sae 979 mon bikes) vend RNa ts wre, er pee ate ae Coreen oe Pane ae ae ey OU Pet as ae pd el ON into ’ Paztsirsirtas z z 2 z ee TITUTION NOILOLILSNI~ NVINOSHLIWS S3IY¥VUGIT LIBRARIES SMITHSONIAN ! 5 - ca ani —= o : = wo — i) = a \ 5 a 5 ka : 2 Ne: 3 > 5 = zw A YAS bo] — w - = \ \S - std a = ” Ne ” 2) z m ~ AS = m =z a, <= 7) < n = a8 yuvusit ~t! B RARI ES SMITHSONIAN INSTITUTION NOILALILSNI _NVINOSHLINS, S34 1uVva wi a = = = < = fy, 3 z a : 3 : Ua < oO uw D B 2 w hip ay © z = 2 ie hie | = AN. > = > = > ” =. 7) 2 77) =z NOILNLILSNI_ NVINOSHLINS LIBRARIES SMITHSONIAN ” 2 % wn > ” as mz) Ba w a un lead = oe = ae Sa si e Fe c < oon ow S cc = role = m ro) : a re) a re} = =z a 2 ae z iINVYHNYSIT LIBRARIES SMITHSONIAN INSTITUTION NOILONLILSN!I NVINOSHLIWS 2 5 = = ZO ae z o = oO = o an 4 x 5 tit GF > 5 2 SN 5 ; GM : : ENY = 2 a U, ras - = = m 2 jl m Z2 m Loe : TITUTION NOILNLILSNI_ NVINOSHLINS S3IYWYSIT_LIBRARIES. SMITHSONIAN INSTITUTE : o” = n Zz ae ” = x. Se ee = = =< G = = Xx . = z = ZWNS 5S =z S : at re) = 59 CR = 6 : 2 2 g ; ? ® \.. 8 2 4 2 = z iE SS = 2 2 S = NOs = _LIBRARIES SMITHSONIAN _ INSTITUTION NOILNLILSNI_ NVINOSHLIWS s3aiuva | a ws Z iw 2 &. U; g aay a6 f Ms, 0G PES pa -_ : Eff 3 : : : re) Fs Ye °o = ro) = 2 J = a z =) TITUTION NOILNLILSNI NVINOSHLINS S3I1¥vVuYdgIT LIBRARIES z ee a Se ~_ ise] = ow — (‘fp wo 5 2 5 2 5 ly, Sa = ee * E Lee e 2 = 2 = OS Ki 2 ” m m n°” m = “ = n < 7) 1 iuvudgd rut B RARI ES SMITHSONIAN INSTITUTION NOILNLILSNI _NVINOSHLIWS S3 1uVvEg , , 7, < ¥ = z ‘ = < = yg Wp = o z x = t z 4 be J 3 iG 8 Nw a id a If = ™. = ae Pa ‘AW r Oo 2 1} > MS fe > = 7 i> = a WS - se ra YF fies = : = WY 2 = ee ae Q / w fart = (27) = 77) = ARIES, SMITHSONIAN INSTITUTION NOILOJILSNI_ NVINOSHLINS, Saluvud 11 LIBRAR = z 5 = < = Vi a 4 Pape = = = z= . = s NAG ig 5 Uefia SX 2 E Lv" 2 iS Z, Ee ON i #2 See ee : : ees ALILSNI. NVINOSHLINS S3IYVYGIT LIBRARIES SMITHSONIAN INSTITUTION NOILNLI ae 7) = ” S w” a = o = G zr : & : < 3 : s : sd = or =: re) - 2 "S 3 S ant = ar . 5 wd RARIES SMITHSONIAN, INSTITUTION NOLLALILSNI NVINOSHLINS Sa1uvudi7_LIBRAR z a & z iB 5 : = = : : 2) ES = E = - z a = = = . = hee Z p ae a JNVINOSHLINS, S31YVY E17 LIBRARIES, SMITHSONIAN _INSTITUTION,, NOILALI Sf. Ne. ”n wD» ” WX - @ am ° SA D ¥f. = 2 EWG 2 EWS 2 Z| a z a . 2B = Fa RARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS Saluvugi7_l!BRAF w” > “” S ee n” = te) SS = ae - . 72 4 pl = e ASS = a a . aa 4 Hd -_ a -~ cS oS a ‘ SA a par aif Z a < 2 WAS < c « Gry = ce ear SS oe 4 a Y rat = = - om 5 Be vii a = < WANT * NOLLAL INLILSNI_NVINOSHLIWS ~°3 tyuvyuag Pte LIBRARI ES_ SMITHSONIAN, INSTITUTION = ! = . (@) =— Oo A = (e) ow ‘ = o = (fp wo eS cad ys 5 a 5 , My, ah 5 > VOW E > = F jp E Ps] 3 YA ra Bs] cay Vf C4. wa ke = AS SS wo par De * = a m “aS z m z= m =z 4) ; = w = 77) = IRARIES SMITHSONIAN INSTITUTION NOILNLILSNI_NVINOSHLINS SaluvadI7_LIBRAI t 2 * AN = Pa X + = =a ws (7 [@) ; NY Ae! rT ~W IANS Oo = SOG FN sy z E XS = = = Ss, } as So = 2 = ee ne AMLULSNI_NVINOSHLINS S3IYVYAIT_LIBRARIES SMITHSONIAN _INSTITUTION Coe z = 4 FA ul “n = Ww Z = c = = YG," = eo + < YY fg Ii A < c c “2° GS fe 5 = 3S = * Y fo 5 m 2 a 2 = z = BRARIES SMITHSONIAN” INSTITUTION NOILNLILSNIT NVINOSHLINS Saluvadi7 LIBRA <1HSON, 5 < auSON, S = NS = v 1150 Sao og Ce So oem a Os 8 CSD © Gan ANNALS OF THE _ ANNALE VAN DIE SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM VOLUME 81 BAND 81 ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM VOLUME 81 BAND THE TRUSTEES OF THE DIE TRUSTEES VAN DIE SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM CAPE TOWN KAAPSTAD 1980 @ SET, PRINTED AND BOUND IN THE REPUBLIC OF SOUTH AFRICA BY THE RUSTICA PRESS (PTY.) LTD., WYNBERG, CAPE 336 LIST OF CONTENTS Page CHALONER, W. G., Forey, P. L., GARDINER, B. G., Hitt, A. J. & YOunNG, V. T. Devonian fish and plants from the Bokkeveld Series of South Africa. (Published February 1980.) aa Ba ae aS ae ie a ras ned 27h GRINE, F. E. see TOLLMAN, S. M. Haun, B. D. see TOLLMAN, S. M. HENDEY, Q. B. Agriotherium (Mammalia, Ursidae) from Langebaanweg, South Africa, and relationships of the genus. (Published February 1980.) .. ae as Me 1 KENNEDY, W. J. see KLINGER, H. C. KLEIN, R. G. Environmental and ecological implications of large mammals from Upper Pleistocene and Holocene sites in southern Africa. (Published June 1980.) .. 223 KLINGER, H. C. & KENNEDY, W. J. The Umzamba Formation at its type section, Umzamba Estuary (Pondoland, Transkei), the ammonite content and Se cee distribution. (Published August 1980.) .. Ais at a ; Ko — ae 207, Louw, E. The South African Museum’s Meiring Naude Cruises. Part 10. Station data 1977, 1978, 1979. (Published April 1980.) oF ; a Si TOLLMAN, S. M., GRINE, F. E. & HAHN, B. D. Ontogeny and sexual dimorphism in Aulacephalodon eae Anomodontia). (Published February 1980.) 56 ae = : ai ae ae tel VAN DEN HEEVER, J. A. On the validity of the Therocephalian oe asc ae paisa (Published February 1980.) ; , 111 ; » : * ie ; 7 1 \ a 4 ; ; 4 a } , . ro * A ) aia ; a 2 7 a % in J * a ‘ : Sass er, oe y - os é 4 . ‘ om ; + a ey ere ee wal Piya ghee a ala NS ; ’ 7 ) b Pear 4 i : Ses i oo. ‘ i it r ¥ a ay 2 , : : ‘ y ’ r a » \ rf ? 3 , zs i 1 _- Ne Dar « : , is ‘ ae oe Uy a” F ; , ‘ es mae 5 NEW GENERIC NAMES PROPOSED IN THIS VOLUME Page Barrydalaspis Chaloner, Forey, Gardiner, Hill & Young, 1980 ts ae a el 29 ‘ 'v ; , “ 7 . i ic A ye ‘ ab, ‘ ‘ 7 r ti 4 * , 7 ‘ i 5 “ iy 5 ? 7 : t a ox ft he q a q en . ru * Re 4 z * 7 P ; y 4 \ roa? te yy On Me ns F i Ri ‘ ‘ F ¢ ‘ ; y i = t j iy j 7 ‘ | f BS a f +t pes Ne é ‘ I» J 5 ‘ 4 ‘ ~ OLUME 81 PART 1 FEBRUARY 1980 ISSN 0303-2515 ANNA OF THE SOUTH AFRICAN | MUSEUM | CAPE TOWN INSTRUCTIONS TO AUTHORS |. MATERIAL should be original and not published elsewhere, in whole or in part. 2. 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For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o, scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BuLLouGn, W. S. 1960. Practical invertebrate anatomy. 2nd ed, London: Macmillan. FIscHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FiscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gén. 74: 627-634. Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320. Konn, A. J. 1960b, Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. Ture, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges, Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 81 Band February 1980 Februarie Part 1 Deel AGRIOTHERIUM (MAMMALIA, URSIDAE) FROM LANGEBAANWEG, SOUTH AFRICA, AND RELATIONSHIPS OF THE GENUS By Q. B. HENDEY Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000 Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na gelang van die beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000 OUT OF PRINT/UIT DRUK 1, 2(1-3, 5-8), 3(1-2, 4-5, 8, t.-p.i.), 5(1-3, 5, 7-9), 6(1, t.—p.i.), 711-4), 8, 9(1-2, 7), 10(1-3), 11(1-2, 5, 7, t.—p.i.), 15(4-5), 24(2), 27, 31(1-3), 32(5), 33 Copyright enquiries to the South African Museum Kopieregnavrae aan die Suid-Afrikaanse Museum ISBN 0 908407 84 X Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap AGRIOTHERIUM (MAMMALIA, URSIDAE) FROM LANGEBAANWEG, SOUTH AFRICA, AND RELATIONSHIPS OF THE GENUS By Q. B. HENDEY South African Museum, Cape Town (With 42 figures and 21 tables) LMS. accepted 11 September 1979] ABSTRACT Agriotherium africanum from the latest Miocene/early Pliocene Varswater Formation at Langebaanweg, and other relevant material indicate that Agriotherium was descended from late Miocene Indarctos. Later Ursidae are divided into the subfamilies Ursavinae (Ursavus spp), Agriotheriinae (tribes Agriotheriini and Ailuropodini), and Ursinae (Ursini and Tre- marctini). ‘Ursavus’ depereti and Ailuropoda melanoleuca constitute the Ailuropodini, and Indarctos and Agriotherium the Agriotheriini. The latter consist of a primary European lineage from which branches in Africa, Asia and North America arose. Either all species of Indarctos, or the later ones only, should perhaps be referred to Agriotherium. CONTENTS PAGE Introduction . : ; 1 The Langebaanweg Agriotherium ; 4 Material . : ; ; : : 4 The skull ‘ ; ‘ 9 The postcranial skeleton : : 29 Discussion . ¥ ‘ ; ‘ 51 Palaeoecology : ; : 53 Relationships of griotherium : ; 72 Nomenclature PUM Gees Soll fei as 98 Summary : 4 3 . 104 Acknowledgements saan . mr 106 References . ; ? : ; . 106 INTRODUCTION Agriotherium africanum from the latest Miocene/early Pliocene Varswater Formation exposed in a phosphate mine (‘E’ Quarry) at Langebaanweg, Cape Province (Fig. 1), was the first member of its genus to be recorded in Africa, and the first bear known from sub-Saharan Africa (Hendey 1972: Wolff et al. 1973). Although the first Agriotherium specimens were found in Europe at least as long ago as 1809 (Stehlin 1907), and others have since been collected at scattered localities through much of the Old World and North America, this genus has remained relatively poorly known, being represented for the most part by fragmentary material belonging to few individuals. Initially this also applied to the Langebaanweg representative of the genus, but later more material came to light and it is remarkable that ‘Agriotherium africanum, the most recently described and most remote record of the genus, is apparently also the best represented’ (Hendey 1977: 112). It is in the latter 1 Ann. S. Afr. Mus. 81 (1), 1980: 1-109, 42 figs, 21 tables. 2 ANNALS OF THE SOUTH AFRICAN MUSEUM LOWER COURSE OF RIVER DURING LATE MIOCENE/ EARLY PLIOCENE ATLANTIC — OCEAN 20 «kilometres Fig. 1. The south-western Cape Province. respect that A. africanum is particularly noteworthy. It provides the best indi- cation yet that Agriotherium, like other bears, was characterized by appreciable variation in its dentition and in size, the latter evidently being due to marked sexual dimorphism. A. africanum specimens are recorded from two stratigraphic horizons in ‘E’ Quarry, namely beds 3aS and 3aN of the Pelletal Phosphorite Member (Hendey 1976). The bed 3aS sample, which includes the holotype, comprises only a small part of the total assemblage. Most of the bed 3aN sample was collected in a restricted area (the fossil accumulation at Locality 5—Dingle et al. 1979, fig. 2). This material is from a river channel lag deposit, which was laid down in the lee of a phosphate rock outcrop, while the remainder of the bed 3aN sample was from lateral extensions of the lag deposit. Since this material was probably accumulated during a very restricted period, the bed 3aN AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 3 A. africanum sample may represent remnants of a single population in a temporal as well as a geographical sense. There is no reason to believe that the character diversity observed in this sample can be ascribed to anything but normal intra- specific variation. On the other hand, there are differences between bed 3aS and bed 3aN specimens which apparently reflect evolutionary changes, suggesting that the time interval between deposition of these beds was of sufficient duration for such changes to have occurred. The differences are, however, insufficient to warrant formal nomenclatural recognition. A. africanum thus joins the growing number of species which indicate that deposition of the Varswater Formation took place over an appreciable period in time (see Hendey 1978; Gentry 1980). Langebaanweg is the only recorded locality where Agriotherium is known to occur in more than one stratigraphic horizon. A. africanum is also of interest since it is one of several species now known from sub-Saharan Africa which indicate that towards the end of the Miocene this region had closer faunal links with Eurasia (Hendey 1978). The present study of Agriotherium has suggested that faunal interchange at this time was between Africa and Europe, presumably by way of an Iberian/north-west African connection, rather than, or in addition to, one between Africa and Asia via the Middle East. Apart from A. africanum, the Langebaanweg car- nivores, Plesiogulo monspessulanus and Dinofelis diastemata, also point to such a connection, as have recent studies on other mammals (e.g. Forstén 1978). Until the recent discovery of an agriotheriine at Sahabi in Libya (Boaz et al. 1979), the Langebaanweg Agriotherium was the only African record of the group, and its location at the southern continental extremity was evidence that it had been widespread on this continent. On a more mundane level, A. africanum has an appeal in that it is still South Africa’s only known bear, living or fossil. The material described below is housed in the South African Museum, and catalogue numbers are prefixed SAM-PQ., which identifies the institution and department concerned. This lettering is omitted from the text, and the locality prefix (L) and serial numbers of specimens only are given. Modern comparative material in this museum is distinguished by the prefix ZM. Other institutional abbreviations used in the text are as follows: BMNH-—British Museum (Natural History), London GSI —Geological Survey of India, Calcutta NMB —Naturhistorisches Museum, Basel NMW —Naturhistorisches Museum, Vienna Originals and casts of Agriotherium and related taxa in, or from, the above institutions were examined in the course of the present study. In addition, skulls and skeletons of extant ursids in the South African Museum and the British Museum (Natural History) were studied. Comparative data were other- wise obtained from the publications cited. 4 ANNALS OF THE SOUTH AFRICAN MUSEUM THE LANGEBAANWEG AGRIOTHERIUM MATERIAL The available Agriotherium africanum specimens are listed below according to the horizon from which they were derived, and, where possible, according to sites or areas within ‘E’ Quarry. Pelletal Phosphorite Member, bed 3aS L2045—Left maxillary fragment with P* (holotype) L1868—Left I,, right P°, fragments of right P* and M? L1844, L3141—Left I, and I,, probably of the same individual (Wolff et al. (1973) were incorrect in identifying L1844 as a right I,) L12637—Incomplete left M! (Hendey (1972) was incorrect in identifying this tooth as M?) L2154—Incomplete proximal left ulna The above specimens were described or discussed by Hendey (1972) and Wolff et al. (1973). L12561—Right M, (see Hendey 1972) L12033, L41270—Right I*’s L40031—Left hemimandible fragment and distal left humerus L40030— Right metatarsal II and 2nd phalanx, probably of same individual as L40031 L40002—Left femur and patella, and a thoracic vertebra, of one individual L3433—Left humerus lacking proximal parts L3994— Shaft of left humerus L40040—Distal right humerus L12383—Proximal right ulna L40003—Left radius lacking distal parts L41702—Left scapholunar L41295— Right cuneiform L12503—Right pisiform L20998, L25862— Right metacarpals III L41575— Right metacarpal IV L40043, L40128— Proximal right and left femora L40028, L40029— Proximal right tibiae L41108—Proximal right metatarsal V L10411—Proximal Ist phalanx L42667—2nd phalanx The above material represents at least three individuals, but since the specimens were collected over a wide area and from different levels within bed 3aS, the actual number is certainly much higher. Unless otherwise stated, each catalogue number could represent a different individual, which makes a total of 28. It is not certain that those specimens in this series with numbers above L40000 are all from bed 3aS (see p. 62). AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 5) Pelletal Phosphorite Member, bed 3aN—excavation LBW-E 1975/1 (site— TCWW Prom) L33160—Incomplete left manus, comprising scapholunar, unciform, magnum, trapezoid and metacarpals II to V, the latter lacking distal ends and some of the carpals incomplete L33341, L34188—Incomplete right and left innominates L30205—Left metatarsal III L33557— Proximal left metatarsal V The above material represents at least 2 individuals. Pelletal Phosphorite Member, bed 3aN—excavation LBW-E 1976/1 (site—RP) L33824—Crown of left C. L33825—Left I, L13826—Proximal right radius L33828— Right metacarpal IV L33830—Distal metapodial fragment The above material represents at least one individual. Pelletal Phosphorite Member, bed 3aN—excavation LBW-E 1976/2 (site—IWRP or Locality 5 of Dingle et al. 1979) L45062—Incomplete skull (partly restored) and mandible (restored), lacking left I, right I°, left and right P', right lower incisors, and left P, Associated postcranial bones, including: parts of at least 7 vertebrae (some numbered L49048 and L49115); fragment of distal right humerus; right ulna lacking distal parts; right scapholunar, cuneiform, pisiform, unciform, magnum and trapezoid; proximal right metacarpal III; proximal left humerus; left ulna with distal parts detached and fragmented; left radius lacking part of shaft; right tibia and proximal fibula; right astragalus, calcaneum, navicular and cuboid; right metatarsals II, IV and V; left astragalus; left metatarsal III and proximal metatarsal II; 11 sesamoids, five Ist phalanges, four 2nd phalanges and four 3rd phalanges L45137—Right I?, C lacking root, and P* to M2; left C and P* to M2; some skull fragments L45114—Right hemimandible lacking ascending ramus, incisors, P,; and M3 The above specimens were briefly discussed by Hendey (1977). 146605, L48564, L48577—Left and right maxillary fragments with M?’s and part of left M1 L46573—Crown of right I? L47758/9—Right and left I°, probably of the same individual L48851— Fragment of left I° L46074—Right P* L47698—Left M? 1L46563—Left M., probably of same individual as L45114 L48742— Fragment of mandibular condyle 6 ANNALS OF THE SOUTH AFRICAN MUSEUM L47449, L47701, L47830—Incomplete cervical vertebrae L45063—Incomplete left forelimb, comprising humerus (partly restored), with proximal end detached; ulna and radius (restored); scapholunar, cuneiform, pisiform, unciform, magnum, trapezoid and trapezium; metacarpal, V, and fragments of metacarpals I, III and IV; three sesamoids; three Ist phalanges L46602— Proximal left humerus L48741, L48747—Proximal left radii L47699 /700— Proximal left and right radii of one individual L46076—Right ulna lacking distal end L46134—Left scapholunar, right pisiform and metapodial fragment, probably of one individual L46132, L48021—Left and right scapholunars L47074— Right magnum L45448—Left metacarpal I and distal metapodial fragment, probably of one individual L48432—Right metacarpal IV L49889— Distal left femur L47533, L47910—Left and right astragali L47387— Proximal right metatarsal I L48572 /3— Right metatarsal V and left metatarsal II, probably of one individual L46133, L46216, L48766—Metapodial fragments L47358, L48533, L48730, L49888—1st phalanges L48213, L48230—2nd phalanges The above material represents at least five individuals. Pelletal Phosphorite Member, bed 3aN—surface finds in the vicinity of LBW-E 1976 |2 L47242—Left premaxilla and right M? of one individual. L50636—Right I? L50981—Left M, L42537—Proximal left ulna and distal femur, lacking epiphyses, of one individual L50635—Left scapholunar L50638— Patella L50637—Distal metapodial L41468— Ist phalanx The above material represents at least two individuals. Pelletal Phosphorite Member, bed 3aN—dump 10 sample from deposits immedi- ately north of LBW-E 1976/2 L55012—Left and right P*’s and right M? of one individual L55015, L55016—Right I*’s L55014—Left I* | | AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA if. L55013—Crown of right C L55017— Four anterior premolars of more than one individual L55029— Right metacarpal I L55019— Distal left fibula L55021— Right navicular L55022/28—Seven metapodial fragments L55020—Sesamoid L55030/35—Six Ist phalanges L55036—2nd phalanx L55037/43—Seven 3rd phalanges The above material represents at least two individuals. Pelletal Phosphorite Member, bed 3aN—dump 9 sample from deposits immedi- ately west of LBW-E 1976/2 L50445—Incomplete left hemimandible with C and P, L50453— Fragment of left P* L50458— Right I, L50446— Right M, L50457— Left magnum L50454— Distal metapodial fragment L51592—2nd phalanx L50455/6—Two 3rd phalanges The above material represents at least one Gadiadnall Pelletal Phosphorite Member, bed 3aN—dump 8 sample from deposits immedi- ately south of LBW-E 1976/2 L50003—Incomplete left hemimandible with P, and M, L50004— Incomplete left hemimandible with P, and associated M, and M, L50903— Fragments of right hemimandible with incomplete C L50008—Incomplete right P* L50005/6— Right and left M,’s L50007— Right M, Unnumbered teeth and tooth fragments, including right I’, I° and I,, a canine and two anterior premolars L50843 /54— Eleven vertebra fragments L50857/8—Incomplete left and right scapulae L50834/5/6/8/9—Humerii fragments of at least two individuals L50806, L50816—Proximal left and right radii L50807— Distal right radius fragment L50777— Distal epiphysis of right radius L50763—Left ulna lacking distal end L50764— Left ulna fragment L50840— Proximal ulna fragment 8 ANNALS OF THE SOUTH AFRICAN MUSEUM L50805, L50808— Distal right ulnae L50767— Right scapholunar L50813, L50819—Fragments of right and left scapholunars, probably of one individual L50786, L50795—Left and right cuneiforms L50772, L50774— Right unciforms L50791, L50794— Right magnums L50775—Left metacarpal I L50783, L50788—Left and right metacarpals II L50810—Proximal right metacarpal III L50855/6/9/60—Femora fragments of at least two individuals L50841, L50833— Proximal and distal left tibia, probably of one individual L50842—Proximal right tibia L50769— Distal epiphysis of right tibia L50815, L50828— Distal left and right fibulae L50765/6—Left astragali L50770, L50789— Right astragali L50768— Right calcaneum lacking tuber calcis epiphysis L50773—Left navicular L50778, L50790— Right and left entocuneiforms L50771, L50787—Left and right metatarsals I L50824— Proximal left metatarsal IV L50812, L50829— Proximal right metatarsals V L50809/11/14/17/18/20/22/23 /25/26/30—Eleven metapodial fragments L50802—Sesamoid L50776/80/81 /84/93 /97—Six 1st phalanges L50821/7—Two incomplete Ist phalanges L50785/96/804—Three 2nd phalanges L50782/98/99 /800/801—Five 3rd phalanges L50831—Incomplete 3rd phalanx The above material represents at least four individuals. Pelletal Phosphorite Member, bed 3aN—carbonaceous deposit south of dump 8 area L41404— Fragmented and incomplete skull, with only the left premaxilla and maxilla largely intact, and with right ?P?, and left C (damaged), P*, M1 and M? (see Hendey 1977) L40044— Mandible fragment with associated right radius and fragments of ulna L43126— Fragments of at least six thoracic and lumbar vertebrae. The above material represents at least two individuals. The minimum number of individuals represented in the combined bed 3aN samples is eleven, although the actual number may be far higher. These samples are from a single horizon of river channel and associated deposits, which were exposed over a linear distance of about 200m, with sample areas either contiguous AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 9 or not far removed from one another. Elements of individual skeletons are likely to be represented in more than one sample unit. THE SKULL Apart from L45062, the only other described skull of Agriotherium is one belonging to A. sivalense from the Siwalik Hills in India (Falconer & Cautley 1836; Lydekker 1884). Both skulls are incomplete, and although L45062 is also slightly distorted in parts, it is perhaps the more informative of the two speci- mens. They are similar in overall size, and, in so far as comparisons are possible, they are also similar in morphology. Another skull which is remarkably like that of the Langebaanweg Agrio- therium is of an Indarctos from Florida which was recently described by Wolff (1978). Although the Langebaanweg and Florida skulls undoubtedly do belong to Agriotherium and Indarctos as these genera are presently conceived, the description of skull characters in the latter (Wolff 1978: 2-4) could, with only slight modification, serve as a description of the Langebaanweg specimen. The significance of this will be discussed later (see p. 93). Other described skulls of Indarctos, of which there are two from Samos (Helbing 1932; Thenius 1949, 1959) and one from Spain (Crusafont & Kurtén 1976), are less like the Langebaanweg specimen because they represent species which are more primitive than that from Florida. A summary account of the skull characters of L45062 has been given elsewhere (Hendey 1977), but they are dealt with in more detail here. The skull of A. africanum (Fig. 2) differs in certain aspects from those of all living bears. Its most striking characteristic is its massive size (Table 1), although in some dimensions it is matched, or even surpassed, by skulls of male Kodiak bears (Ursus arctos middendorffi), which are amongst the largest of living bears (Hendey 1977, table 1). A. africanum is otherwise most readily distinguished from living bears by its relatively short and broad snout, while the braincase, which is surmounted by a very high saggital crest, appears small by comparison. The nuchal crest is also very prominent and dorsally projects well behind the occipital condyles. The zygomatic arches are very stout and strongly arched. They resemble those of the giant panda, Ailuropoda melano- leuca, more than any other living ursid, and, amongst extinct ursids in which the zygomata are known, they are closest to those of the Florida Indarctos and North American Arctodus simus (Kurtén 1967). In all these respects the skull of A. africanum is reminiscent of that of the lion, Panthera leo, although the latter is far smaller, and, of course, very different in detail (Fig. 3). The resemblances between the skulls of Agriotherium and P. leo are probably due to their sharing adaptations to a carnivorous way of life. By curious contrast, the A. africanum skull also resembles that of the most herbivorous of all bears, Ailuropoda, as well as that of Indarctos, another supposed herbivore (Wolff 1978). This may be an indication that the actual nature of the diet is less important than the requirement in certain Carnivora 10 Fig. 2. ANNALS OF THE SOUTH AFRICAN MUSEUM Dorsal, lateral and ventral views of the Langebaanweg Agriotherium skull, L45062. AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 11 , TABLE 1 Dimensions of Langebaanweg Agriotherium skull and mandible. SKULL L45062 141404 Basallength . : ete : ; ; ; ‘ : ; 4 c. 381,0 — Condylobasal length . c. 420,0 — Palate length (posterior alveolar ‘margin of Ps to posterior palatine incisure) ; ak # Ay Sets : : : : : ; ‘ c. 165,0 — Zygomatic width . : . i. : ; : , ¢ : c. 305,0 — Rostral width (over _C’ C's). ‘ 3 ; 5 f ‘ ; ; c. 118,0 — Width over M?’s_. ‘ : ite Re : ; F ; ; ; , 135,0 — Interorbital width . : ¢ . : ; : : : : F c. 125,0 — Width over postorbital processes , OPER. ona oP ROT Se Ss c. 150,0 — Occiput width at base of mastoid processes eg see Woy eees cok icbic Bens c. 130,0 — Condylar width veo, i Gt BES PRR aa ey ee c. 90,0 — C-M? length at alveolar margin : : es : 5 iS Dek : 149.0 154,0 P=M* length at alveolar margin . -. . . . . +. . . 83,5 82,5 MANDIBLE 145062 145114 50003 150004 Length (C to condyle) . ; : : ; j 296,0 — = — Height of ascending ramus . ; ; : , 142,0 = = = Transverse diameter of condyle . : : : 71,0 = os == Depth below M, . é : : g : : 69,0 68,5 c. 74,0 —_ Breadth below M, . : 4 : d : : 26,9 26,8 24,7 — Depth at diastema . 3 5 : 3 : 61,0 66,0 65,3 59,4 C—M,; length at alveolar margin ; : : F 174,0 c.190,0 c.190,0 — P,-M; length at alveolar margin . : F : 104,0 =. 117,0 111,0 — for unusually powerful jaw musculature, with consequent similar modification of the masticatory apparatus. There are also some similarities between the skulls of A. africanum and the polar bear, Thalarctos maritimus, which is the most carnivorous of living Ursinae. Like A. africanum, Thalarctos also has a relatively broad snout and a relatively straight dorsal profile in lateral view. These, and other, resemblances between Thalarctos and A. africanum will be discussed again later. Although Thalarctos has a relatively broad snout, it is simply a modifica- tion of the ‘long-faced’ ursine condition, and it is easily distinguished from ‘short-faced’ Tremarctinae, which in turn bear a greater resemblance to Agrio- therium and Indarctos, and, amongst the latter, particularly the Florida specimen. The relatively short and broad tremarctine snouts are, however, ursine-like in having the posterior palatine incisure well posterior of the M?’s (see Kurtén 1966, 1967). By contrast, in A. africanum the posterior ends of the M?’s are in line with this incisure. The situation in Jndarctos is similar (e.g. I. atticus from Samos—Helbing 1932; Thenius 1959), except that in this instance the M?’s project slightly more posteriorly because, unlike these teeth in Agriotherium, they have a talon and are more anteroposteriorly elongated. Ailuropoda is virtually identical to Indarctos in this respect. The situation of the posterior palatine incisure relative to the M?’s sets Agriotherium, Indarctos and Ailuropoda apart from all other later ursids. It is worth noting here that Davis (1964: 50) believed that the lengthened palate in Ursus relative to that in Ailuropoda ‘is an illusion created by the large Fig. 3. ANNALS OF THE SOUTH AFRICAN MUSEUM Dorsal views of skulls. A. Thalarctos. B. Ailuropoda. C. The Langebaanweg Agriotherium. D. Panthera leo. A-B after Gregory (1936, figs 13-14). —— AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 13 teeth of the latter’ and that relative ‘to the anterior end of the braincase, the palate actually extends farther posteriorly in the panda.’ This is true, but Davis makes no mention here of the great shortening of the posterior parts of the Ailuropoda skull, which undoubtedly has an effect on the position of the palate relative to the braincase. The palate of L45062 is otherwise unremarkable, except that it lacks the minor posterior palatine foramen which is present in all other later ursids. In addition, it has a relatively small anterior median palatine foramen. In the latter respect it resembles Indarctos and Ailuropoda, and differs from the Ursinae and Tremarctinae. The minor posterior palatine foramen has apparently been incorporated with the major one, probably as a result of shortening of the palate. A vestige of the minor foramen is indicated on the posterior part of the major foramen by a shelf of bone projecting from the lateral wall of the latter. Earlier in the history of Agriotherium this shelf may well have extended to the medial walls of the major foramen, thus forming a separate minor foramen posteriorly. The infraorbital foramen is situated above the posterior part of P+, as in Indarctos. This foramen is more posteriorly situated in Ursinae and some Tremarctinae, but is more anteriorly situated in Ailuropoda. It is relatively small compared with that in other Ursidae, and in those A. africanum specimens in which it is preserved (L2045, L45062, L41404) it is in the form of a single opening. In other ursids, such as the A. sivalense skull and some tremarctines, there may be more than one opening. The latter condition is probably of no great significance, since multiple infraorbital foramina have been observed in other carnivores (e.g. Proteles cristatus). There is a marked concavity of the maxilla above the reduced anterior premolars, anteroventrally from the infraorbital foramen, which is caused by the expansion of the maxilla over the massive root of the canine and over the roots of the posterior cheek teeth. In anterior view the bulges over the canine roots completely obscure the infraorbital foramina. Amongst the Ursinae a similar tendency was observed in Thalarctos, although in this instance the infraorbital foramina were only partly obscured. Wolff’s (1978: 2) description of the snout of the Florida Jndarctos shows that it is essentially similar to A. africanum. The latter is distinct only in having features such as the bulge over the canine roots exaggerated, apparently because of the larger size of the Langebaanweg species. Wolff unfortunately provided few measurements of the Florida skull, and consequently most dimensions must be estimated from illustrations. The nasal aperture of L45062 does not recede as markedly towards the nasals as it does in the Ursinae, and A. africanum resembles Indarctos and the Tremarctinae in this respect. This feature is evidently due to relative shortening of the snout. Observations on the frontal region of L45062 are omitted, since there was severe fragmentation of this part of the skull and the restoration is not neces- sarily accurate. For example, the orbit appears smaller and the frontals more 14 ANNALS OF THE SOUTH AFRICAN MUSEUM inflated than in the skull of A. sivalense, in which this region is well preserved and therefore certainly accurate. It is nevertheless clear that the post-orbital processes of A. africanum are relatively less prominent than those of Ursinae, and the former is probably more like Jndarctos (I. atticus) and Ailuropoda in this respect. In his description of the zygomatic arches of Ailuropoda, Davis (1964: 47) stated that in dorsal view they ‘form nearly a perfect circle, compared with the triangular outline in Ursus and other carnivores’. In A. africanum, which has exceedingly stout zygomata, their shape approaches that in Ailuropoda. The glenoid fossa and postglenoid process are also large, and in the postglenoid region a wide shelf of bone extends over the external auditory meatus, linking the zygomatic arch with the nuchal crest. The situation in Jndarctos and Ailuro- poda is similar, although in the latter the shelf of bone is much shorter, owing to the anteroposterior compression of the basicranial region in Ailuropoda. The shelf of bone is less well developed in the Ursinae. Amongst fossil specimens the zygomata which most closely resemble those of L45062 belong to the Florida /ndarctos, and once again Wolff’s (1978: 2) description also applies to the Langebaanweg skull. The latter differs in apparently having zygomata of larger size, and in being more strongly arched and thus more Ailuropoda-like. The sagittal crest of L45062 is extremely well developed, reaching a height of at least 50 mm, and, as in the Florida Indarctos (Wolff 1978: 2, 3), it has ‘a very conspicuous cleft between the parietals’. It is, however, slightly deeper (up to 7 mm) and possibly longer (at least 70 mm) than that of the Florida skull. A similar cleft is found in the sagittal crest of Ailuropoda (Davis 1964). According to Wolff (1978: 3), ‘several large, rather irregularly sized and positioned nutrient foramina appear on either side of the parietals just above the temporal shelves near the posterior of the skull in several agriotheriine specimens’, including the skull described by him. There are two such foramina in L41404, one on either side of the sagittal crest, immediately adjacent to it and close to its posterior limit. The same applies to L45062, but since this region of the skull is incomplete, it is possible that there were more than two such foramina. The large size of the zygomata and enormous sagittal crest, together with various rugosities for muscle attachments similar to those of the Florida Jndarc- tos (Wolff 1978), indicate that the masticatory musculature of A. africanum was exceptionally powerful, and in keeping with the massive canines and posterior cheek teeth of this species. The basicranial regions of L45062, L41404, and L45137 are unfortunately poorly preserved and incomplete, although sufficient remains to show that they are ursid-like. Amongst the living Ursidae there are similarities to both Ursinae and Ailuropoda, while of described fossil specimens they closely resemble, and in several respects are indistinguishable from, the Samos J. atticus (Thenius 1949, 1959) and the Florida Indarctos (Wolff 1978). AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 1S As in Ailuropoda and Indarctos, the foramen rotundum and orbital fissure form a single opening in both L45062 and L41404. There is a well-developed horizontal division separating them a short distance posterior to the common opening as in the Florida Jndarctos. In this respect Ailuropoda differs in that there is at most ‘a paper-thin partition separating them’ (Davis 1964: 49). A. africanum is also like Ailuropoda and Indarctos, and different from most, if not all other Ursidae, in lacking an alisphenoid canal. In addition, A. afri- canum, Indarctos, and Ailuropoda are similar in that the medial edge of the glenoid fossa is closer to the foramen rotundum than is the case in the Ursinae, evidently because of the greater transverse length of this fossa in these three taxa. The same applies in the case of the foramen ovale, which is preserved in L45062, L41404, and L45137. This foramen opens opposite the posterior wall of the glenoid fossa in these specimens, and they are similar to the Samos I. atticus, and probably also the Florida Indarctos, in this respect. In the Ursinae the foramen ovale is more posteriorly situated, while in Ailuropoda it is further forward. The postglenoid foramen in A. africanum is situated between the external auditory meatus and the medial edge of the postglenoid process, but is closer to the latter than is the case with living ursids. Amongst the latter, Ailuropoda is the most distinct, since the postglenoid foramen is ‘more laterally situated than in Ursus’ (Davis 1964: 52). The situation in the Florida Jndarctos and I. atticus is similar to that in A. africanum, although the specimen described by Thenius (1949) differs in having a double opening. The external auditory meatus is incomplete in L45062, but it is evidently similar in position and orientation to that in the Florida /ndarctos. The anterior (squamosal) part of the mastoid process of L41404 is complete. It had not yet fused to the posterior (periotic) portion of this process, nor to the bone which caps this process. In size and orientation it is apparently similar to the mastoid process of the Florida Jndarctos, and appears to have been more laterally directed than that of later ursids. Both the mastoid and paroccipital processes of L45062 are lost. The relative position of the bases of these processes differs from that in later ursids in that the base of the mastoid process is only slightly more laterally situated than that of the paroccipital process. The mastoid process of L45062 apparently differed from that of L41404 in being orientated ventrally. Part of the lateral walls of the stylomastoid foramina, and the posterior margins of the posterior lacerate foramina, are preserved in L45062. Their positions relative to one another are as in the Ursinae and Jndarctos. The same applies to the hypoglossal foramina and the anterior lacerate foramina. Little of the bulla of L45062 remains. The posterior parts appear to have been more inflated than in Ursus arctos, and in this respect the bulla may have been more Jndarctos-like. In overall size the bullae of A. africanum and a European U. arctos (ZM39056) were apparently similar, which is surprising in 16 ANNALS OF THE SOUTH AFRICAN MUSEUM view of the very much larger size of the A. africanum skull. In this respect A, africanum is also Indarctos-like. Ailuropoda is remarkable in that ‘externally there is no indication of a bulla’ (Davis 1964: 319, 320). . This is but one of the peculiarities of the Ailuropoda skull, which has been ‘profoundly modified by the demands of mastication’ (Davis 1964: 46). In less specialized ancestral forms the basicranial and other regions of the skull may well have resembled their counterparts in Indarctos and Agriotherium more closely than those in the Ursinae. The absence of the alisphenoid canal in Indarctos, Agriotherium, and Ailuropoda is probably the single most important basicranial character which distinguishes them from the Ursinae. This canal is present in the Canidae and all other Ursidae (Wolff 1978), including the Amphicyoninae (Ginsburg 1977; Hunt 1977) and, presumably, the Hemicyoninae. The occipital region of A. africanum is known from L45062 (ventral parts) and L41404 (dorsal part and ventral part adjacent to, and including, the squa- mosal part of the mastoid process). In posterior view it is relatively narrow and steeply arched, rather like that of U. arctos and Indarctos. In L41404, which represents a young adult with some sutures unfused, there is an indication of a constriction above the mastoid process. In L45062, an older individual, the nuchal crest is strongly developed, and instead of passing ventrally directly on to the lateral side of the mastoid process, as in most living ursids, it is linked anteriorly with the shelf of bone which projects posteriorly from the zygomatic arch. The base of the mastoid process in L45062 is actually recessed beneath this shelf of bone. In most of the available ursine comparative specimens, the nuchal crest merges ventrally into a ridge of bone on the mastoid process, which is more laterally situated than the shelf extending posteriorly from the zygomatic arch. Only in Thalarctos does the arrangement approach that in A. africanum. It is not clear from illustrations of J. atticus (Thenius 1959) and the Florida Indarctos (Wolff 1978) which arrangement characterizes this genus. The mandible is massive, its size being in keeping with that of the skull (Figs 4-5). It is similar in shape to that of J. atticus and U. arctos, and is dis- tinguished principally by the presence of a premasseteric fossa. This fossa is deep in older individuals (e.g. L45062), but much less pronounced in immature adults (e.g. L45114) (Fig. 33). The only living bear with a premasseteric fossa is the South American spectacled bear, Tremarctos ornatus, while it is otherwise known amongst Ursidae in extinct Tremarctinae, Hemicyoninae and other species of Agriotherium. In tremarctines the masseteric and premasseteric fossae are separated by a prominent ridge of bone, and the latter fossa is deep, exten- sive and more or less circular in shape, with its limits clearly defined (e.g. see Kurtén 1966, plates 8-9). In hemicyonines the ridge separating the two fossae is not as prominent, the premasseteric fossa is less extensive in a vertical sense, but more elongated anteroposteriorly, with the anterior end gradually merging with the buccal surface of the mandible beneath M, (e.g. see Frick 1926, figs 2-3, 12). AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 17 Ee ee Kk Mt A A Fig. 4. Buccal and dorsal views of Langebaanweg Agriotherium hemimandible, L45062. The premasseteric fossa in Agriotherium is closer to the hemicyonine type, and may even be virtually indistinguishable from it (e.g. the A. insigne specimen figured by Viret (1939, fig. 6)). There are, however, Agriotherium specimens in which this fossa does not extend as far anteriorly, terminating instead beneath M, (e.g. the A. schneideri specimen figured by Frick (1926, fig. 36)). There is at least one specimen assigned to Agriotherium in which this fossa is absent (i.e. the A. palaeindicum specimen, GSI-D8, discussed by Pilgrim (1932)). The latter is, however, one of the problematical intermediates between Jndarctos and Agriotherium, which will be discussed in a later section of this report. The premasseteric fossae in A. africanum specimens are generally similar 18 ANNALS OF THE SOUTH AFRICAN MUSEUM vi ov ce eae Fig. 5. Buccal and dorsal views of Langebaanweg Agriotherium hemimandible, L45114. to that in the A. schneideri specimen mentioned above. In the four hemi- mandibles belonging to three individuals in which the entire premasseteric fossa is preserved, the anterior termination is beneath M,. These specimens apparently differ from hemicyonines and other Agriotherium in having a less distinct ridge of bone separating the masseteric and premasseteric fossae. The mandible of A. africanum is otherwise distinguished only by the presence of a distinct ‘chin’ in the symphyseal region, which contrasts with the receding jaw-line in other ursids. The ‘chin’ is formed by an anteroventral expansion of the symphysis, which enlarges the area of the symphysis and presumably strengthened the connection between the two halves of the mandible. With the exception of the nondescript and relatively unimportant [, P, P? and P3, all the teeth of A. africanum are known from at least one complete specimen still in position in a jaw. An incomplete I’ is represented in L45062, while P! and P? are tentatively identified on the basis of isolated specimens. Only P, and P, have not been identified, although they could be represented amongst the six unidentified anterior premolars available. The best represented of the teeth are the larger posterior cheek teeth (P{-M3), which are, fortunately, the most informative in the dentition of Agriotherium (Figs 2, 6-7; Tables 2-3). These teeth are represented by between six and ten specimens belonging to between four and seven individuals. 19 AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA ‘VOVIVT ‘uowsely JNOUS JO SMAIA [eIIUDA pue [Loong “| ‘ZIOSST “uewseIy AXe]PIXCU Wntayjolispy SaMueeqasueyT JO MOIA [eIUSA “W ‘9 ‘BLT A I om 20 ANNALS OF THE SOUTH AFRICAN MUSEUM nl eee PUTA ALL 6 117 118 119 210 211 ; yu 112 1 Fig. 7. Ventral view of Langebaanweg Agriotherium maxillary fragment, L45137. mM 3 1 Mm 5 1 UU 4 1 Apart from their large size, the incisors of A. africanum are unremarkable and are little different from those of living Ursinae. Dimensions of some incisors were given elsewhere (Hendey 1977, tables 2, 5). The I? is distinct in having two lingual cusps projecting from the V-shaped cingulum posterior to the principal (spatulate) cusp. The lateral accessory cusp is the smaller and the more anteriorly situated, being little more than a small projection from the cingulum. The other accessory cusp is much larger, covering much of the lower part of the lingual surface, although it is still much smaller than the principal cusp (Fig. 8). In older individuals (e.g. L45062) the three cusps develop horizontal wear facets in a single plane. The [’ is similar to I*, but smaller, while the I°, which is the largest of the incisors, is morphologically similar to its counterpart in Ursinae. The lower incisors are similar to those of Ursus arctos and other ursines, and are distinguished only by their larger size. 1 Zz ma Lf £ mu if I sil + mint 3 Fig. 8. Medial, posterior and lateral views of Lange- baanweg Agriotherium I*, unnumbered Dump 8 specimen. 21 AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 6 6 rg ste -6 7 —6S7 08st =: LOE GG 0G 6EZ 6ST LLC ae Oe SiGe COS G8. 20 1e CGC. sar be 160 > a 0°8s ae 8°€6 ra 9'Sh a OLY cae 0°87 SO80SsT} £9L0ST ‘OUl[N WNLaYyjolisp SoMueeqosuLyT JO SUOISUSWIIG 1wsrA = Ca) * ssod0i1d projAjs JO “wieIp y104S ssos0i1d projAjs Jo ‘wieip 3u0T * —_ ssed0i1d prouo.109 je wep ysod—juy * —- YO}OU JeUNIIWOs ye ‘WeIpP ‘ysod—juYy ssoooid jeouosue je ‘wieip jsod—juy ; * yoJOU IeUN]IWes Jo “WRId p * yojOU [eIper ye ssoooid plouol0d jo ‘“WieIp “AsueIy, ssosoid jeouooue JO “WIRIpP “ASULIT, : } ; Yj3Ue] [[eI0AO 36 ANNALS OF THE SOUTH AFRICAN MUSEUM Davis 1964: fig. 50), but is less so in A. africanum, especially female specimens such as L45063. Once again, the functional significance of these differences is not known. The semilunar notch differs from those of ursines and Ailuropoda in some respects. In lateral view it is almost perfectly semicircular, with the inner borders of the notch not divergent at their extremities as in ursines, and not showing the beginnings of convergence as in Ailuropoda (see Davis 1964, figs 50-51). In anterior view the dorsal part is nearly parallel-sided, rather than sharply tapering as in ursines. The anconeal process of A. africanum is therefore relatively broad. This, and the relatively narrow olecranon fossa of the humerus, are probably directly related features. When the humerus and ulna of A. afri- canum are articulated and fully extended the olecranon fossa is nearly com- pletely filled by the anconeal process, and little lateral movement of the ulna is possible. In the ursines only about half the olecranon fossa is filled and appreci- able lateral movement is possible. Much the same evidently applies in the case of Ailuropoda, which has an even wider olecranon fossa than ursines, while Davis (1964: 96) noted that there was no protection against lateral shifting of the elbow joint. The significance of these contrasting situations is not known. The shaft of the’ A. africanum ulna is very stout, and although the areas for attachment of the brachialis tendon and interosseous ligament vary both in size and form, they are very pronounced features. Distally the radial articular facet is relatively, and sometimes absolutely, smaller than its counterpart in ursines. Medially between this facet and the styloid process is a deep and almost circular depression, which is directly in line with the prominent ridge on the shaft separating the areas of insertion of the pronator quadratus and the distal part of the flexor digitorum profundus 5 muscles. This depression is absent in available ursine specimens, and apparently also in Ailuropoda (Davis 1964, fig. 50). Its significance is not known. The radius of A. africanum is represented by several proximal and distal fragments, but only one that is complete (L40044), while another has been restored (L45063) (Fig. 17, Table 6). The latter belongs to a female and L40044 to a male. These two specimens differ only in size. They are typically ursid in their characteristics, the proximal end being particularly distinctive, and L40044 is very similar in its proportions to the J. atticus specimen described by Pilgrim (1931). The latter specimen is, however, distinct in having a broad groove on the anterior surface towards the distal end. Pilgrim (1931: 27) thought this noteworthy since ‘a similar structure exists in a corresponding position in the radius referred by Falconer to Agriotherium . . . sivalense’. According to Pilgrim it is also present in Amphicyon, although it is evidently not characteristic of all amphicyonines (see Ginsburg 1977, fig. 25). Since the radius of A. africanum is otherwise unremarkable, it is not described in detail. With one exception, all the carpal bones characteristically present in Ursidae are represented by at least one, and as many as nine, complete speci- mens (Fig. 18, Table 7). The exception is the radial sesamoid, which was evi- AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 37 LLU C fi ; \! | + I 2 1 iafe oon (ieee LF t mn mm wu 6 L hulu t lat Fig. 17. A-B. Medial views of Langebaanweg Agriotherium radii. A. 145063. B. L40044. C. Proximal view of L40044. 38 ANNALS OF THE SOUTH AFRICAN MUSEUM Hui 1 sil tui i = Il 1 Z | nu 1 wut I mn Z th mn - Bs > =) Ul 1 8 mn 1 il y ih 1 S| ma nn ii qn 8 Il ao n i-) 14 il { mn fl il | i mn da H I Fig. 18. Carpal bones of Langebaanweg Agriotherium. A—G. L45063. H. 146134. I. L50767 (reversed). A. Proximal and dorsal views of scapholunar. B. Medial view of cuneiform. C. Medial and dorsal views of unciform. D. Anterior view of pisiform. E. Proximal, dorsal and lateral views of magnum. F. Proximal view of trapezoid. G. Lateral view of trapezium. H.-I. Specimens illustrating size range of scapholunars. 39 AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA e90SV'1 S SET ODE ¥H'8Z THE O6E O'8E 661 90% 677 6€€ OLE SOE y8rp OS ZS £00071 C90SV1 SWNOAISIT vLy L’vEé 918081 e Si8S a es OOS Oe Epes 9c an ne ViL SiC MECC sacOG) OCC Ge YC GIES cs CPEeaS St OSES 686 OTP 6 eV L8G SNNNOVW ‘WIRIP JOYS 4SIq ‘WIRIP SUC] “}SIG ‘WIRIP J1OYS “XOIg ‘WIRIP SUC] *XOIg YSU] [[V10AO SUVYNOATOHdVOS “s[edivo wnideyjolusp SsoMueeqosury] JO SUOISUSWIG L a1avL, L‘€9 == — S‘OL = BS Le Str 9 = =a 0°6r sa 53 mag rsp €‘0s 0'6r 9 L‘0S 8°7S (als y'9S ‘OE S‘LE ‘BE ‘9 oI = 8‘€V Ure O'SHE eS =e o'r0r =a ae: ae €90SVT 9C8EET 669471 vroOVI 790SPT LPL8VI 9080571 ‘UIpel WNayjolisp Somueeqosuey] JO SUOISUSWIG 9 alav “WILIP [PIJUSAOSIOP “xe ‘WIPIp “AsUPI} ‘WIP ‘sod—jue "WUPIP [e1}USAOSIOp “WIeIp “AsuUPIy Jo0vj [eIpel JO ‘WIeIp -jsod—jue : : “WIeIp ysod—"jue pus [eIsIp “weip ‘Asued puod [eISIp “weIp jsod—jue pus jewixoid “wep ‘Asuen puo jeuxoid “werp “jsod—yue “Xe “XPJA “XP “XE “XP “XP “Xe “Xe “XP “XE Yysug] [[eI9AO 40 ANNALS OF THE SOUTH AFRICAN MUSEUM dently absent in A. africanum, since the scapholunar lacks the radial sesamoid facet. In this respect A. africanum is very different from Ailuropoda, in which the radial sesamoid is better developed than in any other arctoid carnivore (Davis 1964: 99, 100). A small radial sesamoid is apparently usually, or always, present in Ursinae, and is larger in Tremarctinae. Otherwise the carpals of A. africanum are essentially similar to their counterparts in Ursinae. The largest carpal, the scapholunar, is also the best represented. As with other A. africanum bones, the scapholunars exhibit appreciable size differences, the smallest (50767) being only about two-thirds the size of the largest (L46134) (Fig. 18H-I). This is a slightly greater size difference than that observed in the scapholunars of North American Tremarctos floridanus, an ursid of similar overall size (Kurtén 1966, table 19), although in both instances sample sizes | are small. There are no significant morphological differences between the ~ smallest and largest scapholunars, and they differ in only minor respects from those of ursines. In the latter the cuneiform facet tends to be clearly distinct from the unciform facet, but in A. africanum they merge and are distinguishable only because the cuneiform facet is flattened and the unciform facet is concave. In some specimens (e.g. L48021) the proximal (radial) articular surface is in contact anteriorly with the articular facet of the trapezium, and in this respect A. africanum is similar to Ailuropoda (see Davis 1964: 99). The proximal articular surface of the A. africanum scapholunar is also Ailuropoda-like, and different from Ursinae, in lacking the lateral depression which receives ‘the saddle on the distal end of the radius’ (Davis 1964: 99). The cuneiform of A. africanum is morphologically similar to that of U. arctos, except that the scapholunar facet is more elongated. In addition, this bone is relatively more flattened than that of U. arctos. Such minor differences in morphology and proportions also distinguish other A. africanum carpals from their U. arctos counterparts. For example, in the A. africanum unciform, the magnum and scapholunar facets are not confluent as in U. arctos, while the pisiform is a considerably stouter bone. At least some of the distinctive features in A. africanum carpals may be due to the large size of the species, but their possible significance in other respects was not investigated. Of the metacarpals of A. africanum only the second is not represented by -a complete specimen, although there is one which lacks only the distal end (L33160). All the metacarpals are similar to their counterparts in ursines in terms of morphology, but are relatively more massive (Fig. 19, Table 8). In spite of this, they are not necessarily much longer than those of the available U. arctos specimen. One of the complete metacarpals I (L45448) is in fact similar in length to that of the U. arctos comparative specimen, although it has a much stouter proximal end and shaft. The A. africanum metacarpals appear to be readily distinguishable from those of Ailuropoda, which are ‘short and stout, relatively considerably shorter than in [other bears] of comparable size’ (Davis 1964: 100). AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 41 > ) So “s[edivovjoul WnlayjoUsp SoeMmueeqosueryT JO SUOIsSUOWIC g ATEV IL, Dene Iaae ‘4SIP “UWIRIpP “ASURI] “xeyAT * worjelnoy1e ‘\SIp “WIeIp “jsod—jue “xey * puo -xoid “uIRIp “AsueI) “xe ies) poids “WeIp ‘jsod—jue -xepy YI3Ug] [[PI0AO Teorey THe 4SIp “WeIp “AsuRI} “xe/[ * uone[nonie *1SIP “WeIp jsod—jue -xey * pus -xoid “‘WieIp ‘Asuei} “xepy ee) Sorel” ‘WeIp “jsod—jue -xey : Yysu9gy [[e10AO AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 43 No complete innominate of A. africanum is known, the best specimen (L33341) comprising only the posterior part of the ilium and the acetabular region. The latter is similar to that of the U. arctos comparative specimen, and is distinguished only by its slightly larger size. There is, however, a greater dorsoventral constriction of the ilium anterior to the acetabulum, while the dorsoventral diameter of the posterior parts of the sacroiliac articulation is actually less than in the comparative specimen. A. africanum is more like Ailuropoda in this respect (see Davis 1964, fig. 59). The iliopectineal eminence is less prominent than in U. arctos, and in this respect it is also Ailuropoda-like. No other important features are observable in L33341, but the fact that it is more like Ailuropoda than U. arctos in at least two respects may be significant. The pelvis of the former is very different from those of other bears (Davis 1964: 113), and it is possible that that of A. africanum was equally distinctive. The femur of A. africanum is represented by several fragmentary specimens, and a complete one (L40002) belonging to an aged and arthritic female (Fig. 20, Table 9). The latter specimen is considerably smaller and more slender than corresponding parts of male specimens, but is morphologically similar to them. It differs from the femur of the U. arctos comparative specimen in being slightly longer and relatively more slender. The head of the femur is hemispherical, as in other ursids, while the neck is distinct and slightly longer than in U. arctos, but similar to that of Asian Euarctos and Ailuropoda. The greater trochanter is lower than the head and it is similar to the U. arctos femur in this respect. The gluteal tuberosity is very prominent in L40002 and terminates well below the level of the lesser trochanter, which is also prominent. The area of attachment of the quadratus femoris muscle between the lesser trochanter and gluteal tuberosity is well marked, as is the area of attachment of the adductor muscle, which extends about three- quarters the length of the very straight shaft. The distal end of the femur is essentially similar to that of U. arctos. The femur of A. africanum is very different from that identified with A. sivalense by Lydekker (1884, pl. 29, fig. 1). This very curious specimen may be pathological. The tibia of A. africanum, of which only one complete specimen (L45062) is known, is also very variable in size (Fig. 21, Table 9). As with the femur, L40002, the tibia is longer than that of the U. arctos comparative specimen, but in this instance it is also much stouter, evidently because L45062 belongs to a male. The proximal end of the tibia differs most markedly from those of ursines in having the tibial tuberosity and crest more prominent. This applies even in the case of specimens belonging to females (e.g. L50842). In addition, the proximal articular facets are relatively longer anteroposteriorly. In Ailuropoda and ursines the lateral edge of the tibia shaft viewed anteriorly is bowed, with the tibial crest paralleling the proximal curvature. This, together with prominent lateral projections at the proximal and distal ends, ‘increases the interosseous space between the tibia and fibula, and the total width across Fig. isabel a I iat sli Ss So ANNALS OF THE SOUTH AFRICAN MUSEUM A-B. Anterior and posterior views of Langebaanweg Agriotherium femora. A. L40002. B. L40128. C. Medial view of L40002. AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA i 5 =z & 5 Fig. 21. Anterior and medial views of Langebaanweg Agriotherium tibia, L45062. 