . . . . . . .. . . - t H . si va le ns is 4 ) . . .. . . . . . . - t H . /a gr el ii 5 ) . . . . .. . . . . - tH . sp . B .M . 16 59 3 . . .. . . . . - tH ·. sp . B .M . 16 59 4 . . .. . . . . - tH . sp . 6 ) • . . . . . .. . . . . - tH . c f. H . a/ ri ca e- au st ra li s (N o. M . 10 00 , B .P .I .) - " " (N o. M . 10 01 , B .P .I .) - " tH . m aj or (N o. M . 10 02 , B .P .I .) ! - " (N o. M . 10 03 , B .P .I .) I - t X e no hy st ri x cr as si de ns (N o. M . 10 04 , B .P .I .) - " (N o. M . 10 05 , B .P .l .) - " " " (N o. M . 10 06 , B .P .I .) - " " (N o. M . 10 07 , B .P .I .) - " " (N o. M . 10 08 , B .P .I .) - " (N o. M . 10 10 , B .P .I .) 1 - " " (N o. M . 10 09 , B .P .I .) - " I W he re L = G re at es t an te ri or -p os te ri or d ia m et er W he re B = G re at es t li ng ua l- la bi al d ia m et er 5· 0 7· 0 6· 0 6· 5 6· 5 - - - - - - - - - - 8· 0 - - - - - - - - \ T A B L E V II - - - - -- p4 M l L . B . L . B. I .- - - 8· 2 6· 0 7· 5 7· 2 10 ·0 8· 0 9· 5 8· 0 9 ·2 7· 3 8· 7 7 ·4 5 10 1o · 5 I 7· 75 8· 5 7· 0 I 9· 5 8· 0 8· 5 7· 5 12 ·0 11 .0 10 ·5 l J . 0 I - - - - - - 10 ·0 7· 5 - - 9 ·0 7· 5 - - - - - - - - - - - - 10 ·5 8· 5 - - - - - - - - - - 13 ·0 9· 5 - 9· 5 - - - - - - - - - - - - - - - - 1 4 ·0 1 1 2 ·0 14 ·0 1 1 . 0 16 ·0 12 ·0 - - 17 ·0 13 ·5 - - - I - - - W he n te et h ar e b u t sl ig ht ly w or n, t he w id th a t its g re at es t pr ox im al ly i s ta ke n. JM 3 I M 2 L . B . I L . B . I - - - - - -- - - - 8· 3 7 ·0 I 8 ·5 6 ·0 10 ·5 8 ·2 1 1 0 ·0 7· 75 S ou th er n A fr ic a 9· 0 7· 7 8· 9 7· 0 9 I 10 N . A fr ic a 9· 0 7· 75 8· 5 7· 5 10 ·0 8· 0 9· 5 8· 0 W . A fr ic a 11 .0 II · 0 10 ·3 9 ·0 Pi ke n1 1i , G re ec e - - - - 9 ·2 8· 5 - - S iw al ik R an ge , In di a - - - - M og ok , B u n n a I - - - - H on an , C hi na I i I O ld uv ai , K en ya I - - - - I - - - O ld uv ai , K en ya - - - - - M og ok , B ur m a - - - - M ak ap an , T ra ns va al - - - - - " " - - - - " " - - - - " " - - - - " " - - - - " " - - - - " " 14 ·5 11 ·5 - 10 ·5 " " - - - - " " - - - - " '' - - - - · " " Discussion of generic and specific characters: In July, 1946, an isolated rodent incisor fragment of large size was recovered from the dumps of Limeworks quarry 1n the Makapan Valley. As the tooth differs only in its robustness from that of the living porcupine, it \vas assumed that further finds vvould bring to light an Hystrix of considerable size. Later 1naterial collected sho,ved that this large rodent differs in several respects from the genus Hystrix. The number and nature of the specimens available at the moment ljmits the de cription to details of tooth and palate struture only. The upper premolar and fir t two molar · of the Syn-type B figure 28B) sho·v a considerable degree of abrasion. Their pattern approximates to that of II ystrix africae-australis at the same stage of vvear vvith the re-entrant enamel folds, prominent in unworn teeth, eparated off to form islands of enamel ' ithin the tooth crown. Further, tho upper premolar bears no trace of an external enamel fold - a feature which per i t in both H. cristata and H. africae-australis. The occlu al urface which ha the appearance of a sn1oothly rounded square, is defined by a continuous band of enamel. The characteristic outline of the occlusal surface in enohystrix crassidens i related to the ubequal antero-posterior hnd transverse dimension of the crown. In Hystrix , on the other hand the occlusal urface i 1nore nearly rectangular with the antero-posterior clearly exceeding the tran verse dimension. A diagno tic character of the genu liystrix is that the upper cheek teeth are rootless or only semi-rooted. In specimens of the living South African and European porcupine examined the premolar and the molar teeth have papilliform non-d1vergent rootlets which are poorly developed in molars M2 and M3 . .L enoh ystrix ha rooted upper m.olar . The matrix above the base of Ml of specimen Syn-type A figure 28A has been removed exposing lingually a single root elliptical in section, and two smaller cylindrical roots on the labial side. These three root are slightly divergent. The enamel sheath of the tooth ends rather abruptly immediately distal to the junction of the root and the crovvn. From the specimen Syn-type B (figure 28B which