45 ANNALS OF THE SOUTH AFRICAN MUSEUM 46 pUo [eISIp “UIP “AsUeN “xe a, = = OS 17S * pus [eISIp “werp ysod—jue ‘xeyy ZT 016 ‘2 0°66 °2 LOOT S801 * puso jeurxoid “ureip *Asueq} “xeyy pes L'S8 €‘L8 1°26 — pus jewrxoid “weip ysod—jue ‘xe a ae a = = peoy jo ‘weip ysod—jue *xeyy — ae x O'8LE = : ; Sug] [[e19AO Cy80ST s8c00VI 6c00VI c90SVI § <€E80ST /T80sT 6886V1 cOO0OVI 8zcl0rT AVIAL], VUOWAT “OEIQN PUL BIOWOJ wniayjoUsp Somueeqosuey JO suoIsuOWIG ACGCKAD AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 47 = mn < Da INIU UH i ni ul INH ht I inn C ni ul a D ll ng A nl Il Itt I Fig. 22. Tarsal bones of Langebaanweg Agriotherium, 145062. A. Proximal, lateral, ventral and distal views of astragalus. B. Lateral and dorsal views of calcaneum. C. Dorsal view of cuboid. D. Proximal view of navicular. 48 ANNALS OF THE SOUTH AFRICAN MUSEUM the leg’ (Davis 1964: 115). The tibia of A. africanum differs in having the proxi- mal and distal lateral projections less prominent, and the shaft less bowed, which gives it a much straighter and more bilaterally symmetrical appearance in anterior view. The distal articular facet, like the proximal ones, has a rela- tively greater anteroposterior diameter than in ursines. The same applies to the proximal and distal fibula facets, while the fibula itself is much like those of ursines. All the tarsal bones of A. africanum, except the mesocuneiform, ecto- cuneiform and tibial sesamoid, are known from at least one, and as many as eight specimens (Fig. 22, Table 10). The tibial sesamoid, like the radial sesamoid, was probably absent in this species. As with the carpals, the tarsals are essentially similar to their counterparts in Ursinae, and they, too, exhibit appreciable sexual dimorphism. The astragalus is the best represented tarsal bone. It is distinguished from those of ursines principally by a longer neck. The available ursine astragali have a variably developed lip of bone projecting posteriorly from the base of the tibial facet, and which is most prominent medially. This lip of bone is absent in A. africanum, although in the specimens L45062 and L47533 there is a ventrally projecting lip of bone in this position. Since it effectively inhibits movement between astragalus and calcaneum, it is probably an abnormality caused by osteo-arthritis. The calcaneum of A. africanum is also ursine-like, differing principally in having a relatively shorter and much stouter tuber calcis. The remaining tarsal bones differ in only minor respects from their counter- parts in U. arctos. All the metatarsals of A. africanum are represented by at least one complete specimen (Fig. 23, Table 11). They are in general much stouter but only slightly longer than their counterparts in U. arctos. An exception is a small metatarsal I (L50771), evidently that of a female, which is of similar length and which is TABLE 10 Dimensions of Langebaanweg Agriotherium tarsals. ASTRAGALI 145062 (R) 150765 147533 L50766 150770 147910 61,9 58,0 63,2 62,1 59,8 c. 56,5 51,2 43,7 Max. ant.—post. diam. . Max. transy. diam. Transv. diam. of tibial facet 44,9 c. 46,0 43,1 Shy 33,5 29,5 Transv. diam. of navicular facet 43,5 44,5 43,0 37,9 32,8 27,0 Max. dorsoventral diam. 40,4 — 37,2 35,3 28,8 27:5 CALCANEUM NAVICULARS 145062 L50773 145062 | L45062 48,7 43,3 38,7 38,2 20,5 17,6 . ant._post. diam. . Max. transy. diam. Max. dorsoventral diam. ; . transy. diam. of tuber calcis (R) = Right AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 49 Fig. 23. Proximal, anterior and lateral or medial views of Langebaanweg Agriotherium metatarsals. A. I, L50787. B-E. II-V, L45062. ANNALS OF THE SOUTH AFRICAN MUSEUM 50 a = TET CET : er ae DONeNoIe [e\sIp “WeIp “ASURI} “Xe = = OI~ L‘6L = A - = wvonrnonse [eisip “werp ysod—jue “xe S*LZ ITE LOE OLZ €ZE : : * puso yecapxoid “UIRIP "ASUBI} “XE ELT 9°67 SOE Ile VE ‘ Sei abe pice al “WieIp “jsod—jue “xe = = S‘SII S‘6IT = é Pa ae al : W13U9y [[eJeAO BOIIy1 6c80ST 790SVE CLS8VI ZI80ST coOSVI = HBOS Vz TC * WONEfNose [e}SIp “UIEIP “ASUI} “Xe y6l S07 uorefnonse [eysip “werp “jsod—yue “xe SC (44 ates pus jeunxoid “urerp “Asue] “XE Ive S‘9E BES [Eanes ‘weIp ‘jsod—jue “xe 6 LOT 9OIT : ysus] [[e19AO SsOcOeT 90ST Til C90SV1 uu ELS8Vv1 ILLOSI 6 L8ELVI = L8 LOST I “S[esIe}e}OU wnidayjousy Zomueegosuey jo suorlsuauid TI S1av.L AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 51 more slender than that of the U. arctos comparative specimen. Morphologically there are no significant differences between the metatarsals of A. africanum and U. arctos. The patella, sesamoids and phalanges of A. africanum are also U. arctos- like, although the Ist and 2nd phalanges of the former are relatively much shorter and stouter (Fig. 24). | Mn hn Fig. 24. Dorsal and lateral or medial views of Langebaanweg Agriotherium phalanges, L45062, possibly of one digit. il DISCUSSION The overall impression gained from the study of the skull and postcranial skeleton of A. africanum is of a large and heavily built animal which was unmis- takably bear-like in its appearance (Fig. 25). It was undoubtedly plantigrade since its postcranial bones exhibit most of the characteristics found by Ginsburg (1961) to be indicative of this condition. The appreciable size variation observed is to be expected of a bear of such large proportions, since marked sexual dimorphism is characteristic of these animals (Kurtén 1955, 1966, 1967). Being typical of large ursids in this respect, it is of interest to compare A. africanum with better known species which belong in this category. They include Ursus spelaeus, Tremarctos floridanus, and Arctodus simus. Of these species A. africanum probably resembled the North American late Pleistocene tremarctine, Arctodus simus, most closely. For example Kurtén (1967: 49, fig. 28) found that the skull of this short-faced bear ‘shows a remark- able convergence with the great cats’, which is also a feature of the skull of A. africanum (see p. 9 and Fig. 3). In overall size the male skull of the latter (L45062) is intermediate between those of female and male A. simus (see Table 1 herein and Kurtén 1967, table 5). In addition, the postcranial bones of A. afri- canum are similar in proportions to those of A. simus, although they differ in 52 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 25. Reconstruction of Agriotherium, adapted from one of North American Arctodus simus by J. Matternes Gin Guthrie 1972). some morphological details. In actual size the bones of A. africanum males compare closely with those of A. simus females (e.g. the Potter Creek Cave sample—see Tables 4-11 herein and Kurtén 1967, tables 10-25). Unfortunately the skeleton of A. africanum is less well represented than that of A. simus and consequently cannot be analysed in as much detail. In view of the marked individual size variation in A. africanum it may be misleading to combine skeletal elements of different individuals in metric analyses. Never- theless, this was done in several instances, one of which is presented here. The isolated metatarsal I, L50787, which is evidently that of a male, was combined with the metatarsals II to V of L45062, also a male, in order to com- pare their relative lengths with the metatarsals of other ursids (Table 12). The similarity to the Potter Creek Cave A. simus sample is striking. On the other hand, the calcaneum length expressed as a percentage of the longest metatarsal length of L45062 is 86,6, a figure which compares closely with the 86,5 of an U. arctos sample, and which is considerably lower than the figures for extinct tremarctines and U. spelaeus, which are over 100 (Kurtén 1966, table 36; 1967, table 27). In spite of such deviations from the Arctodus pattern, A. africanum is like this genus and different from other later ursids in having relatively long legs. This characteristic, together with its specialized skull and dentition, is a highly significant departure from the typical ursid condition. Kurtén (1967: 50) interpreted A. simus as ‘a predominantly carnivorous form’, which may have AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 53 TABLE 12 Relative lengths of the metatarsals of some bears expressed as a percentage of the length of metatarsal V. I I Tl IV Vv Agriotherium africanum . 71 82 96 101 £100 (L50787 + L45062) Arctodus simus* . ; (Potter Creek Cave) Tremarctos floridanus* 70 82 95 99 100 69 80 91 103 100 (males) Ursus spelaeus' . 63 78 88 98 100 (Salzofen) Ursus arctos* 65 79 87 96 100 (Recent) 1 Kurtén 1967, table 26. ‘preyed on large contemporary herbivores’, and although not ‘truly cursorial it may have been capable of bursts of speed exceeding those of U. arctos’. These conclusions presumably apply equally well in the case of A. africanum. The ecology and relationships of this species will be dealt with in more detail in following sections of this report. PALAEOECOLOGY The Langebaanweg Agriotherium assemblage is comprised of over 330 specimens, which represent a minimum of 14 individuals. Females are much less commonly represented than males, and no very young animals are known. Those postcranial bones belonging to immature individuals are probably all of young adults, while those of which teeth are known are all young or prime adults. Some specimens (e.g. L40002) show signs of osteo-arthritis, which suggests an advanced age for the individuals concerned. Bone pathology is otherwise rare, one notable exception being the metacarpal, L45448, which exhibits an osteitis of the proximal end, the cause of which is unknown. All A. africanum specimens were found in, or closely associated with, river channel deposits. They occurred together with a wide variety of terrestrial, freshwater and marine vertebrates, which range in size from shrews to whales. Lists of most associated mammals have been given elsewhere (Hendey 1976, table 4; 1978, table 10). Associated birds will be listed by P.V. Rich (in pre- paration). Lists of lower vertebrates, which include cartilaginous and bony fish, amphibians and reptiles, have yet to be compiled. It is clear that A. afri- canum was an element of a rich and diverse fauna, with resemblances to both late Miocene (Turolian) faunas of Eurasia, and later African faunas. The Varswater Formation fauna includes descendants of taxa typical of the Eurasian late Miocene, with A. africanum included in this category, as well as ancestors of species which are now typically African. A. africanum is one of the Varswater Formation taxa not recorded from the Quartzose Sand Member, the lowest of the three important fossil mammal- bearing units of the succession (Hendey 1976) (Table 13). The Quartzose Sand ANNALS OF THE SOUTH AFRICAN MUSEUM 54 jueurwmopeid So}BIQOVWIOA | RII}Sel1o} i pues onjeydsoyd ul UOUWIWOD sso] Sului09eq jnq ‘“jURpUNQe s]Issoy pues oiyeyd PeprAtp -soyd ojur spieMdn 3ulpeis -qns Se] jIssoy} pue pues zjienb yueq JOA pue jouueys IOAII jou yusseid Ajgeqoid sua][od yng ‘pep1OdeaI $9} ¥1q91.19A AjUO ‘]JJ JO yYInos uoWUWOS SSO] A[PAISSoIs 01d S]ISSOF suoziioy Avid pue pues zj1enb Jepun souilj}oWIoOs “Keo pue pues snosdvuoqgies puod pure ysieur Aejo pue pues sNOsdeUOGIeS O}UI SUIpeIs AI PUe I] UseM19q 9} eIPSUIE} UL Sel jIssoy pue pues zji1enb yueUIWOpeld so}e1q9}19A [eL} -So119} PUL S[eOS f UOUIUOD S]ISsOy yueurmopeid SO} VIQOIOA [P1I}S91I19} pue sjeos ‘juepunqe sjissoy yoor ajeydsoyd jo sa] ur pue jouueys IdATI ‘uo 3x jIsso} pue pues zjrenb UdaNdY ALIYOHdSOHG I1VLATWdg yueulMopeid $9}21Q9}19A [PII}SO1I0} gjees ‘juepunqe s|Issoy yoor ayeydsoyd uo pues Askejo yueq ISAII pue e1el AISA S3}e1QG -O}LIOA ‘S[ISSOJOIOIUI SUIIUI SUIOS poyenucieyipun pues syeqdsoyd [ero] oulTeUr ‘Wdd SAONFAANIIO TISSOFL SLNAWIGAS OILLSIAALOVAVH) SLNANWNOMIANA IVNOILLISOdId yaodai stu} pure 9/6] Aepusyy 6L61 10 ja yIsuld ‘SomuLEgosURT ‘AlIengd .q, “UOMNeUIO, IoJeMSIVA OU} Ul SOOUSIINIDO [ISSOJ PUL SJUSUIIPSs SI}SIIO}OvIeY ‘sJUSWIUOIIAUS [eUOITISOdaqd €[ J1av.L, 55 AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA sjiuin AreJUWIIpas JO soroey oe (AT 01 [ INSO PUe AT O1 I N&E Wedd ‘S}UsTeAINboe I1ey} pue) py 0} Tq pue eq pue ZC SUOISIAIpgns oy, jueulUoperd saye1q -OJIOAUL PUL So}eIQ9}I0A SUTTeUT ‘yuepunge SOoUI}oWIOS s|Issoy uMOUyY JOU yueUIWIOpeld so}e1q -O}IOAUL JoJeMYseIJ pue sUlIe -njso “oulIeur ‘jUepUNge s|Issoy WOUWIUIOS $0} V1IQO}IOA [VII{Se1I0) “UeUTUIOp -o1d suoyjod ‘yuepunge sjissoy yueulwopeid $0}81Q9}19A [PII}S01I9} ‘UOWWIOS SOUIjaWIOS s]ISSOy yueulMopeid SO}B1IQS}IOA [PII}S01I0} ‘yuepunqe sowiiewlos sjissoy pues zj1enb pue odes yoor 9sjeydsoyd uMOUyY j0U wis Appnut Ae pue pues snosovuogieo pues zj1enb puvs zj1enb (¢ “39 “6L61 (72 2 9]8UIC 90S) yoveq Apues pure A011 (Aizenb Jo yjnos pue jsvo-yjnos s}sixo A[qeqoid yng posodxo ou) JouURYS JIOATI { em ysieur JeuUeYd IOATI 0} Josopo ured -pooy g ‘I Jo uorerres o[qissod (jouuPvYyS sseuTeIp IOUTUI ‘puod °3'9) sjusUIUOIAUSOIOIUI poyeioosse yA ‘urefdpooy poyenuorsyipun o ~~ le) Z ECINCI IAN TAAVAD wuasWayY, GNVS dASOZLUVNO N Ta 56 ANNALS OF THE SOUTH AFRICAN MUSEUM Member deposits were laid down mostly on the floodplain of a river which then met the sea to the south or south-west of the existing ‘E’ Quarry. Many of the fossils from this unit are believed to represent the remains of animals which lived in the immediate vicinity (Hendey 1974: 349-353; 1976: 223-226). By contrast, most of the Pelletal Phosphorite Member fossils, including those of A. africanum, were washed into the area by the river, which was then following more northerly courses, first depositing bed 3aS and later bed 3aN (Hendey 1976: 226-230). Consequently, the A. africanum fossils are likely to be out of their natural environmental context, and there is no way of certainly establishing the nature of the preferred habitat of the species. Assuming that A. africanum was a terrestrial species, the number of possibilities is, however, limited, and there is some evidence which favours one of them. The environment in the vicinity of Langebaanweg and in adjacent areas at the time of deposition of the Varswater Formation was clearly very different from that of the present (Hendey 1973). Long-necked giraffes (Giraffa sp.) were common, and, together with other large browsers such as a sivathere (Siva- therium hendeyi), a palaeotragine (Palaeotragus cf. germaini), and primitive proboscideans (Anancus sp., Mammuthus subplanifrons), indicate the presence of trees, probably in substantial numbers, and perhaps in the form of a riverine woodland. On the other hand, grazers such as alcelaphine antelopes (Gentry 1980 in press), an equid (Hipparion cf. baardi), and a _ rhinoceros (Ceratotherium praecox) indicate the presence of grasslands as well. Although there was evidently a variety of micro-environments in the area (Hendey 1976), it is only the major terrestrial habitats of woodlands and grasslands which need be considered in the case of Agriotherium. Of the large herbivores, the one which occurs most commonly in the Quartzose Sand Member and bed 3aS of the Pelletal Phosphorite Member is Ceratotherium praecox, a grazer, while browsing giraffoids are very rare in the Quartzose Sand Member, and only slightly more common in bed 3aS. By contrast, C. praecox is either very rare, or absent, in bed 3aN, while giraffoids are astonishingly well represented. The implication is that either woodlands became a progressively more widespread habitat during deposition of the Varswater Formation, or that taphonomic factors were such that woodland species had their remains incorporated in the deposits with increasing frequency. Either way, the fact that A. africanum is not recorded from the Quartzose Sand Member, is rare in bed 3aS, and is relatively common in bed 3aN, suggests that it was a woodland species. This conclusion has also been reached in respect of Agriotherium elsewhere. Kurtén (1968: 119) suggested that Agriotherium ‘was probably a forest animal like most modern bears’, while Wolff et al. (1973: 226) concluded that ‘it does seem that specimens of /ndarctos and Agriotherium are better represented at localities which have a greater representation of woodland forms’. The possible influence of habitat on the evolution of Agriotherium will be discussed later (see p. 70). =e ER. = = -e ~ —aer AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 57 The habitat preference of Agriotherium may have been a factor which contributed to its comparatively poor fossil record, but this was almost certainly due largely to the habits of the animal. Agriotherium has long been recognized as an atypical ursid because it was apparently carnivorous rather than omni- vorous. Carnivorous species of such gigantic proportions would of necessity have been rare animals, even under the most ideal conditions (Wolff et a/. 1973). The same restriction would not apply in the case of more omnivorous bears, and for later species the ‘fossil record is excellent; indeed, as regards the Pleisto- cene bears of Europe, almost incomparable’ (Kurtén 1968: 119). An added factor in the case of the latter was that extreme climatic conditions during the Pleistocene led bears to use caves as retreats, with consequent concentration of their remains in caves. Such circumstances did not apply to Agriotherium, all remains of which are recorded from open sites dating from a climatically moderate period. Although A. africanum has been said to be a well represented species, it is nevertheless one of the less common elements in the Langebaanweg assemblage, which is now comprised of the remains of many thousands of animals. Pro- portionately, Agriotherium may be no more common at Langebaanweg than it is at localities elsewhere. Many of the Langebaanweg Agriotherium specimens were recovered in the course of mining operations, or by screening of bulk sediment samples mechanically removed from the mine. In these instances the original condition _Of specimens, associations of skeletal elements and body part representations cannot necessarily be determined. More significant from a taphonomic point of view is that material recovered from controlled excavations. Three such excavations yielded Agriotherium remains (Fig. 26, Table 14), and the relevant material is listed on pages 5-6. The deposits in which this material occurred are noteworthy for the almost complete absence of a very coarse lithic fraction. Occasional pebbles of quartz, feldspar and the local phosphate rock do occur, but the sediments are generally made up of medium- to coarse-grade sands. Fine sands and clayey sands are also present. The larger elements of the lag gravels in bed 3aN are almost exclusively bones and teeth of vertebrates (Fig. 27). Consequently, those fossils transported by the river were not subjected to the destructive battering by, and against, cobbles and boulders, which is often a feature of this sedimentary environment. For about 30 km east of Langebaanweg there are few rock out- crops, the area being largely covered by the generally sandy deposits of the ‘Sandveld’ (see Talbot 1947; Visser & Schoch 1973). This means that in its lower reaches the river which was largely responsible for building up the Vars- water Formation (Fig. 1) could have picked up little in the way of a coarse lithic fraction. In addition, its generally sandy bed would have provided a relatively smooth passage for organic materials in its load. In fact the fossils of beds 3aS and 3aN show remarkably few signs of abrasion which could be ascribed to transport in sand-charged water over a 58 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 26. Aerial view of past and present areas of exposure of bed 3aN in ‘E’ Quarry, Lange- baanweg. 1—LBW-E 1975/1; 2—LBW-E 1976/1; 3—Dump 10; 4—LBW-E 1976/2; 5—Dump 9; 6—Dump 8; 7—Carbonaceous deposits; R—Phosphate rock outcrops; W—Wet season river channel; D—Dry season river channel. sandy substratum. What abrasion there is may have developed after deposition rather than during transport (see below). This suggests that many of the fossils reached the vicinity of ‘E’ Quarry still protected by soft tissue, perhaps even as floating carcasses, and that disarticulation and dispersal of skeletal elements took place locally. The bed 3aN Agriotherium specimens were recovered from deposits laid down in three distinct micro-environments. Those deposits exposed in the AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 59 TABLE 14 Controlled excavations in ‘E’ Quarry which yielded Agriotherium remains. EXCAVATION DEPOSITIONAL STRATIGRAPHIC ENVIRONMENT Unit? LBW-E river bank and 1975/1 river channel, with phosphate rock substratum PPM 3aN I river channel, 1976/1 with phosphate rock substratum LBW-E river channel in PPM 3aN 1976/2 lee of phosphate II rock, with unconsolidated sand substratum 1 See Table 13. 2 Locality 5 of Dingle et al. 1979, fig. 2. excavation LBW-E 1975/1 were laid down partly in the river channel and partly on the north bank of the channel. Channel deposits were sampled in both excavations LBW-E 1976/1 and 1976/2, the depositional environments differing only in that the former had a rock substratum and the latter a sandy one. In both the 1975/1 and 1976/1 areas the substratum was a phosphate rock horizon of up to 0,75 m thick. The third micro-environment was not sampled by controlled excavation, but was the source of a few Agriotherium specimens recovered in the course of mining operations. This was an extensive area of carbonaceous (peat-like) deposit over the southern (seaward) limit of the river channel. It was probably the area of accumulation of plant debris washed down by the river during flood times. During the dry season it formed the west bank of the river and was probably a marshy area with appropriate vegetation. The likely positions of the wet and dry season channels are indicated in Figure 26. The bed 3aN deposits are overlain by the thick and extensive commercially exploited phosphatic sand, which was deposited in a marine littoral environ- ment (Tankard 1975), and from which some fossil vertebrates, not including Agriotherium, are known (Hendey 1976: 230). The fossils from LBW-E 1975/1, which were deposited close to or on the north bank of the river channel, were generally better preserved and less frag- mented than those from the other two excavations. This applied particularly in the case of specimens not in direct contact with the phosphate rock sub- stratum. In the area of LBW-E 1976/1 most of the deposit overlying the phosphate rock had been mined away, and the material recovered came from on, or close to the rock surface itself, particularly depressions therein. This material was for the most part very fragmented. The greatest concentration of fossils in bed 3aN was in the area of LBW-E 1976/2. Here the deposits were laid down on an unconsolidated substratum (the Quartzose Sand Member) in the lee (west) of the phosphate rock exposed 60 ANNALS OF THE SOUTH AFRICAN MUSEUM in the other two excavations. Immediately adjacent to the phosphate rock a 60 cm thick horizon of fossils was accumulated (Fig. 27). This thinned out rapidly to as little as 10 cm westwards and southwards. Most of the fossils in this area were highly fragmented and in a poor state of preservation. This applies particularly in the case of the remains of larger species, of which giraffoids, especially Sivatherium, were by far the most commonly represented. By contrast the remains of aquatic vertebrates, with the seal, Prionodelphis capensis, being exceedingly common, tended to be in good condition, although their remains were often fragmented and skeletons disarticulated and dispersed. As this fossil lag deposit thinned westwards and southwards, the clastic matrix became finer-grained, with an increasing clay component. Westwards the fossiliferous horizon terminated abruptly against another phosphate rock outcrop, but southwards it once again thickened and also spread out laterally in a south-westerly direction. There was also a rapid darkening in the colour of the clastic matrix, which coincided with a diminution in the occurrence of vertebrate fossils. These fossils occurred mostly as isolated, and often frag- mented, teeth and bones, but associated parts of skeletons, including at least one of Agriotherium, are recorded. In the case of the latter, it is not known how complete they were, since all were chance discoveries made after disturbance of the deposit by mechanical excavators. The darkening of the deposits southwards was caused by an increasing carbonaceous fraction, which was evidently derived from decomposed plant remains. This deposit was not a pure peat, the carbonaceous material having been mixed with a high proportion of sand and clay, but such peats might well have overlain the remaining carbonaceous deposit (see below). The picture which emerges is that of a river which in times of flood carried in to the area remains of terrestrial vertebrates, sometimes as whole carcasses, depositing some along its banks and others in the channel itself. A major part of the load of vertebrate remains was deposited immediately after the channel passed over the western edge of a southward projecting tongue of phosphate rock. Another such outcrop about 30 m further west then deflected the channel southwards, where it spread out over a wide and flat area. Here it dropped the last of its organic load, this probably being comprised largely of easily trans- portable plant material. Some vertebrate remains also reached this area, but they are recorded only from the more northerly parts, that is, closest to the area where the main vertebrate load was dropped. Although the vertebrate assemblage of bed 3aN is comprised largely of the remains of terrestrial species washed in by the river, marine vertebrates are also represented. In the case of the seal the number of individuals involved is sub- stantial. The marine vertebrate remains are generally better preserved than those of terrestrial species, which suggests, not surprisingly, that they had suffered less transport. The seals, cetaceans, marine birds, bony fish, and sharks may all have been inhabitants of, or visitors to, the river estuary. This probably applied particularly during flood times when the influx of carcasses of terrestrial species *SUOISIAIP WO ¢ Ul BJVOG “JYSII oY} WO UOT}OOIIp Ajioseo-yj10U eB ul spremdn Surdojs doroyno yoo syeydsoyd pure “jJo] pomosse jisodep BxI [Issog “A11end .q, JO vore 7/9/61 A-MAT ‘LZ ‘Stet < 3) ~) fy a =) e) n re] iQ ie} Zz a 5 a fy ; x S q S S N 62 ANNALS OF THE SOUTH AFRICAN MUSEUM provided an abundant source of food for marine scavengers. Whether or not the seal was included in this category is not known. Seals may simply have been attracted by an increase in the numbers of scavengers such as fish and crus- taceans, and would themselves have attracted predators such as sharks. Marine vertebrates were certainly in a position to have their remains caught up in, and dispersed by, the river’s floodwaters. During the dry season when the river was not in flood it is unlikely to have breached the phosphate rock outcrop which underlies the northerly and easterly parts of bed 3aN. Instead it would have been deflected southwards by this outcrop in the direction of bed 3aS, which was laid down during an earlier phase of the Varswater cyclothem. This would account for the fact that bed 3aS was abruptly truncated south of the phosphate rock outcrop. This truncation was not due solely to flood periods in bed 3aN times because bed 3aS terminated ~ at least 25 m east of the first exposure of the carbonaceous deposit of bed 3aN, which is an indication of the width of the dry season channel. There may have been other distributaries of the river north of ‘E’ Quarry. As indicated earlier, that area west of the dry season channel, where floodwaters had dropped their load of organic materials, probably took the form of a marsh. Although the river probably still fed vertebrate remains into the area during the dry season, the number of specimens involved is likely to have been much lower. This raises a problem in connection with specimens collected between the principal exposures of bed 3aS and bed 3aN. There are relatively few such specimens, and their actual source is usually uncertain, since most were collected by mine workers in the course of their activities. This material has been recorded as being from bed 3aS, but it may actually belong with the bed 3aN complex of deposits. Those Agriotherium specimens with numbers between L40000 and L43000 fall into this category. Fortunately there are no doubts about the source of Agriotherium specimens from the bed 3aN excavations, and those from the bed 3aN carbonaceous deposits, the latter being distinctive because of their dark colour. The only other fossils from ‘E’ Quarry in a similar state of preservation are those from the ‘peat bed’ of the Quartzose Sand Member (Hendey 1976: 218, table 2), and there was no possibility of material from these two deposits becoming mixed. Although there can be little or no doubt as to how the bed 3aN Agrio- therium remains reached the ‘E’ Quarry area, it is of interest to consider the nature of some of this material. The partial skeleton, L45062, is interpreted as the remains of an animal which reached the point of its discovery as a complete, or nearly complete carcass. This carcass was deposited about 15 m west of the phosphate rock outcrop in the LBW-E 1976/2 area. It is virtually certain that at least some, and perhaps all, missing parts of L45062 were mined away, the parts recovered having come from an area of 2 to 3 m* immediately adjacent to a vertical face cut by a mechanical excavator. The remains were found in the lag deposit where AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 63 it. was about 20 cm thick, and were centred on a depression on the unconsoli- dated substratum. The preservation of the remains was considerably better than that of the majority of surrounding fossils. Their good preservation, together with the unmistakable characteristics of Agriotherium bones and teeth, facilitated their recovery from a mass of thousands of fragmentary fossils. It appears that after deposition of the carcass of L45062, flowing water scoured the depression beneath it. At the same time the carcass acted as an obstruction to coarse debris, causing it to settle in the immediate vicinity. Disarticulation and slight dispersal of skeletal elements followed decomposition of soft tissue. L45062 is unusual in being one of the few instances in the LBW-E 1976/2 area where parts of one individual were found in a good state of preserva- tion, and with skeletal elements in close association. This was otherwise notice- able mainly amongst the seals, although in these instances the large number of individuals involved made it impossible to separate their skeletons. The Agriotherium forelimb, L45063, was another instance where there was association of skeletal elements of one individual. This material was found in the same depression as L45062, but was readily distinguishable, having belonged to a smaller (female) individual. This forelimb must also have reached the area held together by soft tissue, perhaps with the scapula and most phalanges already detached. Although no two elements of this limb were found in articulation, the individual elements were less dispersed than those of L45062. The incomplete manus, L33160, from LBW-E 1975/1, was a similar occurrence to L45063. Other Agriotherium specimens from the bed 3aN controlled excavations tended to be isolated occurrences of individual skeletal elements. Since they are generally well preserved and show few, if any, signs of abrasion, they, too, are likely to be from carcasses similar to that of L45062. However, in these instances there was greater dispersal of skeletal elements and associations were no longer obvious. The condition of isolated specimens was usually in keeping with the nature of the depositional environment. Long bones were invariably fragmented, but shorter and stouter bones, such as those of the manus and pes, were usually intact. The hemimandible, L45114, had lost the single-rooted teeth (incisors, P; and M3), which are easily separated from the jaw, and the ascending ramus, which is more fragile than the mandibular corpus. The single- rooted canine was still in position, because the root of this tooth is large and slightly bulbous, and impossible to remove from the jaw without breaking the root or the bone enclosing it. Also of interest from a taphonomic point of view is the post-mortem damage to specimens. This was clearly dependent on the micro-environment in which specimens were deposited. The incomplete manus, L33160, was deposited close to, or on the north bank of the river in the LBW-E 1975/1 area. This specimen is remarkable because it provides evidence of the activities of hyaenas in this area. Like many of the fossils from ‘E’ Quarry, L33160 shows signs of having been chewed by 64 ANNALS OF THE SOUTH AFRICAN MUSEUM — — =—_ a thet em nein canal —— ll | < li al ge iS oo Hyaena-gnawed Agriotherium manus, 133160, from Langebaanweg. Arrows indicate areas of major damage. AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 65 a hyaena, and this was done while the bones were still articulated. The phalanges, metacarpal I and distal ends of the metacarpals II to V are lost, probably having been ingested by the hyaena concerned, and there are tooth-marks on some of the metacarpals adjacent to the missing parts. There is also damage on the scapholunar and unciform, but the magnum and trapezoid, which are largely enclosed by the other bones, are intact (Fig. 28). The remains of at least six small hyaenas (Ictitherium preforfex) were recovered from LBW-E 1975/1. This species of hyaena, and perhaps others, probably scavenged the banks of the river for the remains of animals washed into the area. Hyaena-damaged bone was not uncommon in this area, another notable example being the skull of a seal with double punctate marks on the braincase. The lower canines of Ictitherium preforfex fit these punctate marks well. Another specimen from LBW-E 1975/1, the metatarsal L30205, is remark- able in having on its shaft gnaw-marks apparently made by a small rodent (Fig. 29). This type of damage is rare amongst the fossils from ‘E’ Quarry. 110 Mi 12 113 Fig. 29. Rodent-gnawed Agriotherium metatarsal, L30205, from Langebaanweg. Arrow indicates area of damage. Although rodent remains are generally common in the fossiliferous deposits of the Varswater Formation, this was not the case in the LBW-E 1975/1 area, where only bathyergids were represented in moderate numbers. Living bathy- ergids, which are fossorial, are known to gnaw at objects encountered in their tunnels, plastic water-pipes and telephone cables being items recently affected in this way in the south-western Cape Province. It is possible that a bathyergid tunnelling on the river bank was responsible for the damage to L30205. Three distinct types of post-mortem damage are evident on the bones and teeth of the partial skeleton L45062. Most common is simple fracturing of bones, which is ascribed to movement after loss of soft tissue, and to subsequent compression by overlying deposit. Not surprisingly, it was the relatively delicate parts of the skull which were particularly affected by this fracturing. The second type of damage is abrasion, which was probably caused by sand-charged water 66 ANNALS OF THE SOUTH AFRICAN MUSEUM flowing over exposed parts of the partly buried skeleton. This type of damage is not uncommon on the fossils from ‘E’ Quarry, and a good example was mentioned elsewhere (Hendey 1970: 82, fig. 3). The third type of damage was caused by fire, with affected bones and teeth being blackened, more badly fractured and less well preserved than unburnt bone (Fig. 30). This type of damage is also not uncommon on fossils from ‘E’ Quarry, and has been discussed elsewhere (Hendey 1974: 351; 1976: 224). jy Fig. 30. Fire-damaged Agriotherium astragalus, 145062, from Langebaanweg. It was previously assumed that fire-damage was caused by dry-season bush or grass fires. While this may sometimes have been the case, there are instances where the inferred depositional environment and nature of the damage is such that fires of another sort are likely to have been responsible. The fire-damage of L45062 is a case in point. It is clear that this damage was done after the skeleton had been dis- articulated, since severe fire-damage on one bone is not matched by damage to immediately adjacent parts of the skeleton. For example, the symphyseal region of the right hemimandible is fire-damaged and the incisors are lost, whereas the left hemimandible is intact in this region and the incisors are present. Similarly, the distal end and part of the shaft of the left ulna is damaged, but corresponding parts of the left radius are unaffected. AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 67 Assuming that the depositional environment of L45062 has been correctly interpreted, it is highly improbable, if not impossible that the random burning of parts of the skeleton could have been caused by a bush or grass fire. A more plausible explanation is that the fire-damage was caused by peat fires, a phenomenon which has been reported in a North American estuarine swamp by Staub & Cohen (1979). It was indicated earlier that the horizon in which L45062 occurred graded laterally into a peat-like deposit, and that pure peats may have overlain this horizon. Such peats, if they did exist, may have been prone to dry season fires like the American example cited above. Having burnt away they would have left no trace obvious to observers unfamiliar with such phenomena other than some underlying burnt bone. Peat fires may also have led to the formation of hitherto unexplained deposits capping the peat-like sediments immediately south of the LBW-E 1976/2 area. Immediately overlying the ‘peat’ was a horizon of sand a few centimetres thick, which was recorded as being ‘orange-brown’ in colour and in this respect unlike any other sand body recorded before or since in ‘E’ Quarry. The ‘orange-brown sand’ was in turn overlain by a grey clay incorporating scattered sand grains. In the examples of peat fires recorded by Staub & Cohen (1979), and in other similar ones (e.g. Cypert 1961), ponds developed in depres- sions left in the peat after burning, and such ponds become a new and distinct depositional environment in the areas in question. If there was, indeed, a peat fire in the LBW-E 1976/2 area, the orange-brown sand and grey clay may represent sediment accumulated in a resultant pond, while the underlying carbonaceous deposit represents an unburnt residue of the original peat deposit. The carbonaceous deposit may have remained unburnt either because it was waterlogged or because its high mnon-carbonaceous content made it incombustible. The grey clay is finely laminated, suggesting slow accumulation in still- water conditions, with individual sediment particles perhaps having been transported to the pond by wind. The origin of the orange-brown sand is not known. It was initially thought that it may represent the unburnt residue of the peat fire, but the occasional vertebrate fossils incorporated in the sand show no signs of having been burnt. The existence and effects of peat fires at Langebaanweg are largely specu- lative, but they do provide a plausible explanation for hitherto unexplained, or unsatisfactorily explained, aspects of the fossils and the deposits. The ecological role of Agriotherium is also relevant here. It was mentioned earlier that Agriotherium was a carnivorous animal and, judging from its dentition, it was better adapted to carnivory than almost all other later ursids. It is the nature of the posterior cheek teeth which are particularly significant in this respect, and since these teeth are better represented in A. africanum than in previously recorded Agriotherium, it is useful to examine them from a func- tional viewpoint. The P*, M! and M? of A. africanum, and other Agriotherium, resemble one 68 ANNALS OF THE SOUTH AFRICAN MUSEUM another in having a series of prominent buccal cusps and less prominent lingual ones (Fig. 31). In all there are seven buccal cusps, three on P* and two on each of the molars, and they are in the form of a smooth curve, with the convexity directed buccally. In addition to being lower-crowned, the lingual cusps are less distinct and fewer in number, only five excluding the vestigial accessory cusps on the P* protocone lobe. These cusps are more or less in a straight line. TULA LULL mM Tl 6 Fig. 31. Langebaanweg Agriotherium specimens showing prominence of buccal cusps. A. Lingual view of P* and M1. B. Anterior view of M?. Judging from wear facets, the buccal cusps have a combined shearing and crushing function, with the former being predominant. The lingual cusps also have this double function, but in this instance the crushing function is predominant. Much the same applies in the case of P, and the lower molars, in which the more prominent cusps are situated buccally. An exception is the large posterolingual cusp of the M, talonid. This cusp is, however, like the more buccal ones in developing an inclined shearing facet on its buccal side. The other lower teeth or cusps which develop inclined shearing facets buccally are the paraconid, protoconid and hypoconid of M,, the protoconid and hypoconid of M,, and the anterobuccal part of Ms. A most significant development relating to the emphasis of the buccal cusps in A. africanum is the reduction or loss of the anterolingual cusps of the M,.’s, L45114/L46563 and L50007. As far as is known this has not previously been observed in Agriotherium. It creates the impression that the M, of A. afri- canum was tending to become a second lower carnassial. Although smaller than the true carnassial, L45114/L46563 and L 50007 resemble this tooth in having the trigonid large and functioning essentially as a shearing element. To have carried this evolutionary experiment to its logical conclusion AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 69 would have required continued suppression of the lingual cusps of P*, M! and M2, continued emphasis of the buccal cusps of these teeth, and a similar emphasis on the more buccal parts of the lower molars. This would have resulted in a sectorial dentition unique amongst Carnivora, but since no Agriotherium is known in which there was an advance on the A. africanum condition, it evidently served the needs of the genus adequately. It is worth noting in this connection that a similar evolutionary path has been followed by the polar bear, Thalarctos maritimus. In this species the P4 protocone is reduced or absent, the buccal cusps of M! and M? are relatively higher crowned than those of other ursines, while the lingual ones, including the M? talon, are correspondingly reduced. These dental characters reflect ‘the carnivorous habits of the species and diverges from the omnivorous dentition of most other ursids’ (Kurtén 1964: 4). Whether Agriotherium was a predator or a scavenger is not known. Its large and cumbersome build suggests that the former alternative is less likely, although it may have been capable of hunting down at least some of the larger contemporary herbivores, such as the giraffids. The P* of Agriotherium has long been referred to as hyaenid-like (Falconer & Cautley 1836; Wagner 1837), and recently Wolff (1978: 4) has written that this tooth ‘bears a striking resem- blance in robustness to the teeth of the bonecrushing hyaenas’. This implies a belief that Agriotherium may have been a scavenger. However, in the hyaenas it is the more anterior cheek teeth which are enlarged to perform a bone-crushing function, and the enlarged cheek teeth of Agriotherium may have been too posteriorly situated to perform this function efficiently. Irrespective of how its food was acquired, there can be no doubt that an abundance must have been available in the Langebaanweg area when it was inhabited by Agriotherium. There is evidence, however, which suggests that the Varswater Formation fauna dates from the latter part of the period which Kurtén (1971: 152) termed the ‘climax of the Age of Mammals’, and that even in its heyday it was becoming an anachronism. In fact, it is possible that the circumstances which led to the evolution of Agriotherium ultimately also caused its extinction. The late Tertiary was a period of world-wide environmental change, with a general lowering of mean annual temperatures (Butzer 1971, fig. 2), which heralded the glacial-interglacial oscillations of the Pleistocene. Over wide areas of the Old World and the Americas, forests and woodlands were giving way to savannas and grasslands, and these changes had a profound effect on the character and composition of terrestrial vertebrate faunas (Webb 1977). The Varswater Formation fauna provides evidence of these changes. For example, it includes two of the earliest alcelaphine antelopes yet recorded (Gentry 1980), and it is these bovids with their high-crowned teeth which are characteristic of the African savannas today. Similarly, an early ancestor of Africa’s grazing rhinoceros (Ceratotherium simum) is recorded from Langebaan- weg (Hooijer 1972). Palynological evidence from Langebaanweg, and elsewhere 70 ANNALS OF THE SOUTH AFRICAN MUSEUM in the south-western Cape Province, indicates that the modern Cape macchia (fynbos) vegetation was becoming established at this time, and that the climate was changing from ‘Cool Wet’ to ‘Colder Drier’ (Coetzee 1978: 121, fig. 2). There is even some as yet unpublished evidence to suggest that there was a marked fall in local sea temperatures during deposition of the Varswater Formation. The immediate cause of the climatic and environmental deterioration in the Langebaanweg area and adjacent regions was the development of the Benguela Current System off the west coast of southern Africa and the consequent aridification of the adjacent land mass (Siesser 1978; Tankard & Rogers 1978). The development of the Benguela Current was in turn the result of major glaciation in Antarctica later in the Miocene (Kennett et al. 1975). This period of climatic and environmental change must have influenced the Ursidae as much as any other mammalian group. During the Vallesian | the ursids which inhabited Europe were generally relatively small omnivorous animals inhabiting forests and woodlands. That lineage which was to give rise to the Ursinae, and probably also the Tremarctinae, continued to be repre- sented by small species during the Turolian. However, in the Indarctos lineage there was a marked increase in the size of the species during this period. In the following section of this report, evidence will be presented which indicates that Indarctos was the ancestor of Agriotherium, and consequently it is the Indarctos lineage, and the changes it underwent, which are relevant here. By the Turolian (and its equivalents) the environment in mid-latitudes was in the process of change, with woodlands giving way to more open country (Kurtén 1971). Consequently, woodland plant foods such as fruits, nuts and berries, favoured by omnivorous ursids, were probably becoming less readily available. It is the larger species which would have been most adversely affected by this development, and thus Turolian Jndarctos, and its counterparts else- where, may increasingly have been forced to adapt their diets to more abundant food sources. In this instance, the response was evidently in the direction of increased carnivory, with adaptations culminating in the evolution of Agriotherium. During the Pleistocene a similar situation arose in respect of Thalarctos. At the onset of one of the glaciations, populations of Ursus arctos did not retreat southwards in advance of the ice, but instead adapted to the new con- ditions. Amongst other things, this involved a growing reliance on carnivory to replace the plant foods which had previously been part of its diet (Kurtén 1964; Hendey 1972). The early ancestors of the giant panda, Ailuropoda, evidently responded to late Tertiary environmental change in a different way. They adapted their diet to include more readily available plant foods which had not necessarily figured in their diets previously. Thus arose the most herbivorous of all living ursids, whose main diet of bamboo shoots is varied by other plant foods when this is possible (Ewer 1973). : 71 AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA Pe be! Rex : “ff ‘OL6[ Bulidg ‘somuvegesuryT ‘AjIJedo1d ourur py] sojwioyo sul “ze “SI 72 ANNALS OF THE SOUTH AFRICAN MUSEUM By becoming a highly specialized herbivore, Ai/uropoda ensured its survival, although latterly in diminishing numbers and in increasingly remote areas. On the other hand, by adopting carnivory, Agriotherium placed itself in direct competition with smaller and better adapted predators and scavengers for a declining food source (i.e. large woodland browsers), or one which was increas- ingly difficult for a large ambulatory carnivore to acquire (i.e. cursorial grazers of the savannas and grasslands). According to this hypothesis, the fate of Agrio- therium was sealed by the factors which led to its origins. The picture of the Langebaanweg Agriotherium which emerges is of a gigantic carnivore living at a time when vertebrate life flourished in the south- western Cape Province, and when the Langebaanweg area was well-watered and richly vegetated. The semi-arid environment of Langebaanweg today, devoid of indigenous trees and scarred by man’s activities, is a poor reflection ~ of the past (Fig. 32). Only some of the smallest of the carnivorous cousins of Agriotherium, namely foxes, polecats, mongooses and wildcats, still occupy the area and the prospects for their survival are limited. RELATIONSHIPS OF AGRIOTHERIUM The first Agriotherium specimens to be collected and recorded were several isolated teeth of an aged individual found at Montpellier in France early in the nineteenth century. These specimens, which are preserved in the Museum d’Histoire Naturelle, Geneva, were described by Cuvier (1822) under the name of ‘Lophiodon de Montpellier’. Stehlin (1907) gave an account of this historic material, whose true identity remained a mystery for many decades, and pro- vided an indication of the difficulties originally encountered in correctly identi- fying and classifying specimens belonging to Agriotherium. Although its ursid affinities were recognized by Falconer & Cautley (1836) on the basis of material from the Siwalik Hills of India, Agriotherium is in many respects an atypical member of the family. Of all the genera of Ursidae, it is Agriotherium and its herbivorous counterpart, Ailuropoda, whose relationships have been most controversial. The fact that Agriotherium has a poor fossil record, while that of its nearest relatives is not necessarily any better, has aggravated the problem. In addition, there has been a tendency to compare it with the ursine bears, which have long been well known, and this has further obscured matters, since the ursines and Agriotherium are only distantly related. Much the same applies in the case of Ailuropoda. Hyaenarctos Owen, 1845, a junior synonym of Agriotherium Wagner, 1837, was the generic name in common use during the last century and the earlier part of the present one. Prior to the identification of Indarctos by Pilgrim (1913), “Hyaenarctos’ was the name applied to species now referred to both Agrio- therium and Indarctos. This is an indication that these two genera have many characters in common, a point made obvious by studies such as those of Frick (1926) and Matthew (1929). Pilgrim (1931, 1932) did much to clarify the situa- AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 5 tion, and thereafter, apart from occasional lapses (e.g. Viret 1939), the name ‘Hyaenarctos’ fell into disuse, and Pilgrim’s interpretation of the genera Agrio- therium and Indarctos was generally accepted. Pilgrim (1932: 42, 43) listed half a dozen characters which he believed distinguished these genera and concluded that ‘Indarctos appears to represent a distinct line of development from Agriotherium, in some ways more and in others less advanced than the latter’, while ‘J. punjabiensis seems to some extent to bridge over the gulf between [them]’. In spite of the attention given by Pilgrim to the problem of distinguishing Agriotherium from Indarctos, and in determining their relationships to one another, and to other ursids, uncertain- ties have remained. Nevertheless, it has become common practice to include these genera in the subfamily Agriotheriinae, together with ‘the basal genus of the family, Ursavus’ (Kurtén 1966: 7). Since Agriotherium and Indarctos fossils are not common, they, and the question of their relationships, have received only infrequent attention since Pilgrim’s (1932) review. Such attention has been prompted on the one hand by descriptions of new material (e.g. Viret 1939; Kretzoi 1942; Tobien 1955; Thenius 1959; Hendey 1972; Crusafont & Kurtén 1976; Wolff 1978), and on the other by reviews of ursid inter-relationships and phylogeny (e.g. Erdbrink 1953; Thenius & Hofer 1960). On those rare occasions when the relationship between Agriotherium and Indarctos has been discussed, Pilgrim’s (1932) opinion has been favoured. For example, both Erdbrink (1953, fig. 61) and Thenius & Hofer (1960, figs. 34-35) place these genera on separate lineages, and indicate derivation from an unknown common ancestor in the Ursavus group. This theory is based on the belief that Agriotherium is the more ‘primitive’ (i.e. canid-like) of the two genera, which, since it survived later than Indarctos, must represent a distinct lineage. However, there is an alternative hypothesis. In previous studies on the Langebaanweg Agriotherium it was suggested that this genus was directly descended from Jndarctos (Hendey 1972, 1977), an idea which was apparently first conceived by Schlosser (1899). Before examining this alternative in more detail it is worth noting what is known of the temporal ranges of the two genera. In Europe Agriotherium is first recorded from the Ruscinian, while Indarctos is known only from the Vallesian and Turolian (Table 15). In North America Agriotherium is restricted to the late Hemphillian (i.e. 4,5-6,0 Ma) and Indarctos is recorded only from the latter part of the early Hemphillian (i.e. 6-7 Ma) (R. H. Tedford 1979, pers. comm.). In Africa the Langebaanweg Agriotherium is from a Ruscinian-equivalent fauna whereas the cf. Agriotherium from Sahabi (Boaz et al. 1979) is probably an advanced Indarctos in terms of current defi- nitions, and is almost certainly a little older than the Langebaanweg species. The situation in Asia is still obscure, although it is now known that the Dhok Pathan of the Siwaliks, from which important Indarctos and Agriotherium specimens are recorded, spans an appreciable period during the late Miocene 74 ANNALS OF THE SOUTH AFRICAN MUSEUM TABLE 15 The occurrence of Jndarctos and Agriotherium in Europe and the Middle East. APPROX. AGE IN m.y.t TAXON LOCALITY MAMMAL AGE! | MEIN ZONE? Ruscinian Turolian Montpellier Concud Samos Pikermi Maragha Montredon Late Orignac Vallesian — Pfaffstetten Westhofen Can Llobateres Early 12 Can Purull Vallesian Agriotherium insigne . Indarctos atticus Indarctos arctoides Indarctos vireti 1 Berggren & Van Couvering 1974; 7? Mein 1975. and Pliocene (Pilbeam et al. 1977). Consequently, the Dhok Pathan repre- sentatives of these genera, whose taxonomy is controversial (see below), were not necessarily contemporaneous with one another. A similar situation may well exist in the case of Jndarctos and Agriotherium from Chinese late Tertiary localities, with the former known from Localities 30, 31, 43 and 52, while Agriotherium is tentatively identified from Locality 13 (Kurtén 1952). Indications are, therefore, that Agriotherium and Indarctos were not contemporaries, and when their age is known the former appears later in the fossil record. Consequently, their known temporal ranges are in accord with the theory that Agriotherium was descended from Indarctos. While its relationships to Agriotherium may be controversial, the history of Indarctos itself is now reasonably well documented. It apparently had its origins in Europe during the Vallesian, having stemmed from an Ursavus, and subsequently spread through Asia and into North America (Thenius & Hofer 1960), as well as to Africa (see above). Differing interpretations of inter-generic relationships are possible. For example, Crusafont & Kurtén (1976) suggested that J. vireti was ancestral to both I. arctoides and I. atticus, whereas I. arctoides is here regarded as an intermediate between the other two species. Crusafont & Kurtén (1976: 15) further suggested that J. anthracitis ‘may be a precociously specialized form’. This species is, indeed, unusual, perhaps because it evolved in isolation on a Tethyan island, but it is largely irrelevant to present con- siderations. Recorded Asiatic Indarctos are either close to I. atticus or more advanced, while North American /ndarctos is more advanced than J. atticus. Irrespective of the actual inter-relationships of Indarctos species, there is no doubt that J. vireti is a generalized and early form, whereas J. atticus, and Asian and North American Jndarctos are more specialized and younger in age. For example, Crusafont & Kurtén (1976: 15) noted that J. atticus is ‘more advanced [than other European /ndarctos] with larger cheek teeth, more reduced premolars, heavy and powerful jaws, and ursine limb proportions’. a ee — rf -—- ——e > — a J , nn ta ~— eee 6 ee | A oe. AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA US It follows that if Agriotherium were descended from Jndarctos, then the evolutionary trends evident in the /. vireti-J. atticus lineage are likely to have been continued in the hypothetical Indarctos—Agriotherium lineage. This is, indeed, the case, and Crusafont & Kurtén’s comments quoted above apply equally well to Agriotherium relative to J. atticus. Such general trends do not necessarily constitute proof of a direct phylogenetic relationship between the two taxa. Much more convincing evidence comes from an examination of certain tooth and skull characters in advanced Jndarctos (i.e. those between 6 and 9 m.y. old), and early Agriotherium (i.e. those that are, or probably are, about 5 m.y. old). Examples of the latter are A. africanum from Langebaanweg, A. insigne from Europe (Montpellier), and A. palaeindicum from the Siwaliks. Advanced Indarctos includes J. atticus from Europe (Samos) and Iran (Maragha), J. pun- jabiensis from the Siwaliks, and North American Indarctos. All recorded speci- mens of the latter are here referred to the species J. oregonensis (see p. 101). The classification of material referred to some of the above species has long been complicated by specimens which exhibit a combination of characters ‘typical’ of both genera. Pilgrim (1932: 4446) discussed such a problem involv- ing three incomplete mandibles from the Dhok Pathan of the Siwaliks (GSI-D8, D9, D10), and although he found it ‘difficult to give a definite answer’, he decided to reverse the identifications suggested by Lydekker (1884). Pilgrim’s identifications are accepted here, with D8 assigned to Agriotherium palae- indicum, while D9 and D10 are assigned to Jndarctos punjabiensis. The latter has since been recognized as one of the more advanced representatives of Indarctos (Thenius 1959), while A. palaeindicum is one of the more primitive representatives of Agriotherium (Hendey 1977). It is highly likely that more will yet be written on the identity of the Siwaliks’ Indarctos and Agriotherium. Both Pilgrim and Lydekker may have been incorrect in their interpretation of GSI-D8, D9 and D10 (and other specimens) since they could represent a single species which was no more variable than the Langebaanweg Agriotherium. The anomalous situation in respect of J. pun- Jabiensis and A. palaeindicum will be repeatedly evident in the discussions which follow. The study of new and well provenanced material from the Siwaliks may resolve a situation which is beyond satisfactory resolution on the basis of available evidence. Whatever the final outcome of this controversy, the fact that specimens can with some justification be identified with either Indarctos or Agriotherium is here regarded as highly significant, with specimens such as GSI-D8, D9 and D10 being interpreted as ‘intermediates’ between ‘typical’ Jndarctos and ‘typical’ Agriotherium. The present study has shown that such ‘intermediates’ are more common than has hitherto been supposed. The characters which the two genera have in common, coupled with apparent evolutionary trends linking them, provide surprisingly good evidence for the transition of Indarctos to Agrio- therium considering the relatively poor fossil record of the taxa concerned. 76 ANNALS OF THE SOUTH AFRICAN MUSEUM Indarctos—Agriotherium ‘intermediates’ are included in the A. africanum assemblage, while an undescribed A. insigne specimen from Montpellier (NMB-MP549) and the recently described Indarctos skull from Florida (Wolff 1978) are also in this category. The Florida skull is perhaps the best single recorded specimen which is ‘intermediate’ between ‘typical’ Indarctos and ‘typical’ Agriotherium. The age of this specimen is ‘late Hemphillian’ (Wolff 1978: 1), and it probably dates back about 6 m.y. It is certainly younger than the Samos J. atticus skulls described by Helbing (1932) and Thenius (1949, 1959), and it is much younger than the skull of J. vireti from Spain (Crusafont & Kurtén 1976). On the other hand, it is probably a little older than the skull of the Langebaanweg A. africanum. Judging from the few measurements given by Wolff (1978), and from illustrations, the Florida skull is appreciably larger than that of J. atticus, which in turn is larger than that of J. vireti (Crusafont & Kurtén 1976). The trend of increasing size with time in the Jndarctos lineage is clearly illustrated by these specimens. This trend was continued with the evolution of Agriotherium, since the skull of A. africanum is, in some respects at least, still larger than that of the Florida Jndarctos. In respect of size the Florida skull is probably closer to that of A. africanum (and A. sivalense) than the three described skulls of European Jndarctos. The general similarity between the skulls of the Florida Jndarctos and A. africanum was discussed earlier (see pp. 9-16), while the differences between them can all be ascribed to the more advanced condition of the latter. The advances are probably all related to the larger size of the A. africanum skull, and to modifications of its masticatory apparatus. Judging from the Florida Indarctos and A. africanum skulls, the following are the most significant cranial characters shared by advanced Indarctos and Agriotherium: 1. Snouts relatively short and broad. 2. Zygomatic arches very stout and of similar shape. 3. Sagittal crest very high. 4. Overall similarity of the basicranial regions, particularly the absence of the alisphenoid canal, and the positions of the oval, postglenoid and other foramina. In respect of the development of the snout, zygomatic arches and sagittal crests, the Florida Jndarctos is intermediate between J. atticus and A. africanum. The basicranial region of J. atticus is similar to those of the other two skulls, and, according to Crusafont & Kurtén (1976), the basicranial region of /. vireti is similar to that of J. atticus. I. vireti is more primitive, however, in having the ‘facial part of the skull. . . relatively much longer than in J. atticus’ (Crusa- font & Kurtén 1976: 10). The relatively short snout of Agriotherium goes together with a relatively short palate, which is of interest because Kurtén (1964: 22) found that the AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA Ti palate of the polar bear, Thalarctos, is slightly shorter than that of the brown bear, U. arctos, a pattern which he was not ‘able to match... . in any other bear population’. Evidently the Indarctos—Agriotherium example was not taken into account, but it is a parallel to U. arctos-Thalarctos in this respect. Kurtén (1967) subsequently noted that, like Thalarctos, a short and broad snout characterizes Arctodus, another ursid which is convergent with Agriotherium (see p. 51). - Another apparent similarity between the skulls of Agriotherium and Thalarctos is that in lateral view they have a relatively straight profile, with the sagittal crest prominent (Erdbrink 1953). It was on this basis that Wagner (1837) proposed the name Agriotherium for the Ursus sivalensis of Falconer & Cautley (1836) (see Erdbrink 1953: 557). Thalarctos is like Agriotherium and Arctodus in being a carnivorous animal descended from an omnivorous ancestor, and other parallels between them will be mentioned below. The presence of a premasseteric fossa in the mandible of Agriotherium, and its absence in Jndarctos, is an important distinguishing characteristic, the sig- nificance of which has prompted much published and unpublished comment. In one recent account it was erroneously stated that the premasseteric fossa is also characteristic of Ursavus (Hendey 1977), an error stemming from Frick (1926: 99), citing Wegner (1913). In fact, Ursavus, like Indarctos, lacks this fossa, and it is thus peculiar to Agriotherium in the hypothetical Ursavus— Indarctos—Agriotherium lineage. The premasseteric fossa is important from a phylogenetic point of view, since it, like other ‘characteristics’, may not be an invariably diagnostic feature of Agriotherium. For example, it is absent from at least one mandible which has been assigned to Agriotherium, namely, the A. palaeindicum specimen GSI-D8. On the other hand, in the Samos J. atticus specimen described by Thenius (1959) (NMW-Samos 1912, 29), there is a slight depression in the pre- masseteric region which could be an incipient fossa. No such fossa, however, has been reported in more advanced Jndarctos, such as that from North America. Thenius (1959) pointed out that most of the Jndarctos mandibles then known were either incomplete or belonged to immature individuals, and that the premasseteric fossa was either not observable or absent. This fossa clearly is an ontogenetic character, since in the Langebaanweg assemblage it is well developed only in older individuals (e.g. L45062), and is shallow in the mandible of a young adult (L45114) (Fig. 33). A premasseteric fossa is otherwise known amongst ursids in Hemicyoninae and Tremarctinae. Although the early history of tremarctines is not well known (Thenius 1976), there is nothing to indicate that they and the hemicyonines are closely related. Consequently, it is certain that the premasseteric fossa in these two groups was evolved independently. There is thus no reason to suppose that its presence in Agriotherium is indicative of a close relationship with either the Hemicyoninae or the Tremarctinae. Since the significance of the premas- seteric fossa in ursids is not known (Davis 1955), there is no way of knowing why it should develop in some lineages and not in others. 78 ANNALS OF THE SOUTH AFRICAN MUSEUM A B Fig. 33. Dorsoventral cross-sections of Lange- baanweg Agriotherium hemimandibles with arrows indicating premasseteric fossae. A. 145114. B. L45062. The teeth of Indarctos and Agriotherium provide much evidence in support of the theory that they are directly related. As is often the case with carnivore incisors and canines, little of signifi- cance emerged from a study of these teeth in Indarctos and Agriotherium, except that they are essentially similar morphologically The [' and I? are perhaps the most distinctive of the anterior teeth in A. africanum, and they are closely matched by those of the /. atticus specimen from Samos described by Helbing (1932, fig. 2) (NMB-Sam31). The anterior teeth of Agriotherium are distinguished from those of Indarctos principally by their larger size, this being a reflection of the overall size differences between the two genera. It is worth noting in this connection that since the Agriotheriinae, like other ursids, exhibit appreciable sexual dimorphism, it is possible that large males of advanced Indarctos were of similar size to, and perhaps even slightly larger than, small Agriotherium females. For example, in terms of overall size the maxillary fragment of a small A. insigne specimen from Montpellier (NMB-— MP549) is virtually identical in size to corresponding parts of the Vienna I. atticus specimen (NMW-Samos 1912, 29) (Fig. 34). Similarly, the Florida Indarctos skull is in some respects as large as that of the Langebaanweg Agrio- therium, which belongs to a male, and would therefore have been larger than those of A. africanum females. Thus size alone may not necessarily be a reliable criterion for distinguishing the two genera. One of the general trends in ursid evolution has been the emphasis on the development of the posterior cheek teeth, and the. reduction in size or even loss of the anterior premolars (Pj-P3). More reduced premolars is one of the charac- AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 719 ee eee C IT) Fig. 34. Upper cheek tooth rows of Indarctos atticus, NMW-Sam1912/29 (left) and Agriotherium insigne, NMB-MP549 (right). Drawn from casts. ters which distinguishes J. atticus from the earlier J. vireti (Crusafont & Kurtén 1976). In I. atticus P} and P§ are apparently always single-rooted, while P3 usually have two roots, although P, may sometimes have only one root (Thenius 1959). In the Florida Indarctos P! and P? are single-rooted, and P? is double-rooted (Wolff 1978). Judged on the basis of the Langebaanweg sample, the anterior premolars of Agriotherium had undergone even further reduction, since in this instance all are invariably single-rooted and as many as two of a series may be lost (Table 16). This sample does not, however, exhibit one of the extremes in anterior premolar development encountered in Agriotherium. The Montpellier A. insigne specimen, NMB-MP3549, is like advanced Indarctos in having a 80 ANNALS OF THE SOUTH AFRICAN MUSEUM TABLE 16 Anterior premolars in Agriotherium from Langebaanweg. L41404 150003 = = p p p 150445 — == — p p p 140044 — —_— p a p L50004 —_— — p a p 150903 = = p a p p p 145062 p 145114 p = present; a = absent. double-rooted P? set transversely in the jaw. It is especially reminiscent of the Vienna J. atticus specimen in this respect (Fig. 34), the only difference being that the principal cusp in the latter is slightly more distinct. The P? of NMB— MP549 is unlike that of any other recorded Agriotherium. By contrast, the Vienna J. atticus specimen is Agriotherium-like in having the P, to P; reduced to single-rooted teeth. Thus, in respect of anterior premolar development conditions typical of advanced Jndarctos may occasionally be found in Agriotherium, and vice versa. In addition, the anterior premolars of Agriotherium are morphologically similar to those of advanced Indarctos, except for the slightly more distinct principal cusps in the latter. This is a primitive characteristic since the principal cusps of the anterior premolars of /. atticus are in turn less well developed than those of I. vireti. The P,’s of advanced Indarctos and Agriotherium are also almost indis- tinguishable in terms of their basic morphology. Differences in detail are probably no greater than those in the Langebaanweg Agriotherium sample, which includes one specimen (L50445) with the principal cusp configuration resembling that in the J. atticus specimen, NMB-Sam31 (Fig. 35). Of particular significance is the tendency in Indarctos for the development of a postero-internal bulge in the cingular region. This feature is well developed in the Maragha J. atticus specimen (De Mecquenem 1925). In Agriotherium the postero-internal bulge on P, is comparably developed, whereas it is absent or much less pronounced in all other ursids. The principal differences between the P,’s of advanced Indarctos and Agriotherium are that the latter are higher crowned and tend to be larger in AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 81 A Fig. 35. Buccal and occlusal views of P,’s. A. Indarctos atticus, NMB-Sam31 (after Helbing 1932, fig. 3). B. Langebaanweg Agriotherium, L50445. overall size. All the posterior cheek teeth of Agriotherium are higher crowned than their counterparts in Indarctos, although the Florida Indarctos specimen may be an exception in this respect. Apparently the crown height increase in the Indarctos—Agriotherium lineage was more or less in proportion to the overall increase in the size of the taxa concerned. A likely parallel of this situation is that involving Thalarctos, which has relatively higher crowned posterior cheek teeth than Ursus arctos (Kurtén 1964). In the case of the upper carnassial (P*), it is the nature of the protocone lobe and the development of the parastyle which are of particular phylogenetic significance. As a general rule the P* of Agriotherium is distinguished from that of Indarctos by the presence of a parastyle. However, some specimens of Jndarctos do have a P* parastyle, although it is usually less prominent than that of Agrio- therium. For example, this cusp is present in J. atticus from Concud (Crusafont & Kurtén 1976) and Samos (Helbing 1932; Thenius 1959), I. punjabiensis from the Siwaliks (Lydekker 1884), and North American Jndarctos (Merriam et al. 1925; Merriam & Stock 1925; Wolff 1978). The parastyle is particularly well developed in some North American specimens (e.g. the Florida skull), which are younger than European J. atticus. This cusp is absent in the still older and more primitive J. arctoides and I. vireti. There was thus a tendency for the development of a P* parastyle in the Jndarctos lineage, and this was continued in the presumed descendant, Agriotherium, in which this cusp is always present and well developed. As far as is known the only other ursids in which the P? parastyle is present are Ursavus depereti and its possible descendant, the giant panda, Ailuropoda melanoleuca (see p. 96). The situation in respect of the P* protocone lobe is slightly different, although equally informative. According the Crusafont & Kurtén (1976: 8) 82 ANNALS OF THE SOUTH AFRICAN MUSEUM there was a tendency for J/. vireti to develop an antero-internal cusp on the P*, a situation which ‘is not uncommon in J. arctoides and I. atticus’. This cusp is also present in the J. punjabiensis specimen, GSI—D6, although in this instance it is rather small (Lydekker 1884). In the younger and more advanced North American Jndarctos this cusp is either reduced (Wolff 1978) or absent (Merriam & Stock 1925). This indicates that while a double-cusped protocone lobe is characteristic of most /ndarctos, there was a tendency for the antero-internal cusp to be reduced or lost in advanced forms. By contrast, an antero-internal cusp is found only in some of the earlier Old World representatives of Agriotherium, namely, I. insigne from Mont- pellier (NMB-MP549) and the Langebaanweg A. africanum. In the case of the latter, this cusp is best developed in the holotype, L2045, from bed 3aS, which predates those A. africanum P*’s from bed 3aN in which the antero-internal cusp is small or absent. Indications are, therefore, that early forms of European and African Agriotherium still had an antero-internal cusp on P*, but this was soon reduced and lost. On the other hand, no Asian or North American Agriotherium is known to have had this cusp, which had already been reduced and lost in the advanced Indarctos of these continents. This suggests that the Indarctos— Agriotherium lineage may have had at least two geographically separated branches, one in Europe and Africa and the other in Asia and North America, which in respect of their P* protocone lobes evolved at different rates. The impli- cations of this possibility will be discussed later (see pp. 101—4). Reduction of the P* protocone is a characteristic of the Ursus—Thalarctos lineage (Kurtén 1964), and in South American Arctodus (Kurtén 1967). This is another example of a parallel development in Agriotherium, Thalarctos and Arctodus. Once again a double-cusped protocone lobe is otherwise known only in Ursayus depereti and Ailuropoda, although in these taxa the morphology of the lobe is somewhat different (see p. 96). As with the upper carnassial, the lower one (M,) is an important tooth in indicating the origins of Agriotherium. There is, however, a complication with M,, although in the final analysis its phylogenetic significance is not diminished. The complication concerns the identification of the posterolingual cusps of this tooth. The cusp of the Agriotherium M, which has invariably been identified as the metaconid may in reality be the entoconid. If this is, indeed, the case, then the metaconid of the Agriotherium M, is either reduced or absent, usually the , latter. Both the metaconid and entoconid are present and well developed in | Ursavus and Indarctos, although with time the metaconid becomes a progres- sively less prominent feature of My,, since it is reduced in size relative to the entoconid and is increasingly overshadowed by the protoconid. In U. primaevus ~ the metaconid is larger and more prominent than the entoconid, and is only ~ AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 83 slightly less high-crowned than the protoconid (e.g. Crusafont & Kurtén 1976, fig. 13). Much the same applies in the case of J. vireti and J. arctoides, except that in these species the metaconid and entoconid are of comparable size (Crusafont & Kurtén 1976, fig. 2; Tobien 1955, fig. 3). In I. atticus the metaconid is much less prominent than the protoconid, but is still of similar size to the entoconid (Helbing 1932, fig. 3). In the smallest of the A. africanum M,’s (L50006) what is interpreted as a vestigial metaconid is still present. It is completely overshadowed by the proto- conid, and, unlike its counterpart in J. atticus, is smaller than the entoconid. The metaconid is absent in all other M,’s from Langebaanweg, and, apparently also from all other recorded Agriotherium M,’s. The progressive reduction and eventual loss of the M, metaconid in the Ursavus—Indarctos—Agriotherium lineage is illustrated in Figure 36. rs ag Fig. 36. M,’s. A. Ursavus primaevus (after Crusafont & Kurtén 1976, fig. 14). B. Indarctos arctoides (after Tobien 1955, fig. 3). C. Indarctos atticus (after Helbing 1932, fig. 3). D. Lange- baanweg Agriotherium (D1—L50006; D2—L50004; D3—L50446). All are lingual views except C, which is a buccal view with the buccal talonid cusps omitted. e—entoconid; m—metaconid; p—protoconid. If the largest of the posterolingual M, cusps in Agriotherium is indeed the entoconid (i.e. a talonid cusp), rather than the metaconid (i.e. a trigonid cusp), this would account for Tobien’s (1955: 14) observation that the “Metaconid’ of Agriotherium is ‘niedriger und starker zuriickgeschoben’, while in /ndarctos it is 84 ANNALS OF THE SOUTH AFRICAN MUSEUM ‘héher und naher an das Protoconid gestellt’. The cusp which Tobien believed to be the metaconid in Agriotherium may simply be a well-developed entoconid in more or less its usual position, while in Jndarctos the metaconid was correctly identified as such, and it, too, is in its usual position. The metaconid in /ndarctos has a counterpart on the buccal side of the talonid, this being a small cusp situated between the protoconid and hypoconid. It is present in J. vireti (Crusafont & Kurtén 1976), I. atticus (Helbing 1932), I. oregonensis (Dalquest 1969), and other specimens, although traces of its presence may be obliterated by wear. A vestige of this cusp may be represented in the A. palaeindicum specimen, GSI-—D8, by a slightly inclined ridge anterior to the hypoconid. A similar ridge is present in the M, of A. africanum, although in this species it takes the form of an undemarcated horizontal extension of the hypoconid. It meets with the posterobuccally directed keel of the protoconid. L50006 is also unusual in lacking this keel and in having the hypoconid ridge linked directly with the vestigial metaconid. The reduction and loss of the M, metaconid in the Indarctos—Agriotherium lineage thus appears to have been accompanied by reduction and loss of its buccal counterpart. The impression gained is that these cusps were ‘absorbed’ by the protoconid, which is considerably enlarged in Agriotherium. This is a manifestation of the development of the shearing cusps (paraconid and proto- conid) at the expense of the crushing cusps (talonid cusps) in the Indarctos— Agriotherium lineage. It is possible, however, that the traditional interpretation of the large posterolingual M, cusp in Agriotherium as the metaconid is correct. Another of the Langebaanweg specimens, L50446, is unusual in having the largest of the posterolingual cusps flanked posteriorly by two smaller cusps, whereas in all other specimens there is only one such cusp. An almost exact match of L50446 is the M, of the A. palaeindicum specimen, GSI-D8 (Lydekker 1884). If the largest of the posterolingual cusps in these specimens are metaconids, then the smaller cusps immediately posteriorly would be the entoconids. It would, therefore, be the entoconid which is lost in other Agriotherium M,’s, and this, rather than the loss of the metaconid, would have characterized the Indarctos— Agriotherium lineage. This hypothetical link would then be supported by the observation that the entoconid is as well developed in Agriotherium specimens such as L50446 and GSI-D8 as it is in some advanced Jndarctos (e.g. I. oregon- ensis—Dalquest 1969, fig. 4). In the case of the first alternative suggested above, the two smaller postero- lingual cusps in L50466 and GSI—D8 would be interpreted as a duplication of the single cusp in this position in other Agriotherium M,’s. A first-hand examination of all relevant specimens may be necessary before deciding which of the above alternatives is likely to be correct. Irrespective of which applies, a transition from the typical Indarctos condition to that typical of Agriotherium is documented by specimens from the Siwaliks and Langebaanweg. AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 85 Another trend evident in the lower carnassials of Indarctos and Agrio- therium is that of a reduction in their relative lengths with time (Table 17). Only the problematical Siwaliks’ specimens GSI-D8 and D9 are anomalous in terms of their length: breadth ratios. The arrangement and morphology of the M! cusps in advanced Jndarctos and Agriotherium is virtually identical. The M? of A. africanum differs from that of I. atticus only in being relatively shorter and higher crowned, although it may be indistinguishable from more advanced Indarctos (e.g. the Florida specimen) in these respects. In most, and perhaps all, Indarctos M?’s the posterior keels of the meta- cone and hypocone are linked across the posterior end of the tooth by a low ridge of enamel. This feature is present but less obvious in A. africanum, and probably all other Agriotherium, and this region of the tooth also differs in being noticeably shorter than in the corresponding part of the J. atticus M?. In other words, there is a very short ‘talon’ region in the latter, and it is reduction of this feature which contributes to the overall shortening of M! in Agriotherium. TABLE 17 Length : breadth ratios of Indarctos and Agriotherium My,’s. Loca ity and/or TAXON NUMBER LENGTH | BREADTH L:B MEANS Agriotherium africanum .\ { L45062 1 1,82: 1 145062 2 il 150004 oat | L50006 21 150446 Bil 145114 al L50005 Bil Agriotherium insigne .\ Montpellier! oa 17a Montpellier? aI Agriotherium palaeindicum . .| GSI-D8? Balt 2,00 : 1 Indarctos punjabiensis .| GSI-D9? gt 1,88 : 1 Indarctos atticus . .| NMB-Sam31 Bal! 1,90:1 NMW-Saml 912/29? v1 Concud? al Indarctos arctoides* .| Westhofen il 1,93: 1 Pfaffstetten | Montredon ori Indarctos viretit . .| Can Llobateres VP633 $ 2,01 : 1 VP647 VP647 Can Purull Type (pontiensis) Type 1 Viret 1939; * Lydekker 1884; ° Thenius 1959; +4 Crusafont & Kurtén 1976. 86 ANNALS OF THE SOUTH AFRICAN MUSEUM This is a significant difference, because once again a specimen of one genus is known which has the character of the other. An Jndarctos-like post-metacone lengthening is one of the few features visible on the incomplete and badly restored M? of the Montpellier A. insigne specimen, NUB-MP549. The ‘primitive’ M! of this remarkable specimen is yet another of its Indarctos ‘characteristics’, others being its relatively small size, double-rooted and trans- versely orientated P’, and double-cusped P* protocone lobe. This specimen also has the P*, M! and M2? lower crowned than any of their counterparts in the Langebaanweg assemblage. However, it is identified with Agriotherium because its M? is of the Agriotherium type and distinct from that of all specimens referred to Indarctos (see below). Had the M? of this specimen not been preserved, it may well have been referred to Jndarctos. Montpellier could therefore have erroneously acquired the distinction of being the only locality where Indarctos and Agriotherium occurred together. This imaginary situation is mentioned here to indicate the importance of NMB-MP549 as an Indarctos—Agriotherium ‘intermediate’, and to illustrate how easy it is to misidentify specimens belonging to late Indarctos and early Agriotherium. The possibility of similar confusion with certain Siwaliks specimens was mentioned above. Before dealing with the next tooth in the dentition, another parallel between Agriotherium and Thalarctos is mentioned. The M? of the latter is relatively shorter and higher crowned than that of U. arctos, its ‘structural’ ancestor (Kurtén 1964). This also applies in the case of at least some species of Arctodus (Kurtén 1967). The M, of A. africanum, and other Agriotherium, usually consists of two trigonid cusps side by side, flanked posteriorly by two similarly positioned talonid cusps. The M, of Jndarctos differs only in having two lingual talonid cusps, and in being relatively longer and narrower. There are, however, Agrio- therium specimens which are Indarctos-like in both these respects. The M, of the A. palaeindicum specimen GSI—D8 has two lingual talonid cusps, while vestiges of a second cusp are visible in the Langebaanweg specimens L54114/ L46563 and L50007. The length:breadth ratios of the M,’s of GSI-D8 and L45114/L46563 are comparable to those of I. atticus M,’s (Table 18). By contrast, this ratio in the J. punjabiensis specimen GSI-D9 is the same as that of the mean of the A. africanum sample. Curiously, the A. africanum M,’s which are most Jndarctos-like in respect of talonid cusps and proportions are those which are most specialized in terms of their trigonid development (see p. 27). Consequently, these specimens should not be considered as good Jndarctos—Agriotherium intermediates. In addition, it could be argued that Pilgrim (1932) was incorrect in reversing Lydekker’s (1884) identifications of GSI-D8 and D9 and that they, too, are not ‘inter- mediates’ in the sense claimed above. This may, indeed, be the case, but the fact remains that however these (and other) Siwaliks specimens are identified, they exhibit a combination of Indarctos and Agriotherium characters. The dentitions of Indarctos and Agriotherium are perhaps most clearly AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 87 TABLE 18 Length : breadth ratios of Indarctos and Agriotherium M,’s. Locatiry and/or NUMBER LENGTH | BREADTH L:B 145062 145062 150004 150003 150007 145114 146563 Montpellier? Agriotherium africanum . Cie} 'afel (int hele) nie oie pnh | pemed pemek pemek peek feek feed fe Agriotherium insigne Agriotherium palaeindicum . Indarctos punjabiensis Indarctos atticus GSI-D8? GSI-D9? NMB-Sam31 NMW-Sam1 912/298 Montredon Westhofen Pfaffstetten Can Llobateres VP633 VP640/1 VP647 VP647 Can Purull Type Type (pontiensis) 1,39: 1 1,29:1 1,39:1 Indarctos arctoides* 1,41:1 Indarctos vireti* 1 Viret 1939; * Lydekker 1884; * Thenius 1959; 4 Crusafont & Kurtén 1976. distinguished from one another by the fact that the M? of the latter lacks a talon, whereas in Jndarctos this feature, although variably developed, is always present. The postero-internal (talon) region of the Agriotherium M? is also variably developed, the Langebaanweg assemblage being useful in indicating the varia- tion possible in a single population (Fig. 12). Of particular interest is the isolated M?, L47698, in which there is a marked posterior projection of the postero- internal part of the tooth. This is here interpreted as the vestiges of the talon characteristically present in the M? of Indarctos. L47698 is remarkably similar to the M?, GSI-D12, referred to J. punjabiensis by Lydekker (1884, fig. 6), and these two specimens represent an intermediate between the conditions typical of Indarctos and Agriotherium. The progressive shortening and broadening of M”’s in the Jndarctos—Agriotherium lineage is indicated by the data in Table 19. Erdbrink’s (1953: 582) view that there is ‘at best a beginning of a [M?] talon .. . in A. insignis’ is here regarded as the reverse of the true situation. A. insigne, like A. africanum, sometimes has the vestiges of a M? talon. As with M}!, the arrangement and morphology of the four principal cusps of M? in advanced Indarctos and Agriotherium are very similar. The paracones 88 ANNALS OF THE SOUTH AFRICAN MUSEUM TABLE 19 Length : breadth ratios of Indarctos and Agriotherium M*’s. LOCALITY or NUMBER LENGTH} BREADTH L:B Agriotherium africanum .|_ 48577 0,84: 1 148564 0,84 : 1 141404 0,85 : 1 L45062 0,87: 1 145062 0,88 : 1 L45137 0,93 : 1 L45137 0,94 : 1 L47242 1,00: 1 L47698 1,03 : 1 Agriotherium insigne NMB-MP549 0,89 : 1 ail Type* 1,03: 1 Indarctos punjabiensis GSI-D12? 1,06: 1 ou Indarctos atticus NMW-Sam1912/298 1,13:1 cat NMB-Sam31 1,28: 1 Indarctos arctoides Montredon* 1,29:1 al Montredon? 1,43 :1 Orignac* 1,28:1 Gau—Weinheim® 1,37:1 Indarctos vireti VP6334 1,47: 1 | VP646? Sil 1 Frick 1926; *Lydekker 1884; *Thenius 1959; 4 Crusafont & Kurtén 1976; *°Tobien 1955. and metacones are conical, with distinct anterior and posterior keels, the protocone is ridge-like, and, with the small hypocone, is lower than the buccal cusps. The M*’s of Indarctos and Agriotherium illustrate very clearly a progressive development from /. vireti, through I. arctoides, I. atticus, I. punjabiensis to Agriotherium, with A. africanum taken as an example of its genus (Fig. 37). The recorded M?’s of advanced North American Indarctos all have well- developed talons, that of the J. cf. oregonensis specimen recorded by Merriam & Stock (1925) being the most prominent. It is, however, from the same deposits as another J. oregonensis M?, which is a larger tooth but with a relatively smaller talon (Merriam et al. 1925). The latter is here interpreted as belonging to a male, while the smaller specimen with the larger talon belongs to a female. This raises the possibility that the teeth of female Indarctos and Agrio- therium tend to be more ‘primitive’ than those of the larger males. This explana- tion applies in the case of the ‘primitive’ A. insigne specimen, NMB-MP549, discussed earlier, which represents a small individual, apparently a female. Another example is the A. africanum M, with the vestigial metaconid, L50006. The female I. cf. oregonensis is also ‘primitive’ compared with other North American Jndarctos in having a relatively small P* parastyle and a relatively elongated M1. It does not, however, have a double-cusped P* protocone lobe. This, together with the fact that NMB-MP549 does not have a talon on M?, AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 89 A B c D1 D2 D3 Fig. 37. Ms. A. Indarctos arctoides (after Helbing 1932, fig. 7). B. Indarctos atticus (after Helbing 1932, fig. 1). C. Indarctos punjabiensis (after Lydekker 1884, fig. 6). D. Langebaan- weg Agriotherium (D1 —L47698; D2—L45137; D3 —L41404). indicates that females were not necessarily ‘primitive’ in all respects. Never- theless, since increasing size was a characteristic of the Indarctos—Agriotherium lineage, and consequently large size was itself an ‘advanced’ character, it is to be expected that larger individuals would also be ‘advanced’ in other respects, and vice versa. This is a further indication that allowance must be made for appre- ciable variation in characters in studies on Indarctos and Agriotherium. The fact that known North American Indarctos M*’s appear more ‘primitive’ than the I. punjabiensis specimen, GSI-D12, does not necessarily mean that they must be older. There is other evidence to support this opinion. Once again there is a parallel between Agriotherium and Thalarctos, since the latter also has the M? talon reduced, and it may sometimes even be absent. The study by Kurtén (1964) has shown that there is considerable variation in Thalarctos in this respect even within a single population. Particularly remarkable is a specimen in which the right M? talon is reduced, while that of the left M? is completely absent (Kurtén 1964: 17, pl. 4A). Such examples are exceptional, but presumably if samples of Indarctos as large as Kurtén’s Thalarctos sample (n = 113) were available, similar specimens may be found. This suggests that undue reliance may have been placed on the value of the M? talon as a distinguishing charac- teristic of Indarctos. The M, of Indarctos and Agriotherium is less commonly represented than other posterior cheek teeth, evidently because it is single-rooted and easily 90 ANNALS OF THE SOUTH AFRICAN MUSEUM separated from the mandible. It is nevertheless clear that this tooth was reduced in length in the /ndarctos—Agriotherium lineage to a degree comparable to that of M®. In J. vireti the Mg is markedly elongated anteroposteriorly and has a pronounced posterior tapering (Crusafont & Kurtén 1976). The M, of J. atticus is similar, but is less elongated, while that of Agriotherium is usually only as long as it is broad. The latter, however, is still Jndarctos-like in having a pro- nounced posterior tapering. The A. africanum Mg, L50981, is very like those of the Vienna and Basel /. atticus specimens (Thenius 1959; Helbing 1932) in shape. The reduction of Mg is another development shared by Agriotherium and Thalarctos. Although mention has been made of differences in proportions in certain of the posterior cheek teeth discussed above, this matter warrants more detailed attention, since these differences are directly related to changes in tooth mor- phology and are as significant from a phylogenetic point of view. In addition, metrical data have the advantage of being easily represented diagrammatically and can thus give a visual impression of certain evolutionary changes in the hypothetical Ursavus—Indarctos—Agriotherium lineage. Five Old World species were selected to represent successive stages in this lineage. They are Ursavus primaevus, I. vireti, I. arctoides, I. atticus, and A. africanum. Individual species do not necessarily represent the stock from which the next in the series was derived, but are simply structurally suited to such a role. The lengths and breadths of their posterior cheek teeth are plotted in Figures 38—40. It is clear from these figures that U. primaevus is in an isolated position, whereas there are similarities between J. vireti and J. arctoides on the one hand, 10 SE 10 -E p4 BREADTH / p4 LENGTH 2—/ -D 2 -D 1 -C 2 -c 2 -B 2 -B -A 3 -A (ee ee ee a ee ee eee ee ee ee ee SS Sd 8 10 12 14 #16 18 20 22 24 26 1214 16 18 20 22 24 26 28 30 32 34 36 M, LENGTH =) 3 -C 3 -B -A 5 s40 12-4416 8202222 1) 20 22 3A 28 20, S02 34, 36> GU TAUEROEAS Fig. 38. Lengths and breadths of P* and M,. A. Ursavus primaevus. B. Indarctos vireti. C. I. arctoides. D. I. atticus. E. Agriotherium africanum. Sample sizes, ranges and means are indicated. Data from Crusafont & Kurtén (1976), Thenius (1959) and this report. AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 91 6 -E 6 -E mM! BREADTH mM! LENGTH 2 -D -D 3 -C -C -B —B 4 -A 4 -A Le en Se Eee eee ee ee eee (ee ee eS eee ee eee ee 10 12 14 16 18 20 22 24 26 28 30 12 14 16 18 20 22 24 26 28 30 32 -E -E =D -D -c -c —B -B -A —-A ee ee Se ES eS ee es ee ee Ee ee 8 10 12 14 16 18 20 22 24 26 14.16 18 20 22 24 26 28 30 32 34 Fig. 39. Lengths and breadths of M' and M, of some Ursidae, (see Fig. 38 for key). and J. atticus and A. africanum on the other. The isolated position of U. primae- vus is not surprising since it is a contemporary of J. vireti (Crusafont & Kurtén 1976), and its phylogenetic connection with the /ndarctos—Agriotherium lineage must be indirect. The fact that this lineage is divisible into two parts on the basis of cheek tooth size (and other evidence) is significant. The first is comprised of earlier, smaller and more primitive forms (i.e. J. vireti-I. arctoides), and the second of later, larger and more advanced forms (i.e. J. atticus—A. africanum). 9 -E 9 -E M2 BREADTH M> LENGTH 2 -D =D 3 -C -c —B8 -B 3 =A. 3 -A ee ee ee ee eee a SS SS eS ee ee es ee ees | 10 12 14 16 18 20 22 24 26 28 30 32 14.16 #18 20 22 26 26 28 30 32 34 -E -E M, LENGTH =D 2 -D -B 4 -B -A 2 -A (aa) js ts 8 10 12 14 #16 «18 «20 8 10 12 14 16 18 20 22 Fig. 40. Lengths and breadths of M? and M; of some Ursidae, (see Fig. 38 for key). 92 ANNALS OF THE SOUTH AFRICAN MUSEUM It is only in the last segment of the lineage that there are deviations from otherwise general trends in the lineage as a whole. The most obvious of these general trends was the increasing breadth of the posterior cheek teeth. This was probably more or less in proportion to the overall size increase of the taxa concerned. However, the situation in respect of the lengths of these teeth is more complex. Only in the case of P* was there a tendency to increase length throughout the Ursavus—Indarctos—Agriotherium lineage. This is also a reflection of the increasing size of successive members of the lineage. In addition, it may be interpreted as indicating the increasing functional importance of the principal shearing tooth in the upper dentition, with the increased lengths of P* in the last stage of the lineage (i.e. J. atticus—A. africanum) being largely due to the development of the parastyle. It might be expected that there would have been a corresponding increase in the length of the lower carnassial. The fact that there is a barely perceptible increase in the mean length of the A. africanum M, over that of J. atticus may simply be due to the composite nature of this tooth, the increased length of the shearing element (i.e. the trigonid) being obscured by the reduction of the talonid. By contrast, in the case of M? and M, the lengthening trend evident in most of the lineage is reversed in the /. atticus—A. africanum segment. This is a mani- festation of the reduced importance of the crushing function of the most pos- terior cheek teeth in the last stage of the lineage. The situation in respect of the lengths of the intermediate teeth, M! and M,, was itself intermediate, with lengths remaining static once the J. atticus stage was reached. The overall impression gained from the study of the morphology and dimensions of the teeth of successive members of the Ursavus—Indarctos— Agriotherium lineage is that this lineage can be separated into two parts. Up to the I. atticus stage the taxa concerned were generalized ‘omnivorous’ ursids, but thereafter there were modifications of both tooth morphology and pro- portions being manifested which indicate adaptation to a more carnivorous diet. It may thus be more appropriate to think of ‘advanced Jndarctos’ as being “primitive Agriotherium’. Since the transition from Jndarctos to Agriotherium was gradual in the sense that there are specimens which exhibit ‘characteristics’ of both genera, it is far from obvious on what grounds the distinction between these genera is to be made. Earlier attempts to find distinguishing characters are now seen to have been inadequate. For example, lists of such characters given by Pilgrim (1932: 42) and Tobien (1955: 14) can be misleading and should be used with caution, if they are to be used at all. The formulation of mutually exclusive diagnoses for these genera has become difficult, if not impossible. There would, of course, be no problem if these diagnoses were to be based on a primitive Indarctos, such as I. vireti, and any of the species presently identified with Agriotherium. However, when all recorded intermediate forms are taken into AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 93 account, the diagnoses break down under a welter of qualifying statements. The situation could only become more confused if all the Indarctos and Agriotherium assemblages from the critical 5-8 Ma period were as large, or larger, than that from Langebaanweg. Even if this is never the case, the Langebaanweg assem- blage has clearly shown that agriotheriines are as variable as other ursids, and consequently there can no longer be complacency about supposed ‘charac- teristics’ of individual representatives of this group. This situation is here interpreted as indicating an ancestor—descendant relationship between Indarctos and Agriotherium beyond all reasonable doubt. In concluding his study of the Florida Indarctos, Wolff (1978: 11) stated that there are ‘several cranial features [which] may indicate a relatively closer relationship between Ailuropoda and Indarctos than with other bears, although other possibilities exist’. In the light of the preceding discussion and the fact that the Florida Indarctos skull is even more like that of A. africanum than Ailuropoda, the ‘other possibilities’ must certainly include Agriotherium. The similarities between the skulls of advanced Jndarctos, particularly the Florida specimen, and Agriotherium are so great that it is inconceivable that they are only distantly related. Wolff’s concluding remarks refer specifically to ‘great enlargement of the cheek teeth’, ‘expansions of the zygomatic arches and sagittal crests’, and the ‘absence of the alisphenoid canal’ as shared characters indicative of the ‘close relationship’ between Jndarctos and Ailuropoda. The fact that Indarctos and Agriotherium also share them is equally significant, especially taken in conjunction with all the other evidence cited above. It seems superfluous at this stage to consider alternative theories on the origin of Agriotherium, but one is mentioned here since it apparently still has some support. The fact that Agriotherium and the Hemicyoninae have a premasseteric fossa on the mandible in common has contributed to the belief that they are closely related (e.g. Frick 1926). There is also a superficial similarity between the dentitions of Agriotherium and hemicyonines, and both are supposedly canid- like, rather than ursine-like. This is indeed so in the case of hemicyonines, but in terms of the arrangement, morphology and size of individual cusps, the teeth of Agriotherium are far more like those of advanced Indarctos than any hemi- cyonine. It could, of course, be argued that the rather generalized hemicyonines are structurally suitable in both cranial and postcranial characters to be ancestral to Agriotherium. This more tenuous hypothesis is considered less likely than the alternative suggested here. It was indicated earlier that the known temporal ranges of Indarctos and Agriotherium are in accord with the theory that the latter was derived from the former. In the case of the Hemicyoninae—Agriotherium alternative the situation is much less convincing because of an apparent, or actual, gap between the recorded histories of the taxa concerned. In Europe Agriotherium is first recorded from the Ruscinian, whereas hemicyonines were extinct by the end of the Vindobonian (Table 20). This 94 ANNALS OF THE SOUTH AFRICAN MUSEUM TABLE 20 The occurrences of Hemicyoninae and Agriotherium in Europe. LOCALITY MAMMAL AGE! | MEIN ZONE? Ruscinian MeL ka Moc ee Hemicyon sansaniensis, Wintershof- Burdigalian Hemicyon goeriachensis | West, Sansan, to etc.