vvas broken before fossilization it can be seen that the M2 i rooted. The M3 is demonstrably rooted as the ' "ell preserved alveolus of the M3 clearly shows the impressions of a large Inner root and the anterior outer root (Syn-type A o. 1004, B.P.I.). Apart from its large ize, the lower fourth premolar differs little from that of Hystrix. So far as can be ascertained from the material available the degreo of development of the roots in the lo' er cheek teeth is similar to the condition found in that of Hystrix. An i olated P 4 Jo. M1010) with broken root stumps indicates that \vhen perfect the tooth had roots which were si1nilar in proportion to that in the living South African porcupine. The alveolus of the 11 een in the n1andibular fragment o. M.l008 B.P.I. ee figure 28E shows that the tooth had ·hallow root and that it too might be con idered a being emi-rooted although the roots appear to be more cylindrical in hape and are more divergent than tho '"'e of the genu -. Hystrix. For compari on tooth mea urement of orne living and fossil Hystrix specie J.re o-iven in table VI and VII. 84 GENERAL D fSCUSSION Some fossil bones from the Makapan Lime,tVorks Quarry show large rodent incisor marks predating fossilization. However, the remains of porcupines of a size which 'vould have made these marks are extremely rare. Fragmentary skeletal parts see.m to be absent. In X enohystrix crassidens, too skull and tooth specimens only have been identified. The incisors are far greater in diameter than those which gnawed the bones in the same deposit. As regards knowledge of the type of environment which existed at the time these Hystricomorphs were preserved little can be said as living porcupines have such a wide range of environmental tolerance. Ellern1an, in discussing the H ystricoidea, bases his groupings on external characters only, maintaining that cranial and dental characters alone are unreliable. IIowever, in the case of the Makapan fossils , skull and tooth remains only are available. But it must be borne in mind that though generic and specific identifica- 6ons are made in this paper the characters used are those " hich usually do ~ho'v such differences in ancestral and generalized forms of a species. AcKNOWLEDGEME~Ts My thanks are due to Professor R. A. Dart, Head of the Department of Anatomy Medical School , University of the Witwatersrand, for placing the material at my disposal; Dr. V. FitzSimons, Director of the Transvaal Museum, Pretoria for the use of comparative material; to Drs. H.B.S. Cooke A. T. Hopwood, L. S. B. Leakey, and L. H. Wells for their interest and very helpful criticism; to Messrs. J. w-. and B. J. !(itching for their skilful preparation of the specimEns, and to l\1essrs. B. Grobbelaar and A. Pequeux for other technical assistance. . BIBLIOGRAPHY ...:\bel, 0., ( 1922), Lebensbilder au der Tierwelt der Vorzeit II. In der Buschsteppe von Pikermi in Attika zur unteren Pliozanzeit. G~stav Fischer, Jena, pp. 131'133, fig. 116. Bate, D. M. A., ( 1951), Fossil mammals of Africa No. 2, British Museum of Natural History, London. Broom, R. & Scheepers, G. W. H., (1946), The South African fof::sil apemen: the Austra" lopithecinae. Transvaal Museum, Pretoria. Colbert, E. H., ( 1935), Siwalik mammals in the American Museum of Natural History. 'Trans. Amer. Phil. Soc. , N.S., xxvi. Colbert, E. H., ( 1943), Pleistocene vertebrates collected by the American South East Asiatic Expidition. 'Trans. Amer. Phil. Soc. , xxxii, p. 395. Dietrjch, W. 0., ( 1942), .Altestquartare Saugetiere aus drr Siirll ;chen Sercn~c t ~, Deut ~ ch Ostafrika. Palaeontogra.phica, xciv, Abt. A, pp. 43'133, pls. i.-xii. Ellerman, J. R., ( 1940), The families and genera of living rodents. Vol. I, Rodents other than Muridae. London. Gaudry. A., ( 1R6z .. 1867) Animaux fossile et geologie de l'Attique Parif::. (Text not een. Reference only to atlas). Lydekker, R., (1884), Indian Tertiary and post ·Tertiary Vertebrata. Palaeont. Indica, Ser. x. pt. 3, Rodent , Ruminants and a synopsis of Mammalia. Lonnberg, E., ( 1924), On a new fossil porcupine from Honan, with some remark about the development of the Hy tricidae. Palaeont. S inica Serie~ C. i. · r ? "Simpson, G. G. & Roe, A., (1939), Quantitative Zoology. Macgraw.-Hi11. 85