® Goriach, La Vindobonian Grive-St-Alban, APPROX, Agriotherium insigne Steinheim etc® 1Van Couvering 1972, Fahibusch 1976; * Mein 1975, Fahlbusch 1976; *Heizmann 1973. means that there was a period of about 8 m.y. for which no possible inter- mediate between hemicyonines and Agriotherium is known in Europe. Much the same applies in Asia and North America. It is unlikely that this could be due to a defective fossil record, since the period in question covers the Vallesian and Turolian (and their equivalents), of which the faunas are moderately to very well known. It is possible that the Hemicyoninae—Agriotherium transition took place in Africa, where 5-13 Ma faunas are poorly known, but negative evidence is hardly convincing support for a theory. It is much more likely that the characters Hemicyoninae and Agriotherium have in common are due to convergent evolution. Parallel and convergent evolution often complicate studies of relationships, and the Ursidae are a group where this complication definitely exists. Early in the history of the Ursidae, the Amphicyoninae and Hemicyoninae evolved along parallel lines, while later the same applied to the Ursinae and Tremarctinae. Agriotherium is of particular interest in this connection, since not only is it intermediate in age between amphicyonines/hemicyonines and ursines/tremarctines, but it has in certain respects paralleled members of both sets of subfamilies. The superficial similarities between Agriotherium and large amphicyonines and hemicyonines (e.g. Amphicyon major, Dinocyon thenardi) are striking enough to suggest that these animals were ecological vicars. On the other hand, Agriotherium, A. major and D. thenardi (and better known hemicyonines) also differ from one another in certain respects, which can be explained by their having evolved at different times from different ancestors. Amongst the Ursinae, Thalarctos has now been mentioned several times as having evolved characters comparable to some in Agriotherium. In this instance there is no possibility of the shared characteristics being due to a close relationship. They are simply explained by the fact that both Agriotherium and Thalarctos are essentially carnivorous forms which evolved from omnivorous ancestors (Hendey 1972: 122). The relationship between Thalarctos and the Ursus arctos group has been well documented (Thenius 1953; Kurtén 1964), AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 95 and is here regarded as a parallel of the relationship between advanced Indarctos and Agriotherium. A more detailed search for parallelisms between these two sets of taxa may well be worth while. The obvious differences between Agrio- therium and Thalarctos are not unexpected, since their ancestral forms are markedly different from one another. In addition, Thalarctos has as yet had a relatively brief history compared with that of Agriotherium. Given time it would no doubt become increasingly distinct from U. arctos, and perhaps become even more Agriotherium-like. Agriotherium and advanced Indarctos are also paralleled in some respects by tremarctines, especially the large extinct species. For example, several references have been made above to similarities between the skull and skeleton of A. africanum and Arctodus simus. Merriam & Stock (1925: 5) found that there are certain characters ‘in which Tremarctos and [Arctodus] show distinctly closer affinity to [Agriotherium] and its allies of the Pliocene than is seen in Pleistocene and Recent bears of the genus Ursus’. Others have thought this significant. For example, Erdbrink (1953) suggested that Tremarctos was closely related to the ursines, while the larger tremarctines were derived from advanced Jndarctos. Kurtén (1966: 7) disagreed, and concluded that although the ‘earlier history of Arctodus is poorly documented . . . there can be little doubt that it is a tremarctine, and not a member of the Agriotheriinae’. The Indarctos—Agriotherium lineage was but one of several evolutionary developments amongst the Ursidae during the latter part of their history. As indicated above, it was a development which paralleled that undergone earlier by large amphicyonines and hemicyonines, but it, too, was ultimately unsuc- cessful and by the end of the Tertiary Agriotherium was on the verge of extinc- tion, if not already extinct. It had no descendants. Other later ursid lineages, however, were more successful. This applies particularly in the case of the one which gave rise to the Ursinae, which under- went a spectacular radiation beginning in the Pliocene. As with Indarctos— Agriotherium, the Ursinae also stemmed from Ursavus, and Thenius (1977 and earlier papers) has suggested U. ehrenbergi as the likely ancestral form. The Tremarctinae were less successful than the Ursinae, having been con- fined to the Americas, and eventually being supplanted in North America by the Ursinae (Kurtén 1966, 1967). Their origins were recently considered by Thenius (1976), who suggested that they, too, stemmed from an Ursavus. Unfortunately, little is known of early tremarctines, but no doubt more will yet be learnt of the origins and early history of this group. The origin of the giant panda, Ailuropoda melanoleuca, has long been a controversial issue. The earlier views that Ailuropoda is an ursid and not a procyonid, and that it had an agriotheriine ancestor (Hendey 1972), are main- tained here, although it now appears that it may have stemmed from Ursavus and not /ndarctos as previously suggested. As indicated earlier, the skulls of Ailuropoda and advanced Indarctos have many characters in common which suggest that they are more closely related 96 ANNALS OF THE SOUTH AFRICAN MUSEUM to one another than to other bears. However, the latter had already undergone specializations in the direction of Agriotherium which render it structurally unsuitable as a stem form for Ailuropoda. For example, advanced Jndarctos has reduced anterior premolars, while those of Ai/uropoda are unusually large for an ursid. This objection does not apply in the case of earlier, unspecialized Indarctos (see Hendey 1972, table 1), but since such species have much in common with Ursavus, the latter must also be taken into account when con- sidering the origin of Ailuropoda. One of the less well-known species of Ursavus is U. depereti of the European Turolian, which is in some respects atypical of the genus (see Heizmann 1973). This applies particularly to the P*, and it is the unusual characteristics of this tooth which suggest that U. depereti may have a direct phylogenetic connection with Ailuropoda. Casts of upper teeth of U. depereti from Soblay in France were recently examined in the Naturhistorisches Museum, Basel. The originals are in Lyon and were described by Viret (1949) and Viret & Mazenot (1949). Two P?’s are represented and they are remarkable for two reasons. Firstly, unlike the P*’s in other Ursavus species, the Soblay specimens have large parastyles. This cusp is proportionately even larger than those in advanced Indarctos and in Agrio- therium, in which the parastyles are overshadowed by large and high-crowned paracones. The large parastyle and relatively low-crowned paracone of U. depereti are reminiscent of the situation in Ailuropoda, and differ from that in all other ursids. Secondly, the U. depereti P* has an enlarged, double-cusped protocone lobe, which differs from that in Indarctos and Agriotherium in being more anteroposteriorly elongated and regular in occlusal outline. In addition, in the Soblay specimen, AA52 (Viret & Mazenot 1949, pl. 1 (fig. 6)), both cusps are ridge-like rather than pointed and are of more or less equal size. In all these respects the U. depereti P* protocone lobe is Ailuropoda-like, and once again it differs from that in all other ursids. The M? and M? of U. depereti are also basically similar in morphology to their counterparts in Ailuropoda. However, the upper molars are also similar to those of other Ursavus species and Jndarctos, and are thus less significant in indicating a relationship with Ailuropoda. A possibly important charac- teristic of the M? figured by Viret & Mazenot (1949, pl. 1 (fig. 4)) is the rugose enamel of the occlusal surface, which could foreshadow the ‘richly tuberculate’ condition of the Ailuropoda M? (Davis 1964: 127). The M, and M, of U. depereti also have the basic morphology charac- teristic of other Ursavus and Indarctos, although the M, is distinct, and more Ailuropoda-like, in having the metaconid more anteriorly situated (Schlosser 1902, pl. 2 (figs 20, 23)). Slender though the evidence may be, U. depereti appears to be structurally better suited than any other recorded fossil ursid to fill the role of ancestor to Ailuropoda. In addition, it is also temporarily and geographically well suited to this role since it is from the continent on which the early evolution of living AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 97 bears was centred and it dates from a period when the radiation of later bears was just beginning. The absence of any intermediate forms may simply be due to the relatively poor Pliocene record in Asia, the continent to which Ailuropoda melanoleuca is confined (Chorn & Hoffmann 1978). Relevant fossils of Plio- cene age, as well as better knowledge of U. depereti, are required to test this theory of Ailuropoda origins. While the ultimate origins of both Ailuropoda and the Indarctos—Agrio- therium lineage are uncertain, it is evident that these genera are more closely related to one another than to the Ursinae and Tremarctinae. The radiation of the Ursidae, like that of some other mammalian families, was thus characterized by repetitious evolutionary developments, and the correct identification of relationships requires careful study of the fossil record. Ignorance of the details of this record may well lead to misinterpretation of the significance of similar, but independently evolved characteristics in ursid taxa. Kurtén (1967: 5) has found, for example, the analogy between European Miocene Jndarctos and North American Pleistocene Arctodus to be ‘truly astonishing and an example of the déja vu experiences so familiar to the student of fossil bears’. The relationships suggested here are indicated in Figure 41. The named late Tertiary species are all from Europe and consequently the European sub- divisions of this period are used. URSINI TREMARCTINI AILUROPODINI HOLOCENE & (Ursus, Thalarctos, Euarctos, Helarctos. ‘ PLEISTOCENE Melursus) ‘Ursavus’ Indarctos x | depereti atticus TUROLIAN Nas Ursavus sp. Ursavus (Soblay) ehrenbergi , ’ A | as A URSAVINAE Ursavus is Indarctos Indarctos primaevus +: anthracitis arctoides 1 Ursavus 4, Indarctos brevirhinus Z soras- ss ViretL: ee areas VINDOBONIAN ek Ursavus brevirhinus HEMICYONINAE eee elmensis Foueocene | AMPHICYONINAE oN Cephalogale Fig. 41. Suggested relationships of some Ursidae. (Plionarctos. Arctodus, (Ailuropoda) Tremarctos) Agriotherium insigne AGRIOTHERIINAE PLIOCENE MIOCENE 98 ANNALS OF THE SOUTH AFRICAN MUSEUM NOMENCLATURE The suprageneric classification of the Ursidae adopted here (Table 21) is a modification of an arrangement suggested earlier (Hendey 1972: 119), and is based on the phylogenetic relationships indicated in Figure 41. TABLE 21 A classification of the Ursidae. SUBFAMILY TRIBE GENERA, SPECIES Amphicyoninae . : ; Not subdivided t Amphicyon, + Cynelos and tothers Hemicyoninae . ; 4 : Not subdivided {| Hemicyon, | Dinocyon and t others Ursavinae Not subdivided t Ursavus : a Agriotheriini {Indarctos, + Agriotherium Agriotheriinae . . . . : — - - — Ailuropodini t‘Ursavus’ depereti, Ailuropoda Weise Tremarctini {Plionarctos, + Arctodus, Tremarctos Ursini Ursus, Thalarctos, Euarctos, Helarctos, Melursus + = extinct In preceding sections of this report the generally accepted practice of allowing the ursines and tremarctines subfamilial status has been followed. With this as a precedent, it follows that the agriotheriines and Ailuropoda each warrant similar status. Some of those who have included Ailuropoda in the Ursidae have, indeed, placed it in a separate subfamily (e.g. Pilgrim 1932). Similarly, the agriotheriines have been widely recognized as a valid subfamily (e.g. Kurtén 1966). A disadvantage with this arrangement is that it does not indicate the apparently close relationships between the ursines and tremarctines on the one hand (Kurtén 1966, 1967; Thenius 1976), and the agriotheriines and Ailuropoda on the other (Hendey 1972; Wolff 1978; this report). This problem is overcome by reducing the status of the four subfamilies to tribes, and then separating them into two subfamilies, namely, the Agriotheriinae (Agrio- theriini and Ailuropodini) and the Ursinae (Ursini and Tremarctini). Except for the position of the Tremarctini, this was the arrangement suggested earlier (Hendey 1972), and for which there has been some support (e.g. Chorn & Hoffmann 1978). There is also a difference now in the taxa constituting the Agriotheriinae. Firstly, in the classification suggested here, Ursavus depereti is included in the Ailuropodini, since it is regarded as the likely ancestor of Ailuropoda. This means that it can no longer be referred to Ursavus, but must either be given a new generic name, or be referred to Ailuropoda itself. The latter course is favoured here, but in view of the uncertainties still surrounding this species neither of these alternatives is followed, and the doubtful generic identity of ‘Ursavus’ depereti is indicated by quotation marks. AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 99 Secondly, if the Ursini, Tremarctini and Ailuropodini did, indeed, stem from the Ursavus group and are valid tribes, then the Indarctos—Agriotherium group of the Agriotheriini must be given the same status. This means that Ursavus, which is generally regarded as an agriotheriine (e.g. Kurtén 1966; Hendey 1972), has to be excluded from the tribe. The various species of Ursavus, excluding ‘Ursavus’ depereti, are accordingly placed in a new subfamily, the Ursavinae. Since this subfamily is not established on the same criteria as the others, it constitutes an unsatisfactory element in the classification as a whole. When their relationships are better understood, it may be possible to assign the species of Ursavus to other tribes, just as ‘Ursavus’ depereti has been assigned to the Ailuropodini. The need for this additional subfamily will then fall away. The classification of more primitive ursids, or ursid-like carnivores, namely, the hemicyonines and amphicyonines, was not investigated since they are largely irrelevant to the present study. They are mentioned here for the sake of completeness, but the conclusions regarding their status are tentative. The Hemicyoninae are generally regarded as an early off-shoot of primitive ursid stock (probably Cephalogale—see Erdbrink 1953, fig. 61; Hendey 1972, fig. 1), and they are here included in the Ursidae. The connection between the Amphicyoninae and later ursids is more remote, and the classification of this group is more controversial. They have been variously classified as a separate family (e.g. Hunt 1972), a subfamily of the Canidae (e.g. Kuss 1965), and as an ursid subfamily (e.g. Ginsburg 1977). The latter course is followed here. The nomenclature of the species constituting the Agriotheriini is a far more problematical matter than the suprageneric classification of the Ursidae. The conclusion that Agriotherium is directly descended from Jndarctos with certain specimens exhibiting a combination of ‘characteristics’ of both genera, raises the possibility that they are congeneric, with Agriotherium the senior synonym. This situation had been foreseen even before Indarctos became well established in the literature (Pilgrim 1914; Merriam ef al. 1916). It can be considered in relation to a similar situation involving the brown and polar bears. Ewer (1973) and Van Valen (1978) have recently discussed the relative merits of opposing opinions on the generic identity of the polar bear. According to one widely held opinion the polar bear and brown bear are congeneric (i.e. both belong to the genus Ursus), while the opposing view is that the polar bear represents a separate genus (i.e. Thalarctos). Ewer and Van Valen favoured the latter alternative, a view which is supported here. A problem with this arrangement is that by one widely accepted criterion Thalarctos does not merit separate generic status. Thalarctos maritimus and Ursus arctos are known to produce fertile hybrids (see Van Gelder 1977). Van Valen (1978: 292) dismissed this objection in the grounds that ‘lack of intersterility per se’ is of ‘low evolu- tionary importance’ a fact for which Vrba (1979) has found supporting evidence amongst the Bovidae. 100 ANNALS OF THE SOUTH AFRICAN MUSEUM It is clear from their habitats and habits that Thalarctos and U. arctos are set on distinct evolutionary paths, and that given time they may well warrant generic separation by any standards. It is fortuitous that at present their diver- gence is not far advanced, although there is no doubt that this divergence does, in fact, exist. There will always be a problem in classifying species, both living and extinct, which are on separate lineages, but which are still close to a common ancestor. In instances where divergence is certain, and in the case of the brown and polar bears it is, then classification by ‘clade’ rather than ‘grade’ is preferable. As indicated earlier, Thalarctos and Agriotherium are similar in the sense that both are essentially carnivorous forms derived from omnivorous ancestors. There is an apparent difference, however, in the evolutionary histories of these two genera. In the case of Thalarctos there was a divergence from the U. arctos group after which two lineages evolved independently, one with essentially carnivorous elements, and the other continuing with essentially omnivorous ones. In the case of Indarctos—Agriotherium it has been suggested above that the essentially omnivorous ancestor gradually adapted to give rise to the more carnivorous descendant forms and did not itself continue to exist as a separate entity. In other words, successive members of what may be regarded as a single lineage adapted their habits in response to a gradually changing environment (see p. 70). This may, of course, be a misinterpretation of the fossil record and there may have been a dichotomy during the history of Indarctos—Agriotherium similar to that of U. arctos-Thalarctos. For example, Crusafont & Kurtén (1976) may be correct in their interpretation of the early history of Indarctos, and I vireti may have given rise to both I. arctoides and the more Agriotherium-like I. atticus. According to this interpretation J. vireti-I. arctoides wouid be an equivalent of the U. arctos lineage, while J. atticus was the counterpart of Thalarctos. The nomenclatural implication is that J. atticus and other later Indarctos should be referred instead to Agriotherium, leaving only J. vireti, I. arctoides and the aberrant J. anthracitis as representatives of Indarctos. This arrangement is supported by the earlier suggestion that those species referred to throughout this report as ‘advanced Indarctos’, could as well be regarded as ‘primitive Agriotherium’. A minor difficulty which arises is that Indarctos would have to be replaced by another name, because the genotype (J. salmontanus) would then be identified as an Agriotherium. It could also be argued that since J. vireti is readily distinguishable from contemporary Ursavus, the evolutionary changes which were to culminate in Agriotherium were already being manifested early in the Vallesian. Thus J. vireti relative to contemporary Ursavus was the counterpart of the present-day situation involving Thalarctos and U. arctos. According to this interpretation all species presently identified with Jndarctos should be referred instead to Agriotherium. Both the above arrangements are less arbitrary than the existing one in AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 101 which it is only the most advanced agriotheriines which are referred to Agrio- therium. This is unsatisfactory because it does not reflect the realities of the situation. Nevertheless, no changes in this arrangement are proposed here. Undescribed Jndarctos and Agriotherium material from Asia and North America is available, and a decision regarding the status of these genera is best left until it, too, can be taken into account. The same applies in the case of nomenclature at the species level, although some changes which reflect opinions on relationships are proposed here. An appropriate starting point is with the European species, since they are amongst the best known and they represent the primary group from which agriotheriines elsewhere evolved. Four late Miocene species are recognized as valid. They are J. vireti, I. arctoides, I. atticus and I. anthracitis. I. atticus was apparently the first species to spread into Asia. J. maraghanus (De Mecquenem 1925) and J. lagrelii (Zdansky 1924) were apparently broadly contemporaneous with J. atticus and are here regarded as junior synonyms of this species. The situation in respect of Siwaliks species is more problematical. The identifications of Pilgrim (1932), although regarded as unsatisfactory in some respects, are accepted here. Various possibilities will have to be considered in a revision of the Siwaliks material. For example, J. punjabiensis and I. salmontanus may represent a single species which is conspecific with J. atticus. Alternatively this species, or perhaps only J. punjabiensis, may be more advanced. The possi- bility that advanced Siwaliks Indarctos is conspecific with A. palaeindicum will also have to be considered. A. sivalense will be mentioned below. Advanced Jndarctos is also represented in Africa (Sahabi—see p. 73), and in North America, where material has been identified as J. oregonensis (e.g. Dalquest 1969), J. nevadensis (MacDonald 1959), or not identified as to species (e.g. Wolff 1978). There is no reason to believe that recorded North American Indarctos represents more than one species. Whether it should be identified as I. oregonensis, or referred to one of the Old World species, is not certain. The North American species was an immigrant from the Old World during the Hemphillian (Repenning 1967), and must, therefore, be closely related to a contemporary Asian species. It is here regarded as definitely distinct from I. atticus, but if I. punjabiensis is a valid species, it might be conspecific with, and the senior synonym of J. oregonensis. Largely because of the uncertainties surrounding the Siwaliks species, I. oregonensis is here retained as a distinct species. In addition, the possibility was mentioned earlier that in certain respects North American /ndarctos evolved independently of, and at a more rapid rate than its counterparts in Europe and Africa (see p. 82). If this were a development peculiar to North America, rather than both Asia and North America, then it would be another reason for recognizing [. oregonensis as a distinct species. The situation in respect of recorded species of Agriotherium is complex, although the final solution with these species may be very simple. This solution, which is not advocated yet, is for all species of Agriotherium to be regarded as 102 ANNALS OF THE SOUTH AFRICAN MUSEUM junior synonyms of A. sivalense. Judged on the basis of the variation observed in the Langebaanweg Agriotherium, there may be no size or morphological grounds for recognizing more than one species of Agriotherium, However, it was decided to draw at least some distinctions on a geographical basis. European A. insigne, which is here taken to include A. intermedium (Stach 1957), is tentatively regarded as valid. It was suggested earlier that African Agriotherium was an offshoot from the primary European lineage, and although A. africanum is almost indistinguishable from A. insigne, they are not regarded as conspecific, since the former is likely to be a descendant of north African Indarctos and was thus probably only indirectly related to European A. insigne. There are some distinctive features in the dentitions of recorded A. insigne and A. africanum specimens which suggest that their phylogenetic connection may, indeed, have been indirect. For example, the European A. insigne speci- men, NMB-MP549, has a double-rooted P®, a condition unknown in the Langebaanweg sample. On the other hand, no European specimen is known which has an ‘intermediate cusp’ on the P* protocone lobe or a vestigial M, metaconid, while none is known to lack the anterolingual cusp of M,. The possibility that European and African Agriotherium evolved independently will be discussed again below. The situation in respect of Agriotherium in Asia and North America is similar to that with the Indarctos of these continents. A. sivalense is definitely a valid species, while A. palaeindicum is only tentatively regarded as such. Chinese Agriotherium is probably referable to A. sivalense. There is almost certainly only one species of Agriotherium represented in North America, and this may also be referable to A. sivalense. However, the North American species name which has priority, A. schneideri (Sellards 1916), is provisionally retained. It has generally been assumed that North American Agriotherium, like Indarctos, was an immigrant from Asia (e.g. Repenning 1967). This is clearly a simple and logical interpretation of a situation where one genus supersedes another closely related one on a continent known to have received immigrants during the period in question. However, this is not the only interpretation possible. The fact that North American Jndarctos includes some of the most Agriotherium-like specimens known suggests that the Indarctos—Agriotherium transition may have taken place in North America. If this were so then either the subsequent dispersal of Agriotherium started in North America, or Agrio- therium evolved independently from advanced Indarctos in North America as well as in Europe and Africa. The former alternative is unlikely in view of what is known of the Old World history of agriotheriines, whereas the independent evolution theory is a distinct possibility. This alternative appears to be implausible only in that it involves the polyphyletic origin of one genus as it is presently conceived. If, as indicated earlier, ‘advanced Jndarctos’ is interpreted instead as ‘primitive Agriotherium’, it would mean that the Indarctos—Agriotherium transition took place once only (probably in Europe), and that once this evolutionary course 103 AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA sisuauobaJo sojosepu uapleuyos wnueyjouby VOIHAWV HLYON ‘goeds pu SUII} UI TUIMIOYJONIZY JO JUOWOSURIIL SAIIEIUN} YW “Zp “SIT eaceetes hae ee sv snioesyjue sojosepu| | sapiojose sojosepu] ‘ | ears Snowe sojosepuy sisuaigelund sojo1epuy | é wnoipulaejed wnvayjouby ‘ds sojosepul; ANSOOIW ALV1 asuajeais wnueyjouby aubisul wnuayjouby visvana wnueowe wnueyjouby VOldsv ANSOONMd 104 ANNALS OF THE SOUTH AFRICAN MUSEUM had been set the development of Agriotherium (senso stricto) could take place irrespective of the geographical location of the populations concerned. Thus European A. insigne may have been descended from ‘J.’ atticus, A. africanum from the north African Jndarctos-like agriotheriine, A. sivalense from the ‘J.’ punjabiensis/A. palaeindicum complex in Asia, while in North America ‘J.’ oregonensis was ancestral to A. schneideri. It is nevertheless clear that there was a close connection between advanced agriotheriines in Europe and Africa on the one hand, and in Asia and North America on the other, and the above scenario may therefore be unrealistic. Instead A. insigne and A. afri- canum may share a common ancestor and be conspecific, while the same may apply to A. sivalense and A. schneideri. These alternatives are indicated in Figure 42. The possibility that all Agriotherium (senso stricto) evolved from a single source is not favoured here, but it cannot yet be dismissed. The nomenclature of this genus will ultimately have to be established by further testing of the hypotheses presented here, and perhaps by reference to conclusions reached in respect of other taxa with similar wide distributions and generally uniform characteristics. Amongst the latter are the living brown and black bears, as well as a wide variety of other carnivores, notably canids and felids. SUMMARY Available material of Agriotherium africanum from the latest Miocene/early Pliocene Varswater Formation at Langebaanweg is described. This species was found to be as variable as other bears of comparable size, this being in part due to marked sexual dimorphism. The depositional environment and likely habitat and habits of the Lange- baanweg Agriotherium are discussed. It is suggested that it was a large wood- land carnivore, remains of which were transported to their points of discovery by a river which then met the sea in the vicinity of Langebaanweg. The trans- porting agent, and hyaenas, rodents, and fires contributed to the post-mortem damage exhibited by specimens. Agriotherium was evidently descended from late Miocene Jndarctos, the more significant evidence in support of this hypothesis being as follows: 1. There was an increase of size with time in the /ndarctos—Agriotherium lineage, and advanced Indarctos may be little or no different’in size to Agriotherium. 2. The skull characters of advanced Jndarctos (e.g. I. atticus from Samos; Indarctos from Florida) are shared by Agriotherium (e.g. the Langebaanweg specimen L45062). Observable differences reflect further development of trends already evident in the Indarctos lineage. Significant shared characteristics are a relatively short and broad snout, with the posterior palatine incisure in line with the M? metacones, large zygomata and sagittal crest, and similar basi- cranial region, including absence of the alisphenoid canal. AGRIOTHERIUM FROM LANGEBAANWEG, SOUTH AFRICA 105 3. There is little or no difference in the dentitions of advanced Indarctos and Agriotherium. For example, the incisors, canines, P,’s, M?’s, M,’s and M,’s may be morphologically indistinguishable. Differences that do exist reflect trends already evident in the Jndarctos lineage. For example: (i) There was a progressive reduction in the size and number of anterior premolars in the Indarctos—Agriotherium lineage, with their principal cusps becoming progressively lower crowned and indistinct. However, Agrio- therium sometimes has Indarctos-like upper premolars (e.g. A. insigne, NMB-MP549), while Indarctos may have Agriotherium-like lower anterior premolars (e.g. J. atticus, NNUW-Samos 1912, 29). (ii) The shearing elements of the carnassials in the Indarctos—Agriotherium lineage are progressively developed, as are the buccal cusps of M! and M3, which also have a shearing function. There is a corresponding reduction in the crushing elements of the posterior cheek teeth. For example, while the M2? of Indarctos is distinguished from that of Agriotherium by the presence of a talon, this feature is progressively reduced in time. In advanced Indarctos it may be small (e.g. I. punjabiensis, GSI-D12), while vestiges of a talon may be present in Agriotherium (e.g. the Langebaanweg specimen L47698). (iti) As a general rule the P* of Agriotherium is distinguished from that of Indarctos by the presence of a parastyle, but this cusp may be present in advanced Indarctos (e.g. Samos I. atticus; most North American specimens). 4. A double-cusped P* protocone lobe is characteristic of European Indarctos, and is also found in some European and African Agriotherium specimens (e.g. A. insigne, NMB-MP549; Langebaanweg L2045 and others), although the P* antero-internal cusp is sometimes absent (e.g. A. insigne, Gervais (1859) specimen; Langebaanweg, L41404). The fact that this cusp is sometimes absent in A. insigne, and that in those specimens postdating L2045 it is reduced or absent, indicates that there was a tendency in European—African Agriotherium to lose the antero-internal cusp. A similar tendency is evident in advanced Asian and North American Indarctos (e.g. I. punjabiensis, GSI-D6; Florida Jndarctos), while the antero-internal cusp is always absent in the Agriotherium from these continents. This common tendency in Jndarctos and Agriotherium indicates a close relationship, while the examples cited suggest that the European/African and Asian/North American Indarctos—Agriotherium lineages evolved independently and at different rates later in their history. The primary /ndarctos—Agriotherium lineage was European and comprised I. vireti, I. arctoides, I. atticus and A. insigne, with I. anthracitis as an aberrant, probably island-dwelling offshoot. Indarctos emigrated to Africa, probably late in the Miocene, and an Agriotherium (A. africanum) very similar to the European A. insigne was evolved on this continent. Advanced Indarctos also migrated eastwards and the Asian/North American lineage is comprised of species of 106 ANNALS OF THE SOUTH AFRICAN MUSEUM Indarctos and Agriotherium postdating J. atticus. Pending further studies, most existing species names of these genera are retained. It is suggested that either all /ndarctos, or all later Indarctos (I. atticus and younger species) should be referred instead to Agriotherium, although the status quo is maintained for the present. A modified suprageneric classification of later Ursidae is proposed. The species of Ursavus are included in a new subfamily, the Ursavinae. The Agrio- theriinae are divided into two tribes, namely, Agriotheriini (Indarctos—Agrio- therium) and the Ailuropodini (‘Ursavus’ depereti—Ailuropoda melanoleuca). The Ursinae and Tremarctinae are reduced to the status of tribes within a reconstituted subfamily Ursinae. ACKNOWLEDGEMENTS I am greatly indebted to the many persons who assisted directly and indirectly in the study of the Langebaanweg Agriotherium. They include Drs G. de Beaumont (Geneva), B. Engesser (Basel), A. W. Gentry (London), L. Ginsburg (Paris), J. Jewell (London), C. A. Repenning (Menlo Park, Cali- fornia) and E. Thenius (Vienna) who provided access to, and/or casts of, specimens in their care. Drs A. J. Tankard (Knoxville, Tennessee) and R. H. Tedford (New York) provided information on geological and palaeontological matters respectively. Almost all the photographs of specimens were made by Miss J. Nolte, while Mr A. Byron printed Figures 27 and 32. Miss L. Scott prepared Figure 34, Mrs P. Eedes typed the manuscript, and the South African Air Force provided the aerial photograph for Figure 26. Drs R. G. Klein (Chicago) and B. Kurtén (Helsinki) reviewed early drafts of the manuscript. The Langebaanweg Research Project is supported by Chemfos Ltd, the South African Council for Scientific and Industrial Research, and the Wenner— Gren Foundation for Anthropological Research (Grant no. 2752-1834), and the assistance of these organizations is gratefully acknowledged. REFERENCES BERGGREN, W. A. & VAN COUVERING, J. A. 1974. The Late Neogene. Amsterdam: Elsevier. Boaz, N. T., GAziry, A. W. & EL-ARNAUTI, A. 1979. New fossil finds from the Libyan Upper Neogene Site of Sahabi. Nature, Lond. 280: 137-139. BuTzer, K. W. 1971. Environment and archaeology. Chicago: Aldine. Cuorn, J. & HOFFMANN, R. S. 1978. Ailuropoda melanoleuca, Mammalian Species 110: 1-6. CoETZEE, J. A. 1978. Late Cainozoic palaeoenvironments of southern Africa. In: VAN ZINDEREN BAKKER, E. M. ed. Antarctic glacial history and world palaeoenvironments: 115-127. Rotterdam: Balkema. CRUSAFONT, M. & KurTEN, B. 1976. 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A history of savanna vertebrates in the new world. Part 1. North America. Ann. Rey. Ecol. Syst. 8: 355-380. Wo ttr, R. G. 1978. Function and phylogenetic significance of cranial anatomy of an early bear (UIndarctos) from Pleistocene sediments of Florida. Carnivore 1 (3/4): 1-12. Wo rr, R. G., SINGER, R. & BisHop, W. W. 1973. Fossil bear (Agriotherium Wagner 1837) from Langebaanweg, Cape Province, South Africa. Quaternaria 17: 209-236. ZDANSKY, O. 1924. Jungtertiare Carnivoren Chinas. Palaeont. sinica (C) 2: 1-149. | 6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature | (particularly Articles 22 and 51). : | Names of new taxa, combinations, synonyms, etc., when used for the first time, must be | followed by the appropriate Latin (not English) abbreviation, e.g. gen. noy., sp. nov., comb. | novy., Syn. NOv., etc. An author’s name when cited must follow the name of the taxon without intervening | punctuation and not be abbreviated; if the year is added, a comma must separate author’s | name and year. The author’s name (and date, if cited) must be placed in parentheses if a | species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, i.e. all published | scientific names by which the species previously has been designated are listed in chronological | order, with all references to that name following in chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845) | Figs 14-1SA | Nucula (Leda) bicuspidata Gould, 1845: 37. Leda plicifera A. Adams, 1856: 50. Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b), Nucula largillierti Philippi, 1861: 87. | Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. Note punctuation in the above example: comma separates author’s name and year “semicolon separates more than one reference by the same author full stop separates references by different authors figures of plates are enclosed in parentheses to distinguish them from text-figures dash, not comma, separates consecutive numbers Synonymy arrangement according to chronology of bibliographic references, whereby the year is placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable. In describing new species, one specimen must be designated as the holotype; other speci- mens mentioned in the original description are to be designated paratypes; additional material | not regarded as paratypes should be listed separately. The complete data (registration number, depository, description of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g. : | Holotype | SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach | Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973. 3 Note standard form of writing South African Museum registration numbers and date. |7. SPECIAL HOUSE RULES Capital initial letters (a) The Figures, Maps and Tables of the paper when referred to in the text | e.g. *... the Figure depicting C. namacolus...’; *. . . in C. namacolus (Fig. 10)...’ |(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by initials or full names e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene |(c) Scientific names, but not their vernacular derivatives | e.g. Therocephalia, but therocephalian | Punctuation should be loose, omitting all not strictly necessary | Reference to the author should be expressed in the third person | Roman numerals should be conyerted to arabic, except when forming part of the title of a | book or article, such as ‘Revision of the Crustacea. Part VIII. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: “Smith (1969) describes .. .’ ‘Smith (1969: 36, fig. 16) describes .. .’ “As described (Smith 1969a, 19696; Jones 1971)’ ‘As described (Haughton & Broom 1927)...’ ‘As described (Haughton et al. 1927)...’ Note: no comma separating name and year Dagination indicated by colon, not p. names of joint authors connected by ampersand - et al. in text for more than two joint authors, but names of all authors given in dist of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list a, scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FiscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gén. 74: 627-634. Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320. Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 81 Band February 1980 Februarie Rant 2 Deel ON THE VALIDITY OF THE THEROCEPHALIAN FAMILY LYCOSUCHIDAE (REPTILIA, THERAPSIDA) By J.A. VAN DEN HEEVER Cape Town _. Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000 Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na gelang van die beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000 OUT OF PRINT/UIT DRUK 1, 2(1-3;,5=8), 3(1-2, 4-5, 8, tpi), 5(-3,.4, 7-9); 6(1, t.-p.i.), 7(1-4), 8, 9(1-2, 7), 10(1-3), 11(1-2, 5, 7, t-p.i.), 15(4-5), 24(2), 27, 31(1-3), 32(5), 33 Copyright enquiries to the South African Museum Kopieregnavrae aan die Suid-Afrikaanse Museum ISBN 0 908407 85 8 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., . Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap ON THE VALIDITY OF THE THEROCEPHALIAN FAMILY LYCOSUCHIDAE (REPTILIA, THERAPSIDA) By J. A. VAN DEN HEEVER South African Museum, Cape Town (With 9 figures) LMS. accepted 29 November 1979] ABSTRACT The taxonomic position of the therocephalian family Lycosuchidae is discussed in the light of published accounts and a re-examination of most of the type material, together with additional information from undescribed specimens of early therocephalians. It is shown that the primary distinguishing characteristic of the Lycosuchidae which separates it from the Pristerognathidae, i.e. two simultaneously functional canines in each maxilla, is based on a misinterpretation. It is therefore concluded that the family consists of an unnatural grouping of members of the Pristerognathidae and should consequently be regarded as invalid. CONTENTS PAGE Introduction : , Papell Historical review F ; is exalt Material and techniques . oe ey alily/ Description and discussion . 118 Conclusions . : ; ae P23 Acknowledgements . : ay el23 References . , : : oo 124 Abbreviations . pe Baits S- FALZS INTRODUCTION The early Therocephalia of the upper Permian Tapinocephalus Zone (Dinocephalian and Pristerognathus/Diictodon Assemblage Zones of Keyser & Smith 1979) of the Beaufort Series of the South African Karoo are generally poorly understood in comparison with other therapsid groups such as the Dicynodontia and the Cynodontia, mainly as a result of the intractable matrix in which the material is usually found. Since the dentition of an unprepared specimen is often its most distinctive feature the number and position of the teeth have been predominantly used in the past to distinguish between the various taxa of the group. Consequently, serious doubts have only recently been raised about the naturalness of taxa which have existed in the literature, e.g. the Lycosuchidae (Haughton & Brink 1955). Since the family Lycosuchidae was established its relatively rare members, identified principally by the posses- sion of two maxillary canines, have always been regarded as closely allied to 111 Ann: S. Afr. Mus. 81 (2), 1980: 111-125, 9 figs. 112 ANNALS OF THE SOUTH AFRICAN MUSEUM the more abundant family Pristerognathidae, contemporary therocephalians with one maxillary canine. Most authors (Haughton & Brink 1955; Kermack 1956; Romer 1956; Watson & Romer 1956; Boonstra 1969) who have discussed the early Therocephalia have placed great taxonomic weight on the number of canines and consequently the concept of double-canined therocephalians is widespread in the literature. Illustrations of early therocephalians usually present the double-canined Lycosuchus as a general representative of the group (Du Toit 1954; Romer 1956, 1966). Kermack (1956) demonstrated in the Therocephalia and the Gorgonopsia the existence of two upper canine positions which alternate in housing a single functional canine. According to Hopson (1964) this is also the case in the cynodont Thrinaxodon liorhinus and probably most other cynodonts as well. From this it is to be expected that while the functional canine was being replaced, the animal would have two canines of different ages in each maxilla, super- ficially similar to the condition frequently observed in living mammals when the permanent canine is in the process of replacing the milk canine (Fig. 1). Kermack also described a lycosuchid Trochosaurus major with two erupted canines and states that the possession of two simultaneously functional canines was primitive for Therocephalia. This idea probably stems from the view that a similar condition was thought to typify sphenacodont pelyccsaurs, the pre- sumed ancestors of therapsids. In an important paper Mendrez (1972) established the existence of an incipient crista choanalis in the pristerognathids Pristerognathus polyodon and Ptomalestes ayidus, situated on the inner surface of the maxilla medial to the canines. She interpreted this structure as the first step on the way to the develop- ment of a bony secondary palate as in mammals. Since the gorgonopsian maxilla is completely smooth in this area (Kemp 1969), this structure makes it possible to distinguish readily between the otherwise very similar snout frag- ments or isolated maxillae of therocephalians and gorgonopsians. Mendrez Fig. 1. Stereophotograph of the left maxilla of Felis caracal (SAM-ZM38191) to show the eruption of the permanent canine anterolingual to the milk canine. Scale =10mm. — ON THE VALIDITY OF THE LYCOSUCHIDAE 113 (1972) also noted the presence of two canine positions in pristerognathids (but did not cite Kermack’s prior discovery of this fact) and states at page 2961: ‘Pristerognatus polyodon ainsi que Ptomalestes avidus possédent également une autere caractéristique qui, selon les descriptions classiques, était, parmi les Pristerosauria de la zone a Tapinocephalus, \a propriété exclusive des Lyco- suchidae, a savoir la présence de deux canines de chaque cété de la téte. Ceci diminue le nombre déja faible des caractéres opposant ces deux familles. Il est fort probable que le Pristerognathidae décrits comme présentant un diastéme entre la canine et les postcanines possédaient a cette place une seconde canine.’ From this she concluded that the Pristerognathidae and the Lycosuchidae probably form a single family. However, from her statement it appears as if the Pristerognathidae possessed, like the Lycosuchidae, two functional canines in each maxilla and thus that the accepted definition of the Lycosuchidae should include the Pristerognathidae as well. In summary, the only distinguishing characteristic of the family Lyco- suchidae that at present still appears to separate it from the Pristerognathidae is the presence of two functional canines in each maxilla. In an effort to deter- mine the validity of this morphological distinction, and thus of the family Lycosuchidae, a detailed study of the mode of replacement of the upper canines was undertaken. This study is intended to resolve the question of whether the two canines were fully mature and remained simultaneously functional for a long period of time (as assumed by most authors), or whether the condition represents a short-lived phenomenon in the replacement process, representing a stage during which the new canine is well erupted but the old functional canine has not yet been shed. The latter interpretation implies that the double- canined condition is a short segment of the normal replacing cycle of all early therocephalians and that there is no valid basis for taxonomically separating the double-canined forms (Lycosuchidae) from the Pristerognathidae. HISTORICAL REVIEW The first early therocephalian possessing two maxillary canines was described by Broom (1903a) as Lycosuchus vanderrieti (Figs 2-3). According to Broom the only other theriodont known at that time which possessed two canines in each maxilla was the Albany Museum specimen of the cynodont ? Cynognathus leptorhinus Seeley (currently placed in Cynognathus cratero- notus). However, in an addendum to the description of Lycosuchus vanderrieti, Broom (1903a) notes that ? Cynognathus leptorhinus is similar to Cynognathus platyceps and that the other known species of Cynognathus all had only one canine; therefore, the double-canined condition in this specimen was regarded by him as temporary. Broom also drew attention to the type of Trirachodon kannemeyeri Seeley which on one side of the snout, in front of the canine, shows the tip of a second canine similar to that in both Cynognathus and Lycosuchus. Broom (1903a) felt that the anterior canine in all these genera is the morphological equivalent of the permanent mammalian canine and the 114 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 2. Stereophotograph of the right maxilla of the type skull of Lycosuchus vanderrieti (Stellenbosch D173) to show the canines. The specimen is covered with polymethylmethacrylate for preparation in acid. Scale = 10 mm. Fig. 3. Stereophotograph of the left maxilla of the type skull of Lycosuchus vanderrieti to show the canines. The specimen is covered with polymethylmethacrylate for preparation in acid. Scale = 10mm. posterior canine is the equivalent of the deciduous canine of mammals. He also stated that both teeth may, however, have been functional for some time in Lycosuchus and the higher theriodonts because the posterior canine which developed first is more powerful and the anterior canine is ‘peculiarly specialized’ as if developed for a different function. The suggestion of separate functions was due to his observation that both the anterior and the posterior borders of the anterior tooth are serrated, whereas only the posterior border of the posterior tooth appeared to be serrated. a wg ON THE VALIDITY OF THE LYCOSUCHIDAE 115 Fig. 4. Stereo photograph of the left maxilla of the type specimen Trochosuchus acutus (SAM-— 1076) to show the canines. Scale = 10mm. Broom (19035) described the isolated maxilla of a second therocephalian possessing two canines as Lycosuchus mackayi. Not until five years later (Broom 1908), however, when describing the double-canined Hyaenasuchus whaitsi, does he mention the fact that he now regards both canines in these early thero- cephalians as being simultaneously functional with neither of them being a replacement tooth. In the same article Broom described the anterior part of a small therocephalian skull as Trochosuchus acutus, noting the presence of two maxillary canines, the anterior being the smaller (Fig. 4). Broom (1915) described Trochosuchus major specifically, stating that neither of the two canines in the maxilla is a replacement tooth and that in the light of the descriptions of Lycosuchus, Hyaenasuchus and Trochosuchus he regards these genera as having two large canines functioning simultaneously in each maxilla. Haughton (1915) in his description of Trochosaurus intermedius followed Broom in interpreting the two canines present in each maxilla as being simul- taneously functional, notwithstanding the fact that they differed in size and that a replacement tooth was situated medial to the anterior canine in each maxilla. In his book on the mammal-like reptiles of South Africa, Broom (1932) redescribed all of the species with two canines in each maxilla and stated that both teeth are simultaneously functional because more than a dozen specimens were then known to possess this arrangement of teeth. He regarded them as a 116 ANNALS OF THE SOUTH AFRICAN MUSEUM group separate from the Pristerognathidae but did not formally establish a new family for them. (Lycosaurus mackayi at his p. 50 is an error and should read Lycosuchus mackayi.) He also synonymized Trochosaurus intermedius (Haughton, 1915) with Trochosuchus major (Broom, 1915) under the name Trochosaurus major. He retained the genus Trochosuchus for the single specimen of Trochosuchus acutus (Broom, 1908). A third specimen of Trochosaurus major was described by Boonstra (1934) who indicated in the text as well as in the figures that the canines were under- going replacement. He also noted the relatively broad epipterygoid which is narrowest in the middle and expanded dorsally and ventrally. Broom (1936a) described Trochorhinus vanhoepeni as closely allied to Trochosaurus major and possessing two canines of unequal size, the larger being anterior. Broom (1936b) described Trochosaurus dirus as having two large functional » canines in each maxilla. However, the canines are at different stages of develop- ment and the roots of their eventual replacements are visible medially. He notes that both canines are functional and situated so close together that they probably functioned as one tooth. Furthermore he states that: *. . . each anterior canine has a very young replacing tooth; but the posterior canine on the left is being replaced by an already well-developed successor. On the right side the specimen is imperfect but the inner canine is of large size and apparently functional. Probably the outer canine is shed or being absorbed.’ No new early therocephalian specimens showing double upper canines were described after 1936. Romer (1945) included all of the above-mentioned genera in the Pristerognathidae. Although Broom (1932) developed the rather loose concept of double- canined therocephalians, it was actually Houghton & Brink (1955) who estab- lished the family Lycosuchidae, for which they gave the following diagnosis: ‘Medium-sized therocephalians with two large functional canines in each maxilla.’ They listed the species as: Hyaenasuchus whaitsi Broom, 1908; Lyco- suchus vanderrieti Broom, 1903 (not 1902 as given by Haughton & Brink); Lycosuchus mackayi Broom, 1903; Trochorhinus vanhoepeni Broom, 1936; Trochosaurus major (Broom, 1915), and Trochosaurus dirus Broom, 1936. The single specimen of Trochosuchus acutus was referred by them to the family Akidnognathidae. Tatarinov (1974) ascribes the establishment of the family Lycosuchidae to Haughton (1924). The references in Tatarinov’s article reveal that the paper in question was actually published in 1925; however, in this paper Haughton retained the double-canined forms in the family Pristerognathidae. Shortly thereafter, Watson & Romer (1956) followed Romer (1956) who independently established the family Trochosuchidae comprising the same genera as those placed by Haughton & Brink (1955) in the Lycosuchidae. They also synonymized Trochosaurus Haughton, 1915, with Trochosuchus Broom, 1908. Watson & Romer (1956) diagnosed the family Trochosuchidae as: ‘Large ON THE VALIDITY OF THE LYCOSUCHIDAE 117 therocephalians which resemble the Pristerognathidae in fundamental features of their structure but differ in having a much lower skull with a broad and rather flattened snout, a sagittal crest never elevated and the occiput trans- versely widened. They may have six incisors and normally two canines, each separately replaced.’ Since their classification is predated by that of Haughton & Brink (1955) the name Lycosuchidae has precedence and has been used by nearly all subsequent authors, e.g. Boonstra (1969, 1971, 1972), Mendrez (1972), and Tatarinov (1974); Lehman (1961 : 232), however, incorrectly follows Watson & Romer (1956). Von Huene (1956) retains the members of the Lycosuchidae within the Pristerognathidae. In spite of having synonymized Trochosaurus with Trochosuchus (Watson & Romer 1956), Romer (1966), synonymized the Lycosuchidae of Haughton & Brink (1955)-with a new family, the Trochosauridae. This was apparently done to facilitate the inclusion of Trochosuchus in another family, the Alopeco- dontidae (Romer 1966). However, Haughton & Brink (1955) had placed Trochosuchus in the Akidnognathidae (defined as having one small canine in front of the large functional canine) a family not recognized by Romer (1966), and they described the Alopecodontidae as therocephalians with two small canines in front of the large functional canine in the maxilla. The weathered type specimen of Trochosuchus acutus (SAM-—1076) in the South African Museum has one canine in the right maxilla and two canines in the left maxilla, the anterior being the smaller (Fig. 4). However, the last incisor appears to lie within the maxilla when viewed laterally and may have been mistaken for a small canine by Romer. The genera included by Watson & Romer (1956) in the Trochosuchidae (Lycosuchidae) do not have larger skulls than those early therocephalians possessing a single maxillary canine, and an examination of the available material indicates that the other diagnostic differences of the family can be attributed to post-mortem distortion. Consequently, in a later description of the Lycosuchidae, Boonstra (1969) characterizes the family as: ‘Early fairly large Therocephalia with fairly broad flattened skulls with two functional canines in the maxilla, advanced broadened epipterygoid and low sagittal crest. Otherwise very similar to pristerognathids. With four monotypic genera.’ However, as Mendrez (1972) quite rightly points out, the so-called broad epipterygoid of the Lycosuchidae is actually known in one specimen only, Trochosaurus major (BMNH RS5747), and it is, in fact, no broader than that of the pristerognathid Ptomalestes avidus. MATERIAL AND TECHNIQUES The type material of Tapinocephalus Zone therocephalians at the South African Museum was examined. In addition a complete therocephalian skull (G.S. C60) with lower jaw, lacking only the occipital bones, was borrowed from the Geological Survey. The medial aspect of the right maxilla of this specimen was carefully prepared by mechanical means to show the canines. 118 ANNALS OF THE SOUTH AFRICAN MUSEUM The type skull of Lycosuchus vanderrieti (D173), on loan from the University of Stellenbosch, is currently being prepared in an 11 per cent solution of formic acid owing to the extreme hardness of the matrix. It is at present still covered with polymethylmethacrylate (Figs 2-3), but the double canines are well pre- served and have been examined. Other material was prepared mechanically where necessary. In addition, the right maxilla of an unidentified species of therocephalian, SAM—K317 (identified as therocephalian according to the method of Mendrez (1972)), was sectioned frontally on a Beuhler Isomet Low Speed Saw at intervals of 2 mm. One section was stained with Alizarin Red S in a 4 per cent solution of potassium hydroxide to show the resorption of the canine root. All photographs were taken on Kodak Panatomic-X film with a stereo apparatus built by N. J. Eden of the South African Museum. DESCRIPTION AND DISCUSSION The inner surface of the left maxilla of therocephalian SAM-—K317 shows a distinct canine boss which contains the two canine alveoli (Fig. 5). The relative positions of the roots are visible as two smaller bulges separated by a shallow vertical sulcus. This condition can also be seen in the type of the pristerognathid Ptomalestes avidus, SAM-11942. The functional canine lies in the anterior alveolus and is broken off at the alveolar border. No tooth is externally visible in the posterior alveolus. The right maxilla of SAM—K317 shows the same features as does the left side, but a frontal section through the posterior alveolus shows the root of an old canine being resorbed from the alveolar border upwards (Figs 5-6). The skull of Geological Survey specimen C60 has been compressed laterally, but in lingual view the maxilla clearly shows the canine boss with the functional canine in the anterior alveolus (Fig. 7). A replacement canine of which the tip is serrated both anteriorly and posteriorly is erupting from the posterior alveolus. This condition is identical to that in the left maxilla of Lycosuchus Fig. 5. Stereophotograph of the medial surface of the left maxilla of an unidenti- fied pristerognathid (SAM-—K317) showing the boss containing the canine alveoli. Anterior is to the right. Scale = 10 mm. ON THE VALIDITY OF THE LYCOSUCHIDAE 119 Fig. 6. Frontal section through the posterior canine alveolus of the right maxilla of an unidentified pristerognathid (SAM-—K317) to show the resorption of the old canine root and its replacement by spongy bone. Scale = 10 mm. vanderrieti except that in the latter the younger tooth lies in the anterior position (Fig. 3). See externally only, G.S. C60 would have to be classified as a lycosuchid according to the accepted definition of the family. However, the diameter of the posterior alveolus is the same as that of the anterior and much larger than that of the erupting canine. This suggests that the posterior alveolus was probably occupied previously by a large canine and that the immature tooth is not the first to have erupted in that position. Medial to the functional canine (Figs 7-8) an unerupted replacement canine is visible where the bone of the 120 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 7. Stereophotographs of the medial surface of the right maxilla of an unidentified pris- terognathid (G.S. C60) showing the boss containing the canine alveoli and the sequence of canine replacement. Scale = 10 mm. Fig. 8. Stereophotographs of a ventral view of the right maxilla of an unidentified pristerognathid (G.S. C60) to show the sequence of canine replacement. Scale = 10 mm. ON THE VALIDITY OF THE LYCOSUCHIDAE 121 medial wall is damaged. In this ‘lycosuchid’, then, there is direct evidence that the sequence of canine eruption alternates between the two alveoli in such a way that the time lapse between the eruption of teeth in the two alveoli produces a single functional canine at a time. This is also the most likely interpretation of the condition in the type of Lycosuchus vanderrieti. Specimens with two large canines in the same maxilla represent the terminal stages of the older tooth of the pair. Since the second canine is older than the first in Lycosuchus vanderrieti, it would naturally be more powerful than the immature tooth, and since both tips of the posterior canines in Lycosuchus are damaged, Broom (1932) had no grounds for stating that the anterior canines are ‘peculiarly specialized’ for a different function. In fact, specimens of therocephalians which have complete canines show serrations at the tips of these teeth both in front and behind, regardless of whether the tooth is in the anterior or posterior position. Kermack (1956) regards the single lycosuchid specimen (Trochosaurus major, BMNH R5747) described in his paper as one of the most primitive of the therocephalians because it has two functional canines in each maxilla (Fig. 9). By his own description (Kermack 1956: 114) the roots of replacement canines can be seen in a fracture lying lingual to each of the canines in the right maxilla. The anterior of this pair is in a more advanced stage of development than the posterior, which strongly suggests that the two large functional teeth are also of different ages. On the left, the fracture is such that the replacement teeth cannot be seen but the large canines are clearly also of differing ages since the anterior canine was still in the process of erupting and has a wide-open pulp cavity. This indicates not that both teeth were functional at the same time but rather that replacement was taking place at the time of death. However, Kermack (1956: 115) states: “This specimen compares closely with the two specimens of Ae/urosaurus (R339 and R855a). The only essential difference is that, in the two gorgonopsids the pair of alveoli in the maxilla each alternately bears the functional canine, while in the therocephalian each bears a functional tooth simultaneously. The difference is one of timing only.’ Kermack (1956: 121) notes further that in sphenacodont pelycosaurs such as Dimetrodon, as well as in Trochosaurus, two functional canines were present in each maxilla and elsewhere (Kermack 1956: 130) he states: ‘As in Dimetrodon there was a pair of functional upper canines on each side in these primitive therocephalia, and they were replaced alternately. The functional replacement for each of the pair was the next number of its own tooth family. Like Dimetro- don when one of the upper canines was being replaced, these Therocephalia must have had but one functional canine on that side of the jaw.’ Kermack (1956) apparently implies here that in Dimetrodon and Trocho- saurus-like primitive therocephalians the double canine condition was the prevailing one, whilst the period during which only one canine was functional was, in fact, of a comparatively short duration, i.e. there were normally two upper canines functioning simultaneously. 122 ANNALS OF THE SOUTH AFRICAN MUSEUM R L Fig. 9. Diagram from Kermack (1956) to show the canines and canine replacements of Trocho- saurus major (BMNH R5747). R = right, L = left. The South African Museum holds at least 112 specimens of early thero- cephalian skulls and skull fragments in which the canines can be seen. Of these, fourteen specimens possess two canines in either one or both of the maxillae, including the above-mentioned types. This ratio of roughly one specimen with double canines for every seven with a single canine per maxilla illustrates the relative scarcity of the two-canine condition and indicates that the period during which two canines were externally visible was probably of relatively short duration. In no South African Museum specimen with double canines are there any indications that the teeth are of the same age and, from the literature cited above, it is also clear that in all described specimens of Lycosuchidae the canines are also staggered in age. It is highly improbable that in carnivores such as the Therocephalia, in which the tips of the canines are serrated both anteriorly and posteriorly, these teeth would have functioned optimally as a closely packed unit. Not only would the efficiency of penetration be impaired ON THE VALIDITY OF THE LYCOSUCHIDAE 123 by the bulky ‘unit’ compsed of two large teeth, but also, since some of the serrations would be obscured, the teeth would tear less efficiently as well. Therefore, it seems more likely that these animals possessed a single piercing canine of long functional duration and that the period of replacement, during which two canines were externally visible in each maxilla, was as short as possible. This is indicated by the relatively few specimens actually showing this condition. The functional replacement for each canine would then not be the next tooth of its own family (i.e. in the same tooth position), but the next tooth erupting from the other canine alveolus. This model of canine tooth replace- ment is supported by the work of Edmund (1960) who, contrary to the observa- tion of Kermack (1956), found that in Dimetrodon the pair of canines in each maxilla were only occasionally functional at the same time and usually alternated so that only one tooth was functional at a time. CONCLUSIONS Kermack (1956) is correct in stating that in the Therocephalia the two canine alveoli each bore the functional canine alternately, but he is incorrect in assuming that in the Lycosuchidae, e.g. Trochosaurus, both alveoli normally bore functional canines simultaneously. In view of the importance of canines in carnivore dentitions it is to be expected that the replacement of any fang will develop at such a time and replace the mature canine in such a way that the animal is never without at least one functional canine in each maxilla. This necessitates a period of time when the erupting replacement coexists with the old functional tooth. Because of the distinct advantages of the single over the double functional canine condition, the actual period of time in which the two canines were externally visible was probably kept as short as possible. The Lycosuchidae is therefore not a separate primitive therocephalian family but consists of members of the Pristerognathidae in which death occurred while the erupting replacing canine was visible externally. Lycosuchus van- derrieti (Figs 2-3) is an especially good example of this condition. Therefore the family Lycosuchidae (—Trochosauridae) represents an unnatural grouping of members of the Pristerognathidae, and it is suggested here that it be invalidated. ACKNOWLEDGEMENTS I wish to thank the following persons: Mrs K. Rial and Mr N. J. Eden, both of the Department of Palaeontology at the South African Museum for, respectively, preparation and photography, Professor W. J. Verwoerd of the Department of Geology at the University of Stellenbosch, and Dr A. W. Keyser of the Geological Survey in Pretoria for the loan of specimens, Dr J. A. Hopson of the University of Chicago for critically reading the manuscript, and Mrs P. Eedes and Mrs G. E. Blaeske, of the South African Museum, for typing. 124 ANNALS OF THE SOUTH AFRICAN MUSEUM REFERENCES BoonstrA, L. D. 1934. A contribution to the morphology of the mammal-like reptiles of the suborder Therocephalia. Ann. S. Afr. Mus. 31: 215-267. Boonstra, L. D. 1969. The fauna of the Tapinocephalus zone (Beaufort beds of the Karoo). Ann. S. Afr. Mus. 56: 1-73. ‘ BoonstrA, L. D. 1971. The early therapsids. Ann. S. Afr. Mus. 59: 17-46. BoonstraA, L. D. 1972. Discard the names Theriodontia and Anomodontia: a new classi- fication of the Therapsida. Amn. S. Afr. Mus. 59: 315-338. Broom, R. 1903a. On an almost perfect skull of a new primitive theriodont (Lycosuchus vanderrieti). Trans. S. Afr. phil. Soc. 14: 197-205. Broom, R. 19036. On some new primitive theriodonts in the South African Museum. Ann. S. Afr. Mus. 4: 147-156. Broom, R. 1908. On some new therocephalian reptiles. Ann. S. Afr. Mus. 4: 361-367. Broom, R. 1915. Catalogue of types and figured specimens of fossil vertebrates in the American Museum of Natural History. II. Permian, Triassic and Jurassic reptiles of South Africa. Bull. Am. Mus. nat. Hist. 25: 105-164. Broom, R. 1932. The mammal-like reptiles of South Africa and the origin of mammals. London: Witherby. Broom, R. 1936a. On some new genera and species of Karroo fossil reptiles, with notes on some others. Ann. Transy. Mus. 18: 349-386. Broom, R. 19365. On the structure of the skull in the mammal-like reptiles of the suborder Therocephalia. Phil Trans. R. Soc. 226: 1-42. Du Torr, A. L. 1954. The geology of South Africa. 3rd ed. S. H. Haughton ed. Edinburgh; London: Oliver & Boyd. Epmunp, A. G. 1960. Tooth replacement phenomena in the lower vertebrates. Life Sci. Contr. R. Ont. Mus. 52: 1-190. HauGutTon, S. H. 1915. On two new therocephalians from the Gouph. Ann. S. Afr. Mus. 12: 55-57. HauGuton, S. H. 1925. A bibliographical list of pre-Stormberg Karroo Reptilia, with a table of horizons. Trans. R. Soc. S. Afr. 12: 51-104. HAUGHTON, S. H. & BRINK, A. S. 1955. A bibliographical list of Reptilia from the Karroo beds of Africa. Palaeont. afr. 2: 1-187. Hopson, J. A. 1964. Tooth replacement in cynodont, dicynodont and therodephalian reptiles. Proc. zool. Soc. Lond. 142: 625-654. Kemp, T. S. 1969. On the functional morphology of the gorgonopsid skull. Phil. Trans. R. Soc. (B) 256: 1-83. Keyser, A. W. & Smit, R. M. H. 1979. Vertebrate biozonation of the Beaufort Group with _ Special reference to the western Karoo basin. Ann. geol. Surv. Pretoria 12: 1-35. Kermack, K. A. 1956. Tooth replacement in mammal-like reptiles of the suborders Gorgonopsia and Therocephalia. Phil. Trans. R. Soc. (B) 240: 95-133. LEHMAN, J. P. 1961. Therocephalia. In: Traite de Paleontologie 6: 224-245. J. Piveteau, ed. MENDREZ, C. 1972. Premiéres ébauches d’un palais secondaire osseux chez les reptiles mam- maliens. C. r. Acad. Sci. Paris 274: 2960-2961. Romer, A. S. 1945. Vertebrate paleontology. 2nd ed. Chicago: University of Chicago Press. Romer, A. S. 1956. The osteology of the reptiles. 1st ed. Chicago: University of Chicago Press. Romer, A. S. 1966. Vertebrate paleontology. 3rd ed. Chicago: University of Chicago Press. TATARINOV, L. P. 1974. Theriodonts of the USSR. Moscow: NAUCA. Von HEUNE, F. 1956. Paldontologie und Phylogenie der niederen Tetrapoden. Jena: VEB, Gustav Fischer Verlag. WATSON, D. M. S. & Romer, A. S. 1956. A classification of therapsid reptiles. Bull. Mus. comp. Zool. Harv. 114: 37-89. ON THE VALIDITY OF THE LYCOSUCHIDAE ABBREVIATIONS anterior canine canine boss crista choanalis canine root functional canine incisor lower postcanine maxilla milk canine old alveolus postcanine position of anterior canine posterior canine permanent canine position of posterior canine replacing canine sulcus unerupted canine tip of unerupted canine upper postcanines British Museum of Natural History Geological Survey, Pretoria South African Museum. 125 i} — We Hy 6. SYSTEMATIC papers must conform to the International code of zoological nomenclature (particularly Articles 22 and 51). Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov., etc. An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific namé must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, i.e. all published scientific names by which the species previously has been designated are listed in chronological order, with all references to that name following in chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845) Figs 14-15A Nucula (Leda) bicuspidata Gould, 1845: 37. Leda plicifera A. Adams, 1856: 50. Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b). Nucula largillierti Philippi, 1861: 87. Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. Note punctuation in the above example: comma separates author’s name and year “semicolon separates more than one reference by the same author full stop separates references by different authors figures of plates are enclosed in parentheses to distinguish them from text-figures dash, not comma, separates consecutive numbers Synonymy arrangement according to chronology of bibliographic references, whereby the year is placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable. In describing new species, one specimen must be designated as the holotype; other speci- mens mentioned in the original description are to be designated paratypes; additional material not regarded as paratypes should be listed separately. The complete data (registration number, depository, description of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.: Holotype SAM-A13535 in the South African Museum, Cape Town. Adult female from mid- tide region, King’s Beach Port Elizabeth (33°51’S 25°39’E), collected by A. ‘Smith, 15 January 1973, Note standard form of writing South African Museum registration numbers and date. 7. SPECIAL HOUSE RULES Capital initial letters (a) The Figures, Maps and Tables of the paper when referred to in the text e.g. “... the Figure depicting C. namacolus ...’; *. . . in C. namacolus (Fig. 10)...’ ' (b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by initials or full names e.g. DuToit but A.L.du Toit; Von Huene but F. von Huene (c) Scientific names, but not their vernacular derivatives e.g. Therocephalia, but therocephalian Punctuation should be loose, omitting all not strictly necessary Reference to the author should be expressed in the third person Roman’ numerals should be converted to arabic, except when forming part of the title of a book or article, such as ‘Revision of the Crustacea. Part VIII. The Amphipoda.’ Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial capital letter, provided the same generic name is used consecutively. Name of new genus or species is not to be included in the title: it should be included in the abstract, counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts. J. A. VAN DEN HEEVER ON THE VALIDITY OF THE THEROCEPHALIAN FAMILY LYCOSUCHIDAE (REPTILIA, THERAPSIDA) VOLUME 81 PART 3 FEBRUARY 1980 j ANNALS OF THE SOUTH AFRICAN MUSEUM |CAPE TOWN ca INSTRUCTIONS TO AUTHORS 1. MATERIAL should be original and not published elsewhere, in whole or in part. 2. LAYOUT should be as follows: (a) Centred masthead to consist of Title; informative but concise, without abbreviations and not including the names of new genera or species Author's(s') name(s) Address(es) of author(s) (institution where work was carried out) Number of illustrations (figures, enumerated maps and tables, in this order) (b) Abstract of not more than 200 words, intelligible to the reader without reference to the text (c) Table of contents giving hierarchy of headings and subheadings (d) Introduction (e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents (f) Summary, if paper is lengthy (g) Acknowledgements (h) References (i) Abbreviations, where these are numerous 3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with 2,5 cm margins all round. First lines of paragraphs should be indented. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes .. .’ ‘Smith (1969: 36, fig. 16) describes...” “As described (Smith 1969a, 19696; Jones 1971)’ “As described (Haughton & Broom 1927)...’ “As described (Haughton et al. 1927)...’ Note: no comma separating name and year Dagination indicated by colon, not p. names of joint authors connected by ampersand et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, 5, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o, scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BuLLouGn, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140, FiscHer, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gén. 74: 627-634. ‘ Konn, A. J. 19604. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon, Ann. Mag. nat. Hist. (13) 2: 309-320. Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. , THIELE, J. 1910, Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 81 Band February 1980 Februarie Part 3 Deel DEVONIAN FISH AND PLANTS FROM THE BOKKEVELD SERIES OF SOUTH AFRICA By W. G. CHALONER E.R. Pk. FOREY B. G. GARDINER ALS; BILL V. T. YOUNG Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000 Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na gelang van die beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000 OUT OF PRINT/UIT DRUK 1, 2(1-3, 5-8), 3(1-2, 4-5, 8, t.—p.i.), 5(1-3, 5, 7-9), 6(1, t.-p.i.), 711-4), 8, 91-2, 7), 10(1-3), 11(1-2, 5, 7, t.—p.i.), 15(4-5), 24(2), 27, 31(1-3), 32(S), 33 Copyright enquiries to the South African Museum Kopieregnavrae aan die Suid-Afrikaanse Museum ISBN 0 908407 88 2 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap DEVONIAN FISH AND PLANTS FROM THE BOKKEVELD SERIES OF SOUTH AFRICA By W. G. CHALONER F.R.S. Birkbeck College, University of London P. L. FoREY British Museum (Natural History) B. G. GARDINER Queen Elizabeth College, University of London A. J. HILL Birkbeck College, University of London & V. T. YOUNG British Museum (Natural History) (With 14 figures and 1 addendum) [MS. accepted 20 September 1979] ABSTRACT Fossil fish and plants from upper strata of the Bokkeveld Series of Barrydale, Cape Province, Republic of South Africa, are described. The fishes are represented by arthrodire placoderms, Barrydalaspis theroni gen. et sp. noy., a phlyctaenaspid arthrodire and, possibly, Groenlandaspis; an acanthodian spine indistinguishable from Gyracanthides warreni White; and an egg case. The plants are represented by lycopods: Archaeosigillaria plumsteadiae sp. noy., A. cf. picosensis Kraéusel & Dolianiti, and two different types of lycopod endocortical cast. The fishes and plants are compared with those in Australia, Antarctica, North America and South America, and it is concluded that these Bokkeveld fossils are either Middle or Upper Devonian. CONTENTS PAGE Introduction Sine a) HCE py re ei ee ene ms cert 2%.) The fish remains , : j ; ‘ : 3 : Z = 28 Arthrodires ; f : : ; : : . ; B28 ?Chimaeroid . ; 3 F ; ; : { ‘ . 140 Acanthodian ; , : : : : : : : . 140 The plant remains. : : : : ; . 141 Archaeosigillaria plumsteadiae sp. nov. . 5 Teas Archaeosigillaria sp. cf. A. picosensis Krausel & Dolianiti =). 149 Lycopod ?endocortical cast . : j : : ; . 150 Discussion . : P 3 : : : : : E : =) £t50 Acknowledgements . : : : . : : : ; Ch Pd4: References . ah ee : 4 E : Se : : . 154 127 Ann. S. Afr. Mus. 81 (3), 1980: 127-157, 14 figs, 1 addendum. 128 ANNALS OF THE SOUTH AFRICAN MUSEUM INTRODUCTION In 1974 a small collection of fossil fish and plants was sent to one of the authors (B. G.) for identification by Dr J. N. Theron of the Geological Survey of South Africa. The material comes from the upper strata of the Bokkeveld Beds at Barrydale, Cape Province, Republic of South Africa. Precise locality information is not available. The Bokkeveld Series is reputed to be of Devonian age (Du Toit 1939) and therefore the presence of fish is of significance because, with one possible exception, fish have not previously been recorded from the Devonian of South Africa. This one possible exception is a reference to the occurrence of Machae- racanthus in the Bokkeveld Beds which is given without detail by Du Toit (1939: 222). It is possible that Du Toit extracted this record from Schwarz (1900) who referred to a fossil fish spine. At the very best, therefore, Devonian fish from southern Africa are very poorly known and this collection adds a new dimension to our rapidly increasing knowledge of Devonian fish from the Southern hemisphere. The primary purpose of this paper is to describe these fishes. The fish are associated with a few plant remains and these are described and discussed (W. G. C. & A. J. H.). A few notes are added concerning the age of the Bokke- veld Beds based on comparisons of the fish and plants with those preserved in presumed contemporaneous strata in North and South America, Antarctica, Australia and west Africa. The fish are represented mainly by placoderms, but an acanthodian and the impression of a large egg case are also present. This last is particularly interesting since the form of the egg case is similar to those attributed to chimae- roids which are not known prior to the Jurassic. The plants are represented by several types of lycopods. All the fossils are poorly preserved, the majority being represented as impression or internal casts. The matrix is a texturally variable and poorly bedded micaceous siltstone. Because of this and the type and condition of the contained fossils, it appears as if the sediment was deposited in a freshwater environment and that the fossils lived in the place of sedimentation. The specimens referred to in this paper come from the collections of the South African Museum, Cape Town and from the Geological Survey, Cape Office. The former are prefixed by SAM;; the latter are left unprefixed, simply being quoted by the field number. THE FISH REMAINS ARTHRODIRES A number of facts have made the study of the arthrodires difficult; only impressions of the plates remain; with few exceptions the plates are isolated and sometimes broken, making it difficult to associate parts of animals; there DEVONIAN FISH AND PLANTS FROM SOUTH AFRICA 129 is some degree of post-mortem deformation with resulting distortion. Never- theless, the margins of the plates are perfectly angular, ruling out the possibility of post-mortem/pre-depositional transportation. Among the specimens several show associated ventral thoracic armour and on the basis of these two types of arthrodire can be recognized. Some isolated plates of the lateral aspect of the trunk shield can be referred to one or other of these types. It has been decided to name one of these because it is distinctive from other named arthrodires. The second arthrodire type shows a general similarity to named arthrodires from elsewhere. To avoid cluttering the already burdensome literature with yet another name, this is referred to as a ‘phlyctaenaspid arthrodire’ with the implication that what little is known of this form is similar to Neophlyctaenius and presumed close relatives. Additionally, specimens of a head and a piece of a large arthrodire are briefly mentioned. Order ARTHRODIRA Suborder PHLYCTAENIOIDE! Miles 1973 Infraorder and Family incertae sedis Genus Barrydalaspis gen. nov. (Figs 1-4, 6A) Etymology From Barrydale, South Africa, the area in which these fossils are found. Diagnosis Phlyctaenioid with short, broad ventral thoracic armour; spinal long with denticles along the medial edge, spinals set at a very divergent angle; sub- pectoral emargination broad; interolateral with prominent transverse groove; anterior ventrolateral of either side meeting its partner in the mid-line; anterior dorsolateral with a prominent groove on the lateral face of the anteroventral corner; ornament consisting of small, simple tubercles; body behind thoracic shield at least partially covered with scales. Type species “@ Barrydalaspis theroni sp. nov. Barrydalaspis theroni sp. nov. Etymology Named after Dr J. N. Theron who brought this material to the attention of the authors. Holotype SAM-K4647, the impression of the ventral thoracic armour from the Bokkeveld Series of Barrydale, Cape Province, South Africa. 130 ANNALS OF THE SOUTH AFRICAN MUSEUM Other material SAM-K 4648-50, K4770-73, K4766, K4779, K4785, K4789, K4791, and an anterior dorsolateral plate preserved on K4798. Diagnosis As for genus, only species. Remarks The ventral thoracic armour shows a superficial resemblance to that of the petalichthyid Lunaspis (see Gross 1961); the armour is short and broad and the spinals are set at a divergent angle. However, unlike Lunaspis, Barrydalaspis shows well-differentiated posterior ventrolaterals. (Gross (1961) describes two pairs of plates lying behind the anterior ventrolaterals in Lunaspis, the posterior ventrolaterals and the postero-ventrals. These are interpreted as body scales by Miles & Young (1977) and their interpretation is accepted here.) Furthermore, the dorsal part of the armour is similar to that of a phlyctaenioid and unlike that of a petalichthyid in showing a ‘ball and socket’ dermal neck joint and a large posterior dorsolateral. The anterior ventrolateral (AVL) is short and broad, being 1,35 times as wide as long in the smallest individuals and 1,25 times as wide as long in the 20mm (= [oe Fig. 1. Barrydalaspis theroni gen. et sp. nov. Silicone cast of holotype showing ventral thoracic armour in external view. DEVONIAN FISH AND PLANTS FROM SOUTH AFRICA 131 a.t.s.c Fig. 2. Barrydalaspis theroni gen. et sp. nov. Restoration of the ventral thoracic armour in ventral view. Based on SAM-K4647 and K4649. AVL —anterior ventrolateral, 1L—intero- lateral, PVL—posterior ventrolateral, SP—spinal, a.t.s.c—anterior transverse sulcus. larger (presumably older) individuals. With this change in linear dimensions there is also a change in shape, the posterior area of the AVL becoming squarer with increasing size (cf. Figs 1 and 2). The length of the spinal margin, expressed as a percentage of the maximum width of the plate, also varies from 48 per cent in the small individuals to 44 per cent in the large specimens. A similar decrease in the relative length of the spinal margin has been recorded for Coccosteus by Miles & Westoll (1968: 433-434). There is a well-marked ‘Ventrolateralkante’ (Gross 1933) running longitudinally on the posterior part of the AVL and across the posterior ventrolateral. This implies that there was a narrow subpectoral wall of the flank armour and the development of a postbrachial lamina. The centre of radiation of the AVL lies remarkably far forwards, more so than in any of the ‘dolichothoracid’ types figured by Denison (1958, fig. 112). The ornament on the AVL consists of simple tubercles which are very small at the radiation centre but become larger along the anterior, median and posterior margins. Ornament is absent from that part of the plate adjacent to the sub- pectoral emargination. A similar lack of ornament is noted by Miles & Westoll (1968) in the corresponding area of Coccosteus. The visceral surface of the AVL shows that perichondral bone lined the scapulocoracoid. Impressions left by this perichondral layer suggest that the abdominal division and the coracoid process of the scapulocoracoid (termi- nology of Stensi6 1959) were both very broad, as in most ‘dolichothoracids’. In some specimens the contact faces with the anterolateral can be seen, sug- gesting that the width of the pectoral fenestra is equal to about half the length of the subpectoral emargination. The posterior ventrolateral (PVL) is a little longer than broad and, as usual, the left PVL overlaps the right. As mentioned above, there is a strongly 132 ANNALS OF THE SOUTH AFRICAN MUSEUM IL MRDORRT POC ne x Sta I ANA ON RRARAL AAG a KAS An anann ananene 20mm Fig. 3. Barrydalaspis theroni gen. et sp. nov. Restoration of anterior ventrolateral, intero- lateral and spinal of the left side in visceral view. Based on SAM—-K4779, K4770, K4791 and K4771. Abbreviations as in Fig. 1. developed ‘Ventrolateralkante’ which divides the plate into lateral (vertical) and ventral (horizontal) laminae. The lateral lamina, which is relatively long, suggests that the postbrachial lamina was also long, as in phlyctaenaspids (Denison 1958: 534). The ornament is similar to that on the AVL. The medial margin of the left PVL is ‘S’-shaped where it overlaps the right but there is no indication of the complex overlap relations seen in Tiaraspis and Romundina. The spinal (SP) is relatively long and the proportions agree more with those of Denison’s phlyctaenaspid genera than with those of any other arthrodire group. The spinals are relatively longer in smaller individuals. A suture between the SP and AVL can be traced except anteriorly. As mentioned in the diagnosis, the SP is set at a very divergent angle, this being approximately 50°. Phlyctaenius acadica also shows a spinal set at a high angle (about 48° from the restoration given by Heintz 1934), but in this species the SP is much shorter. The ornament consists of well-developed tubercles along the lateral edge of the anterior two- thirds. Smaller tubercles are also present over half of the ventral surface and about one-fifth of the dorsal surface. Posteriorly, the medial edge bears six to eight recurved denticles. DEVONIAN FISH AND PLANTS FROM SOUTH AFRICA 133 The interolateral (IL) is orientated almost transversely and has ventral (external) and dorsal (internal or postbrachial) laminae which both become wider laterally. On the ventral lamina there is a shallow sulcus between two rows of tubercles. This sulcus, which has been variously named in arthrodires (‘anterior ventral sulcus’—Miles & Westoll 1968; anterior transverse sensory canal— Mark-Kurik 1973; Orvig 1975), implies that there was a neuromast line as in Actinolepis and several brachythoracids. The ventral surface of the IL is covered with tubercles, similar to those on the edges of the AVL. The tubercles on the dorsal surface are regularly arranged into four or five rows. The presence of an anterior median ventral is seen in SAM-—K4649 where there is the impression of tubercles between the IL and the AVL of either side. How- ever, the shape of this plate and the existence of a posterior median ventral cannot be demonstrated in this material. An anterolateral (AL) has not been found associated with the ventral armour, but two specimens showing most of this plate can be referred to Barry- dalaspis because they show ornament similar to that on the AVL. A note of caution must be introduced when associating plates by using similarities in ornament. White (1969: 303) has pointed out that in Heightingtonaspis anglica Traquair the ornament on the AL may differ considerably from that on the AVL. However, in the South African material there are only two types of AL present, each with ornament which matches that on one or the other of the two types of ventral armour. Thus, the criterion of association by ornament seems the most reasonable with the available material. The AL is tall and relatively narrow, similar proportions being seen only among ‘dolichothoracids’ in the arctolepid described by Miles (1965). As usual the bone is raised to a focal point, which in this case is centrally placed, and from this four ridges run to the corners to divide the bone into quadrants. The posterodorsal corner is produced as in Arctolepis decipiens Woodward and, to some extent, in Neophlyctaenius sherwoodi (Denison). The pectoral emargi- nation is very wide, matching that of the AVL. Tubercles are present on the dorsal, anterior and posterior quadrants but they are very sparse on the ventral quadrant. The anterior dorsolateral (ADL) is known from two specimens, one of which (SAM-—K4648) shows part of the trunk armour preserved in lateral view. This specimen can be associated with the holotype because of the similarity of ornament and the fact that there is evidence of scales on the body. Among the ‘dolichothoracids’ the ADL of Barrydalaspis resembles that of Tiaraspis. Both are tall and narrow and have a dorsal margin which slopes postero- dorsally and have the anteroventral angle produced. This last feature is also seen in Neophlyctaenius sherwoodi (Denison 1950, fig. 2). The bone is divided into lateral and dorsolateral faces by a prominent ridge below which runs the lateral line canal. A small trochlear is developed on the anteromesial edge, immediately in front of where the ridge and lateral line converge. Beneath the trochlear the anterior margin is swollen to resemble an obstantic process, but 134 ANNALS OF THE SOUTH AFRICAN MUSEUM o. MD 10mm 0. AL Fig. 4. Barrydalaspis theroni gen. et sp. nov. Restoration of anterior dorsolateral of left side. Based on SAM-K4648. 1.1—lateral line, o. AL—area overlapped by anterolateral, o. MD— area overlapped by median dorsal, tr—trochlear. it is impossible to determine if an articular face was present. The lateral face of the anteroventral corner is marked by a prominent groove. This is a feature usually found in brachythoracid arthrodires where it receives the dorsal part of the postbranchial lamina. The presence of this groove thus suggests that the dorsal part of the AL may have been inturned. The surface of the ADL is ornamented with tubercles which become larger along the posterior margin and along the crest of the ridge. The centre of radiation is found at the base of the trochlear. One specimen (SAM-K4648) shows evidence of a posterior dorsolateral but, apart from noting the fact that it is a large plate, it is too incomplete to merit further comment. A relatively large plate is significant in showing that the lateral face of the trunk armour is not ‘reduced’ as it is in many brachythoracids. The holotype and SAM-—4643 show impressions of scales behind the trunk armour. The latter specimen shows that the scales are moderately large, deeper than long and completely cover at least 10 cm of the body. The scales immedi- DEVONIAN FISH AND PLANTS FROM SOUTH AFRICA 135 ately behind the posterior dorsolateral plate are deeper than those above or below and in this respect the squamation is similar to that seen in Sigaspis Goujet (1973, fig. 3a). There are impressions of dorsal ridge scales but no further details of these or the flank scales can be established. Relationships of Barrydalaspis The interrelationships of the arthrodire groups have recently been reviewed by Miles (1973) and Miles & Young (1977) and some rational outline of arthro- dire phylogeny has been proposed. Within the cladistic framework provided by these authors, Barrydalaspis is to be regarded as a phlyctaenioid arthrodire by virtue of the possession of a ‘ball and socket’ dermal neck joint. Unfortunately, the relationships of Barrydalaspis cannot be considered further due to lack of information about the head and the median dorsal plate. The suborder Phlyctae- nioidei of Miles contains the collateral infraorders Phlyctaenii and Brachy- thoracii. The latter can be shown to be monophyletic (Miles 1973) based on synapomorphies in features of the head and the median dorsal plate. The former, as Miles admits, is possibly a grade group. In other words monophyly has not yet been demonstrated for the Phlyctaenii (families Tiaraspidae, Groenland- aspididae, Phlyctaenaspidae, Williamaspididae and the genus Aggeraspis) and more rigorous analysis of the species included within the Phlyctaenii is necessary. Some recent work suggests that Tiaraspis and Groenlandaspis may be sister groups based on the synapomorphies of a high median dorsal plate and the fact that the lateral line, in crossing the posterior dorsolateral describes a sharp dorsal flexure. These two genera, plus an unnamed form from the Middle Devonian of Australia, are included in the Groenlandaspididae by Ritchie (1975). Thus to place Barrydalaspis as Suborder Phlyctaenioidei incertae sedis means that it shows the synapomorphy of that Suborder but that the material does not allow us to specify its position within that group. Barrydalaspis resembles some members of the Brachythoracii in showing a prominent groove on the ADL to receive the AL but it is not yet clear whether this should be regarded as a synapomorphy of the brachythoracids or as a feature primitive for the Phlyctaenioidei. ‘Phlyctaenaspid arthrodire’ The second type of placoderm which is recognizable in the present collec- tion is known from ventral views of the trunk armour and a partial AL which is associated with the ventral plate because of the similarity of the ornament. This arthrodire is distinct from Barrydalaspis in a number of respects: the ventral armour of the trunk is much longer and narrower; the SP is set at a much lower angle (i.e. it is more nearly parallel to the sagittal plane) and lacks the medial denticles; the SP appears to be fused with the AVL throughout its length; the lateral end of the IL is swollen to produce what is here termed an ‘elbow’; the ornament consists of coarse tubercles along the outer edge of the SP and IL 136 ANNALS OF THE SOUTH AFRICAN MUSEUM 20mm Fig. 5. ‘Phlyctaenaspid arthrodire’. Restoration of ventral thoracic armour in ventral view. Based on SAM-K4640. Abbreviations as in Fig. 1. but elsewhere is represented by minute tubercles. The proportions and shape of the ventral armour are similar to both Neophlyctaenius sherwoodi (Denison 1950) and Gaspeaspis Pageau (1969). The former is from the late Middle or early Upper Devonian of New York State, the latter from the early Middle Devonian of Gaspé Peninsula, Quebec. As in Gaspeaspis, the South African form shows AVLs which meet one another in the mid-line, leaving only small areas for the anterior and posterior median ventrals (these were not seen in the South African form). The ‘Ventrolateral- kante’ is well developed and there was probably a long postbrachial lamina. The South African phlyctaenaspid differs from both N. sherwoodi and Gaspeaspis in the relatively longer AVL and the fusion of that plate with the SP. The division between the IL and SP is recognized as a deep groove. The prominent elbow of the ILis matched elsewhere in the Lower Devonian phlyctaenaspid Dicksonosteus (see Goujet 1975, pl. 4 (fig. 1)). An incomplete AL (SAM-K4775) which can be referred to the ventral armour is tall with a wide pectoral emargination and a focal point which is situated ventral to the centre of the bone. Distinctively the dorsal margin slopes anteroventrally towards the front. This last feature is ‘SLLYA-INVS JO SUIMeIp eplony e1oweD ~s11poiyjie prdseuoryoAyyd, Jo oyejd jesrojeoINUY “g OS9PA-INVS JO SurMesp epion] eroweD “Aou “ds jo ‘usd iWosay) sidspjopdadpg JO opis 1YBII Jo o1eId [eIo\e[OINUY “WY °9 ‘Shy 137 WW O} DEVONIAN FISH AND PLANTS FROM SOUTH AFRICA aN I a a ER ASE I LE LS IIL TRISTE SB ait 138 ANNALS OF THE SOUTH AFRICAN MUSEUM rare among arthrodires but is seen in species currently referred to Phlyctaenaspis and in Arctolepis decipiens (see illustrations in White 1969, figs 2-21). In summary, this South African form is considered to be a phlyctaenaspid because it lacks the anteroventrals (a derived feature of actinolepoids (Miles & Young 1977)) and because it shows phenetic resemblances in the proportions of the ventral thoracic armour and constituent plates to certain phlyctaenaspid genera. Arthrodire head One specimen (SAM-—K4748) shows the matrix impression of the under- surface of a partial skull roof. It cannot be associated with either of the two types described above although it is of a size that would match either. Further- Cael Sere Fig. 7. Arthrodire head. Camera lucida drawing of internal cast of a partial skull roof, SAM-K4748. Ce—central, M—marginal, Nu—nuchal, Po—postorbital, Pro—preorbital, Sc.c—impression left by semicircular canals, Ce.s.c.—central sensory canal, Cl.l—cephalic division of the main lateral line. DEVONIAN FISH AND PLANTS FROM SOUTH AFRICA 139 more, it cannot be assumed that the pattern of sutures visible on the under- surface of the skull roof corresponds faithfully to that on the upper surface, which is the surface most frequently studied. For these reasons our remarks about this specimen must be limited. The anterior end of the nuchal is narrow, gently rounded and reaches a considerable distance between the centrals. The postorbital is short and broad with the posterior margin orientated transversely. These features are seen in combination in Gaspeaspis (Pageau 1969, fig. 20). The marginal has a broad area of contact with the central, a feature rarely seen in actinolepoids but common in phlyctaenaspids, Groenlandaspis and Aggeraspis. What little that can be seen of the paths of the sensory canals agrees with that expected in a ‘dolichothoracid’. Undetermined arthrodire In the collection there is one specimen (SAM-K4646, Fig. 8A) of a large arthrodire which shows coarse ornamentation unlike the forms already described Fig. 8. A. Silicone mould of the right side of part of the trunk armour of a large arthrodire, SAM-K4646. ADL—anterior dorsolateral, PDL—posterior dorsolateral, PIL—postero- lateral, 1.1—lateral line, o.AL—overlap area for anterolateral. B. ?Chimaeroid egg case. Silicone mould of SAM-K4814. 140 ANNALS OF THE SOUTH AFRICAN MUSEUM from this locality. If we have interpreted this specimen correctly, this represents an impression of the posterior part of the flank of trunk armour displaying parts of the anterior dorsolateral, posterior dorsolateral and posterolateral. The anteroventral margin of the ADL and the anterior margin of the PL are similar to those in Groenlandaspis antarcticus Ritchie (1975). Gavin Young (pers. comm.) suggests that the size and pattern of ornament shown in this specimen is similar to the ornamentation seen in the Antarctic Groenlandaspis. However, it is to be admitted that similarity in ornament is a weak basis for associating this single South African specimen with the Antarctic Groenlandaspis and in consequence this specimen is left unnamed. ?CHIMAEROID There is, in the collection, a specimen of an egg capsule (Fig. 8B), 160 mm in length and with approximately thirty-two unbranched narrow transverse ridges in each lateral flange. The egg capsules of living chimaeroids are leathery, bilaterally symmetrical and elliptical in outline. They possess a membranous lateral web, which may be strengthened by simple or branched, rib-like thickenings and the margin of which can be fimbriate or entire. Presumably by analogy with these Recent types, some eleven fossil chimaeroid eggs have been described. The earliest descriptions are of forms from the Middle Jurassic of Germany (Bessels 1869; Jaekel 1901) while the majority of later finds have been those from the Upper Cretaceous (Gill 1905; Dean 1909; Brown 1946; Voronets 1952; Vakhrameey & Pushcharovskii 1954; Obruchev 1967). Additional material has been described from the Jurassic of Canada (Warren 1947) and the Oligocene of the U.S.A. (Brown 1946). A rather differently shaped form, almost butterfly-like, has been recorded from the Triassic of Connecticut (Bock 1949) but this is possibly a pteropod. Thus the find reported here takes the known occurrence of these supposed egg capsules back into the Devonian. In this respect it is interesting to note that the Chimaeroidei extend back only to the Jurassic (Toarcian) although members of the Menaspoidei first occur in the Upper Devonian (Patterson 1967). If, however, pelvic claspers are a primitive feature of the elasmobranchiomorphs, as is suggested by their presence in ptyctodonts, then it is reasonable to assume that placoderms also produced egg cases (Patterson 1965) and that the egg case reported above may be that of a placoderm. ACANTHODIAN A specimen of an incomplete pectoral spine of Gyracanthides (Fig. 9) is present in the collection. The spine is flattened and is deeply grooved along the medial edge of the posterior half. The upper and lower surfaces are marked with tuberculated ribs which pass obliquely across the surface and meet in a chevron pattern along the free edge. A prominent ridge runs the length of this free edge. DEVONIAN FISH AND PLANTS FROM SOUTH AFRICA 141 30mm Fig. 9. Gyracanthides sp. Pectoral spine. Freehand sketch of rubber latex cast. Gyracanthides is known by two species, G. murrayi (Woodward 1906), from the Mansfield Slate of Victoria, Australia, and G. warreni White (1968) from the Aztec Siltstone of Victoria Land, Antarctica. The latter is known only from two specimens representing part of the base of a pectoral spine, perhaps of the same individual (White 1968). The South African Gyracanthides is similar to G. warreni in a number of biometric details. In both, the ribs and alternating grooves are of equal width; the ribs are almost straight; there are nine to ten ribs per cm at the base of the insertion area and thirteen to fifteen just behind this level (these counts are taken by placing a cm scale at 90° to the direction of ribbing) and the tubercles on the ribs are very closely packed. Thus, the South African Gyracanthides and G. warreni are similar in all features in which the two can be compared and there is every reason to regard them as being conspecific. Gyracanthides murrayi differs from G. warreni in that the pectoral spine and the ribs are more strongly curved and the ribs are spaced further apart, as are the tubercles upon the ribs. THE PLANT REMAINS The plants associated with the fish fauna consist of various types of lyco- pods, preserved either as ‘external moulds’ (‘impressions’), or in some cases as matrix infillings of the cortical cavity within the stem (‘endocortical casts’). Before describing the fossils, this form of preservation must be briefly reviewed. Available for study were a number of latex casts prepared from moulds in the original matrix in South Africa, and sent to London. There was also a rather smaller number of specimens on the rock matrix, where this had been sent to London for study of the associated fish. From these, additional casts were made in latex or silicone rubber. The lycopods represented evidently had a relatively tough cortical cylinder (Fig. 10A) within which (by analogy with Palaeozoic lycopods preserved uncompressed, as petrifications) a small stele (circle in that figure) was sur- rounded by a broader middle cortical cavity. As the fragmented stems became buried in matrix, this cavity became filled with mud (with or without the stelar woody cylinder at the centre). 142 ANNALS OF THE SOUTH AFRICAN MUSEUM we wege+vweeeeweewiwivif 2 + ef ef + ~~ = '@@@e@00800060006 0 06600606 0 @ e e ee e eee e d’@ Fig. 10. Compression of stem of a lycopod such as Archaeosigillaria, producing the two types of fossil, endocortical cast (EC) and the external mould (EM). A. Stem lying horizontally in matrix (M). Matrix both surrounds it and fills the cortical cavity. The stem outer surface shows protruding leaf cushions; the only features on the cortical inner surface are indentations corresponding to the position of the passage of a vascular trace into the cortex. The positions of these indentations will correspond to the position of the leaf cushion. B. Compressed matrix (M) and coaly matter of the plant (Co). (The lower half only is shown.) On the outer surface protrusion of the leaf cushions is reduced with little distortion of their horizontal dimensions. The greater compressibility of plant tissue as against matrix caused collapse of DEVONIAN FISH AND PLANTS FROM SOUTH AFRICA 143 On compression, with resulting collapse of the plant tissue, the matrix filling the cortical cavity became compressed to a rod of matrix of ellipsoidal cross-section (Fig. 10B). In all cases studied, the plant material was missing, being represented either by a gap (white region in Fig. 10C), or by a dark brown (iron-rich?) porous mineral substance. The surface of an endocortical cast (EC in Fig. 10C) of such a fossil is revealed by a fracture plane passing between the cast and the enclosing matrix, i.e. the external mould. It generally shows (Fig. 14E) a topography dependent partly on any indentations on the inner surface of the cortex (e.g. passage of leaf trace, etc.) and partly on the collapse of the plant material on compression into features (e.g. leaf cushions) on the stem outer surface. Endocortical casts in this material typically show several series of longitudinal rows of bosses (Figs 12E, 14E) which correspond to the positions of leaf cushions on the original stem outer surface. They may show a small central protrusion which was a depression on the inner face of the cortex (EC in Fig. 10C). Latex ‘moulds’ were prepared (La in Fig. 10D) from such endocortical casts. It must be emphasized that the topography of these endocortical casts corresponds only in the broadest way to the original external appearance of the stem. The external mould (or impression in the matrix) of the original outer surface shows a closer approximation to the original appearance of the stem. External features (e.g. leaf cushions, represented symbolically by ridges in Fig. 10A) appear in somewhat reduced topography, on such an external mould (EM in Fig. 10E). Latex casts (La in Fig. 10E), approximating to the original outer surface of the plant, may be prepared from such a mould (e.g. Fig. 13A—B). Where leaves were still attached to such a stem, evidence of their presence is normally seen on an external mould. Where a leaf has a broad expanded base (leaf cushion) the leaf plus cushion became compressed on the upper and lower surfaces of the cylindrical stem. Such a leaf is shown in Figure 11A as though lying on the lower surface of a stem. On compression and subsequent removal (by weathering or diagenesis) of the plant material the leaf cushion is repre- sented in the external mould by a depression in the matrix (Fig. 11C). At the bottom of this depression a narrow slit extends into the rock matrix representing the site of the lamina of the leaf. When latex is poured on to such a mould, the shape and topography of the leaf cushion are shown faithfully, but usually the latex (La in Fig. 11D), does not penetrate into the narrow mould of the the matrix filling the endocortical cavity into the area behind the leaf cushions. The matrix forming the endocortical cast now also shows raised bosses, corresponding to the positions of leaf cushions. C. Subsequently some, or all, of the coaly matter is removed leaving the two matrix surfaces, the endocortical cast (EC) and the external mould (EM). D. Upper half of the endocortical cast (EC) exposed by fracture plane passing over cast surface, together with latex mould (La) prepared from it, a negative version of the cortical cavity (a plaster cast taken from this mould will correspond to the original endocortical cast). E. Latex cast (La) of the external mould (EM), corresponding to the original (compressed) stem surface with protruding leaf cushions. 144 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 11. Archaeosigillaria plumsteadiae sp. nov. Compression of leaf and leaf cushion resulting in the production, in the latex mould, of a leaf cushion showing only the ‘false leaf scar’. A. Original shape of leaf and cushion (as seen in profile at edge of stem in Figs 12C, 13B), surrounded by uncompressed matrix (M) which also fills the endocortical cavity; the leaf lies in the matrix as on the lower surface of a horizontal steni. PT —plant tissue. B. Compression distorts the shape of both leaf base and leaf. The leaf cushion becomes less protruding, the thickness of the leaf lamina is reduced and its angle of emergence is decreased. Co—coaly matter. C. Removal of coaly matter (by subsequent diagenesis or weathering) gives a negative impression of the leaf cushion with a narrow mould of the leaf lamina going down into the matrix. D. Latex (La) applied to this negative mould fails to penetrate the narrow mould of the leaf lamina, leaving a ‘false leaf scar’ on the (positive) cast of the leaf cushion (Fig. 13A—B, f.l.s. in Fig. 12D). leaf itself. As a result, the leaf cushion, seen (as a protruding ‘positive’ feature) on the latex cast prepared from this mould, does not show the leaf but merely a transverse marking—a kind of ‘false leaf scar’—in the middle of the cushion (f.l.s. in Fig. 12D) where the latex failed to flow into the narrow space repre- senting the leaf. In such a specimen the leaf may be seen in profile at one or both margins of the compressed stem (Fig. 12C; see left side of Fig. 13B). In one specimen of Archaeosigillaria cf. picosensis the narrow mould of the leaf itself was wide enough to allow latex to enter, so producing a somewhat flattened replica of the original leaves attached to the leaf cushion surface (Figs 12A-B, 14A, F). Although, in what is said above, the distinction is made between an endo- cortical cast and the external mould, the former may be encountered lying, in effect, within the latter (Fig. 14B). In this case, the visible surface features are those of the endocortical cast, but the leaves may be seen in profile at the margin (Fig. 14C-D). Unfortunately, there were no cases of ‘part-and-counterpart’ specimens, where external mould and endocortical cast could be seen on opposed, fractured, faces of matrix. . Two well-defined taxa of lycopods may be recognized in this material; DEVONIAN FISH AND PLANTS FROM SOUTH AFRICA 145 one is made the basis of a new species and the other is compared with a South American species. Two further kinds of rather less satisfactory lycopod fossils are also described. Division TRACHEOPHYTA Class LYCOPsIDA Order PROTOLEPIDODENDRALES Family Archaeosigillariaceae Genus Archaeosigillaria Kidston Archaeosigillaria plumsteadiae sp. nov. (Figs 12C-D, 13A-B) Etymology Named after Dr Edna Plumstead who has contributed so much to our knowledge of South African Palaeozoic plants. Holotype 185B, external mould (Fig. 13A). Paratypes 161, 190A, 192A, 175. Silicone and rubber casts from the type material, two natural external moulds, are shown in Figure 13A-B. Diagnosis Fragments of lycopod leafy stems up to 1,5 cm diameter and 11 cm in length, represented by external moulds. Stem surface completely covered by hexagonal cushions, each typically 6,5 mm wide by 4 mm high, upper and lower edges of leaf cushions flat and in contact with cushions above and below. Cushions arranged in vertical ranks with corresponding orthostichies in alter- nating series. Leaves seen only in profile at margin of flattened stem mould; free part of leaf typically 6 mm long, leaving stem at about 45° and diverging from it, the apical part of the leaf being almost perpendicular to the stem. Shape of lamina otherwise unknown, but evidently not thicker than 1 mm. No evidence of leaf abscission nor of ligule or ligule pit. Remarks As indicated above, the shape of the leaf lamina may be seen in profile at the edges of the compressed stem (Fig. 13A, left-hand side) but over the stem surface the leaves (represented by cavities in the matrix of the fossil) cannot be seen (Fig. 11D). The latex poured into this natural mould evidently failed to flow into these cavities. Thus the only clear feature on each hexagonal leaf cushion is a transverse line at the widest part of the cushion (f.l.s. in Fig. 12D). This is rather comparable to the ‘false leaf scar’ of a lycopod compression 146 ANNALS OF THE SOUTH AFRICAN MUSEUM fossil when a fracture plane has detached the leaf in the counterpart fossil (cf. Chaloner & Boureau 1967: 533). It must be borne in mind that whatever is seen of the leaf lamina in profile at the flattened stem margin (Figs 12C, 13B) is only a minimum length. It may actually have been longer, depending on the shape and taper of the leaf (cf. Lacey 1962, fig. 12A, D). There is no evidence of Fig. 12. Archaeosigillaria and ‘lycopod endocortical cast . A-D. Leaves and leaf bases of both species of Archaeosigillaria. Drawn from photographs of the latex cast. A-B. Archaeosigillaria sp. cf. A. picosensis Krausel & Dolianiti. A. Rhomboidal leaf cushion with leaf attached, 2 mm long, probably complete, lying parallel to the stem sur- face, with its sides tapering abruptly to form a spatulate tip; SAM-—K4785. B. Another leaf cushion, with a small tab-like and probably incomplete leaf emerging from the centre of the leaf cushion; SAM-—K4785. C—D. Archaeosigillaria plumsteadiae sp. nov. C. Leaf seen in profile at side of stem, showing uncompressed dimensions of leaf cushion, and thickness of leaf lamina (as in Fig. 11A). Leaf emerging at 45° and diverging from the stem to become nearly perpendicular to it; 190A. D. Leaf cushion of holotype (as in Fig. 13A). Hexagonal, strongly protruding leaf cushion, featureless apart from the false leaf scar (f.l.s.), a transverse line at the widest part of the cushion; 185B. E. ‘Lycopod endocortical cast’ with bosses on the surface produced by processes of compression and collapse explained in Figure 10. The bulge in the centre of each boss is interpreted as a feature produced by the site of passage of the leaf trace into the cortex. DEVONIAN FISH AND PLANTS FROM SOUTH AFRICA 147 Fig. 13. A-B. Archaeosigillaria plumsteadiae sp. nov. Latex casts coated with ammonium chloride and illuminated from top left. A. Holotype, 185B. Stem showing three rows of con- tiguous hexagonal leaf cushions. x 3. B. Paratype, 161. x 3. Both specimens showing leaves in profile on the left. C_D. Lycopod ?endocortical cast, cf. Haplostigma irregulare Seward. Illuminated from top left. SAM-K4744. C. Plaster cast prepared from latex mould, representing the original rock surface, showing circular raised features corresponding to positions of leaf bases. x 3. D. Latex mould showing positions of leaf bases as depressions. x 3. 148 ANNALS OF THE SOUTH AFRICAN MUSEUM leaf shedding (abscission) in these specimens; none shows a leaf scar, and hence no parichnos or vascular scar could have been represented. There is no evidence of a ligule. One specimen shows several missing leaf bases (Fig. 13A, bottom and right of specimen). The most probable explanation of this is that secondary growth of the cortex caused eventual sloughing off of the whole leaf cushions, as in the case of Sigillaria (Chaloner & Collinson 1975). Grierson & Banks (1963), in their emended generic diagnosis of Archaeo- sigillaria (which the authors generally follow), do not regard this genus as having leaf cushions, and refer only to ‘enlarged leaf bases becoming hexagonal on larger stems’. Their figure of the leaf of A. vanuxemi in profile (their pl. 35 (fig. 4)) conforms closely with that seen in our specimens. They further state (Grierson & Banks 1963: 239) that the leaves of their plant ‘were persistent and that the six-sided leaf bases cannot be regarded as true cushions from which the leaf abscissed but rather as merely the enlarged base of the leaf’. The authors prefer to follow the broader concept of a leaf cushion developed by Meyen (1976) and regard the swollen leaf bases of their plant as constituting cushions even though the leaf was not abscissed, and would emphasize that they differ from Grierson & Banks only in terminology, and not in interpretation of their plant. The broad leaf cushions of the present species with flat upper and lower faces in contact with cushions above and below are reminiscent of some Upper Carboniferous Sigillaria species, particularly those of the Favularia group (e.g. S. elegans Brongniart, particularly the specimens figured as S. hexagona Brongniart, a synonym of the former species). However, of course, Archaeo- sigillaria plumsteadiae differs from all Sigillaria species in having no indication of leaf abscission. The appearance of the leaf cushions and the profile view of the leaves in the present species are most closely matched in Archaeosigillaria kidstoni where the much smaller leafy shoots show comparable hexagonal leaf cushions (Lacey 1962, fig. 12B—C; Chaloner & Boureau 1967). Archaeosigillaria plumsteadiae shows good general agreement with several species of Archaeosigillaria including A. vanuxemi, A. kidstoni, and with the Ghanaian A. essiponensis (Mensah & Chaloner 1971). It is noteworthy that these are all Lower Carboniferous species. Comparison with A. caespitosa (Schwartz) Plumstead from the Witteberg is limited, as the holotype of this species is apparently an endocortical cast (Plumstead 1967, pl. 11 (fig. 2)), and none of the specimens she assigns to that species shows details of leaf shape. A. plumsteadiae differs from all other species of the genus with hexagonal leaf cushions in their being broader than long. It must be accepted that this distinction is relatively trivial, but on available information this separates the present species consistently from earlier described ones. The authors endorse Meyen’s (1976) emphasis on the need for as wide a range of specimens as possible in order to establish the extent and variability in fragments of lycopod stems. Unfortunately, as here, such a range is not always available. DEVONIAN FISH AND PLANTS FROM SOUTH AFRICA 149 Archaeosigillaria sp. cf. A. picosensis Krausel & Dolianiti (Figs 12A-B, 14A, F—G) Material SAM-K4785 (Fig. 12A, B; Fig. 14A, F), ‘D’ (Fig. 14G), SAM-K4650. Description This species is represented by several specimens showing fragments of stem outer surface, seen only as a natural mould (i.e. a negative version of the original stem surface) in the matrix. The stem surface is formed of contiguous rhomboidal to rounded-rhomboidal leaf cushions (expanded leaf bases), typi- cally 4 mm wide by 2,5 mm high, arranged in prominent alternating vertical series. Arrangement of the cushions must have been either in alternating whorls or a very low angle spiral. The leaves were still in attachment, the free portion being about 1 mm wide and 2 mm long. Remarks No leaves are seen in profile at the edges of the specimens as in the last species, perhaps because they represent fragments of a larger stem rather than parts of a complete cylinder. However, latex evidently penetrated the leaf cavities in the mould more freely than in A. plumsteadiae (possibly due to greater thickness of the leaves) so that something of the form of the leaves is seen in the latex cast of the stem prepared from the mould (Figs 12A—B, 14A, F). This can be compared with the situation in A. conferta (Menendez 1965) where leaves are seen on the surface because the fracture plane exposing the fossil followed the plane of the leaf laminae rather than the stem surface. It is possible that these small tab-like leaves (seen in the latex casts) represent incomplete infill by the latex of a larger leaf cavity, or possibly the original leaf shape was somewhat eroded before fossilization. It is interesting to compare the situation here with that in A. plumsteadiae, where the latex did not penetrate the leaf cavity. On one of the specimens (Fig. 14G) there is evidence of secondary cortical growth resulting in the lateral separation of the leaf bases revealing (?) cortical tissue between them. This is, of course, a common phenomenon in many Lepidodendron species (Thomas 1966). In the leaf shape and arrangement this material agrees well with those species of Archaeosigillaria seen to have short tab-like leaves turning abruptly from the leaf cushion to parallel the stem surface or lie obliquely to it; these include A. vanuxemi, A. kidstoni, and A. essiponensis, with leaves showing at the side of the stem, and more particularly A. picosensis. This plant, from the Picos member in Brazil (Lower Devonian according to Krausel & Dolianiti 1957), is preserved like the specimens here as a natural mould with leaves showing on the surface and is the closest species to the Bokkeveld plant. Those authors describe their specimen (free translation from the German summary) as ‘small stems, [covered with] leaf cushions pressed together, rounded-angular to rhomboidal, also in part hexagonal, bearing in their upper part a small 150 ANNALS OF THE SOUTH AFRICAN MUSEUM thin leaf’ (literally, leaflet). It should be noted that the age of Krausel & Doli- aniti’s specimen, cited by them as Lower Devonian, may well be much younger. Sampaio & Northfleet (1973) offer an age correlation for the Picos member (of the Pimenteiras Formation) as ranging between Emsian and Eifelian; Bar & Riegel (1974) favour ‘Middle Devonian Age’. The age of A. picosensis is probably best placed only within a broad bracket between Emsian and Frasnian (Brito 1971, fig. 10). LYCOPOD ENDOCORTICAL CASTS (Fig. 14B-E) Material SAM-K4790, K4798a. Remarks The specimen illustrated in Figure 14E is one of several in the latex casts available in which the topography of the fossil appears to represent only a very blurred version of the lycopod stem. This cannot be reconciled with leaf bases or remains of attached leaves, and this fossil is interpreted as representing an endocortical cast. This shows the positions of leaves in the form of bulges or bosses on the endocortical cast, which simulate leaf bases or cushions (EC in Figs 10C—D, 12E, 14E). One of these casts K4798a (Fig. 14B-D), besides showing these vertically seriated bosses, also shows leaves in profile along the margin. The leaves appear to be preserved in a brown mineral substance which must have come to occupy the site of the original plant tissue. This mineral matter is a thin layer on the outside of the cast, representing the cortical tissue and leaf bases of the stem. The leaves are somewhat similar to those of some species of Archaeosigillaria, such as A. essiponensis (Mensah & Chaloner 1971) and A. kidstoni (Lacey 1962; Chaloner & Boureau 1967), being short with a broad base and tapering towards the apex. On the other cast SAM-K4790 (Fig. 14E) this mineral matter is not present, and on the left-hand side a cavity can be seen representing the gap between the endocortical cast and the external mould, i.e. the site of cortical Fig. 14. A, F, G. Archaeosigillaria sp. cf. A. picosensis Krausel and Dolianiti. A. Latex cast of a fragment of an external mould showing contiguous rhomboidal leaf bases with attached leaves, one of which, near top left, is more complete than the others; SAM—K4785. F. Scanning Electron Micrograph photograph of an area of the same specimen, taken on a Cambridge S600 at a stub angle of 10°. (Horizontal axis only, x 12.) G. Latex cast showing separation of leaf bases, SAM ‘D’ x 7,5. B-E. Indeterminable lycopod endocortical casts. Photographs of original rock surface. B. Part of cast immersed in alcohol, photographed to show vertical files of raised bosses on the surface. At both sides there is a thin layer of mineral matter, representing the site of original plant tissue, and showing leaves in profile, one on the left-hand edge, three on the right. SAM-K4798a. x 3. C-D. Detail of leaves (immersed in alcohol). Short leaves with a broad base tapering to a thin lamina, at about 45° to the stem surface. x 5. E. Cast photographed dry, lighting from top left, showing seriated raised bosses; SAM-K4790. x Bb 151 DEVONIAN FISH AND PLANTS FROM SOUTH AFRICA 152 ANNALS OF THE SOUTH AFRICAN MUSEUM tissue. These endocortical casts are generally comparable to those formed on the matrix infill inside A. essiponensis (cf. Plumstead 1967, pl. 11 (fig. 2); Mensah & Chaloner 1971, pl. 64 (fig. 7)), but other genera can produce similar casts (e.g. Plumstead 1967, pl. 15 (fig. 3) attributed to Haplostigma). The authors do not believe that such fossils can be assigned to genera based on charac- teristics of leaf cushion shape and prefer to leave these present specimens unassigned. Lycopod ?endocortical cast, cf. Haplostigma irregulare Seward (Fig. 13C-D) Description The specimen figured in Figure 13D (and a plaster cast prepared from that latex mould, so representing the original rock surface, Fig. 13C) shows vertically seriated round markings on a grooved stem surface, the grooves apparently separating elongated leaf cushions. This specimen is preserved as a ‘positive’ cast apparently showing detail of the surface topography, rather than the blurred bosses described above. It is believed that this is due to the plant posses- sing only a narrow zone of cortical tissue, which in compression would collapse to a uniformly thin layer more or less conforming to the stem’s original external topography on both surfaces. No leaves are in evidence in the material, and it is not clear whether they were abscissed. It is accordingly regarded as generically indeterminable but is figured since it shows some resemblance to the holotype of Haplostigma irregulare Seward, 1903, from the Bokkeveld (as refigured by Plumstead 1967, pl. 13 (fig. 3), pl. 14 (fig. 5)). It is also comparable to the speci- mens attributed to the same species by Krausel (1960, fig. 88) from the Ponta Grossa Formation of Brazil. DISCUSSION The Bokkeveld Series consists of alternating bands of sandstone and shale which vary in number and thickness over wide areas of the Cape Province (Plumstead 1967). This is overlain conformably by the Witteberg Series. Most authors divide the Bokkeveld Series into lower and upper beds which, by reason of the contained fossils, are thought to represent marine and shallow marine/ freshwater deposits respectively. One author, Swart (1950), suggests that in at least one locality shallow marine conditions persisted throughout the Bokkeveld Series. In the Barrydale area the more usual freshwater beds are clearly seen, and the fish are found in these upper beds of the Series. The interpretation of the lower beds as representing marine conditions is well founded. A large number of marine species have been described (Lake 1904; Reed 1925 and refs; Haughton 1969) including lamellibranchs, brachiopods, trilobites, gastropods, cephalopods, corals and crinoids. A consensus of opinion holds that the lower marine beds of the Bokkeveld Series are of Lower Devonian age, and Boucot ef al. (1967) are more precise in suggesting an Emsian age. DEVONIAN FISH AND PLANTS FROM SOUTH AFRICA 153 Further, several authors (Du Toit 1939; Doumani 1965; Haughton 1969) note the close similarity of the marine faunas of the Bokkeveld with the presumed contemporaneous strata in the Falkland Islands, Bolivia, Argentina, southern Brazil and Antarctica. Therefore, on the strength of the evidence of the under- lying marine sequence, the fish- and plant-bearing beds of the Bokkeveld Series cannot be older than Middle Devonian. The only information available here (J. N. Theron, pers. comm.) on the position of the fish fauna is that it is some ‘6 000 ft’ below the fish zone of the Upper Witteberg (Gardiner 1969) and as such may be anything from Middle Devonian to Lower Carboniferous. At first sight the fact that the fauna con- tains two dolichothoracid arthrodires suggests a Lower/Middle Devonian age since the dolichothoracids reached their acme in the Emsian/Eifelian (Miles 1969). Nevertheless, Groenlandaspis is a widespread late Devonian representa- tive of the dolichothoracids, while Neophlyctaenius survived into the Frasnian in the eastern United States (Denison 1950). Since one of the dolichothoracids is a completely new form and the other closely resembles Phlyctaenius, their stratigraphic significance is not apparent. Similarly, the occurrence of large arthrodire plates resembling Groenlandaspis does no more than confirm a Middle or Upper Devonian age. However, from the same general area as the other specimens, but not, unfortunately, as accurately located within the sequence, occurs a spine of the large acanthodian Gyracanthides. Elsewhere Gyracanthides is recorded from the Upper Devonian of Victorialand, Ant- arctica (White 1968) and the Lower Carboniferous of Mansfield, Australia (Woodward 1906). The Upper Devonian Antarctic fish fauna from Victorialand is charac- terized by the presence of Bothriolepis, Phyllolepis, Groenlandaspis, holopty- chiids and various acanthodians and sharks (Gavin Young pers. comm.) whereas the Australian Lower Carboniferous fauna from Mansfield has Strep- sodus, Ctenodus, Elonichthys and three acanthodian genera but no placoderm genera. There can be little doubt that the Antarctic fauna is Upper Devonian (Young 1974) and if this South African fauna is to be interpreted as being of similar age then the absence of more typical Upper Devonian forms such as Bothriolepis, Phyllolepis and Holoptychius from the Bokkeveld is difficult to understand. Nevertheless the presence of a typical Lower Carboniferous fish fauna in the overlying Witteberg Series, some 1 800 m (6000 ft) above the fish-bearing layer of the Bokkeveld convinces the authors that the fauna under discussion must be at least of Upper Devonian age and the only safe con- clusion that can be drawn is that the fish fauna is Middle/Upper Devonian. The only plants in this flora which may be of significance in dating these rocks are the two species of Archaeosigillaria, and in particular the very distinc- tive A. plumsteadiae. Plants belonging to this genus range from Middle Devonian to Upper Carboniferous (Banks 1960; Grierson & Banks 1963; Lejal 1970; Mensah & Chaloner 1971). Recently Lejal-Nicol (1975) has described a number of typically Middle or Upper Devonian and Lower Carboniferous lycopod 154 ANNALS OF THE SOUTH AFRICAN MUSEUM genera (including Protolepidodendron, Lepidodendropsis, Lepidosigillaria and Archaeosigillaria) from Libya. Lejal-Nicol maintains that these deposits are of Lower Devonian age, and a further flora is described containing A. kidstoni from rocks believed to be of Pre-Siegenian age (either Gedinnian or Siluro- Devonian). These genera, which are characterized by various peculiarities of their leaf cushion shape and arrangement, do not appear in Europe and North America until the Middle or Upper Devonian (Grierson & Banks 1963; Chaloner & Boureau 1967). Typical lycopods of the Lower Devonian from continents other than Africa (e.g. Drepanophycus and Baragwanathia) show no significant development of leaf base expansion comparable to the cushions of the later arborescent lycopods. Even in the Middle Devonian lycopods with a leaf cushion or cushion-like feature (e.g. Protolepidodendron, Leclercqia, and Colpodexylon) this feature is poorly developed compared with that seen in the arborescent lycopods of the late Devonian and early Carboniferous. On this basis an extreme age bracket is put on these Bokkeveld lycopods as Middle Devonian to Lower Carboniferous, with the strongest possibility of their being Upper Devonian (Frasnian—Fammenian). It may be useful to note that shoots of lycopods such as Archaeosigillaria were evidently among the more robust of plant remains occurring in the Upper Palaeozoic; they commonly survived when no, or few, other plants were repre- sented in coarse non-marine lithologies, or even in marine environments. Archaeosigillaria kidstoni occurs in the coral/brachiopod-rich Carboniferous Limestone in Britain (Chaloner & Boureau 1967) and the holotype of A. vanuxemi is closely associated with a brachiopod fauna. The present association of lycopods with fish remains is therefore not surprising. The evidence of both the fish and the plants suggests, therefore, that these fossils come from an horizon within the Bokkeveld Series that is either Middle or Upper Devonian. Hopefully, further collecting in these strata will yield fossils giving a more precise stratigraphic position. ACKNOWLEDGEMENTS We should like to thank Dr J. N. Theron for bringing this material to our attention. Our thanks are also due to Drs R. S. Miles and G. C. Young for comments on the fishes, and to Dr O. Rosler and Professors H. P. Banks and J. D. Grierson for comments on South American stratigraphy and the lycopods. Finally, we thank the authorities of the South African Museum for allowing us to comment on this material. REFERENCES BANKS, H. P. 1960. Notes on Devonian Lycopods. Senckenberg. leth. 41: 59-88. Mi BAr, P. & RiEGEL, W. 1974. Les microflores des séries paleozoiques du Ghana (Afrique | occidentale) et leurs relations paléofloristiques. Bull. Sci. nat. Géol. 27 (1-2): 39-58. BESSELS, R. 1869. Ueber fossile Selachier-Eier. Jh. Ver. vaterl. Naturk. Wiirtt. 25: 152-155. Bock, W. 1949. Triassic chimaeroid egg capsules from the Connecticut valley. J. Paleont. 23: 515-517. 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A new arthrodire from the New York State Devonian. Am. J. Sci. 248: 565-580. DENISON, R. H. 1958. Early Devonian fishes from Utah. Part III. Arthrodira. Fieldiana, Geol. 11: 461-551. Doumanl, G. A. 1965. Geologic Setting. Lower Devonian fauna of the Horlick Formation, Ohio Range, Antarctica. Antarctic Res. Ser. Washington 6: 241-248. Du Torr, A. L. 1939. Geology of South Africa. 2nd ed. London: Oliver & Boyd. GARDINER, B. G. 1969. New palaeoniscid fishes from the Witteberg Series of South Africa. Zool. J. Linn. Soc. 48: 423-452. Git, T. 1905. An interesting Cretaceous chimaeroid egg-case. Science, Washington 22: 601-602. GouseT, D. 1973. Sigaspis, un nouvel arthrodire du dévonian inférieur du Spitzbergen. Palae- ontographica (A) 143: 73-88. GouseT, D. 1975. Dicksonosteus, un nouvel arthrodire du dévonian du Spitsberg: remarques sur le squelette visceral des dolicothoraci. Colloques int. Cent. natn. Rech. scient. 218: 81-99. GRIERSON, J. D. & Banks, H. P. 1963. 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DEVONIAN FISH AND PLANTS FROM: SOUTH AFRICA 157 ADDENDUM ADDED IN PRESS After preparing this manuscript for publication, the authors received (August 1979) an offprint of Plumstead’s (1977) account of Zosterophyllum de-vriesii and Z. bokkeveldensis. (The former name is here hyphenated to give a single epithet, in accordance with the International Code of Botanical Nomen- clature, Art. 23.) It is immediately evident from Plumstead’s figures and speci- men citations that the species described above as Archaeosigillaria plumsteadiae is based on the same fossil assemblage (and in part the same specimens ?), from the same locality, as her Zosterophyllum de-vriesii. No basis in her paper is found for revising the views expressed here, that these cylindrical structures covered with closely spaced hexagonal leaf cushions represent a lycopod vege- tative axis and not a zosterophyll fructification. Her photographs and her text-figure 3 (central figure) clearly show what has been here interpreted as the free tips of the leaves, seen in profile at the stem margin (her “bisected empty sporangial sacs’). The clear validity and priority of Plumstead’s specific name is acknowledged, and accordingly it is reassigned: Archaeosigillaria de-vriesii (Plumstead) comb. nov. Synonyms: Zosterophyllum de vriesii Plumstead, 1977: 270, text-fig. 3, pl. 1 (figs 1-10). Archaeosigillaria plumsteadiae Chaloner et al. 1979 (this paper): figs 12C—D, 13A-B. Plumstead’s Zosterophyllum bokkeveldensis does not appear to be strikingly distinct from Z. de-vriesii, but her view that they are distinct species is not challenged. It is agreed that they are congeneric (i.e. may both be placed in Archaeosigillaria), but no formal reassignment of the former species is suggested. The age implication of the authors’ systematic assignment of these plant fossils is, of course, at variance with Plumstead’s. In rejecting assignment to Zosterophyllum, the suggestion of a Middle or Upper Devonian horizon rather than the Lower Devonian which was implicit in attributing these fossils to Zosterophyllum, is sustained. REFERENCE PLUMSTEAD, E. P. 1977. A new Phytostratigraphical Devonian Zone in southern Africa which includes the first record of Zosterophyllum. Trans. geol. Soc. S. Afr. 80: 267-277. W. G. CHALONER London, September 1979 | % Ci on atl aa srt Nan 6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature (particularly Articles 22 and 51). : Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. noy., comb. nov., syn. Nov., etc. ‘ An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a - species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, i.e. all published scientific names by which the species previously has been designated are listed in chronological order, with all references to that name following in chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845) Figs 14-1SA Nucula (Leda) bicuspidata Gould, 1845: 37. Leda plicifera A. Adams, 1856: 50. Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b). Nucula largillierti Philippi, 1861: 87. Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. 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Adult female from mid-tide region, King’s Beach Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973. Note standard form of writing South African Museum registration numbers and date. 7. SPECIAL HOUSE RULES Capital initial letters (a) The Figures, Maps and Tables of the paper when referred to in the text e.g. ‘... the Figure depicting C. namacolus ...’; *. . . in C. namacolus (Fig. 10)...’ (b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by initials or full names e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene (c) Scientific names, but not their vernacular derivatives e.g. Therocephalia, but therocephalian Punctuation should be loose, omitting all not strictly necessary Reference to the author should be expressed in the third person Roman numerals should be converted to arabic, except when forming part of the title of a book or article, such as ‘Revision of the Crustacea. Part VIII. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes .. .” ‘Smith (1969: 36, fig. 16) describes . ‘As described (Smith 1969a, 19695; ates sh ‘As described (Haughton & Broom eo “As described (Haughton et al. 1927) . : Note: no comma separating name anit year Dagination indicated by colon, not p. names of joint authors connected by ampersand et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o, scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BuLLouGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FiscHer, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FiscHer, P.-H., Dvuvat, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gen. 74: 627-634. Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Ann, Mag. nat. Hist. (13) 2: 309-320. Konn, A. J. 19606. Spawning behaviour, SRE masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4) os TuHreELeE, J. 1910. Mollusca: B. Cobirlncenbaes Gastropoda marina, Bivalvia. In; SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 81 Band February 1980 Februarie Part 4 _ Deel ONTOGENY AND SEXUAL DIMORPHISM IN AULACEPHALODON (REPTILIA, ANOMODONTIA) By S.M. TOLLMAN EF. E. GRINE & B. D. HAHN Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na gelang van die beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad OUT OF PRINT/UIT DRUK 1, 2(1-3, 5-8), 3(1-2, 4-5, 8, t-p.i.), 5(1-3, 5, 7-9), 6(1, t-p.i.), 70-4), 8, 9(1-2, 7), 10(1-3), 11(1-2, 5, 7, t—p.i.), 15(4-5), 24(2), 27, 31(1-3), 32(5), 33 Copyright enquiries to the South African Museum Kopieregnavrae aan die Suid-Afrikaanse Museum ISBN 0 908407 87 4 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., , Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap ONTOGENY AND SEXUAL DIMORPHISM IN AULACEPHALODON (REPTILIA, ANOMODONTIA) By S. M. TOLLMAN F. E. GRINE* Department of Anatomy, Medical School, University of the Witwatersrand, Johannesburg & B. D. HAHN Department of Applied Mathematics, University of the Witwatersrand, Johannesburg (With 10 figures and 6 tables) [MS. accepted 11 October 1979] ABSTRACT A large number of Aulacephalodon crania have been examined by means of biometrical (allometric) methods. The results of this investigation indicate that this sample represents a morphometrically homogeneous group, and that probably only a single species of Aulacepha- lodon, viz. A. baini, is represented in the Permian sediments of South Africa and Zambia. A qualitative analysis of nasal boss and temporal arch morphology indicates that these features are related to sexual dimorphism. Sexual dimorphism of the cranium appears to have been expressed in individuals with a basal skull length of between 190 and 245 mm. The geographical distribution of these fossils has been examined also. CONTENTS PAGE Introduction . 4 : 4 ; 3 All i) Material and methods . ; ; ‘ . 162 Cranial growth and variability . ; : . 165 Sexual dimorphism . ; : ‘ ‘ . 169 Distribution . : : ; : 3 a Ze) Discussion . : ; : : . . 180 Summary and conclusions p : ; = 83 Acknowledgements . : ; ; : . 184 References . F : ; : ; Ze l84: INTRODUCTION Numerous anomodont fossils have been recovered from lower Beaufort Group sediments of South Africa. Well over 100 anomodont species have been described from Cistecephalus Zone strata alone. The Cistecephalus Zone, as defined by Kitching (1970, 1977), includes both Broom’s (1906) Endothiodon Zone and the lower and middle portions of his Cistecephalus Zone. Kitching (1970, 1971) has classified the upper horizons of Broom’s (1906) Cistecephalus Zone as the Daptocephalus Zone. 159 *Present address: South African Musem, Cape Town. Ann. S. Afr. Mus. 81 (4), 1980: 159-186, 10 figs, 6 tables. 160 ANNALS OF THE SOUTH AFRICAN MUSEUM One of the most commonly occurring forms in these sediments is a rather homogeneous group of anomodonts which have been referred to the genus Aulacephalodon by Haughton & Brink (1954) and Keyser (1969). The fossil remains of this reptile appear to have a rather limited stratigraphic range; © in the Cistecephalus Zone, they occur in sediments with a vertical thickness of some 330 m, but they are concentrated in a relatively thin horizon (about 80 m) in which the zone fossil, Cistecephalus, occurs abundantly (Keyser 1969). Aulacephalodon has been recovered, albeit rarely, from within the first 3 m of the overlying Daptocephalus Zone (Kitching 1977), and fossils of Aula- cephalodon have been found in comparatively large numbers at the northern (Chikonta) localities of the upper Member of the Upper Madumabisa Mudstone ‘Formation’, Luangwa Valley, Zambia (Drysdall & Kitching 1963). A number of genera and species of aulacephalodonts have been described, but only a few have been based on more than fragmented fossils, and none has been diagnosed adequately. In very few instances has either ontogenetic growth or sexual dimorphism been considered in the description of a new taxon. Owen (1844) described the first species of this group, on the basis of a single specimen, as Dicynodon baini. He later referred a second specimen to D. baini because it showed “the same character of the tusk’ as the type-species (Owen 1876). A second species, D. tigriceps, was described also by Owen (1855). Seeley (1898) divided Dicynodon into two subgenera. He proposed that those forms of Dicynodon which exhibit a short snout and a wide cranium be included in the subgenus Aulacephalodon. Broom described two further species of Dicynodon, viz. D. laticeps (Broom 1912) and D. moschops (Broom 1913). In 1921 he proposed a new genus, Bainia, for the ‘tusked specimens of || Dicynodon’; he included the species D. baini, D. tigriceps and D. laticeps in the genus Bainia and named two more species, B. peavoti and B. haughtoni | (Broom 1921). Later, Broom (1932) recognized the validity of Seeley’s (1898) subgeneric name Aulacephalodon but, as pointed out by Keyser (1969), he altered the spelling, probably as a mistake from the spelling of Seeley’s (1898) other subgenus Aulacocephalus, to Aulacocephalodon. Broom considered that, ‘Aulacocephalodon ... ought to be accepted for the group of broad skulled anomodonts typified by Dicynodon baini if we regard them as worthy of separate generic rank. ... Certainly they must be placed in at least a subgenus, and I think we can quite safely regard Aulacocephalodon as a distinct genus. Some years ago I proposed the name Bainia for the large broad-headed types not fully recognising the claims of Seeley’s name’ (Broom 1932: 191-192). He referred six species, namely Dicynodon baini, D. tigriceps, his own D. laticeps, D. moschops, Bainia peavoti, and B. haughtoni to the genus ‘ Aulaco- cephalodon’; and he described a new species, A. Jatissimus (Broom 1932). In the same work Broom (1932, fig. 65A—B) figured the dorsal and lateral views of an apparently nearly complete cranium with the legend, ‘view of skull of ~ Aulacocephalodon whaitsi, Broom’ ; however, there is no accompanying descrip- tion which serves to define or differentiate that taxon. Accordingly, the name ONTOGENY AND SEXUAL DIMORPHISM IN AULACEPHALO DON 161 Aulacocephalodon whaitsi does not satisfy Article 13 of the International Code of Zoological Nomenclature (1964) and it must, therefore, be considered as a nomen nudum. The generic name ‘Aulacocephalodon’ was used subsequently by Broom (1936, 1937, 1940, 1941, 1948), Broom & George (1950), Van Hoepen (1934), and Haughton & Brink (1954). However, Keyser (1969) has pointed out that the name ‘Aulacocephalodon’ is probably an incorrect subsequent spelling of Seeley’s name Aulacephalodon and thus ‘Aulacocephalodon’ has no status in nomenclature; the correct generic name is Aulacephalodon Seeley. Broom (1928) described the species Dicynodon milletti, and Van Hoepen (1934) placed this species in Aulacephalodon; Keyser (1969) has referred it to Oudenodon, as a junior synonym of O. baini, and he has noted that Broom’s (1913) Dicynodon moschops (which Broom referred to Aulacephalodon as a valid species in 1932) probably represents a valid species of Pelanomodon. To date some seventeen species which have been referred to Aulacephalodon have been described (Table 1). Keyser (1969) has examined the supposed features that have been used in the diagnosis of various Aulacephalodon species and has concluded that, for most of these characters, their variability and susceptibility to diagenetic distortion makes them highly questionable as taxonomic criteria. He made the important observation that, since many of the features used to distinguish the various species are to at least some extent size-dependent, the possibility that these species are all synonyms deserves consideration. TABLE 1 List of suggested synonyms of Aulacephalodon baini (Owen). Dicynodon baini. : ‘ Owen, 1844 Dicynodon tigriceps : 5 Owen, 1844 Dicynodon laticeps . : j Broom, 1912 Bainia peavoti : ; ‘ Broom, 1921 Bainia haughtoni . : ; Broom, 1921 Aulacephalodon latissimus ; Broom, 1932 Aulacephalodon nesamanni : Broom, 1936 Aulacephalodon nodosus . : Van Hoepen, 1934 Aulacephalodon luckhoffi . : Broom, 1937 Aulacephalodon hartzenbergi . Broom, 1937 Aulacephalodon coatoni . : Broom, 1941 Aulacephalodon brodiei_ . : Broom, 1941 Aulacephalodon cadlei_ : Broom, 1948 Aulacephalodon pricei ; : Broom & George, 1950 Aulacephalodon vanderhorsti . Broom & George, 1950 Consequent upon Keyser’s suggestion, the hypothesis entertained in this study was that many, if not all, of the aulacephalodont specimens which have been described possibly constitute an ontogenetic series of a single species of Aulacephalodon. A number of aulocephalodont crania were biometrically analysed in an attempt to ascertain whether an ontogenetic growth series could be demonstrated for this anomodont, and also whether the phenomenon of allometric growth could explain the supposed morphological differences between 162 ANNALS OF THE SOUTH AFRICAN MUSEUM the various proposed taxa. The possibility that sexual dimorphism was expressed in the cranium of Aulacephalodon (as suggested by Broom 1937, 1948; so 1969) has been examined. MATERIAL AND METHODS Thirty-three specimens were examined. The material ranged from relatively undistorted crania to portions of the skull. The principles of relative (allometric) growth were applied to 18 of the crania (Table 2, Nos. 1-18) whilst the remainder of the specimens received less rigorous biometric treatment. Some 31 different measurements were defined (Fig. 1), but, because of the often fragmentary and distorted nature of the fossils, there were only 4 specimens for which all 31 TABLE 2 List of all specimens of Aulacephalodon examined in this study. Previous taxonomic Specimen Number designation Description Vv Sex 1 SAM-3328 A. haughtoni (T) cranium 12 indet. 2 SAM-8747 A. latissimus (T) cranium 28 female 3 SAM-K1221 A. luckhoffi (1) cranium 25 female 4 BPI.FN. 1207 cranium 31 indet. 5 BPI.FN. 806 A. pricei (T) cranium 31 male 6 BPI.FN. 300 A. baini cranium 31 ? female 7 BPI.FN. 904 cranium 26 indet. 8 BPI.FN. 4087 A. baini cranium 30 female 9 BPI.FN. 4124 A. cf. baini cranium 29 female 10 BPI.FN. 642 A, cf. baini cranium 7 male 11 : BPI.FN. 2594 A sp. cranium 14 male 12 BPI.FN. 634 A. vanderhorsti (T) cranium 28 male 13 BPI.FN. 2460 A, sp. cranium 31 male 14 BPI.FN. 3950 A. cf. baini cranium 20 female 15 BPI.FN. 2983 A. baini cranium 29 male 16 BPI.FN. 4106 A, baini cranium 12 female 17 BPI.FN. 493 A. baini skull 23 male 18 BPI.FN. 304 A. baini cranium 29 female 19 TM 287 A. hartzenbergi (T) snout = indet. 20 T™ 1494 A. brodiei (T) skull ae female 21 T 4043 Aulacephalodon cranium * female 22 + TM 4118 A. sp. cranium na female 23 + T™ 953 ? Aulacephalodon occiput a 24 + TM 2043 A. sp. snout a female 25 + T™ 4471 A. sp. cranium = indet. 26 + T 1506 A. sp. skull ~ indet. 27 + T™ 4468 A, sp. snout = indet. 28 ++ TM, 4452 A. sp. cranium * indet. 29 + TM 4467 A. sp. cranium * indet. 30 + T™< 4469 A. sp. cranium x 31 GS K30 A. baini snout = female 32 GS R550 A, baini cranium = male 33 GS RS415 A. baini cranium = male V = number of mensurable parameters recorded; * = specimens not included in the allo- metric analysis; -- = specimens for which no locality data are available (thus, not included ; in Table 6); (1) = holotype; SAM = South African Museum, Cape Town; BPI.FN. = Bernard Price Institute for Palaeonotological Research, Johannesburg; TM = Transvaal © Museum, Pretoria; GS = Geological Survey, Pretoria. ONTOGENY AND SEXUAL DIMORPHISM IN AULACEPHALO DON 163 Fig 1. Aulacephalodon cranial measurements. A. Dorsal. B. Basal C. Occipital. D. Lateral. 1. Basal cranial length. 2. Basal temporal length. 3. Basal snout length. 4. Palatal length. 5. Pterygoid fossa length. 6. Total cranial length. 7. Temporal cranial length. 8. Bicanine breadth. 9. Least prespenoid breadth. 10. Interquadrate distance. 11. Canine-snout length. 12. Temporal fossa length. 13. Temporal fossa breadth. 14. Buccolingual diameter of canine (or socket). 15. Breadth of caniniform process. 16. Width between premaxillary ridges. 17. Least squamosal breadth. 18. Intermediate temporal breadth. 19. Greatest cranial width. 20. Breadth of occipital condyle. 21. Least post-temporal fossa breadth. 22. Post-pineal length. 23. Pre-pineal length. 24. Least interorbital width. 25. Greatest width between nasal bosses. 26. Snout width. 27. Orbital length. 28. Greatest width between prefrontal bosses. 29. Intertemporal width across pineal foramen. 30. Greatest snout length. 31. Length of nasal bosses. 164 ANNALS OF THE SOUTH AFRICAN MUSEUM metrical features could be recorded. The cranium only was measured, as the mandible is missing from most of the specimens. The variables selected for measurement were designed to reflect the overall shape of the cranium and its various parts rather than the configuration of individual bones. All measurements were taken with either a sliding vernier caliper or the top segment of an anthropometer and were recorded to the nearest millimetre. In a number of instances, owing to either distortion or breakage of the specimen, some variables could not be measured directly. In these cases estimates of the diameters were made on the basis of symmetry in order to correct for distortion and missing fragments of bone. For those crania which were too badly distorted or broken to permit reasonable estimates, the affected measurements were not recorded. The study of relative growth has been characterized by Gould (1966) as the analysis of size and its consequences. Allometric growth refers simply to the changes in proportion that occur as an organism increases in size. Such growth can be assessed and described quantitatively when measurements are fitted to the biparametric power function Wy Sede where y is a variable whose increase relative to that of another parameter, x, is considered; a is a numerical constant and f is the slope of the rectilinear plot, or, simply, the ratio of the specific growth rates of variables x and y (Huxley 1932). This approach rests on the observation that the size of an organism, and not its rate of growth, is important when one determines the proportions of its parts (Dodson 1975a). If this equation is converted to logarithms, the problem is reduced to the fitting of a straight line: log y = loga + Blog x. This may be rewritten as Y=a-+ BX where Y = log y, X = log x and « = log a. The allometric coefficient, f, is the slope of the ‘best straight line’ through the data, and may be considered as the value of Y when X = 0. Thus, f is the ratio of the specific growth rates of Y and X;; it serves as an indication of the intensity of differential size increase. Values of 8 greater than | (positive allometry) imply a differential increase of Y relative to X; when f is less than 1 (negative allometry) the Y/X ratio decreases with an increase of the absolute magnitude of X. Isometry, when 8B = 1, repre- sents the maintenance of geometrical similarity with size increase. In determining the slope of the ‘best straight line’ through the data, Kermack & Haldane (1950) and Kermack (1954) have cautioned against the use of regression models which assume error to be related to only a single variable. Consequently, the fitting procedure utilized here was that of Bartlett’s (1949) ‘best fit’. This method was found to be the preferred procedure in a computer simulation of ten different methods by Kidwell & Chase (1967), because it (i) is highly accurate, (ii) has a small variance, (iii) has a simple procedure for setting confidence limits on the estimates, and (iv) includes a ONTOGENY AND SEXUAL DIMORPHISM IN AULACEPHALO DON 165 simple linearity test. Bartlett’s (1949) method has been recommended for analysis of allometric growth by Simpson et al. (1960) and has been so used by Dodson (1976) and Grine et al. (1978). A description of Bartlett’s method has been provided by Grine et al. (1978) and will not be repeated here. Two diameters, basal cranial length (variable 1) and basal snout length (variable 3) were utilized as comparative (x) measurements in the present study. Basal snout length was found to have a nearly isometric relationship to basal cranial length, and use of the former permitted specimens, for which the latter length was not obtainable, to be included in the analysis of sexual dimorphism. In addition to a bivariate quantitative assessment, several qualitative features were examined in an attempt to define sexual dimorphism in Aula- cephalodon crania. CRANIAL GROWTH AND VARIABILITY The basal cranial length of those fossils in the present series for which this parameter could be recorded ranged from 135 mm (BPI.FN. 904) to 410 mm (BPI.FN. 806). This range, if basal cranial length is accepted as an indicator of the general size of the cranium, can be accommodated comfortably within the ontogenetic size range for skulls of modern large reptiles (e.g. Alligator and Crocodylus). And, as mentioned previously, specimens of Aulacephalodon have been recovered from localities within a rather limited horizontal and vertical range. Aulacephalodon appears therefore to satisfy the requirements for a possible growth series as established by Olson & Miller (1951). Coefficients of allometry and other relevant data for bivariate plots against basal cranial length are presented in Table 3. Selected bivariate plots of cranial measurements against basal cranial length are shown in Figure 2. In no instance could linearity be rejected, by analysis of the t statistic, in favour of a possible parabolic (quadratic) relationship (Table 3). Correlation between variable sets is rather high; most correlation coefficients are greater than 0,95 (Table 3). The high correlation coefficients and the degree of clustering of the points about the slope lines suggest that the specimens studied here represent a morphologically homogeneous group of animals in various stages of ontogenetic development. This series includes the types of Aulacephalodon haughtoni, A. latissimus, A. luckhoffi, A. pricei, and A. vanderhorsti. In several cases the allometric coefficients indicate isometric or nearly isometric growth relative to basal cranial length (e.g. variables 2, 3, 6, 8, 16, 21, 26), but in each instance the confidence intervals for 8 range from well below to well above isometry. Dodson (1975a) has. noted that for Alligator missis- sipiensis coefficients as close to isometry as 0,98 or 1,02 can be shown to differ from 1,00 at p = 0,02 or even p = 0,001, an indication of the high degree of correlation between variables, which is conditioned, in part, by the magnitude of the ontogenetic size range of specimens of Alligator. In three instances—palatal length (4), greatest cranial width (19), and the greatest width between the nasal bosses (25) (Fig. 1)—positive allometric 166 ANNALS OF THE SOUTH AFRICAN MUSEUM Log 4 3,80 Log |4 3,26 3,10 = 490 5,06 522 536 5,54 570 586 6,02 490 5,06 522 5,38 554 5/70 5,86 6,02 3,9 : 490 5,06 522 538 554 5,70 586 602 4,90 5,06 522-5,38 5,54 5,70 5,86 6,02 ONTOGENY AND SEXUAL DIMORPHISM IN AULACEPHALO DON 167 Log 25 4,20 3,80 3,40 3,00 490 5,06 5,22 536 5,54 5/70 586 6,02 Log 28 3,00 4,90 5,06 5,22 538 554 5,70 5,86 602 Log 31 3,90 4,90 5,06 5,22 5,38 5,54 570 586 6,02 490 5,06 5,22 538 554 570 586 602 Fig. 2. A-L. Selected bivariate plots of Aulacephalodon crania. The scale on both axes is logarithmic. In each case the X-axis is basal skull length (variable 1). The solid line represents the calculated line of Bartlett’s “best fit’. 168 ANNALS OF THE SOUTH AFRICAN MUSEUM TABLE 3 Summary of data concerning relative growth and variability in Aulacephalodon (X = basal cranial length). y Nhe a B-Cl a-Cl r rede t RL 2. 13 0,99 - 0,46 . 0,875 1,11 0/50; =043 099) io) 0713 eee 3 13. 0,99 —0,84 0,79; 1,19 —0,90; —0,78 0,97 10 0,444 No 410 4,15 =1569' 1,045. 1,26> 1,725! 165 91.00), aca 5 10 0,82 0,52 0,63: 1,23. 0,623) 0,42" 094) 7) Gage 6 12. 1,00 0,18 0,85; 1,14 0,13; 0,22 10,9959 Ogepene 7 ie 091 0,19 0:63;, 1,16. 0,11; 0,28 0,95 9, "=o 554nuenm 8 12 1,01 —O78 0,59: 1,29 —0,89; =067 095 9 ~l o.0ggummnm 9 12 1,10 —2,64 0,78: 1,44 —2,74 2053 005 9) Odeo 10 6 5-°1,20 ~=1,49 3 11,66) 1,80; 117 90198 9 92) oo ee 11, 13. 1,17 2,13. 0,86; 1,51. 2,23; 2/03), (0:95 910; —0lgnp aan 12 41 1,12, —1,35 0,87: 1,53 —1,43; =1,26-. 096 6 ose 13 12 0,94 0,67 0:59; 1,16° —0,76; —0)59' 0,96 9° = 1197 14.13 087 —1)67 <0,54; 117 .—1,76; "= 157") O91 10) nie 15 10- 0,81. —0,80. 0,60; 1,01 .—0)87; 9-078" 0977.07 =Onaee 16 12 1,05 —295 0,643 1,37 - —3,07; 283, 0.94 9 =0aumuee 17° 12 091° —0,05 0,43: 1,39 —O11; “00 086 9) ons eee 18 11 1,145 —0,89 091: 145 —0l98: ogi, 0196 3) = 2c neem 19 11° . 1,21.» 1,01 1,04: 1,33. =1,06; —096 (099 8” (NO asauem 20 10 0,89 1,33. 03227413 .°=1,42; 124 70:96 S78 eso nee 21 10 0198 | S09 0.71. 1.03 0,97; —0,84 098 7 —0558 NE 22°13. 126° ' 2,60 0,80; 1.90 - °—2,77;°=—2,43 086 10 9 20, gueaee 23. 13. 0,84 . 0,49 0,58; 1,04. . 0,42; 0:56..0:96 40) s=i:705 ene %” 12 1,31 °—294 0,95; 1,64 —305; =—293 006 "9, —S0sRunemem 25° 13 1,33 22,73 1,02: 1,57 ° 2182" 2165 0.97" 108 no coo 26 13 1,01 —1,70 0,56; 1,34 1/82; —1,59" (0935 10" | “Onesmeamm 27.12. 0,87. —0,69 0,64; 1,10 _-—0,76; —0,61 096 9 —1,488.0eNe 2% 12 1,39 3,08 O81:°1,76 —3.20; —296 095 9) —enqugz ame 29 13° 059 0:80 - O01 1,07 0,63; 0,96 0,72 10 0,363 No 30 11 0,86 0,76 0,312 1,21 —091: 062% 090, 8 Soars ame 31.13 1l1_ = 2,33 091; 1.27 22395 2:98 0.08) Oe SOc enn Y = variable number; N = number of specimens; a—-CI = 95% confidence interval for a; B-CI = 95% confidence interval for 8B; r = correlation coefficient; d.f. = degrees of freedom; t = Bartlett’s test statistic; RL = rejection of linearity in favour of a parabolic relationship. growth was indicated. On the other hand, whilst some eleven variables showed values of B less than 1,00, in all of these cases the 95 per cent confidence intervals ranged above isometry; variable 29, intertemporal width, showed the lowest value (0,59) but even here the upper end of the confidence limit ranged above isometry to 1,07. This parameter shows a particularly high degree of variability in terms of the magnitude of its allometric confidence limits and its low corre- lation coefficient (Table 3). It is interesting to note that the diameters of both the tusks (variable 14) and the caniniform processes (variable 15) show negative allometric growth (Table 3), although the upper limits of the confidence intervals for each range to above isometry. It appears that the eruption, or replacement of the tusks, by which increasingly larger teeth were introduced, did not keep pace with cranial enlargement. Also, the size of the orbits, as judged by their length (variable 27), enlarged in a negative allometric manner compared to basal | | | | ONTOGENY AND SEXUAL DIMORPHISM IN AULACEPHALODON 169 cranial length (Table 3). This is generally characteristic of higher vertebrates, where the eyes are disproportionately large in young individuals. Gould (1968) has noted that increase in size subjects organisms to different orders of forces and, thus, new morphological configurations to cope with these new forces are required. In his discussion of the allomorphism of species, Hennig (1966) maintained that knowledge of allometric correlation is important, because it permits recognition of linkages between different series of trans- formations that might otherwise not be recognized. New shapes produced by continuation of an ontogenetic allometric relationship into new size ranges are not independent taxonomic criteria (Gould 1966). For the most part, the features which have been used variously in the diagnosis of new aulacephalodont species have been (i) the size of the nasal bosses, (ii) the shapes and sutural relationships of the individual bones of the cranium and (iii) the length—breadth ratio of the cranium. Although several Aulacephalodon specimens have been described as possible ‘juveniles’ (Haughton 1917; Broom 1921), no description of a new species has taken into consideration the phenomenon of allometry. Cluver (1971), in a detailed study of another anomodont, Lystrosaurus, has recorded that minor changes in skull morphology, such as a change in the sutural relationships of bones, may be attributed to the size and consequently the ontogenetic age of individual specimens. Keyser (1972) has postulated that the characters which have been used to distinguish the various species of Aulacephalodon depend greatly upon the size of the individual. It is evident that in Aulacephalodon rather marked changes in the shape of the cranium and its various parts occurred with increasing size. In general, the relative width of the skull across the zygomatic arches and across the pre- frontal and nasal bosses, as well as the robusticity and sculpture of the temporal arches and other parts, show a rather dramatic increase with ontogenetic age (Fig. 3). The results of the present investigation indicate that at least five of the type specimens of Aulacephalodon species could be interpreted as constituting a homogeneous ontogenetic growth series. Two other types—A. hartzenbergi and A. brodei—have been examined also, and whilst these specimens were not subjected to allometric analysis, it is evident that they, too, form part of the same growth series. Although not every type and available specimen has been studied, the results of this preliminary investigation appear to support the con- tention that perhaps all the specimens, which have been regarded previously as belonging to different Aulacephalodon species, constitute a taxonomically homogeneous ontogenetic growth series of but a single species, Aulacephalodon baini (Owen). SEXUAL DIMORPHISM Sexual differences of both living and fossil reptiles are generally poorly expressed in the skeleton, and, as such, they are difficult to identify (Olson ANNALS OF THE SOUTH AFRICAN MUSEUM 170 171 "WD Ul SoTeOg “WU Op U)SUT [erUeIO [eseq :(1a014d “py Jo adAjoOjoY) 908 “NA'Idd ‘¢ ‘Ou “UoUTOeds OBIE] AIOA “D “WUT 76] YISUOT [erueIo [eseq ‘OO “NAId dl “9 “Ou uouNoedg “gq “WIUI get YISUD] [eIURIO [eseq ‘1071 "NA'Id@ “py ‘ou ‘uoumIoeds [[eUIg “YW “eIUeID wopojnYydaoDjnp JO solIas d1joUed0jUO poysossng ‘¢ “31.7 ONTOGENY AND SEXUAL DIMORPHISM IN AULACEPHALO DON 172 ANNALS OF THE SOUTH AFRICAN MUSEUM 1969). Dodson (1976) has used successfully two complementary approaches to define quantitatively sexual dimorphism in skulls of a small ceratopsian dinosaur. His first approach made use of bivariate allometric plots, a use of ratios, whilst the second was based on multivariate statistical analysis. In the present study an attempt was made to assess quantitatively sexual dimorphism in a growth series of Aulacephalodon crania. Dodson’s (1976) bivariate technique was utilized but application of the multivariate approach was not attempted owing to the small sample size of relatively complete crania. Bivariate plots of all thirty-one parameters were examined. For each case, individual values that lay above the line of Bartlett’s “best fit’ were assigned a score of —1, those below the line a score of +1, and values on the line were accorded a score of 0. Scores of 0 were very infrequent and were treated as indeterminate; thus they were not included in the final total of a score. The lower limit for the acceptance of either ‘maleness’ or ‘femaleness’ was set arbitrarily at 75 per cent, that is, at least three-quarters of the plotted points for an individual specimen lay to one side of the line. Only those specimens for which at least fifteen of the thirty possible variable plots were recorded were included in this part of the study (Table 4). Those fossils which fell above the line for 75 per cent of characters were considered to be male, whilst those that lay below the line for 75 per cent of traits were deemed to be female (see Dodson 1976). Bivariate analysis of sexual dimorphism revealed that no single specimen exhibited a consistently male or female pattern. Two specimens, however, were consistent in expression in at least 74 per cent of the traits (specimens 2, SAM— 8747, and 3, SAM-K1221), whilst a third (specimen 12, BPI.FN. 634) was consistent in 72 per cent of the characters. Accordingly, two of the specimens (2 and 12) may be accorded male status, whilst specimen 3 could be considered to be a female. Two of the fossils were completely indeterminate (7 and 8) and the others were only vaguely consistent in expression. In this analysis all available characters were considered; no selection of sexually distinctive features of the cranium was made. Thus, the technique utilized here differed from Dodson’s (1976) in that he selected characters which were believed to be dimorphic for Protoceratops. It is possible that, in the bivariate analysis of sexual dimorphism in Aulacephalodon, those features which are related to sexual dimorphism could have been ‘swamped’ by a larger number of variables which are not dimorphic. Furthermore, as not all the specimens possessed all possible measurable features, it is possible that those traits which may be sexually related were not present in some of the fossils. The principal cranial feature of Aulacephalodon which has been postulated to show sexual dimorphism is the relative size of the nasal bosses. Broom (1937) noted considerable variation in the size of the nasal bosses. He considered that the larger bosses were shown by male skulls, whilst females evinced more weakly developed bosses. He compared briefly two skulls which he considered represented a male and a female, and concluded also that the ‘canine’ tusks in hE a IS PY TS es ONTOGENY AND SEXUAL DIMORPHISM IN AULACEPHALODON Wie) TABLE 4 Bivariate sexual dimorphism scores for Aulacephalodon crania. Specimen Variable no. ae i | +O ba) Plesk ee [ee Ho Se eae te ld es |S +] [4+ ]+4/& | +++0 | | ae | lbsese ae a Wes ae | eae ett | Labs Peet | apap (ae [LaF | | Woe sires eeatesant Mar | Ste [etely (eater late stag eater Nee \ ate tet eetle toto 7 | eer J+) ++ I] tt | eeet] + +) 4+o4+14+4+!]4+4+4+11 11 1400 Wate eel alfa elites peli | |; ++] +++] t+) ) +) +444) 4) 4+4+4+4+4+4 I te ee tott |] F4+4+t] [++] ) ++) 4+] 44+) 444+] 4+] 4a. ed | aesest se fk se ji tese arc | se pel se se hse eset at ++4++++++] + ++|lo++++| He ie ML +++] 1] 1] 31 Dominant sign = SE Se ies pa Ochnts ae ae Score 74 74 65 59 63 50 50 59 72 63 68 58 Score = percentage frequency of occurrence of the dominant sign. -++ = individual plot below the line of Bartlett’s ‘best fit’; — = individual plot above the line of Bartlett’s ‘best fit’; 0 = individual plot on the line. See text for explanation. ++ males are longer and further apart than in females. In his description of A. cadlei he noted the nasal bosses of the type to be smaller than those in the type of A. rubidgei, and he postulated that the former specimen was a female whilst the latter was a male (Broom 1948). The question of sexual dimorphism in dicynodont crania has been considered by several workers (Owen 1860, 1876; Broom 1912, 1932; Barry 1957; Tripathi & Satangi 1963). Cluver (1971) dis- cussed the possibility that Lystrosaurus crania exhibited pronounced sexual dimorphism, but he concluded that the dimorphic skulls of that genus probably represent two groups of species. Keyser (1969) recorded that the nasal bosses of Aulacephalodon appear to increase in prominence concomitantly with an increase in skull size; he maintained also that the bosses were related to sexual dimorphism. 174 ANNALS OF THE SOUTH AFRICAN MUSEUM In light of the previous speculations which have related nasal boss size and shape variation to sexual dimorphism, these structures were examined both metrically and osteoscopically in the present series of crania. As an indication of the relative size of the nasal bosses the length of the protuberances (variable 31), as well as the greatest width between them (variable 25), were considered. When these two parameters are compared to basal cranial length (Table 3), both are found to increase in a positive allometric fashion, although the confidence interval for nasal boss length ranges to below isometry. When these variables are considered against basal snout length (variable 3), the positive allometric nature of their growth rates is slightly more pronounced (Table 5) and the 95 per cent confidence limit for nasal boss length ranges as low as isometry only. The considerable degree of morphological variability which is evinced by the nasal bosses may be related to their rates of growth. Dodson (1975a, 1975b) found that in two living reptiles (Alligator and TABLE 5 Summary of data concerning relative growth and variability in Aulacephalodon (X = basal snout length). Ve INI B a B-CI a-CI rt df t RL 1S) aor 0,85 0,84; 1,27 0,79; 0,91 0,97 10 —0,444 No 2 13 ~~ 1,00 0,38 0,73; 1,41 0,25; 0,48 093 10 #—0,550 No 4 10) tie) >) OF 1,09; 1,26 —0,74; —0,68 1,00 7 —0,912 No 5 10 0,83 0,18 0,60; 1,35 0,06; 0,30 0,93 7 -—0,599 No 6 13° 0:93 1,39 0,73; 1,16 1,32; 1,46 0,97 10 —0,548 No Wy (83 0,93 0,67; 1,28 0,83; 1,03 0,93 10 0,480 No 8 14 1,01 0,07 0,69; 1,34 —0,04; 0,18 0,93 11 —0,113 No Oe Sila lil —1,71 0,69; 1,77 —1,87; —1,54 0,88 9 —0,140 No 10 6 0,85 1,07 0,70; 2,18 0,61; 1,52 0,83 3 —0,747 No 11 15 1,13 —O,88 0,84; 1,51 —0,99; —0,77 0,92 12 -—1,418 No 12) 12%) 1307) 0313) 10} 8i 47, —0,22; —0,05 0,93 9 -—2,404 Yes 13 13 0,99 -—0O,07 0,75; 1,22 -—0,14; 0,01 0,97 10 0,353 No 14 15 0,88 —0,95 0,63; 1,16 —1,03; —0,86 0,92 12 1,186 No 15 10 0,80 -—0,03 0,56; 1,09 —0,12; 0,06 0,95 7 —0,352 No 16S ssc Ol —1,88 0,70; 1,31 —1,98; —1,78 0,95 10 —0,220 No 17 13 0,86 1,09 0,43; 1,26 0,95; 1,22 0,88 10 —0,264 No 182 1.08 0,38 0,85; 1,42 0,29; 0,48 0,96 9 1,344 No LO aD ile 0,46 0,85; 1,42 0,36; 0,55 0,97 9 —0,082 No 20 10 0,92 -—0,67 0,35; 1,18 —0,77;- —0,58 0,95 7 —1,849 No Zl eelil 1,03 —O,32 0,86; 1,41 —0,39; —0,25 0,96 8) 2 1425 NG Py M8) PAT Ile) OLT2 DAU 2s — 134 0583)" 10) O6Siaeeeine 2B) als 0182: 1,32 0,60; 1,05 1,25; . 1,40 0,94. 12 . =1,572 No 24 14 1,24 \—1,45 0,94; 1,57 S1s63 iss) OS — i 0,910 No 25) ale —2,44 1,02; 2,13 —2,63; —2,26 0,89 12 -—0,705 No 26 15. 0,97 —O0)58 0,59; 1,34 0,70; —0,45 0,89 12 -—0,114 No 27 14 0,89 —0,03 0,75: 1,06 —O0,08; 0,03 0,97 11 —1,449 No 28 13 1,20 —0,98 0,44; 1,69 —1,13; —0,82 0,90 10 —0,076 No 29 IS O68 0,86 0,25; 1,06 0,73; 1,00 0,80 12 —0,078 No 30 13 1,12 —1,36 0,75; 1,60 —1,46; —1,16 0,90 10 0,651 No SE Si LS S25 00S 77 —1,69; —1,56 0,98 12 -—1,507 No Y = variable number; N = number of specimens; «-CI = 95% confidence interval for «; B-CI = 95% confidence interval for 8; r = correlation coefficient; d.f. = degrees of free- dom; t = Bartlett’s test statistic; RL = rejection of linearity in favour of a parabolic relationship. | ) ONTOGENY AND SEXUAL DIMORPHISM IN AULACEPHALO DON 17/5) Sceloporus) the greatest variability is associated with the most strongly allo- metric variables. In Aulacephalodon the greatest width between the bosses (which includes the breadth of both these structures as well as the rest of the nasal bones) shows the strongest allometric growth compared to basal snout length (Table 5). The allometric nature of these protuberances is not quite so marked when basal skull length is used for comparison, but in the latter case (Table 3) fewer specimens were included. The width across the nasal bosses shows both stronger allometric growth and more variability than the length of these structures when the two are compared to either basal cranial (Table 3) or basal snout (Table 5) lengths. Three rather distinct types of nasal boss morphology, as assessed quali- tatively, were discerned in the present cranial series (Figs 4-5). In crania with ‘Type 1’ boss development (Figs 4A, 5A), the lateral margins of the nasal bones above the external nasal apertures show very little, if any, form of swelling; when viewed from above, the dorsal surface of the snout tapers anteriorly from the orbits. Nasal bosses of the second type (Figs 4B, 5B) project laterally to various degrees from the edges of the nasals above the external nares. The bosses project laterally and form a nearly flat (horizontal) surface with the nasal bones. Bosses of “Type 3’ (Figs 4C, 5C) also project laterally above the external nasal apertures, but are more swollen in appearance than those of the second type; they project dorsally as well as laterally. The third type of nasal pro- tuberance projects above the dorsal surface of the nasal bones, and in several specimens a longitudinal ‘gutter’ is present between the midline of the nasal bones and the boss on each side. The first type of nasal boss was found in small crania, whilst the second and third types appear in larger skulls. Nasal bosses of ‘Type 1’ are very slightly developed (Fig. 6). The second and third types of nasal bosses appear to increase in size with cranial length, but the two types can be distinguished in nearly all cases throughout a broadly overlapping range of individual size (Fig. 6). In the present sample, skulls which show “Type 3’ nasal bosses achieve a larger size than those with bosses of “Type 2’ morphology. This apparent size difference, however, may be an artefact of sampling. Several specimens (e.g. 19, 25-29) show a nasal boss configuration that appears to be intermediate between the second and third types. Although the morphological configuration of the nasal bosses may be altered through burial deformation (e.g. a ‘Type 3’ could be changed to a “Type 2’ through vertical diagenetic pressures), in the vast majority of cases these structures seem to have suffered very little from such deformation. In one specimen (15, BPI.FN. 2983) the nasal bosses had been pushed ventrally so that they form a nearly horizontal transverse plane with the nasal bones, but the bilateral shallow longitudinal troughs are still discernible owing to the presence of the median sagittal nasal ridge. Inasmuch as the present sample of Aulacephalodon crania exhibits a rather high degree of biometric homogeneity, it seems reasonable to suggest that the morphological differences shown by the nasal bosses may be sexual in nature. 176 ANNALS OF THE SOUTH AFRICAN MUSEUM = A showing the three basic types. A. ‘Type 1’. B. ‘Type 2’. C. ‘Type 3’. See text for explanation. i 1 1 1 Fig. 4. Schematic representation of a coronal section through the nasal bosses 1 t t 1 Various cranial sizes, or inferred ontogenetic stages, are present for both the second and third types of nasal bosses, whilst the smallest, and presumably youngest, skulls in the present sample evince a ‘Type 1’ nasal boss. It is possible that the nasal bosses in Aulacephalodon could have served as a sexually selective feature for mate recognition and/or agonistic display mechanisms, or for both. Furthermore, the several relatively large crania (specimens 19, 25-26 and 29) which show nasal bosses intermediate in form between “Types 2 and 3’ may represent the antimodal overlap of the bimodal distribution of a discontinuous feature such as sexual dimorphism. However, a number of the crania in which the nasal bosses are indeterminate are broken and poorly preserved. It is possible that those crania with the more robust, “Type 3’ nasal bosses represent the male condition. The same has been postulated obliquely by pre- vious workers (Broom 1937, 1948; Keyser 1969). In the present series, it appears that, once a certain cranial size had been reached, the nasal bosses assumed one of two rather distinct configurations (Fig. 7). It is suggested that the smaller crania, in which the nasal bosses are only slightly expressed (‘Type 1’), represent relatively young, sexually immature individuals. Specimen 6 (BPI.FN. 300) possesses rather small nasal bosses of the ‘Type 2’ (female) configuration (Fig. 8). This fossil has been identified tentatively as female (Figs 6-7). It is possible that with growth the nasal bosses of both sexes were of a “Type 2’ configuration before sexual maturity (or dimorphism) had been achieved; the smallest positively identifiable male cranium is somewhat larger than the specimen in question. If this sample is at all representative of Aulacephalodon, then it seems that the expression of ONTOGENY AND SEXUAL DIMORPHISM IN AULACEPHALO DON 177 Fig. 5. Stereophotos of the three basic nasal boss configurations. A. ‘Type 1’, specimen no. 7, BPI.FN. 904. B. ‘Type 2’, specimen no. 18, BPI.FN. 304. C. ‘Type 3’, specimen no. 17, BPI.FN. 493. See text for explanation. Scales in cm. 178 ANNALS OF THE SOUTH AFRICAN MUSEUM Type Size very large 3 large § 3 medium S very large g large Ss 2 8 medium small 40 60 80 100 120 140 I60 180 200 220 Basal Snout Length (mm) Fig. 6. Relationship of nasal boss size and morphotype to cranial (snout length) size. sexual dimorphism was achieved by individuals when snout length was between about 80 and 95 mm and basal cranial length approximately 190 to 245 mm. The nasal bosses enlarged in a positive allometric manner compared to snout and cranial lengths, but it appears that the general morphology of these structures remained relatively constant with an increase in individual size. Another feature of the cranium also appears to be related to sexual dimorphism. In large presumptive male crania (e.g. specimens 5, 13, 15 and 17), the lateral border of the zygomatic arch shows a thickening of the squamosal. This body thickening is in the form of a gently curved, ventrally directed convexity, the lateral margin of which shows a rugose texture (Fig. 9). Large female crania, on the other hand, show no such thickening of the zygomatic arch; in these specimens the lateral margin of the squamosal is thinner and the arch has parallel dorsal and ventral surfaces. In smaller male crania the zygo- matic arch is similar to those of females in that there is no sign of squamosai thickening. The significance of this thickening of the anterior portion of the zygomatic arch in larger male crania is enigmatic. Keyser (1969) noted that these squamosal ‘bosses’ may have been related to the jaw adductor muscu- lature; but if this were the case it is puzzling why only the older males developed these structures. Of the three crania sexed by bivariate analysis, the metrical diagnosis of two of them (specimens 3 and 12) agreed with the non-metrical assignment; however, whilst the third skull (specimen 2) was male according to the bivariate sexing technique, it shows a ‘Type 2’ nasal boss configuration, and although it is a fairly large specimen the zygomatic arches are not thickened. I ONTOGENY AND SEXUAL DIMORPHISM IN AULACEPHALO DON 179 Male 3 (sa A AaA A A Immature 4 ~-" a a (en o> 0 a Nasal Boss Type 40 60 80 100 |20 |40 I60 180 200 220 Basal Snout Length (mm) Fig. 7. Graphic representation of nasal boss morphotype compared to cranial (snout length) size. The questionable individual is specimen no. 6, BPI.FN. 300 (see Figs 3B, 8). Fig. 8. Stereoview of nasal boss size and morphology of specimen no. 6, BPI.FN. 300. Scale in cm. DISTRIBUTION An important factor to be considered in an analysis of the biological implications of the Type 1, 2 and 3 nasal boss configurations of Aulacephalodon is the geographic and temporal distribution of these various types. The geo- graphical distribution of the localities which have yielded the crania examined in this study is shown in Figure 10. Although the Madumabisa Mudstone ‘Formation’ is situated a considerable distance from the concentration of lower Beaufort localities in the Cape Province, the Aulacephalodon specimens found at the Zambian sites do not appear to be morphologically distinct from those recovered from the Cistecephalus and lower Daptocephalus zone sediments of South Africa (Drysdall & Kitching 1963). The specimens examined in this study were recovered from localities over a rather limited geographical (Fig. 10) as well as vertical (Keyser 1969; Kitching 180 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 9. Lateral view of a large Aulacephalodon skull showing the thickening of the squamosal at the anterior end of the zygomatic arch. Specimen no. 17, BPI.FN. 493. Scale in cm. 1977) range. There is no horizontal separation by either size or nasal boss } morphology of the specimens; furthermore, two specimens (4 and 17) from the ! same site (Fig. 1OR, Table 6) show considerable difference in both size and ¢ nasal boss configuration. One (BPI.FN. 1207) is a small individual with only | minimal nasal boss development, whilst the other specimen (BPI.FN. 493) | represents the largest skull.in the sample and it posseses well-developed “Type | 3’ nasal protuberances. Specimens which show variously Type 2 and 3 nasal , bosses have been recovered from at least one other locality (Fig. 10I, Table 6). The facts that specimens of various sizes with different nasal boss con- | figurations have been recovered from single localities, and that there is no | geographic or stratigraphic separation of specimens evincing different sizesand | shapes, appear to support the hypothesis that the Aulacephalodon specimens |} studied here do, indeed, represent an ontogenetic growth series of a single , species which evinced sexual dimorphism. DISCUSSION During the deposition of the Cistecephalus Zone sediments the terrain appears to have been characterized by broad, low gradient floodplains traversed by numerous, intermittent streams (Keyser 1970). The climate throughout © this time, as inferred from the abundance of calcareous concretions and of | ‘desert roses’, composed of pseudomorphs of calcite after gypsum (Keyser 1966), may have been sufficiently dry to warrant the term arid. The flora of the lower Beaufort seems to lack variety, the most commonly occurring plants being the equisetaleans, Schizoneura and Phyllotheca (Keyser 1970). Both ~ Keyser (1969, 1970) and Kitching (1977) have commented on the improbability q ONTOGENY AND SEXUAL DIMORPHISM IN AULACEPHALODON 181 RICHMOND VICTORA® Db) @ EST IDDELBURG 500km * @ -RASERBURG RY GRAAFF-REINET e f@ Fig. 10. Distribution of the localities from which the specimens examined in this study were recovered. A. Murraysburg commonage. B. Swaelkrans. C. Leeufontein. H. Ringsfontein. I. Graaff-Reinet commonage. J. Katbosch. K. Vleiplaats. L. Bultfontein. M. Leeuriviers- berg. N. Houd Constant. O. Roodebloem. P. Matjiesfontein. Q. Petersburg. R. Hoekplaas. S. Ferndale. of the synchronous and apparently sympatric coexistence of a large number of anomodont species under the ecological conditions envisaged for Cistecephalus Zone times. Keyser has noted the possibility that only a single species of Aulacephalodon, a \arge dicynodont, is represented in these strata. Biometric (allometric) analysis of a fairly large sample of Aulacephalodon crania, which included seven of the seventeen type specimens, indicates that at least this group of crania constitutes a homogeneous and probably species- specific, ontogenetic growth series. The ecological roles of at least two large extant reptiles, Alligator and Crocodylus, have been observed to change continuously throughout the life of the animal (Cott 1961; Dodson 1975a), the size and type of food obtained being a function of the size of the individual. Analogous food changes probably occurred with the Triassic cynodont Diademodon (Grine et al. 1978). Similar changes in the ecological role of Aulacephalodon baini might have taken place as a function of ontogenetically increasing body size, and thus increased the breadth of the niche occupied by this reptile; but this is by no means certain. Keyser (1972) has pointed out that Aulacephalodon possessed a relatively short, broad snout, reinforced by anterior longitudinal palatal ridges, and he concluded that biting was restricted to the tips of the jaws. Aulacephalodon probably had a horny beak at the tip of the snout, and it is possible that the changes in the 182 ANNALS OF THE SOUTH AFRICAN MUSEUM TABLE 6 Tabulation of locality data. The code corresponds to the legend for Figure 10. Nasal boss Code Locality Specimen morphotype Sex A Murraysburg commonage 7 1 indet. B Swaelkrans 5 3 male C Leeurivierspoort 33 3 male D Modderfontein 9 2) female E Bloukop 8 2 female F Oudeplaas 13 3 male G Leeufontein 12 3 male H Ringsfontein 11 3 male 15 3 male I Graaff-Reinet commonage 10 3 male 18 D female 21 2 female J Katbosch 6 2 ? female K Vleiplaats 3 2 female L Bultfontein 14 2 female M Leeuriviersberg 32 3 male N Houd Constant 20 2 female 31 2 female oO Roodebloem 1 1 indet. P Matjiesfontein 16 2 female Q Petersburg 19 23 ? male R Hoeksplaas 4 1 indet. 17 3 male S Ferndale 2 2 female shape of the snout that occurred during growth may have been further accen- tuated by the beak. Although sexual differences of both extant and fossil reptiles are generally poorly expressed in the skeleton, Dodson (1976) has provided convincing evidence of sexual dimorphism in skulls of the late Cretaceous dinosaur, Protoceratops andrewsi. He showed sexual dimorphism to be an important source of variability in this ceratopsian, and found that it was reflected in many characters of the skull apart from the obvious ones. A preliminary and limited biometric (bivariate) analysis of Aulacephalodon skulls indicated that only three of the specimens in the sample could be assigned even possibly to a sex: two specimens were provisionally accorded male status whilst a third was regarded as possibly female. The nasal bosses of Aulacepha- lodon have been regarded by several workers as being related possibly to sexual dimorphism e.g. Broom 1937, 1948; Keyser 1969). A simple size and shape analysis of these structures throughout an ontogenetic growth series has revealed that they do, indeed, appear to be related to sexual differences (Figs 6-7). The smallest crania exhibit only slightly developed nasal bosses (‘Type 1’), whilst larger skulls evince either a somewhat flattened, laterally projecting boss (‘Type 2’) or a more bulbous, expanded structure (‘Type 3’). The sex of the smaller crania could not be determined although Haughton (1917) considered SAM-—3328 (specimen 1, described by Broom in 1921 as the type of Bainia ONTOGENY AND SEXUAL DIMORPHISM IN AULACEPHALODON 183 haughtoni) to be a ‘young male’. It was not possible in this study to refute or substantiate Haughton’s claim. Those crania which showed a “Type 2’ nasal boss are believed to represent females, whilst it is suggested that males evinced a more strongly developed (‘Type 3’) protuberance. Although the bivariate sexing technique used here proved to be generally inconclusive, as discussed above, three specimens showed relatively consistent biplot patterns (SAM-8747, BPI.FN. 634—males; SAM-K1221—female). The qualitative determination of sex, based solely upon nasal boss configuration, agreed with the biometric determination in two instances; but in the case of SAM-8747, a relatively large specimen, whilst the biparametric technique indicated ‘maleness’, it was deemed to be a female on the shape of its nasal bosses. It is suggested that males possessed relatively more strongly developed bosses than females, and that these structures might have served some sexual selective function. They might have served as important visual signs in a mate recognition system and/or as agonistic display mechanisms. Keyser (1969) has suggested that the bosses could have protected the eyes (from thorny vegetation or during fighting), supported horny growths, and possibly even served a thermoregulatory function. While the size of the nasal bosses is related in an allometrically positive fashion to the growth of the cranium (as represented by either basal snout or basal cranial lengths), the morphotypic expression of their general configuration appears to have been established at the attainment of sexual dimorphism and to have remained recognizably distinct throughout continued ontogenetic development. Thus, the male and female configurations are discernible over a rather large size range, whilst the smaller specimens show only minimal boss development. If nasal boss morphology is sexually related, as indeed it appears to be, and if the present sample is representative of Aulacephalodon, then it seems that sexual dimorphism was first expressed when the base of the cranium attained a length of between 190 and 245 mm. The locality data appear to support the hypothesis that the present sample represents a morphometrically homogeneous, ontogenetic growth series of a single species of Aulacephalodon and that with age this species exhibited sexual dimorphism of the nasal bosses and of the anterior portion of the temporal arch. The confirmation or refutation of these hypotheses must, however, await a more detailed biometrical and osteoscopic analysis of a larger series of specimens which includes all the type specimens of Aulacephalodon. SUMMARY AND CONCLUSIONS A biometrical (allometric) analysis of a number of Aulacephalodon crania was undertaken. The sample included representatives of various supposed species as well as the holotypes of a number of these. The results of this investi- gation indicate that this sample represents a morphometrically homogeneous, 184 ANNALS OF THE SOUTH AFRICAN MUSEUM species specific, ontogenetic growth series. The features utilized previously as * taxonomic criteria can be understood as one result of allometric growth. A qualitative analysis indicates that nasal boss morphology is related to sexual dimorphism. It is apparent that the smallest (ontogenetically youngest) individuals showed no or minimal nasal boss development, and that sexual maturity or at least the earliest expression of sexual dimorphism may have occurred in individuals with a basal cranial length of between 190 and 245 mm. The larger male crania show a thickening of the lateral margin of the temporal arches, and this is not present in small male or any female skulls. Large male crania show a thickening of the anterior portion of the temporal arches. The locality data for the specimens in this sample support the inference that this group of fossils represents a species—specific ontogenetic growth series, and that sexual dimorphism was expressed in the cranium of Aulace- phalodon baini. ACKNOWLEDGEMENTS The fossil material was made available to us through the generous offices of the Bernard Price Institute for Palaeontological Research, Johannesburg, the Geological Survey, Pretoria, the South African Museum, Cape Town, and the Transvaal Museum, Pretoria. Discussions with Drs M. A. Cluver, J. W. Kitching, and A. W. Keyser, who helped in the mapping of the locality data, were beneficial to this paper. We thank Dr M. A. Cluver, and Professors J. A. Hopson, M. Raath, and P. V. Tobias for reading and constructively criticizing this paper. We thank Miss J. Groom for drawing the text figures and Mr H. Thackwray for photography. This work was supported in part by a grant to F. E. Grine by the University of the Witwatersrand Senate Research Committee. REFERENCES Barry, T. H. 1957. On the significance of tuskless specimens of Dicynodon grimbeeki Broom. Palaeont. afr. 5: 57-66. BarTLeETT, M. S. 1949. Fitting a straight line when both variables are subject to error. Biometrika 5: 207-212. Broom, R. 1906. The classification of the Karroo beds of South Africa. Geol. Mag. (5)3: 36. Broom, R. 1912. On some new fossil reptiles from the Permian and Triassic beds of South Africa. Proc. zool. Soc. Lond. 1912: 859-876. Broom, R. 1913. On some new genera and species of dicynodont reptiles, with notes on a few others. Bull. Am. Mus. nat. Hist. 32: 441-457. Broom, R. 1921. On some new genera and species of anomodont reptiles from the Karroo beds of South Africa. Proc. zool. Soc. Lond. 1921: 647-674. Broom, R. 1928. On three new species of Dicynodon. Ann. S. Afr. Mus. 22: 421-425. Broom, R. 1932. The mammal-like reptiles of South Africa. London: Witherby. Broom, R. 1936. On some new genera and species of Karroo fossil reptiles, with notes on some others. Ann. Transy. Mus. 18: 349-386. Broom, R. 1937. A further contribution to our knowledge of the fossil reptiles of the Karroo. Proc. zool. Soc. Lond. 107: 299-318. Broom, R. 1940. Some new Karroo reptiles from the Graaff-Reinet district. Ann. Transv. Mus. 20: 71-87. | ONTOGENY AND SEXUAL DIMORPHISM IN AULACEPHALO DON 185 Broom, R. 1941. Some new Karroo reptiles with notes on a few others. Ann. Transy. Mus. 20: 193-213. Broom, R. 1948. A contribution to our knowledge of the vertebrates of the Karroo beds of South Africa. Trans. R. Soc. Edinb. 61: 577-629. Broom, R. & Georce, M. 1950. Some new anomodont reptiles in the Bernard Price collection. S. Afr. J. Sci. 46: 275-278. Ciuver, M. A. 1971. The cranial morphology of the dicynodont genus Lystrosaurus. Ann. S. Afr. Mus. 56: 155-274. Cort, H. B. 1961. Scientific results of an inquiry into the ecology and economic status of the Nile crocodile (Crocodilus niloticus) in Uganda and Northern Rhodesia. Trans. zool. Soc. Lond. 29: 211-356. Dopson, P. 1975a. Functional and ecological significance of relative growth in Alligator, J. Zool., Lond. 175: 315-355. Dopson, P. 1975b. Relative growth in two sympatric species of Sceloporus. Am. Midl. Natur. 94: 121-150. Dopson, P. 1976. Quantitative aspects of relative growth and sexual dimorphism in Proto- ceratops. J. Paleont. 50: 929-940. DRYSDALL, A. R. & KITCHING, J. W. 1963. A re-examination of the Karroo succession and fossil localities of part of the upper Luangwa Valley. Mem. Surv. Nth. Rhod. 1: 1-55. Gow p, S. J. 1966. Allometry and size in ontogeny and phylogeny. Biol. Rev. 41: 587-670. GouLp, S. J. 1968. Ontogeny and the explanation of form: an allometric analysis. J. Paleont. 42: 81-98. Grine, F. E., HAHN, B. D. & Gow, C. E. 1978. Aspects of relative growth and variability in Diademodon (Reptilia: Therapsida). S. Afr. J. Sci. 74: 50-58. HaucGuton, S. H. 1917. Investigations in South African fossil reptiles and Amphibia, Part 10. Descriptive catalogue of the Anomodontia, with especial reference to the examples in the South African Museum. Ann. S. Afr. Mus. 12: 127-174. Haucuton, S. H. & Brink, A. S. 1954. A bibliographical list of Reptilia from the Karroo beds of Africa. Palaeont. afr. 2: 1-187. HeEnnic, W. 1966. Phylogenetic systematics. Urbana: University of Illinois Press. Houxtey, J. S. 1932. Problems of relative growth. New York: Dial Press. KerMack, K. A. 1954. A biometrical study of Micraster coranyuinum and M. (Isomicraster) senonensis. Phil. Trans. R. Soc. (8) 237: 375-428. Kermack, K. A. & HALDANE, J. B. S. 1950. Organic correlation and allometry. Biometrika 37: 3041. Keyser, A. W. 1966. Some indications of arid climate during the deposition of the Beaufort series. Ann. geol. Surv. S. Afr. 5: 77-79. Keyser, A. W. 1969. A re-evaluation of the systematics and morphology of certain anomodont Therapsida. Unpublished Ph.D. Thesis, University of the Witwatersrand. Keyser, A. W. 1970. Some ecological aspects of the Cistecephalus zone of the Beaufort series of South Africa. In: I.U.G.S. 2nd Symposium Gondwana Stratigraphy and Palaeontology: 653-657. Cape Town and Johannesburg. Keyser, A. W. 1972. ’A re-evaluation of the systematics and morphology of certain anomodont Therapsida. Palaeont. afr. 14: 15-16. Kwwe Lt, J. K. & Case, H. B. 1967. Fitting the allometric equation—a comparison of ten methods by computer simulation. Growth 31: 165-179. Kircaine, J. W. 1970. A short review of the Beaufort zoning in South Africa. In: I.U.G.S. 2nd Symposium Gondwana Stratigraphy and Palaeontology: 309-312. Cape Town and Johannesburg. KircuinG, J. W. 1977. The distribution of the Karroo vertebrate fauna. Mem. Bernard Price Inst. palaeont. Res. 1: 1-131. Oxson, E. C. 1969. Sexual dimorphism in extinct amphibians and reptiles. In: WESTERMANN, G. ed. Sexual dimorphism in fossil metazoa and taxonomic implications. 1.U.G.S. series A (1): 223-225. Oxson, E. C. & Miter, R. L. 1951. Relative growth in paleontological studies. J. Paleont. 25: 212-223. Owen, R. 1844. Description of certain fossil crania, discovered by A. G. Bain, Esq., in sand- stone rocks at the south-eastern extremity of Africa, referable to different species of an 186 ANNALS OF THE SOUTH AFRICAN MUSEUM extinct genus of Reptilia (Dicynodon), and indicative of a new tribe or suborder of Sauria. Proc. geol. Soc. 4: 500-504 Owen, R. 1855. Description of certain fossil crania, discovered by A. G. Bain, Esq., in sand- stone rocks at the south-eastern extremity of Africa, referable to different species of an extinct genus of Reptilia (Dicynodon), and indicative of a new tribe or suborder of Sauria. Trans. geol. Soc. Lond. 7: 59-84. Owen, R. 1856. Report on the reptilian fossils of South Africa. Part II. Description of the skull of a large species of Dicynodon (D. tigriceps, Ow.), transmitted from South Africa by A. G. Bain, Esq. Trans. geol. Soc. Lond. 7: 233-240. Owen, R. 1860. On some reptilian fossils from South Africa. Q. JI. geol. Soc. Lond. 16: 49-54. Owen, R. 1876. Descriptive and illustrated catalogue of the fossil Reptilia of South Africa in the collection of the British Museum. London: British Museum (Natural History). SEELEY, H. G. 1898. On Oudenodon (Aulacocephalus) pithecops from the Dicynodon beds of East London, Cape Colony. Geol. Mag. 5: 107-110. Simpson, G. G., RoE, A. & LEwWonrtIN, R. C. 1960. Quantitative zoology. 2nd ed. New York: Harcourt, World and Brace. TRIPATHI, C. & SATSANGI, P. P. 1963. The Lystrosaurus fauna of the Panchet Series of Raniganj coalfield. Mem. geol. Surv. India Palaeont. indica 37: 1-49. VAN HOoePEN, E. C. N. 1934. Oor die indeling van die Dicynodontidae na aanleiding van nuwe vorme. Paleont. Navors. nas. Mus., Bloemfontein 2: 67-101. ' 6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature (particularly Articles 22 and 51). ; Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov., etc. , An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: “Smith (1969) describes...’ ‘Smith (1969: 36, fig. 16) describes . “As described (Smith 1969a, 19696; peas ay “As described (Haughton & Broom xa): ‘As described (Haughton ef al. 1927) . Note: no comma separating name acl year Dagination indicated by colon, not p. names of joint authors connected by ampersand ' et al. in text for more than two joint authors, but names of all authors given in ‘list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, 5, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o, scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BuLtouGu, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FISCHER, P. —H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FiscHer, P.-H., Dovat, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. ‘gen. 74: 627-634. Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Ann, Mag. nat. Hist. (13) 2: 309-320. Konn, A. J. 19606. Spawning behaviour, cee. masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. THEELE, J. 1910. Mollusca: B. Pabulccrteal Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-. Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 81 Band April 1980 April Part 5 Deel THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES PART 10 STATION DATA 1977, 1978, 1979 Compiled by ELIZABETH LOUW Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na gelang van die beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad OUT OF PRINT/UIT DRUK 1, 201-3, 5-8), 3(1-2, 4-5, 8, t.—p.i.), 5(1-3, 5, 7-9), 6(1, t.—p.i.), 7(1-4), 8, 9(1-2, 7), 10(1-3), 11(1-2, 5, 7, t.—p.i.), 15(4—5), 24(2), 27, 31(1-3), 32(5), 33 Copyright enquiries to the South African Museum Kopieregnayrae aan die Suid-Afrikaanse Museum ISBN 0 908407 90 4 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape ' Courtweg, Wynberg, Kaap THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES PART 10 STATION DATA 1977, 1978, 1979 Compiled by ELIZABETH Louw South African Museum, Cape Town (With 1 figure and 6 tables) LMS. accepted 6 December 1979] CONTENTS PAGE Introduction : F ; ; : : 187 The cruises . F : , ‘ : Be el ee/ Acknowledgements . : : 2 e205) References . herec 5 5 ; . 205 INTRODUCTION In 1975 the South African Museum’s Department of Marine Biology embarked on the first of a series of cruises in deeper waters (about 500 m and more) off the east coast of southern Africa. Station data for the 1975 and 1976 cruises have been published (Louw 1977). During the period 1977-9 three additional cruises were carried out in areas south of the region sampled in 1975 and 1976. Publications dealing with collections from the 1977, 1978 and 1979 cruises will in most instances give station numbers only, and refer to the present paper for further data. THE CRUISES Grants from the South African National Council for Oceanographic Research (SANCOR) and assistance from the South African Museum enabled the Department of Marine Biology to undertake the three cruises aboard the R.V. Meiring Naude. From 9 to 18 May 1977 work was carried out south of Durban, Natal, in the area 30°01’S to 31°30’S and 30°03’E to 32°02’E with bottom depths ranging from 690 m to 2 600 m. In 1978 the area from 32°40’S to 34°11’S and 28°50’E to 27°08’E was worked during the period 24 May to 1 June, in depths ranging from 80 m to 2 880 m. In 1979 permission was obtained to fish off the coast of the Repubtic of Transkei and from 18 to 28 June the area 30°46’S to 32°37’S and 30°39’E to 28°54’E was sampled in depths of 82 m to 2 820 m. In both 1978 and 1979 some stations that were worked were in relatively shallow water due to adverse weather and sea conditions which prevented 187 Ann. S. Afr. Mus. 81 (5) 1980: 187-205, 1 fig., 6 tables. | ‘ q | 188 ANNALS OF THE SOUTH AFRICAN MUSEUM | r 26° 29° 30° 3° ' \sv. DURBAN 52 E51 53 | ih Ly NZ 1476 | | SOUTH AFRICA we Ng y6 1 3° fag 959, *254 2008 * 53 243 207 25009249 0 $28 3F sue 2 246. Ile 0210 ais 022 Caer Be : 200 30235 237 2215 : 225 223 gv 22848226 220 231-8 °230 2)9- $0218 ©6224 227 © $221 223” 0999 2159 #159J-60 i 162, 161 4 E.LONDON « | 3 4 = 4169 3 o { oe 169 0. 466-7 | 171 168 i 0174 el715 | VEIT 7 "2 179 176 178, aja } 184-5 77 4182 ! | 773 j ; ep (83 D 187 j 188: 3 189 Y d #190 i 191 | Pr we 29° 30° 31° vig Fig. 1. Chart of the area sampled during the 1977 (stations SM 116-158), 1978 (stations i SM 159-191), and 1979 (stations SM 192-256) cruises of. the South African Museum on the R.V. Meiring Naude. ‘i THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 189 sampling further off shore. In all 141 stations were occupied during the three cruises and station data are presented in Tables 1, 3 and 5. Relevant temperature data are provided in Tables 2, 4 and 6. The gear used was basically the same as for the 1975 and 1976 cruises, but with the following changes: Use of the IOSN and IKMT was discontinued The small square-framed neuston net (Neustong) used in 1976 was replaced during the 1977 cruise with a larger oval-framed net with floats (Neuston,) In addition to the RMT with uniform anchovy mesh (RMT),), a similar net with stepped-mesh (RMT,) was used at some mid- water stations A double beam trawl (Menzies 1964: 103, fig. 14; Belyaev 1966: 18, fig. 17) replaced the 18 ft beam trawl on the 1979 cruise The material collected is housed in the South African Museum collections, with duplicate fish material in the J. L. B. Smith Institute of Ichthyology, Grahamstown, and duplicate decapod and isopod Crustacea in the U.S. National Museum of Natural History, Washington, D.C. Some animal groups are still available for study by interested biologists. These include Gasteropoda, Pelecypoda, Brachiopoda, Anthozoa, Mysidacea, Tanaidacea and Euphausiacea. 77 NOTES FOR TABLES S, D and N following SM numbers, e.g. SM 124S, SM 124D and SM 124N, indicate shallow Bongo, deep Bongo and Neuston net hauls respectively, at the same station. J following SM numbers, e.g. SM 159J, indicates surface fishing with jiggers or dip-nets at night lights after completion of the haul at the station of the same number. The beam trawl was lost at SM 137 (during the third cruise), at SM 165 (during the fourth cruise) and the double beam trawl was lost at SM 256 (at the end of the fifth cruise). Bathythermograph column in Tables 1, 3 and 5 indicates the number (BT-) of the bathythermograph profile to which one should refer for temperature data relevant to that station, and for which temperatures at selected depths are given in Tables 2, 4 and 6, respectively. In Table 1 ** SM 115 Neuston net was towed during the RMT haul * SM 126 An Expendable Bathythermograph (XBT) was used instead of a standard bathythermograph ~ SM 119 RMT, was on the bottom for part of the haul In Tables 4, 6 * Temperatures at Om were obtained from the readings on the Multipoint chart for sea surface temperature. In Table 6 ** There are no temperature readings for BT—50 and BT-—51 due to equipment failure. ee oemer 44° OU Sito Oser Oree 41S 62° OR Oe 1? Oto Ve b FE o 5: a — www ell we ew LL'S01 [T8I-So8t TOSOE S'TEOE Sb?I WS aspoip 0 Pl‘97 AACOH 069 069 LL'S Ot SSSI-ScSI LOSI /9°8V.0E VEE.0E €cl WS See aS hod ¢—La osuog CIT LL Ol 80rI-9reEl 0°SP.0E O9E.0€ AZZI WS 68°T L1‘97@ €-—Ld osu0g os LL’S ‘Ot OvVEI-EET /0°Sv.0E 0°9€.0€ Szcl WS [Mey 98°T £6°S7 uwuvoq ¢c9-006 ¢¢9-006 LL 01 OfIT OPOI-TE60 O80 /8°7S0E /CTEOE IZl WS IL‘T L8°Sz C-Ld osu0g CIT LL'S OT TS90-T£90 0°8S.0€ L'6T.0E G07 WS z €£0 06°SZ c-Ld osu0g 6S LLS Ot ¥c90-L190 /0°8S.0€ /L‘6TOE socl WS a 5 89'T 69ST I“ Ld WIWY OSL OSL LLS 6 CSIt EIT HOIT EV6r GO ELLE VVTOE 6Il WS a Z est 86°S7Z I-Ld osu0g CI? LLS'6 T@8I-8SLT /V800TE /681.0€ d8II WS S 6ET 00°97 Ld osu0g os LLS'6 CSLI-9PLI /V 8001 6 81.0€ S8IT WS 2 [Med] y8'T 00°97 uvoq 0¢8 078 LL 6 SCLI OS9T-OZ9T ESST 0 OTTE /SLToO€ LIT WS SB 9 aspeip 2 0 66 °SZ AAvoH 078 078 LL 6 OISt LvvI-LOvI Let VSTONE 0°ST.O€ SIT WS & = s}OUy ays wi wi Ao So a “dua “ON puq suysiy jaADIS. apnjigsuoT — apnjijvT s paads aavfans ydoss dpay yjdaq yidaq 210Q——_ ‘ON a S.diys pas -OWABY] SULYSL wojjog Out, uouisog uouvis ~AYIOg “LLOI ABW 81-6 “SINIO paryy OY} Joy wep MONIES | a1av.L 190 191 THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 09'T 8Z'SZ ZI-LE osuo0g TIZ LL €1 S€90-7190 /VLIOE OTIoTE G9ET IWS £9°T ZEST 7I-La osuog = 0S LL €1 8090-2090 LIE OO UTOTE S9ET NS €9'T €9°S7Z TI-La osuog = ZIZ LL ZI ISTI-62Z1 /CVTOE VEOTE ASEl WS 09°T 79°SZ lI-La@ osuog 0S LL ZI O@ZI-VIZI /CVTOOE EO TE SSET WS [Med} 0 Or‘SZ weog (006 006 LLS ZI OIII O£€0I-0€60 6580 /CLTOE 00001 vel WS 6r'T 6S5°SZ = TuojsnoN 86 LLS TI UIUL ¢ S970 COTE NEEL WS 6r' I 6S‘STZ OI-La osuog = ZIZ LLS TI 0’90-8190 /S‘9TOE /TIOTE AEE WS VET 1S‘*Sz OI-La osuog = 0S LL'S TI 7190-9090 /S‘9TOE /CIOTE SEETWS 69° 1197 6-Ld WLINU 0-0£8 OSLI LL I 6E2~ 6rIT-8P0Z 7007 /C CHOOE VSVo0E ZET NS [Mery 0 €0°9Z weog O8/ O8Z LL SILI ZP9I-OI9T SPST /8‘0V.0E iC EVE Tel WS 06'T v8°SZ = TuojsnaN =O) LLS TI UIW OT /80€00€ /CSS0E NOEI WS 06'T P8‘SZ 8-—Ld osuog = ZZ LL ST 60EI-8r7ZI /80€.00€ /7SS.0€ COET WS 88‘T 8L‘°SZ 8-—Ld osuog 0s LUST SYTI-6ETI /80€00€ (SSO SOET WS espoip 0 8L°SZ ACOH 0s8 0s8 LLS IT SIII-ZIOl 760 /LTEOE VES .0E 6cI WS Mey 0 08‘SZ weog O£6-0£8 Of6-0£8 LLS II 6260 6v80-EI80 OFLO ,L‘SE.0€ 1 6V00€ 8zI WS SLT 79°SZ = SuojsnaN =O LUSH “UI OT 9 Tvo0E Lvvo.0€ NLZ7I WS SLT Z9S7Z L-Ld osuog OST LLS TI Z£90-1 190 WOE ZLbyO0E ALZI WS orl 19°Sz L-14 osuog 0s LLS'T1 4090-1090 9 THO Lvvo0E SLZI WS €L‘€ S1‘9Z +9- LE WIN O-19r ~— O81 LL'S ‘01 SIIO OSEZ-OS7%Z LIZZ ,9°6S.0€ ,9°6£.0E 971 WS. 061 TABLE 1 Station data for the third cruise, 9-18 May 1977. Se ee ee eee fi eas 5 Bathy- Br Position Time Bottom Fishing Meni: Sea Ship’s aS D Latitude Longitude Start Fishing End “s Ze nee ee oN. : pice ee : : Ss °E m m 1c knots oO 14 ° Ld SM 116 30°15,0 31°15,4 1347 1407-1447 1516 9.5.77 820 820 Heavy 25,99 0 dredge : oO SM 117 30°17,5’ 31°10,0’ 9.5.77 1725 1553 1620-1650 trawl a a ee eee SM 118S 30°18,9’ 31°08,1’ 1746-1752 9.5.77 50 Bon teas cae 2. go BT-1 26,00 1,39 SM 118D 30°18,9 31°08,1 1758-1821 9.5.77 212 Bongo BT-1 25,98 1,83 SM 119 30°14,4’ 31°13,9’ 1943 2104-2134 2152 9.5.77 750 750 RMT BT-1 25,65 1,68 SM 120S 30°29,7’ 30°58,0’ 0617-0624 10.5.77 59 Bon cell Bel Laks FX0) BT-2 25,90 0,33 SM 120D 30°29,7 30°58,0 0631-0651 10.5.77 212 Bongo BT-2 25,87 1,71 SM 121 30°32,2’ 30°52,8’ 0830 0931-1040 1130 10.5.77 900-625 900-625 Beam 25,93 1,86 trawl SM 122S 30°36,0’ 30°45,0’ 1334-1340 10.5.77 50 Bon PS at 5. go BT-3 26,17 1,89 SM 122D 30°36,0 30°45,0 1346-1408 10.5.77 212 Bongo BT-3 26,15 1,78 SM 123 30°33,4’ 30°48,6’ 1507 1525-1555 10.5.77 690 690 Heavy 26,14 0 dredge ; SM 124S 30°32,5’ 30°59,2’ 1805-18 5. SM 124D 30°32,5’ 30°59,2’ 1817-1 par ieee es Hee eae Ee oe i 4 . ' rast 5M 124m 30°32,5’ _ 30°SO.27 12. min. 10.5.77 ad ° Ne rmdir: 26,24. La 2h a SM 125 30°32,2’ 30°S7,5” 1909 1929-2029 2050 10.5.77 1280 415-0 . £RMtTtm BY-s 2618 Ann ‘SM 126 30°39,6’ 30°59,67 2217 2250—2350 0115 10.5.77 1820 464-0 RMTm BT-6* 26,15 3,73 SM 127S 30°44,7’ 30°41,6’ 0601-0607 11.5.77 50 Bongo BTI-7 25,61 1,46 SM 127D 30°44,7’ 30°41,6’ 0611-0632 11.5.77 150 Bongo BT-7 25,62 1,78 SM 127N 30°44,7’ 30°41,6’ 10 min. 11.5.77 0 Neustons — 25,62 1,78 SM 128 30°49,1’ 30°35,7’ 0740 0813-0849 0929 11.5.77 830-930 830-930 Beam 25,80 0 trawl SM 129 30°53,4’ 30°31,7’ 0942 1012-1115 11.5.77 850 850 Heavy 25,78 0 dredge SM 130S 30°55,2’ 30°30,8’ 1239-1245 11.5.77 50 Bongo BT-8 25,78 1,88 SM 130D 30°55,2’ 30°30,8’ 1248-1309 11.5.77 212 Bongo BT-8 25,84 1,90 SM 130N 30°55,2’ 30°30,8’ 10 min 11.5.77 0 Neuston. — 25,84 1,90 SM 131 30°43,2’ 30°40,8’ 1545 1610-1642 1715 11.5.77 780 780 Beam 26,03 0 trawl SM 132 30°45,4’ 30°42,2’ 2002 2048-2149 2239 11.5.77 1750 830-0 RMTm BT-9 26,11 3,65 SM 133S 31°01,2’ 30°26,5’ 0606-0612 12.5.77 50 Bongo BT-10 25,51 1,34 SM 133D 31°01,2’ 30°26,5’ 0618-0640 12.5.77 212 Bongo BT-10 25,59 1,49 SM 133N 31°01,2’ 30°26,5’ 5 min 12.5.77 0 Neustony —_ 25,59 1,49 SM 134 31°00,0’ 30°27,2’ 0859 0930-1030 1110 12.5.77 900 900 Beam 25,40 0 trawl SM 1358S 31°03,4’ 30°24,2’ 1214-1220 12.5.77 50 Bongo BT-11 25,64 1,60 SM 135D 31°03,4’ 30°24,2’ 1229-1251 2ST 212 Bongo BT-11 25,63 1,63 a a ee erg a Se a Oe ee Ce Pe en SM 1368S 31°11,0’ 30°17,1’ 0602-0608 13.5.77 50 Bongo BT-12 25,32 1,63 SM 136D 31°11,0’ 30°17,1’ 0614-0635 13.5.77 212 Bongo BT-12 25,28 1,60 WngasnW NVOrddv HLNOS AHL AO STVNNV SASINUO FGAVN ONIVIAW SIWNASQW NVOIdAVY HLNOS AHL 161 ANNALS OF THE SOUTH AFRICAN MUSEUM 192 RS ETI 6st 00o% sI-ld $ cBuog os LU 91 vIZI-6071 EOE 98Z1E SOPI WS SO‘e ¥6°STZ LI-La WIN 0-6cT1 0007 LLS 91 PEO! S760-S780 SELO 0°P0.0€ POETE Svl WS 19‘T 00°92 LI-La osuo0g CIT LLS 91 $£90-F190 /S°9000€ N9ZTE bbl WS 9ET 00°97 LI-La@ osu0g os LL’S91 0190-7090 /S°90.0€ MOTTE Spr WS 19‘T 18°Sz 9I-La osuog CIZ LL'S’ST OrsI-9I8I LYLE LYTLE aetril WS Z9'T €8°S7Z SI-La osuog os LLS SI 7181-9081 LVI OE LYTLE Sevl WS es pep 0 69°S7Z Kaeo O0vT OOrT LLS'S1 SO9l CcSI-csvl SIvI 6 91.0€ O'VTolE Chl WS rT 90°97 ST-—La osuo0g CIC LLS'SI STELVScl /£VTOE YOToIE Albl WS 6r'T €0°97 SI-—Ld osu0g os LL SI 6rCI-EvCl /£ VTE IO OVAE STIvI WS LI‘€ £0°97 vI-La WIWY O-OZIT o9sT LL ST 6071 6011-6001 SI60 6 0C0E VPToIE Ort WS VE CHV €I-La WIW4a 0-0S¢ oort LLS VI Sv00 IZ00-OZET VOET TOTAE /8°TTOE 6€I WS 97E 78°EZ €I-La@ WIWY 0-0£8 OcET LLS v1 8vIZ COIT-6S61 ST6l LST OE /£ 1T0E 8€I WS [Mey ae =a weog ate OL8 LLS €1 -LE80 S080 Cele JESToE Let WS sjouy Do uw uw Ao So “duiay ‘ON puq SUIYSIY idDJIS apnyisuoT apnjijoT paads aavfains ydois iva) yidaq yidaq | ¢ (a ‘ON 5, ays pag -OUWls2YI SUIYSLT wojylog aul, uoljIsog uolDnis ~Ayj0g (panuljuod) | aTaV 193 THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 6L‘I 10°97 S7-La osuog = ZZ LL'S'81 9ETI-BITI OTE 880.0 Assi WS 68'T 00°97 ~=s-« S7-Ll osuog = 0S LL'S'81 vITI-6071 OTO.TE —-8°80.0€ S8ST WS vee 86°SZ S7-Ld WINU 0-0SL O0EI LLS81 OvOI 660-680 6080 O'LS.1E S$ °SO0€ LSI WS y9'T 76°SZ = TuojsneaN =) LL'S'81 UI OT TLSoLE S100€ NIST INS v9'T 76'S7 b7-La osuog = ZIZ LL 8 v£90-€ 190 TLSLE iS TO0€ d9St WS 6£'1 $6°S7 pc-La osuog = 0S LL'S 81 6090-7090 TLSNE S10.0€ S9ST WS 67E Ol‘vZ TUO}sNeN«x 0 JES Ph IU Q] IS CVE — 4S VCE SST WS 67€ 1) 4 €c—La WINU 0-005 0097 LAS HEN! 8000-80EZ 6E7Z = SM 250 31°59,3’ 29°22,5’ 2132 2140-2142 2202 27.6.79 cc. 150-200 c.150-200 Heavy BT-62 20,85 0,00 2 Dredge 5 a SM 251 31°56,0’ 29°24,6’ — 2300-0100 — 27.6.79 137 137 JDNNL -- 21,31 0,00 g SM 2528S 31°44,8’ 29°37,1’ — 0600-0605 — 28.6.79 650 53 Bongo BT-63 21,35 1,68 © SM 252D 31°44,8° 29°37,1’ — 0612-0626 — 28.6.79 650 212 Bongo BT-63 21,35 1,63 & SM 252N 31°44,8’ 29°37,1’ — 0635-0646 — 28.6.79 650 0 Neustony, — 21,34 2,68 $ ro) SM 253 31°44,6’ 29°41 ,2’ 0739 0809-0828 0907 28.6.79 1010 1010 Heavy BT-63 21,23 0,00 = Dredge a 5 SM 254 31°42,6’ 29°40,4’ 0928 1016-1047 1136 28.6.79 860-850 860-850 Dbl. Beam BT-63 21,18 0,23 A Trawl na ¢ an SM 255 31°37,8’ 29°40,8’ 1257 1302-1305 1311 28.6.79 125 125 Heavy — 21,55 0,00 bY Dredge SM 256 S1°37,2" 29°41,7’ 1325 1330—- — — 28.6.79 105 os Dbl. Beam _— _ — Trawl £07 ET TENS Hew Ec Ree REC aay “9ESL OLOL OLE CVLi OLLI OF 6l PLIT SLIT LLIt LL It OV It €9-Le er eal =a a Pan a — St ILO OOLT OTLT SZ7LI OLE LOTT 891% 391% 89% O707 Z9-LA tb 8S8b 6(9ObS = BLD dIZ8s«OSG SOT ISIT €LEl OOST 99%ST E8‘ST OSOT €8°Or OL6I OL6I LOT SL‘6I O8'61 I19-LE = = = =e — 006 £801 ZLIL POEL IOP] ILL O6LYI Z6PT B8I'St OOO! 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SEC. 6rd Zz [Sve LV S909) OY) S8Ree Ol Seis VEE oot) MOSEL OLS) Ceol SS0% 651% OS:%e ‘POC 6O'ee OLee ESce SPALal o = ae uA re ee SOLO ee ia Dem OPC Cl ami Ol OSS CrOle SGGLmnChOCMmbONG —LUCCe Shc Shee Cate irae s OV CS 069/81 OGG 0S Ol OS IT 17S cl OS She TOGls G0G cO0G 16a | 0S6C chSe SES pesc Ores Peer SPE : OS OS VLG OS TGS MONON 61s ba nec Iason EDO aSivelamOP len ticaSile COOL utrOce 9 IommcSee. ES:CG OS CC SPALel B Rive SOS 985 60/8 OG mse OL (SPIE IOC OS El LaLy . POG Siok ison SG0C Lice. O8te sec Ocse Te tc Pre 9 6fr Ssh £65 OSL IL'8 €L6 LBOL B8LIT ZOPIT LIT 99°91 P69T O8LT O96I Or07 680% LOIZ O6S‘%% EL’? €b-Ld = coy BGY SCO OS CSS IOOl ae) 11) 00 Clee TL CU moron TLE I COLT wiGLSI) (S0\0GesGbicG OSG CSCC cece. OLice “chale E Leb 6b 679 69%L SL'S PL LEOL OIL HEHE 8991 ILE 7ST 890% 8zIZ EEC 96°C B67TC B8OCC PEC? 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GE-LE zZ on ae Be as a ae ee ats = me — = a= — — — = = _ OF Ces SESE < ON Do Do Oo Jo Do Do Do Jo Oo Oo Oo Do Do Do Oo Oo Jo Do Do ydeis ‘dwej ‘duwiey ‘dwoy ‘dwoy ‘dwoy ‘dwoy ‘dwoy ‘dwoy ‘dwoy ‘dwoy ‘dway ‘dwoy ‘dwoy ‘dwoy ‘dwoy ‘dwoy -duay ‘dwoy “dwoy, -owoyy -Ayeg wOOdT 006 “O08 WOOL W009 WOODS WOOP WOOE WOOT WOOT WSL WOL WOOD WoOS WOp Woe WOT WOT WOx Wdeq ‘9SInIo Uy oy} SuLINp seyyoid ydessoureyAyjeq Wor, pourejgo se ‘syjdep pojoojes ye Sornje1oduwa], 204 9 ATaVL THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 205 ACKNOWLEDGEMENTS The South African Museum thanks the following organizations and individuals for their help during this programme: Government of Republic of Transkei for permission to fish in territorial waters; South African National Council for Oceanographic Research for financial assistance; Sea Fisheries Branch, Oceanographic Research Institute, East London Museum, Captain G. Foulis and crew of the Meiring Naude, Messrs P. Slevin and D. M. Pim (National Research Institute of Oceanology), Prof. M. M. Smith and Dr. P. C. Heemstra (J. L. B. Smith Institute of Ichthyology), Mr G. B. Ross (Port Elizabeth Museum), and Dr B. Kensley (U.S. National Museum of Natural History). Mr V. Branco (South African Museum) prepared Figure 1. REFERENCES BELYAEV, G. M. 1966. Hadal bottom fauna of the world ocean. Trans. by A. Mercado. Jerusalem: Israel Program for Scientific Translations, 1972. Louw, E. 1977. The South African Museum’s Meiring Naude cruises. Part 1. Station data 1975, 1976. Ann. S. Afr. Mus. 72: 147-159. Menzies, R. J. 1964. Improved techniques for benthic trawling at depths greater than 2 000 metres. Antarctic Res. Ser. Washington 1: 93-109. 6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature (particularly Articles 22 and 51). : Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. Nnov., syn. nov., etc. An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, i.e. all published scientific names by which the species previously has been designated are listed in chronological order, with all references to that name following in chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845) Figs 14-15SA Nucula (Leda) bicuspidata Gould, 1845: 37. Leda plicifera A. Adams, 1856: 50. Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b). Nucula largillierti Philippi, 1861: 87. Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. Note punctuation in the above example: comma separates author’s name and year “semicolon separates more than one reference by the same author full stop separates references by different authors figures of plates are enclosed in parentheses to distinguish them from text-figures dash, not comma, separates consecutive numbers Synonymy arrangement according to chronology of bibliographic references, whereby the year is placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable. In describing new species, one specimen must be designated as the holotype; other speci- mens mentioned in the original description are to be designated paratypes; additional material not regarded as paratypes should be listed separately. The complete data (registration number, depository, description of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.: Holotype SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973. , Note standard form of writing South African Museum registration numbers and date. 7. SPECIAL HOUSE RULES Capital initial letters (a) The Figures, Maps and Tables of the paper when referred to in the text e.g. ‘.... the Figure depicting C. namacolus ...’; *. . . in C. namacolus (Fig. 10)...’ ’ (b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by initials or full names e.g. DuToit but A.L.du Toit; Von Huene but F. von Huene (c) Scientific names, but not their vernacular derivatives e.g. Therocephalia, but therocephalian Punctuation should be loose, omitting all not strictly necessary Reference to the author should be expressed in the third person Roman numerals should be converted to arabic, except when forming part of the title of a book or article, such as “Revision of the Crustacea. Part VIII. The Amphipoda.’ D Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial capital letter, provided the same generic name is used consecutively. Name of new genus or species is not to be included in the title: it should be included in the abstract, counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts. ELIZABETH LOUW | THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES ~ PART 10 STATION DATA 1977, 1978, 1979 — ISSM 0303-2515 OF THE SOUTH AFRICAN MUSEUM CAPE TOWN INSTRUCTIONS TO AUTHORS 1. MATERIAL should be original and not published elsewhere, in whole or in part. 2. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes...” ‘Smith (1969: 36, fig. 16) describes . “As described (Smith 1969a, 19695; ros ihe ‘As described (Haughton & Broom ee) “As described (Haughton ef al. 1927) . Note: no comma separating name ard year Dagination indicated by colon, not p. names of joint authors connected by ampersand et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o, scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BuLLouGn, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FiscHer, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gen. 74: 627-634. Konn, A. J. 19602. Ecological Fabre on Conus (Mollusca; Gastropoda) in the Trincomalee region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320. Konn, A. J. 19606. Spawning behaviour, pe masses and larval development in Conus from the Indian Ocean, Bull. Bingham oceanogr. Coll. 17 (4): 1-51. Tureve, J. 1910. Mollusca: B. ate eee Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Wahimea R1 Band August 1980 Augustus Part 6 Deel THE UMZAMBA FORMATION AT ITS TYPE SECTION, UMZAMBA ESTUARY (PONDOLAND, TRANSKEI), THE AMMONITE CONTENT AND PALAEOGEOGRAPHICAL DISTRIBUTION By HERBERT CHRISTIAN KLINGER & WILLIAM JAMES KENNEDY Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000 Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na gelang van die beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000 OUT OF PRINT/UIT DRUK 1, 2(1-3, 5-8), 3(1-2, 4-5, 8, t—p.i.), 5(1-3, 5, 7-9), 6(1, t.-p.i.), 711-4), 8, 9(1-2, 7), 10(1-3), 11(1-2, 5, 7, t.-p.i.), 15(4-5), 24(2), 27, 31(1-3), 32(5), 33 Copyright enquiries to the South African Museum Kopieregnavrae aan die Suid-Afrikaanse Museum ISBN 0 908407 93 9 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap THE UMZAMBA FORMATION AT ITS TYPE SECTION, UMZAMBA ESTUARY (PONDOLAND, TRANSKED, THE AMMONITE CONTENT AND PALAEOGEOGRAPHICAL DISTRIBUTION By HERBERT CHRISTIAN KLINGER South African Museum, Cape Town & WILLIAM JAMES KENNEDY Geological Collections, University Museum, Oxford (With 5 figures and 3 tables) [MS. accepted 6 December 1979} ABSTRACT Detailed collecting at the type section was undertaken to determine the exact age of the Formation and to determine whether more than one faunal zone is present. Faunal analysis based on ammonites shows that the age of the Formation at the type section ranges from Middle Santonian to Lower Campanian. The ammonoid faunas at the type section of the Umzamba Formation in Pondoland were compared with those of the False Bay region of Zululand, and it was found that certain morphotypes, e.g. oxycones, compressed evolute forms, and serpenticones are more dominant in the shallower water transgressive environment of Pondoland, presumably as a reflection of different living conditions in the two areas. CONTENTS PAGE Introduction j : : : : : : : 3 207 Description of the exposures ; j ; : ; - : sy > 208 The age of the Umzamba Formation . : Sees a aac 209 The base of the section . - : : 2 ; : = 209 Top of the succession 5 : ; Be ee ; : > \216 Subdivision of the sequence . e216 Comparison of Pondoland and Zululand ammonoid faunas ney 21S References . : : ; : ; ; F ? ; : 5 PPR INTRODUCTION Although the presence of Cretaceous sediments in the vicinity of the Umzamba River Estuary (Pondoland, Transkei) has been known since at least 1824, the exact age and the question of whether or not a succession of distinct faunal associations is recognizable have long been disputed (Baily 1855; Griesbach 1871; De Grossouvre 1901; Rogers & Schwarz 1902; Woods 1906; Du Toit 1912, 1920, 1954; Van Hoepen 1920, 1921, 1965; Plows 1921; Spath 1921b, 1922, 1953; Rennie 1930; Haughton 1963, 1969; Kennedy & Klinger 207 Ann. S. Afr. Mus. 81 (6), 1980: 207-222, 5 figs, 3 tables. 208 ANNALS OF THE SOUTH AFRICAN MUSEUM 1975). On the basis of the fauna and flora described by Baily (1855), Griesbach (1871), Chapman (1904, 1923), Woods (1906), Lang (1906), Broom (1907), Spath (1921la, 1921b), Van Hoepen (1920, 1921, 1965), Rennie (1930, 1935), Smitter (1956), Little (1957), Madel (1960), Miiller-Stoll & Méadel (1962), Dingle (1969), and Kennedy & Klinger (1977a, 19776 1979), ages varying from Albian to Maastrichtian were postulated, the prevalent idea being that the Formation was of Senonian age (i.e. Coniacian to Campanian). Failure to arrive at a decisive conclusion may be ascribed mainly to lack of precise stratigraphic control, and/or lack of current knowledge of the systematic affinities and stratigraphic ranges of some of the fossil taxa. Aided by the extensive monographical studies of the Madagascar ammonoid faunas by Collignon (1928 onwards), his series Atlas des fossiles caracteristiques de Madagascar (ammonites), and by the authors’ preliminary biozonation for | southern Africa (Kennedy & Klinger 1975), a new attempt was made to fill this gap in our current knowledge. For this purpose a detailed sampling programme was undertaken by one of the authors (H.C.K.) in 1974. However, due to circumstances beyond the authors’ control, results of this investigation could not be published earlier and consequently parts thereof were disclosed elsewhere (Klinger & Kennedy 1977: Cooper 1977). Finally, the apparent faunal differences between northern Zululand and Pondoland are examined briefly to determine their extent and probable causes. DESCRIPTION OF THE EXPOSURES Descriptions of the exposures at and near the Umzamba Estuary are provided in varying degrees of detail by Garden (1855), Griesbach (1871), Rogers & Schwarz (1902), Du Toit (1912, 1920), Plows (1921), Gevers (in Rennie 1930) and Kennedy & Klinger (1975), that of Plows (1921) being the most complete. Apart from details of the ammonoid succession at the type section, little else can be added to these general accounts. The lithologies encountered may be described briefly as follows: The Cretaceous strata overlie Ordovician? quartzites belonging te the Table Mountain Group unconformably with a slight seaward dip of the order of two to four degrees. The basal beds are conglomeratic, consisting of abundant sandstone and lydianite pebbles set in an arenaceous matrix. Large logs, chelonian scutes and other reptilian remains, comminuted shell material, selachian teeth, and baculitid ammonites are locally common in these basal beds. Higher up in the sequence, lithologies become finer-grained, consisting essen- tially of alternating layers of grayish-green, fine-grained silts, and coarser- grained, shelly or sandy concretionary horizons. Some of the concretionary layers show traces of cross-bedding and scouring. Both silts and concretionary layers yield abundant fossils, and sections of gigantic inoceramids, more than a metre in diameter, are conspicuous. Fossils in the silty horizons are generally preserved as internal moulds, whereas those in the concretionary horizons retain THE UMZAMBA FORMATION AT ITS TYPE SECTION 209 the original shell material. The majority of ammonites extracted from the concretionary horizons were embedded horizontally. The most complete exposure is on the northern side of the estuary (Figs 1-2), named the Umzamba Cliff by Plows (1921, pl. 8 (fig. 3)). This is Gevers’s (in Rennie 1930) ‘first locality’ and the authors’ (Kennedy & Klinger 1975) locality 1. Griesbach’s (1871, fig. 5) locality was probably taken about 100 m north-east of here (Fig. 3). Details of the section here vary considerably due to landslides and heavy surf action at the base of the cliff (see Rogers & Schwarz 1902: 40; Plows 1921: 60). The Umzamba Cliff is here referred to as locality A (see Figs 3-4). Beds A3 and A7 are the levels of the prominent caves remarked upon by the early workers. These result from the collapse of the soft, silty beds between hard, concretionary layers. Horizons below Bed A8 can be easily reached from the base of the cliff by climbing on rubble from landslips. Higher horizons can be reached by scaling the cliffs or by taking a footpath over the top of the hill and then climbing down the cliff (see also Gevers 1977 for anecdotal details). During low tide, foreshore platforms are exposed north-east of locality A for more than a kilometre along the beach. Due to the abundance of silicified tree trunks, this locality is known as the ‘Petrified Forest’ and is indicated as such on tourist and topographic maps. Most of the larger logs appear to be orientated in an east-north-east direction, presumably paralleling the Cretaceous shoreline. The exposures are in horizons below those seen at locality A, and are here referred to as locality B. Locality C is situated on the southern side of the estuary (Plow’s 1921 Right Bank), and extends for some distance along the coast, but the latter is only poorly exposed at low tide (see Rogers & Schwarz 1902: 40). Strata even lower down in the succession than those found at locality A and B are exposed here, but the actual contact with the underlying basement rocks was not exposed during the authors’ visits. Large boulders of quartzite derived from the Ordovician? Table Mountain Group are exposed on the NW side of the estuary, but one or two metres of sand covered the actual contact. Approxi- mately 10 m of sediment are exposed at locality C. The highest bed, Bed C11, probably corresponds to Bed 3 at locality A. THE AGE OF THE UMZAMBA FORMATION In determining the age of the Umzamba Formation at the type section, the ammonoid zonation compiled for Madagascar by Collignon (1966, 1969) and the provisional one compiled for Natal and Zululand by the authors in a slightly modified form (Kennedy & Klinger 1975) (Tables 1 and 2 respectively) are employed. The base of the section The lowermost fossiliferous units are Bed C4 and the foreshore outcrops in the northern part of locality B. Occasional Baculites capensis Woods, rare ANNALS OF THE SOUTH AFRICAN MUSEUM *A[COS IOF JJl[9 OY} JO JOOF Oy} ye SI OUBAUONY SoUULYOL ITA] "PL6 Ul paydessojoyd “WD equiezwi~ ‘equiezuip 3e sInsodxo ule oY], *[ “SI “p INSLEY Ul PUL 4X9} OY} UI 0} PodIajol Se s}TUN o}eIISN]II OF YO equiezw jo Zumelg ‘7 ‘3Ly oN THE UMZAMBA FORMATION AT ITS TYPE SECTION 212 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 3. Section at Umzamba about 100 m to the north of the main exposure shown in Figure 1. This appears to be the site on which Griesbach (1871 (fig. 5)) based his section. Gaudryceras varicostatum Van Hoepen, Texanites umzambiense Klinger & Kennedy and Scaphites sp. occur here. The authors had originally considered (Kennedy & Klinger 1971, 1975) that ’ the base of the Umzamba Formation was of Coniacian age on the basis of the reference of Muniericeras cricki Spath (1921b, 1922) (of which Barroisiceras umzambiensis van Hoepen 1965 (Fig. 7A) is a synonym) to the Coniacian collignoniceratid genus Subprionotropis. Since most of the above listed species are typically Santonian forms, this determination became open to question, and the authors would now suggest that this species is better referred to the homoeo- morphic genus Lehmaniceras Collignon, 1966 (see also Klinger & Kennedy 1977: 103; Cooper 1977: 16). Van Hoepen’s specimen was collected from an unrecorded horizon on the southern side of the Umzamba Estuary at locality C. Apart from Pondoland, Lehmaniceras is known only from Madagascar where it is relatively abundant in the Middle Santonian. The association of Texanites umzambiense Klinger & Kennedy with Baculites capensis Woods is related to that of Plesiotexanites olivetiforme Klinger & Kennedy and Baculites capensis occurring in the first division of the Santonian in Zululand. In Madagascar, Baculites capensis occurs in the Lower THE UMZAMBA FORMATION AT ITS TYPE SECTION 213 EXPOSURE AT UMZAMBA ESTUARY PROBABLE AGE CORRELATION MADAGASCAR ZULULAND STAGE SUBSTAGE ZONE CAMPANIAN II Baculites sulcatus Hauericeras madagascariense Anapachidiscus arrialoorensis Menabites boulei Baculites sulcatus Glyptoxoceras subcompressum Scaphites cf. aquisgranensiformis CAMPANIAN CAMPANIAN |! Submortoniceras condamyi Eulophoceras spp Hauericeras gardeni (abundant) Pseudoschloenbachia umbulazi (var. spinifera abundant) Damesites compactus Heteroceras amapondense ts 3 Submortoniceras woodsi Hauericéras gardeni Plesiotexanites stangeri P. ‘seudoschloenbachia By: tee Saghalinites nuperus Madrasites afri icanitm. Sore Pseudophyllites indra Texanites soutoni 5 Hauericeras gardeni Hauericeras gardeni cae ee Phylloceras woodsi P>umbulazi umbulazi Ye Echinoids and inoceramids ae x conspicuous Texanites soutoni Es Submortoniceras woodsi Pseudoschloenbathia sp. Texanites soutoni SN P. umbulazi Hauericeras gardeni Anapachydiscus wittekindi Eulophocerus jacobi Pseudoschloenbachia umbulazi SANTONIAN III SANTONIAN Madrasites similis Baculites capensis Texanites umzambiense Mean sea level Texanites hourcq! Baculites capensis Gaudryceras varicostata Scaphites sp. LOCALITY B SANTONIAN II Interminate molluscs Table Mountain Group quartzite boulders LOCALITY C Fig. 4. Stratigraphic section at Umzamba, including Umzamba Cliff, here marked A, the foreshore beach exposures north of A, here marked as B, and the section on the southern side of the estuary, here marked C. 214 ANNALS OF THE SOUTH AFRICAN MUSEUM TABLE 1 Ammonoid zonation for Madagascar as compiled by Collignon; after Collignon 1966 and 1969. Z = wm Z — < Basal zone of Pachydiscus lamberti RS a< ae) Ss Subzone of SJ : i Termierella s Zone of Menabites boulei lenticilnnis § and Subzone of SS Anapachydiscus arrialoorensis Rabeiella z & SS orthogonia < = 2 y =| may Subzone of Z 3 Scaphites < coat & 2 8 Thee GH reesidei < = Karapadites karapadensis Subzone of ~) mls Maorites Fa 3| 3 aemilii = S R Subzone of S 2 Zone of | Hourcquiella ke} ere is S = Anapachydiscus wittekindi WERE, S a8 and Subzone of | S 28 Eulophoceras jacobi Besareita xz s = P J besairici e | Q Basal unit with Neogauthiericeras zafimahovai Zz Ss x A Zone of Pseudoschloenbachia umbulazi Oe - Pn Z < — is A Zone of Texanites hourcqi Qe =F =e.) ZA < Zz, Zone of Texanites oliveti [a4 i BS B4 Hn SEE OED ts a THE UMZAMBA FORMATION AT ITS TYPE SECTION 215 TABLE 2 Provisional ammonoid succession for Natal and Zululand as compiled by the authors (1975), here presented in slightly modified form to accommodate new data. Menabites (Australiclla) abundant in lower part. Some appear to range throughout together with Bevahites spp. Baculites sulcatus is abundant throughout while pachydiscids become common in the higher part, e.g. A. wittekindi, A. arrialoorensis CAMPANIAN II Submortoniceras woodsi and related forms are common: other ammonites include Bevahites and Menabites, Haucericeras gardeni, Pseudoschloenbachia, Bostrychoceras and diplomoceratids. CAMPANIAN I The local base of the stage is drawn below the level of abundant Submortoniccras Hauericeras gardeni is abundant; the remainder of the fauna is as in Santonian II and is relatively scarce. SANTONIAN Ii Abundant Plesiotexanites stangeri and varieties, Texanites soutoni, Texanites spp., Hauericeras and Pseudoschloenbachia occur, as do Eupachydiscus? Hyphantoceras and diplomoceratids SANTONIAN II Texanites oliveti, Plesiotexanites stangeri densicosta and sparsicosta, Hauericeras gardeni, Pseudoschloenbachia sp Pseudophyllites indra, Karapadites?, Eupachydiscus? sp.. Gaudryceras spp., Hyphantoccras sp. and diplomoceratids. The base of the stage is drawn at the level of the appearance of Texanites sensu strictu in numbers SANTONIAN I Abundant baculitids ornamented only by growth striae. Also ammonites resembling Pseudoschloenbachia primitiva Collignon and Scaphites. CONIACIAN Vv 216 ANNALS OF THE SOUTH AFRICAN MUSEUM | and Middle Santonian. Gaudryceras varicostatum occurs as low as the second division of the Coniacian of Zululand, but is also recorded from the Lower Santonian of Madagascar (Collignon 1966: 3). On the basis of these ranges, it seems quite certain that the basal beds are of Santonian age, and probably uppermost Lower to Middle Santonianinthesense of Collignon (1966) and equivalent to the authors’ first or second divisions, probably the latter, of the Zululand Santonian. Top of the succession The uppermost ammonite-bearing beds exposed, Beds Al4 and A15 (probably the equivalent of Gevers’s (in Rennie 1930) Bed T) yield Baculites sulcatus (Baily), Hauericeras madagascariense Collignon (Fig. 5B), Glyptoxoceras subcompressum (Forbes) and Scaphites cf. aquisgranensiformis Collignon. Hauericeras madagascariense has a very restricted range in Madagascar, occurring only in the upper part of the Lower Campanian zone of Menabites boulei and Apapachydiscus arrialoorensis (see Collignon 1961, 1969; and Table 1 herein). Comparisons with Zululand at this interval are tenuous, but Baculites sulcatus is comparable with Baculites vanhoepeni Venzo, as discussed. earlier (Klinger & Kennedy 1977: 73-74) and is indicative of the second division of the Campanian in Zululand. The absence of Menabites s.1. species which characterize this horizon in Zululand, however, is puzzling. The uppermost exposed beds at the Umzamba Cliff are thus provisionally dated as uppermost Lower Campanian in the sense of Collignon (1969) and tentatively the second division of the Zululand Campanian sensu Kennedy & Klinger (1975). Subdivision of the sequence The ammonites collected in situ from the remainder of the sequence exposed at the Umzamba Estuary are shown in Figure 4. Apart from a thin zone of rolled and encrusted clay pebbles and hiatus concretions in Bed A5, which may represent a very short break in deposition, no evidence could be found of a major sedimentological interruption within the Umzamba Formation. It may thus be assumed that deposition was virtually continuous. Bed A7 yields abundant Pseudoschloenbachia umbulazi umbulazi (Baily), P. umbulazi (Baily) griesbachi van Hoepen, and P. umbulazi (Baily) spinifera van Hoepen (all probably conspecific). All three ‘subspecies’ occur together, but P. umbulazi spinifera appears to become more abundant towards the top of the Bed. Hauericeras gardeni (Baily) is also very abundant. This association corre- sponds to the third division of the Santonian in Zululand, and the Upper Santonian zone of Pseudoschloenbachia umbulazi in Madagascar. Eulophoceras tenue van Hoepen, FE. umzambiense van Hoepen and other Eulophoceras species (all probably conspecific) occur together with Submortoniceras condamyi near the contact of Beds A7 and A8. In Madagascar the base of the Campanian is drawn immediately below the first occurrence of Eulophoceras, whereas it is THE UMZAMBA FORMATION AT ITS TYPE SECTION 217 Fig. 5. A. Lehmaniceras cricki (Spath, 1921). (= The holotype of Van Hoepen’s 1965: 161, pl. 6 and text-fig. 2a Barroisiceras umzambiensis. From the southern side of the Umzamba Estuary (herein locality C) at an unknown horizon (see Van Hoepen 1965: 162). Geological Survey SAS-P1093. x1. B. Hauericeras madagascariense Collignon, 1961. This is the specimen collected by Gevers from his horizon ‘T’ at the Umzamba Cliff (herein locality A), probably horizon A15, being associated with specimens of Baculites sulcatus Baily on reverse side. South African Museum SAM-7043. x0,75. 218 ANNALS OF THE SOUTH AFRICAN MUSEUM drawn in Zululand below the first occurrence of abundant Submortoniceras. It seems reasonable to draw the contact between the Santonian and Campanian Stages at the contact between Beds A7 and A8. It should be pointed out, however, that Submortoniceras appears slightly earlier in Pondoland than in Madagascar. The paucity of ammonites from above Bed A7 at the type section appears to be due more to physical difficulties encountered in collecting in higher sections of the cliff rather than real differences. Scaphites cf. aquisgranensiformis in Bed A14 is comparable with S. aquis- granensiformis Collignon, which occurs in the subzone of Scaphites reesidei at the boundary between the Zone of Menabites boulei and Anapachydiscus arrialoorensis and the Zone of Karapadites karapadensis of the Lower Campanian of Madagascar. COMPARISON OF THE PONDOLAND AND ZULULAND AMMONOID FAUNAS The authors (Klinger & Kennedy 1977: 104; 1980) and Spath (19216: 53) had previously pointed to the apparent differences between the ammonoid faunas of Pondoland and those of biostratigraphically equivalent sediments further north at Umkwelane Hill and in the False Bay region of Zululand. A detailed comparison of the faunas of Pondoland with those of Durban, Richards Bay, Umkwelane Hill, and Zululand must await a complete revision of all the ammonoid faunas, but preliminary results are given here in Table 3. The paucity of the faunas in the Durban and Richards Bay and to a lesser extent the Umkwelane Hill regions as shown in the table are primarily due to lack of sufficient exposures, rather than real differences. Furthermore, the absence of some Pondoland species in Zululand may be partially due to our inability to recognize the generally small Pondoland species as the nuclei of larger species occurring in Zululand. This latter discrepancy is not applicable to the Texanitinae, which generally grow to very large size, especially in Pondoland. Despite these shortcomings, the picture that emerges is not one of total geographic separation, but rather one of concentration of certain shell morpho- types in specific areas. Very good examples of this are the oxyconic pseudo- schloenbachiids and Eulophoceras spp., the compressed evolute Hauericeras gardeni (Baily) and also the serpenticonic rounded Plesiotexanites stangeri (Baily). Examination of the Van Hoepen Pondoland collections in the Transvaal Museum (Pretoria) and personal collection at the type section, shows that Pseudoschloenbachia and Hauericeras can be counted by the thousands, as compared to numbers well below the hundreds in biostratigraphically equivalent strata in Zululand. Known specimens of Eulophoceras from Pondoland number about fifty, whereas the number from Zululand is well below ten. Similarly, only two tentative specimens of Plesiotexanites stangeri (Baily) are known from the False Bay region of Zululand, whereas it is relatively common in Pondoland. THE UMZAMBA FORMATION AT ITS TYPE SECTION 219 TABLE 3 Distribution of ammonoid taxa at Umzamba, Durban, Richards Bay, Umkwelane Hill and northern Zululand. Abbreviations used in Table Absent Mabeiistieas Yale) Common =). 9. (©! —10=50 1-5 Abundant . ee (AN) 21100 1 Occasional : ‘ . (O) = 5-10 SPECIES RICHARDS BAY UMK WELANE PONDOLAND HILL DURBAN ZULULAND Phylloceras (H.) woodsi woodsi van Hoepen Partschiceras umzambiense (van Hoepen) . Tetragonites superstes van Hoepen Saghalinites cala (Forbes) Saghalinites nuperus (van Hoepen) Pseudophyllites indra (Forbes) Pseudophyllites teres (van Hoepen) Gaudryceras varicostatum van Hoepen (= G. cinctum Spath 1922) ‘Gaudryceras’ sigcau van Hoepen Anagaudryceras subsacya (Marshall) . Anagaudryceras subtilineatum (Kossmat) Verte brites kayei (Forbes) Gaudryceras denseplicatum (Jimbo) (= amapondense van Hoepen) “Heteroceras’ amapondense van Hoepen Pseudoxybeloceras amapondense van Hoepen Glyptoxoceras compressum (Forbes) Hoploscaphites spp Baculites capensis Woods Baculites bailyi Woods . Baculites sulcatus Baily Damesites compactum van Hoepen Desmophyliites simplex van Hoepen . Desmophyllites crassa (van Hoepen) . Hauericeras gardeni (Baily) . Hauericeras madagascariense Collignon Natalites spp. (including N. atalensis Spath, N. goutico= status Spath, N. faku van Hoepen, N. africanus van Hoepen, N. similis Spath) Parapuzosia haughtoni Spath Pachydiscus simplex van Hoepen Pachydiscus umtafunensis Spath . Pachydiscus antecursor van ae ; Eulophoceras spp. . ‘ O-C IR — (including Pulephoceras atalense Fiyatt, “Spheniscoceras’ africanum Spath, ‘S’ tenue Spath, ‘S’ minor Spath, ‘Pelecodiscus’ ama- pondense van MHoepen, ‘P’ umzambiense van Hoepen) Pseudoschloenbachia spp _.. A — O O O-C (including P. umbulazi (Baily), P. pee fournieri Spath, P. papillata Spath, P. gries- bachi Spath) Texanites umzambiense Klinger & Kennedy Texanites presoutoni Klinger & Kennedy . Texanites soutoni (Baily) : Texanites texanus s.1. . : Submortoniceras woodsi (Spath) . Submortoniceras condamyi Collignon . Plesiotexanites stangeri (Baily) Plesiotexanites matsumotoi Klinger & Kennedy Lehmaniceras cricki (Spath) soe . | | | mw] | | | | 7 | | ° on badass Pr| WR PRORORO | Ae alae |S Oo | 1 O oO out ea eles | Rwy | | | | cel) WRRARZRZXARA Lee Le ee LP eS cee | | | BOooRnaF| \ewassaltWworiseit)26 Oh owe asckwsuntsr.hlCO Ke Wceneren wes: ae 220 ANNALS OF THE SOUTH AFRICAN MUSEUM Other species, such as Texanites soutoni (Baily), Submortoniceras woodsi (Spath) and S. condamyi (Collignon) are common to both areas, though subtle differences exist as discussed earlier (Klinger & Kennedy 1980) meriting separation at sub- specific level. In the smooth lytoceratid forms the picture is somewhat obscure due to limited numbers (Kennedy & Klinger 1977b). Saghalinites nuperus (van Hoepen) and Pseudophyllites indra (Forbes) are common to both Zululand and Pondoland, but are numerically superior in Pondoland. Pseudophyllites teres (Marshall) is poorly known and has so far been recorded from Pondoland only. Phylloceratids are also restricted in numbers in Pondoland, but both known species, Phylloceras (Hypophylloceras) woodsi woodsi van Hoepen and — Partschiceras umzambiense van Hoepen have so far not been recorded from the False Bay region of Zululand (Kennedy & Klinger 1977a). Details on the pachydiscids are still in preparation but show no distinct trends, neither do the gaudryceratids (Kennedy & Klinger 1979). Amongst the heteromorphs, Baculites capensis Woods and B. bailyi Woods are common to both Zululand and Pondoland, whereas the Pondoland species, B. sulcatus (Baily), has a possible equivalent in Zululand in B. vanhoepeni Venzo (Klinger & Kennedy 1977). ‘Heteroceras’ amapondense van Hoepen (= Anaklinoceras stephensoni Collignon 1966) is relatively rare, but is known from both areas, as also, apparently are species of Pseudoxybeloceras and Glyptoxoceras. The scaphitids have not been studied sufficiently for detailed analysis. The fact that species such as Texanites soutoni, Submortoniceras woodsi, S. condamyi, Pseudoschloenbachia umbulazi and Hauericeras gardeni occur in all major outcrop areas, ranging from Umzamba, through Durban, Richards Bay and Umkwelane Hill to the False Bay region of Zululand, clearly precludes the presence of an impenetrable physical barrier, and supports the authors’ previous views (Klinger & Kennedy 1977: 104) of open marine connection between the areas. One of their previous views, however, that the biostrati- graphically equivalent strata in Zululand were probably not well exposed (Klinger & Kennedy 1977: 104) now appears erroneous in view of the above data. It has been suggested recently (see Cooper 1977: 32) that trophic resources increase during transgressions, which in turn leads to population explosions of certain favoured species. In the case of the diachronous southwards-extending Umzamba Formation Santonian transgression (Klinger & Kennedy 1977), these favoured species appear to be the oxyconic pseudoschloenbachiids and Eulo- phoceras spp, the compressed evolute Hauericeras gardeni and the serpenti- conic Plesiotexanites stangeri. Due to the abundance of specimens, a high degree of splitting into various morphotypes was applied to species (see Van Hoepen 1921; Spath 1922). The authors would suggest that it will be possible to reduce the various ‘species’ or variants of Pseudoschloenbachia and Eulophoceras to a single species each, as has been done in the case of Plesiotexanites stangeri (Baily) (Klinger & Kennedy 1980). | . THE UMZAMBA FORMATION AT ITS TYPE SECTION 221 (A comparable situation to that at Umzamba occurs in Zululand following the Lower Coniacian transgression (Kennedy & Klinger 1971). Here the sediments are dominated by the oxyconic Proplacenticeras ‘species’ P. umkwelanense (Etheridge), P. subkaffrarium (Spath) and P. kaffrarium (Etheridge) and very evolute serpenticonic peroniceratids belonging to the groups of Peroniceras tridorsatum (Schliter), P. dravidicum (Kossmat), P. westphalicum (Schliiter) etc. Klinger, et al. (in prep.) suggest that the Proplacenticeras ‘species’ ranging from the completely smooth P. umkwelanense through P. subkaffrarium with undulating flanks to the distinctly umbilically spinose P. kaffrarium all belong to one variable species. A similar simplification of peroniceratid systematics is also envisaged.) The question which now arises, and for which the authors can find no satisfactory answer, is why these particular morphotypes (oxycones, e.g. Pseudoschloenbachia; compressed evolute, e.g. Hauericeras; and serpenticone, e.g. Plesiotexanites stangeri) proved to be so successful in terms of numbers in a shallow-water, transgressive environment and tended towards a certain degree of endemism and great intraspecific variation. The wide global distribution of Plesiotexanites stangeri (Klinger & Kennedy 1980) clearly shows that these forms were not restricted to transgres- sive habitats only and were capable of substantial dispersal, but apparently preferred a shallower water transgressive milieu. It may be suggested that the shell types were hydrodynamically suited to this particular type of environment, but again specific explanations are lacking. REFERENCES Baty, W. H. 1855. Description of some Cretaceous fossils, South Africa. Q. J/ geol. Soc. Lond. 11: 454465. Broom, R. 1907. On some reptilian remains from the Cretaceous Beds at the mouth of the Umpenyati River. Rep. geol. Surv. Natal Zululand. 3: 95. CHAPMAN, F. 1904. Foraminifera and Ostracoda from the Cretaceous of East Pondoland, South Africa. Ann. S. Afr. Mus. 4: 221-237. CHAPMAN, F. 1923. On some Foraminifera and Ostracoda from the Cretaceous of the Umzamba River, Pondoland. Trans. geol. Soc. S. Afr. 26: 221-237. CoLLIGNon, M. 1961. Ammonites néocrétacées du Menabe (Madagascar) VII. Les Desmo- ceratidae. Annls. géol. Serv. Mines Madagascar 31: 1-115. CoLLiGNon, M. 1966. Atlas des fossiles caractéristiques de Madagascar (Ammonites). XIV (Santonien). Tananarive: Service Geologique. COLLIGNON, M. 1969. Atlas des fossiles caractéristiques de Madagascar (Ammonites). XV (Campanien inférieur). Tananarive: Service Geologique. Cooper, M. R. 1977. Eustacy during the Cretaceous: its implications and importance. Palaeogeography, Palaeoclimatol., Palaeoecol. 22: 1-60. DINGLE, R. V. 1969. Upper Senonian Ostracods from the coast of Pondoland, South Africa. Trans. R. Soc. S. Afr. 38: 347-385. Du Toit, A. L. 1912. The geology of Pondoland. Ann. Rep. geol. Surv. U. S. Afr. 1912: 153-180. Du Torr, A. L. 1920. The geology of Pondoland and portions of Alfred and lower Umzimkulu Counties, Natal. An explanation of Cape sheet 28 (Pondoland). Pretoria: Geological Survey. Du Torr, A. L. 1954. Geology of South Africa. London: Oliver & Boyd. 222 ANNALS OF THE SOUTH AFRICAN MUSEUM GarbkN, R. J. 1855. Notice of some Cretaceous rocks near Natal, South Africa. Q. JI geol. Soc. Lond. 11: 453-454. Gevers, T. 1977. Fossils and dynamite. Quart. News Bull. geol. Soc. S. Afr. 19: 10-11. GrigsBACH, C. L. 1871. On the geology of Natal in South Africa. Q. JI geol. Soc. Lond. 27: 53-72. GrossouvrE, A. DE 1901. Réchérches sur la Craie Supérieure. 1. Stratigraphie générale. Mém. serv. Carte géol. dét. Fr. 1013 pp. HaucutTon, S. H. 1963. The stratigraphic history of Africa south of the Sahara. Edinburgh, London: Oliver & Boyd. HauGutTon, S. H. 1969. Geological history of southern Africa, Cape Town: Geological Society of South Africa. Kennepy, W. J. & Kuincer, H. C. 1971. A major intra-Cretaceous unconformity in eastern South Africa. J. geol. Soc. Lond. 127: 183-186. KENNEDY, W. J. & K1LinGer, H. C. 1975. Cretaceous faunas from Zululand and Natal, South Africa. Introduction, Stratigraphy. Bull. Br. Mus. nat. Hist. (Geol.) 25: 265-315. KENNEDY, W. J. & KLINGER, H. C. 1977a. Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family Phylloceratidae. Bull. Br. Mus. nat. Hist. (Geol.) 27: — 349-380. KENNEDY, W. J. & KLINGER, H. C. 1977b. Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family Tetragonitidae Hyatt, 1900. Ann. S. Afr. Mus. 73: 149-197. KENNEDY, W. J. & KLINGER, H. C. 1979. Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family Gaudryceratidae. Bull. Br. Mus. nat. Hist. (Geol.) 31: 121-174. K.inGer, H. C. & KENNEDY, W. J. 1977. Upper Cretaceous ammonites from a borehole near Richards Bay, South Africa. Ann. S. Afr. Mus. 72: 69-107. K.LinceErR, H. C. & KENNEDY, W. J. 1980. Cretaceous faunas from Zululand and Natal, South Africa. The ammonite subfamily Texanitinae Collignon, 1948. Ann. S. Afr. Mus. 80. Lane, W. D. 1906. Polyzoa. In: Woops, H. The Cretaceous faunas of Pondoland. Ann. S. Afr. Mus. 4: 275-350. LitTLe, J. DE V. 1957. A new species of Trigonia from the Upper Cretaceous beds near the Itongazi River, Natal. Palaeont. afr. 4: 117-122. Maéper, E. 1960. Monimiaceen-HOlzer aus den Oberkretazischen Umzambaschichten yon Ost-Pondoland, S. Afrika. Senck. leth. 41: 331-391. MULLER-STOLL, W. R. & MADEL, E. 1972. Fossil woods of Monimiacea and Euphorbiacea from the Upper Cretaceous Umzamba Beds of East Pondoland, C.P. Trans. geol. Soc. S. Afr. 65: 93-104. PiLows, W. J. 1921. The Cretaceous rocks of Pondoland. Ann. Durban Mus. 3: 58-66. RENNIE, J. V. L. 1930. New Lamellibranchia and Gasteropoda from the Upper Cretaceous of Pondoland (with an appendix on some species from the Cretaceous of Zululand). Ann. S. Afr. Mus. 28: 159-260. Rocers, A. W. & SCHWARZ, E. H. L. 1902. General Survey of the rocks in the southern parts of the Transkei and Pondoland including a description of the Cretaceous rocks of eastern Pondoland. Rep. geol. Commn Cape Good Hope 1901: 25-46. SMITTER, Y. H. 1956. Foraminifera from the Upper Cretaceous beds occurring near the Itongazi River, Natal. Palaeont. afr. 3: 103-107. SPATH, L. F. 1921a. On Cretaceous Cephalopoda from Zululand. Ann. S. Afr. Mus. 12: 217-321. SPATH, L. F. 19216. On Upper Cretaceous Ammonoidea from Pondoland. Ann. Durban Mus. 3: 39-57. SPATH, L. F. 1922. On the Senonian ammonite fauna of Pondoland. Trans. R. Soc. S. Afr. 10: 113-147. SpATH, L. F. 1953. The Upper Cretaceous Cephalopod fauna of Grahamland. Scient. Rep. Falkld. Isl. Dep. Surv. 3: 1-60. VAN Hoepen, E. C. N. 1920. Description of some Cretaceous ammonites from Pondoland. Ann. Transy. Mus. 7: 142-147. VAN Hoepen, E. C. N. 1921. Cretaceous Cephalopoda from Pondoland. Ann. Transy. Mus. 8: 1-48. VAN Hoepen, E. C. N. 1965. New and little known Zululand and Pondoland ammonites. Ann. geol. Sury. S. Afr. 4: 158-172. Woops, H. 1906. The Cretaceous fauna of Pondoland. Ann. S. Afr. Mus. 4: 275-350. rd SF AR A mS OI I NA item A? rn ne 6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature (particularly Articles 22 and 51). 4 Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov., etc. An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, i.e. all published scientific names by which the species previously has been designated are listed in chronological order, with all references to that name following in chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845) Figs 14-15A Nucula (Leda) bicuspidata Gould, 1845: 37. Leda plicifera A. Adams, 1856: 50. Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b). Nucula largillierti Philippi, 1861: 87. Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110, Barnard, 1964: 234, figs 8-9. Note punctuation in the above example: comma separates author’s name and year semicolon separates more than one reference by the same author full stop separates references by different authors figures of plates are enclosed in parentheses to distinguish them from text-figures dash, not comma, separates consecutive numbers Synonymy arrangement according to chronology of bibliographic references, whereby the year is placed in front of each entry, and the synonym repeated in full for each entry, is “not acceptable. In describing new species, one specimen must be designated as the holotype; other speci- mens mentioned in the original description are to be designated paratypes; additional material not regarded as paratypes should be listed separately. The complete data (registration number, depository, description of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.: Holotype SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973. Note standard form of writing South African Museum registration numbers and date. 7. SPECIAL HOUSE RULES Capital initial letters (a) The Figures, Maps and Tables of the paper when referred to in the text e.g. ‘... the Figure depicting C. namacolus ...’; *. . . in C. namacolus (Fig. 10)...’ (b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by initials or full names e.g. DuToit but A.L.du Toit; Von Huene but F. von Huene (c) Scientific names, but not their vernacular derivatives e.g. Therocephalia, but therocephalian Punctuation should be loose, omitting all not strictly necessary Reference to the author should be expressed in the third person Roman numerals should be converted to arabic, except when forming part of the title of a ' book or article, such as ‘Revision of the Crustacea. Part VIII. 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REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: “Smith (1969) describes...’ ‘Smith (1969: 36, fig. 16) describes . “As described (Smith 1969a, 19696; oe me “As described (Haughton & Broom ie Vi ‘As described (Haughton ef al. 1927) . Note: no comma separating name BEN year Dagination indicated by colon, not p. names of joint authors connected by ampersand - et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc. to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (abbreviated according to the World list o, scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number (only if independently paged) in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) BuLLouGu, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan. FISCHER, P. —H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140. FiscHer, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs Zool. exp. gén. 74: 627-634. Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320. Konan, A. J. 19606. Spawning behaviour, cee. masses and larval development in Conus from the Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): THIELE, J. 1910. Mollusca: B. Bele ahbee: Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270, Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. (continued inside back cover) ANNALS OF THE SOUTH AFRICAN MUSEUM ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM Volume 81 Band June 1980 Junie Part dl Deel ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS FROM UPPER PLEISTOCENE AND HOLOCENE SITES IN SOUTHERN AFRICA By RICHARD G. KLEIN Cape Town Kaapstad The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000 Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na gelang van die heskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000 OUT OF PRINT/UIT DRUK 1, 2(1-3, 5-8), 3(1-2, 4-5, 8, t.—p.i.), 5(1-3, 5, 7-9), 6(1, t.—p.i.), 7(1-4), 8, 9(1-2, 7), 10(1-3), 11(1-2, 5, 7, t.—p.i.), 15(4-5), 24(2), 27, 31(1-3), 32(5), 33 Copyright enquiries to the South African Museum Kopieregnavrae aan die Suid-Afrikaanse Museum ISBN 0 908407 91 2 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press, Pty., Ltd., ' Die Rustica-pers, Edms., Bpk., Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS FROM UPPER PLEISTOCENE AND HOLOCENE SITES IN SOUTHERN AFRICA By RICHARD G. KLEIN Department of Anthropology, University of Chicago (With 5 figures, 7 tables and 1 appendix) [MS. accepted 13 December 1979] ABSTRACT There are now more than seventy-five Upper Pleistocene and Holocene localities in southern Africa that have provided analysable remains of large mammals. The purpose of this paper is to summarize the information these remains have provided on past environments, on the evolution of man-environment relationships, and on the ecology and demise of extinct species. For the purposes of discussion, the fossiliferous Upper Pleistocene and Holocene sites are divided among six modern southern African ecozones, distinguished from one another on climatic, phytogeographic, and zoogeographic grounds. Changes in large mammal distributions or in species frequencies that probably reflect Upper Pleistocene and Holocene environmental change can be demonstrated in all six ecozones, but a well-defined pattern of change in mammalian faunas that can be correlated with a pattern of long-term environmental change established on other grounds can be demonstrated in only one ecozone. This is the Cape Zone, where cooler intervals during the Upper Pleistocene repeatedly witnessed an increase in grazing ungulates relative to browsers. In part, the failure of comparable patterns to emerge in other zones may reflect the fact that Upper Pleistocene environmental and faunal change was greater in the Cape Zone than elsewhere, but in large part it almost certainly reflects the better overall quality of data from the Cape—more well-dated sites and more relatively large faunal assemblages for which detailed numerical data are available. The greater quantity and higher quality of data from the Cape also make it the only ecozone in which there is a substantive basis to discuss: (i) long-term changes in human ability to obtain large mammals; and (ii) the reasons for the disappearance of several large mammal species which were common in various parts of southern Africa during the Upper Pleistocene. The Cape data suggest: (i) that Middle Stone Age people were less proficient hunter—gatherers than their Later Stone Age successors; and (ii) that a combination of environmental change and the greater hunting proficiency of Later Stone Age peoples was responsible for the large mammal extinctions. CONTENTS PAGE Aims and basic definitions : : 2 : F 5 5 Hs Materials and problems . : renee Cuaeaice! nay 47) Mammalian evidence for enviropmental change : . BPI The Zambesian Ecozone . i : : ‘ i an) eP2BY2 The Transvaalian Ecozone ; ; : j ; 236 The Kalaharian Ecozone . : d ; : : a) (BAY The Basutolian Ecozone . ‘ : 5 ‘ fo DAS The Karoo—Namaqualian Ecozone : : ‘ . 246 The Cape Ecozone . : 5 : ; ; : . 250 Conclusions. : : ‘ : > 259 Mammalian evidence for cultural change : ; 5 . 260 223 Ann. S. Afr. Mus. 81 (7), 1980: 223-283, 5 figs, 7 tables, 1 appendix. i) i) SG ANNALS OF THE SOUTH AFRICAN MUSEUM PAGE The ecology and demise of extinct species . . . . 263 Equus capensis . . . . ee TE Lae gtk 264, Metridiochoerus sp... Sib he aah a ae cae OS Pelorovis antiquus . j ‘ - ; : 3 . 268 Megalotragus priscus ; a : ‘ ‘ ‘ . 269 Antidorcas bondi j ; i i ; : ; “1 PATA) Antidorcas australis . ; . ; ‘ ; : nu AL The causes of extinction . F 4 5 ‘ ; se era Conclusions Sealed cate | ana es Bt eters ae Acknowledgements . é , i : ‘ 3 , . 2714 References . i : i ‘ ; i : ; : rue 2 Appendix. Vernacular and scientific names of the extant mammalian species mentioned in the text . 5 Be Aye AIMS AND BASIC DEFINITIONS As a result of research undertaken mainly in the 1960s and 1970s, there are now more than seventy-five Upper Pleistocene and Holocene localities in southern Africa that have provided analysable remains of large mammals. This paper aims to summarize the information these remains have provided on past environments, on the evolution of man—environment relationships, and on the ecology and demise of extinct species. For the purposes of this paper, southern Africa is defined relatively broadly to include the Zambesian, Transvaalian, Kalaharian, Basutolian, Karoo— Namaqualian, and Cape Ecozones of Devred, as presented in De Vos (1975) (Fig. 1 here). The ecozones themselves are defined on the basis of climatic, phytogeographic, and zoogeographic features which are summarized below (based mainly on information in De Vos (1975) and Brown (1965)). The zones are of course abstractions in the sense that they grade into one another rather than being sharply bounded, and within each there is important climatic, phytogeographic, and zoogeographic variation, as determined for example by great differences in altitude, subsurface drainage, or proximity to permanent water. However, the zones clearly reflect gross differences in the historic distri- bution and relative abundance of large mammal species and therefore have definite value as a first basis for gauging the palaeoenvironmental significance of fossil mammal faunas. In modern political terms, southern Africa, as defined here, comprises especially the countries of Angola, Zambia, Malawi, Mozambique, Zimbabwe Rhodesia, Botswana, South West Africa (Namibia), South Africa, Lesotho, and Swaziland. The approximate locations of the principal fossiliferous Upper Pleistocene and Holocene sites within these countries are shown in Figure 2, from which it is clear that the overwhelming majority are located in South Africa, South West Africa, Zimbabwe Rhodesia, and Zambia. In general, blank spots on the map reflect the absence of interested researchers as much or more than any other factor. Undoubtedly, many pertinent sites especially wait to be discovered or excavated in Angola, Botswana, Mozambique, and Malawi. For the purposes of this paper, the Upper Pleistocene and Holocene have been taken to comprise the time interval represented by stages 1 through 5 of the \ Burchell's eland oes ees ay zebra Pr Burcnel ‘s ‘ he } ere Ae, me KALAHARIAN is ; gemsbok gf 3] we ~~ ee KAROO- i MAN } eas Ted Fa ingbok blesbok Xx hartebeest if southern roedbuck : Se a melt bonte- elond bie red antelope artebeest blue Ryle Lied ie bac) a lgrysbok ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS t t ; hy xy GUINEAN \ - ea Tho a= okap/ wo (aN ee bushbuck ae ‘ ae a eo -—- {7-~~ Seer \ \ eed \ h H Wer Spee grate ay \/ So { —/ | \ \ ZAMBESIAN \ greater | kudu 7 / / { / Sharpe's . / grysbok i | | pola /echwe Lichtenstein's harteboest 4D il bushbuck i Wee TS, eland iy) Burchall's | TRANSVAALIAN ee Se ae N eland ~~ = =~ Isessebe | ape a gre) ue wie beest “Bing? wilde duiker == Goering } YN we Ny 5 jhe worthog \ eest | (OMe seater \ oN) ye a EE os \ Ce) buffalo / yo ( stoenbok worer- buck myn hs EN zebra pay ne a te ae buffalo 225 EASTERN Grant's gazelle mm son’s gorelle blue wildebeest buffalo waterbuck ZAMBESIAN ortb/ Fig. 1. Southern African ecozones, with the most prominent suid, bovid, and equid species found in each. The outlines and names of the zones are from De Vos (1975). Information on species distributions was obtained primarily from Ansell (1971a, 19715), Ellerman er al. (1953), Joubert & Mostert (1975), Smithers (1966, 1971), and Smithers & Lobao Tello (1976). 226 ANNALS OF THE SOUTH AFRICAN MUSEUM deep-sea core oxygen-isotope stratigraphy, that is, approximately the last 130 000 years. These stages are listed in Table 1, along with the temperature conditions they are believed to reflect and their dates, in so far as these have been established. In conventional terms, stage 1 may be equated with the Holocene, stages 2-4 with the ‘Last Glacial’, and stage 5 with the ‘Last Interglacial’, though some authorities would prefer to restrict the Last Interglacial to sub- stage Se, the only part of stage 5 that compared in overall warmth with the Holocene. In so far as it is possible below, the oxygen-isotope stage numbers are used in preference to the terms ‘Last Interglacial’ and ‘Last Glacial’, since the stages more fully reflect the true complexity of Upper Pleistocene and Holocene climatic change, particularly in the latitudes of southern Africa (roughly 10°S to 35°S). ‘Large mammal’ has been defined very loosely to include all mammalian species in which adults weigh at least 0,7-0,9 kg. The principal creatures excluded by this definition are bats, insectivores (except hedgehogs), and rodents (except primarily springhare, cane rats, the largest of the mole rats, and, of course, porcupines). The rationale for excluding ‘small mammals’ is that this paper is mainly concerned with mammalian remains as reflections of past human ecology, and it seems unlikely that people have ever systematically exploited mammals weighing less than 0,7-0,9 kg. Circumstantially, this proposition is supported by the fact that where small mammal remains have been found in large quantities in southern African sites (for example, at Redcliff Cave, Wonderwerk Cave, Die Kelders Cave 1, Byneskranskop Cave 1, Boomplaas Cave A, and Nelson Bay Cave), they are clearly concentrated in layers where artefacts and bones of large mammals are rare. This suggests that the small mammals were TABLE 1 Upper Pleistocene oxygen-isotope stages defined in deep-sea cores (based mainly on Shackleton & Opdyke (1973, 1976)). Approximate years B.P. Oxygen-isotope stage Inferred world climate (0 ee ence eer FS ie 1 very warm 12000 - - - -- -- - --- = -- ---------- 2 very cold 32000 - - - -- - -- —- -- -- -- --------- 3 cold with warmer oscillations 64000 —- - -- - - ------ ------------ 4 very cold 75000 - - ---------------------- Sa warm 92 000 5b cold Se warm 109 000 5d cold Se very warm ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS 227 brought in mainly by predatory birds who occupied the sites when people were absent. Of course, even though (or in part because) small mammal remains are generally not a product of human activity, they constitute a potentially valuable source of information about past environments, and their value in this regard is enhanced by the fact that it is often possible to obtain very large samples from relatively small excavations. Pertinent examples of palaeoenvironmentally oriented small mammal studies in southern Africa are those of Brain (1974a; Brain & Brain 1977) in the Namib Desert, and of Avery (1977 and in prep.) in the southern Cape Province. Brain has pointed out that fluctuations in the abundance of the principal species represented in the relatively simple micro- faunas of the Namib can be used to monitor past changes in the amount of grass cover and of sand v. gravel in the vicinity of a site. In the more complex situation of the southern Cape, with a wider variety of well-represented microfaunal species, Avery is using sophisticated statistical procedures to detect relatively subtle changes in microfaunal communities, with the goal of checking and amplifying inferences on Upper Pleistocene and Holocene environmental change drawn from parallel studies of large mammal bones, palaeobotanical remains, and sediments. MATERIALS AND PROBLEMS The basic materials on which this paper is based are lists of large mammal species reported from Upper Pleistocene and Holocene sites in southern Africa. Both archaeological ana uon-archaeological sites have been surveyed, but among the archaeological ones, the focus is almost exclusively on sites occupied by Stone Age (v. Iron Age) people. People making stone artefacts were the only human occupants of southern Africa during the Pleistocene and most of the Holocene, and they persisted into the historic period over much of the sub- continent, especially in the Cape, Karoo—Namaqualian, and large parts of the Basutolian and Kalaharian Ecozones. In the Zambesian and Transvaalian Zones, Stone Age people were pro- gressively displaced by Iron Age agriculturists, beginning in the first centuries A.D. The Iron Age people were immigrants from further north, who subsequently also penetrated into those parts of the Basutolian and Kalaharian regions where their system of mixed farming was practicable. Iron Age faunas have been analysed from sites in Zambia by Fagan (1967; also Fagan et al. 1969), in Malawi by Voigt (1970, 1973, 1977), in Rhodesia by Brain (19745) and Huffman (1974, 1975), in Botswana by Welbourne (1975), in the Transvaal by Voigt (1978) and Welbourne (1971, 1973), in Natal by Klein (as reported in Maggs & Michael 1976), and in the Orange Free State by Maggs (1975), but for the purposes of this paper the utility of the samples is limited, because most of them are small and they tend to be dominated by introduced domesticates (cattle, sheep and/or goats). However, the Iron Age lists have been scanned carefully for evidence they may contain on the distribution of indigenous mammals in relatively recent 228 ANNALS OF THE SOUTH AFRICAN MUSEUM \ \ 1 K “ on cant -—— Secor } Peas Vex \ / Se \ RSs H \ Sst N | \ *Fingira \ ! ? 1 ) H ( t “ ia NY ar ( { eer eae oe nN ~ \ \ U 1 ~~/\ \ eNachikufu | 1 1 Sa, NO / \ 1 ~ ¢ » \ 1 Kalemba , ‘ BeeeF \-s) Thandwe | 4 Bee \ Makwee* S. Chencherare . - \ *Broken Hill ==" amy \ | Mumbwa ae | | | Leopard's Hill | } | | Twin Riverse ever ( 7 1 a ‘i a 3 Se \ 2 i Gwisho W = a ny 1 ~ 2 =X oa 1 PS \ Vi \ AS Sa yn eee 7 / SS SQ Ssoe= Sess Inyangae ‘ er ~ I c N 7 NI . le \ *Redcliff 1 gi \ i a SSK Chelmer | By /awayo Waterworks J 1 \ Pomongwe / 1 Ney lA H 1 / e@Frandberg Sites 1 AS Se HH eFackeltrager ool YS ye G . Ald BEEN Gig Elephant & Striped | -- A Giraffe Shelters 2 BA Kalkbank. \ O \ Gobabis 1 Ve Fs \ ! 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These are the creatures which were predominant in most parts of the Zambesian Zone historically, suggesting definite limits to overall Pleistocene and Holocene environmental change, particularly as compared to the Cape Zone (discussed below) in which historically abundant mammals disappeared or became very rare on several occasions during the same interval. Where there is deviation from the general pattern of dominance by zebra, warthog, and alcelaphines, it is toward an emphasis on other large grazing ungulates whose local abundance was predictable from the location and dating of a site. Thus, lechwe and buffalo are especially well represented at the late mid-Holocene (‘isotope stage 1, late’) spring sites at Gwisho, located on the margin of the Kafue floodplain on which these creatures were quite common historically (large herds of lechwe still occur near by). Zebra, warthog, blue wildebeest, and impala are also well represented in the Gwisho sites, presumably reflecting their prominence in the typical miombo woodland that flanks the Kafue floodplain at Gwisho. The long-term persistence of adjoining Kafue floodplain and miombo woodland habitats is clearly suggested by the much earlier (Middle Stone Age) fauna from Twin Rivers Kopje, which, like the Gwisho sites, is located on the margin of the Kafue floodplain, and which has also provided a fauna in which lechwe, wildebeest, and zebra are abundant, although precise numerical estimates are not available. A further site in which creatures favouring near-water situations are very common is Kalemba Rock Shelter which has provided a relatively large number of bushpig and waterbuck remains. It is not clear that these creatures (at least waterbuck) were so abundant near Kalemba historically, and since their remains come principally from layers that probably belong in isotope stage 3 or 4, they may reflect once moister conditions near by. The best evidence for environmental change in the Zambesian Zone, as reflected in mammalian fauna, comes from Redcliff Cave, where three species are present which did not occur in the Zambesian region in historic times, and which are so far unknown in any fauna clearly postdating isotope stage 2. These species are the blesbok, the common springbok, and the mountain reedbuck. In an earlier publication on the Redcliff fauna, the writer suggested that the ‘Tshangula’ industry at the site was a late Middle Stone Age manifestation, similar perhaps to the Howieson’s Poort variant of the Middle Stone Age further south (Klein 1978a). However, with the appearance now of a more complete report on the artefacts (C. K. Cooke 1978), the writer believes the ‘Tshangula’ industry is more likely to be an early Later Stone Age variant, dating to between 30 000 and 20 000 years B.P., as is in fact suggested by the single available radiocarbon date. This would place the ‘Tshangula’ industry and as- sociated fauna in isotope stage 2. The underlying Bambata levels, which are undoubtedly Middle Stone Age, are clearly beyond the range of radiocarbon dating, and their placement in one 236 ANNALS OF THE SOUTH AFRICAN MUSEUM or another isotope stage is not straightforward. However, Brain’s (1969a) analysis of the Redcliff sediments is helpful in this regard. This shows that CaCO, concentration is relatively low and the matrix is relatively coarse in the earlier Bambata and especially in the Tshangula levels, perhaps reflecting stronger water flushing of the deposits, in turn reflecting moister climate. During accumulation of the intervening later Bambata levels, with a higher CaCO, content and finer matrix, conditions may have been generally drier. If the Tshangula horizons are properly placed in isotope stage 2 and this was a relatively moist time near Redcliff, it seems most reasonable to place the earlier Bambata horizons, indicating comparable moistness, in the next oldest isotope stage that was generally comparable to ‘2’ world-wide. This would be ‘4’ (= early ‘Last Glacial’). The intervening later Bambata levels, with their sedimentologic evidence for relative dryness, would then date from stage 3, — during which world climates were generally less different from present ones than during stages 2 and 4. It is interesting in this context that the three ‘exotic’ species found at Redcliff are significantly more common in the Tshangula and earlier Bambata levels than in the later Bambata ones, suggesting that these species were locally most abundant during relatively moist intervals. All three presumably extended their ranges to Redcliff from regions much further south where they were common historically. Range extension Equatorwards (pre- sumably as a result of vegetational change) might itself be taken as evidence for cooler conditions, moister conditions, or both. The presence of common springbok in the Zambesian Zone is confirmed for stage 2 at Leopard’s Hill Cave, where both radiocarbon dates and well-described associated (early Later Stone Age) artefacts leave no doubt about stage place- ment, although the overall Leopard’s Hill faunal sample is too small for detailed palaeoenvironmental interpretation. There is no evidence that springbok survived into stage 1 (the Holocene) in the Zambesian region, but during this interval, a close east African relative of the springbok, the Thomson’s gazelle, apparently penetrated the Zambesian Zone at least as far south as Kalemba Rock Shelter and perhaps also into Malawi, where a possible gazelle has been recorded in Iron Age sites (Voigt 1973) and in the broadly contemporaneous Later Stone Age deposits of Chencherere Rock Shelter II (Crader n.d.). In fact, the ‘tommie’ may have extended through the eastern portion of the Zambesian Zone into the Transvaalian one, if the writer’s tentative identification of material from the Iron Age site of Ntshekane in the Tugela Basin of Natal is correct (Maggs & Michael 1976). Why the species did not occur in the Zambesian and Transvaalian Zones historically is not clear, but perhaps its absence is related in some way to the introduction and proliferation of domestic stock in Iron Age times. THE TRANSVAALIAN ECOZONE On the southern margin of the Zambesian Zone, mopane woodland tends to give way to shrubby acacia steppe or bushveld in a semi-arid setting. This ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS 237) semi-arid country, running more or less across the continent, constitutes the Transvaalian Ecozone. As in the Zambesian Zone, rainfall is restricted almost entirely to summer, but the average is generally less than 500 mm/a, and there are great differences in total amount from year to year. Of all the ecozones considered here, the Transvaalian one is the least satisfactory as a unit. It might well be better to consider it as three zones: (i) a western one, comprising the northern Transvaal, southern Zimbabwe Rhodesia, northern Botswana, northern South West Africa, and southern Angola; (ii) an eastern one, comprising the eastern Transvaal, adjacent south- western Mozambique, and eastern Swaziland; and (iii) a narrow southern extension comprising the Natal and south-eastern Cape coastal strips and their immediate hinterlands as far south-west as Port Elizabeth. The western portion of the Transvaalian Zone, from the northern Transvaal westwards, is the driest part. The vegetation is typically grassland with inter- spersed shrubby trees, among which acacias are often most prominent. In a sense, the area is not so much a distinct ecozone as a transitional region between the Zambesian Zone to the north and the Kalaharian one to the south. Grazers (including blue wildebeest, Cape hartebeest, tsessebe, Cape buffalo, springbok, gemsbok, Burchell’s zebra, and warthog) are most common, but browsers (greater kudu, bushbuck, giraffe, black rhinoceros ef al.) and mixed feeders (eland and impala) are also prominent. Some of the large grazers incorporate parts of the Kalaharian Zone in their seasonal movements, and the transition to the Kalaharian region is clearly indicated by the presence of both gemsbok and springbok. The eastern portion of the Transvaalian Zone, known in South Africa as the ‘eastern Lowveld’, is moister than the western. Bush and tree cover is much denser, and the term ‘bushveld’ is clearly appropriate. The overall variety of large mammal species is basically the same as to the west, but browsers and mixed feeders are more prominent numerically, as are grazers that prefer more wooded country (roan antelope, sable antelope, and buffalo). Springbok and gemsbok are absent. The southern extension of the zone is covered by subtropical thornbush and scrub-forest in a subhumid rather than semi-arid setting. In the north, the fauna is very similar to that of the eastern Lowveld. Further to the south, beyond Zululand (KwaZulu), many of the Lowveld species drop out, and the fauna becomes essentially indistinguishable from that of the adjacent part of the Cape Ecozone. Geomorphic evidence of former very large lakes in the Makarikari and Makgadikgadi Depressions (Street & Grove 1976; Grey & Cooke 1977), as well as interstratified evaporites and aeolian sands in a cave in the Kwihabe Hills of northern Botswana (Grey & Cooke 1977) demonstrate alternation between wetter and drier periods during the Upper Pleistocene and Holocene in the western part of the Transvaalian Zone. Data presented by Heine (1978) suggest that conditions were particularly wet 30 000 to 18 000 and again 11 000 years 238 ANNALS OF THE SOUTH AFRICAN MUSEUM ago, with arid conditions in between. If this is correct, then long-term pre- cipitation trends in the western part of the Transvaalian Zone may have been. out of phase with those immediately to the south in the Kalaharian Zone, where much of the interval between 18 000 and 11 000 B.P. appears to have been very wet (see Heine 1978 and below). Levels of éboulis secs formed by frost weathering in Bushman Rock Shelter and Border Cave in the eastern part of the Transvaalian Zone (Butzer, Beaumont & Vogel 1978) document the intervals of much reduced Upper Pleistocene temperatures apparent in the deep-sea record. The sedimentary fills are not so informative about past precipitation changes, but it appears that there was no one-to-one correspondence between cooler and wetter periods. Sedimentation rates extrapolated from the radiocarbon-dated portion of the Border Cave sequence provide a basis for correlating the temperature fluctuations it records with ones established in the global marine record. The extent of temperature depression involved has not been established, but to the north, at Wolkberg Cave in the north-central Transvaal, the oxygen-isotope ratios of Upper Pleistocene cave carbonates have been used to suggest very tentatively that average tempera- tures during cold episodes were as much as 8,5-9°C below present ones (Talma et al. 1974). Pollen recovered from peat deposits at Wonderkrater near Naboomspruit in the central Transvaal, indicates that, during at least some cooler phases, Transvaalian bushveld was replaced by open grassveld (Scott & Vogel 1978). It was presumably this kind of vegetational change which encouraged the spread of springbok and blesbok to the Cave of Hearths and Kalkbank, as discussed below. The Upper Pleistocene and Holocene sites in the Transvaalian Zone that have provided remains of large mammals are listed in Table 3, along with the probable or suggested correlation of sites (or levels within them) with various oxygen-isotope stages. The Transvaalian Zone is second only to the Cape Zone in the number of sites correlated with various stages, but the available faunal samples are mainly small, incompletely described, or both. The most dramatic changes through time in large mammal species frequencies have been recorded at Melkhoutboom Cave, located at the extreme south-western margin of the Transvaalian Zone, in an area that is transitional to the Cape Zone. Historically, the vegetation near Melkhoutboom was dominated by forest, closed bush, and sclerophyllous scrub, and the most common large mammals were various browsing ungulates—notably bushbuck, kudu, blue duiker, grysbok, and bushpig. The only prominent grazer was the Cape buffalo, which, in spite of its dietary preferences, is known to be very much at home in closed, bushy environmental settings. The deposits at Melkhoutboom have provided a semi-continuous series of large mammal assemblages dating from approximately 15 400 to 2 000 years ago. The principal species in deposits dated to younger than 7600 B.P. are the historically prominent browsers and the buffalo. However, in deposits older than ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS 239 7 600 years, and particularly in ones older than 10 500 years, the browsers and buffalo are rare or absent, and the fauna is dominated by alcelaphine antelopes (black wildebeest, Cape hartebeest, blesbok/bontebok) and equid (either mountain zebra or quagga or both). This assemblage is more reminiscent of the fauna of the Basutolian Zone to the north than of the Transvaalian one in which Melkhoutboom is presently located. More generally, the pre-10 000 B.P. fauna from Melkhoutboom clearly suggests that the environs of the site were grassier in the terminal Pleistocene (late stage 2) than during most of the Holocene (stage 1). The same sort of vegetational change—from grassier to bushier or scrubbier—is reflected in faunal change within late Pleistocene to Holocene sequences at several sites in the near-by Cape Zone, as discussed below. The fact that the faunal change at Melkhoutboom and in the Cape Zone sites is so clear-cut probably reflects their position near the southern margin of the continent where Upper Pleistocene and Holocene environmental changes were perhaps more dramatic than in many areas nearer the Equator. The basal levels at both Wilton and Uniondale, located not far east of Melkhoutboom, also perhaps date from a time (very early Holocene or terminal Pleistocene) when large grazers were relatively common near-by, but the faunal samples are far too small to document this. At both sites, the bulk of the fauna comes from mid to late Holocene levels, and the principal species represented are the same ones that dominate the contemporaneous deposits at Melkhoutboom— bushpig, bushbuck, duiker, grysbok or steenbok, and Cape buffalo. An earlier Upper Pleistocene interval broadly comparable to the terminal Pleistocene at Melkhoutboom may be reflected in the fauna from Aloes, also in the transitional area between the Transvaalian and Cape Zones. Land-snail shells associated with the bones at Aloes yielded a radiocarbon age of greater than 37 000 years. The relatively large sample has provided no identifiable bones of browsers, though large grazers (quagga, ‘giant Cape horse’, wildebeest, common springbok, and warthog) are well represented. In the heartland of the Transvaalian Zone, far to the north, the available evidence suggests that faunal change related to more general environmental change was more subtle than at Melkhoutboom, but such change did occur. This is particularly clear at Border Cave, which has provided analysable faunal remains from deposits correlated with deep-sea isotope stages 5d through 3. Using the number of squares in which bones of a species occur as an index of its abundance in each Border Cave level, the writer has been able to show that levels in which bushpig, Cape buffalo, tragelaphine antelopes (kudu, nyala, bushbuck, and eland), and impala are relatively common, alternate with levels in which warthog, Burchell’s zebra, and alcelaphine antelopes are more prominent. Warthog, zebra, and alcelaphines were probably more common near by in historic times, and the writer has suggested that levels in which bushpig and buffalo are relatively abundant reflect Upper Pleistocene intervals in which the vegetation contained more bush than in recent times. This conclusion is supported by analysis of the sediments (Butzer, Beaumont & Vogel 1978), VSW pue VST VST VST VST VSI VST VSI VST VSI VST VSI VST VST VSI VSI VSI VST VST ANNALS OF THE SOUTH AFRICAN MUSEUM SNOLLVIDOSSY TVYOIDOTOAVHOUV (LL61) UPI “DU (LL61) UIA “DU uPplTy “Da We) “Td Y UPI Da Bury “f up “Oa (vL61) uosdwieg ul poiodal sv ulelg yO up OD a urelg “IO uP OD a SUIYOIIY “f Bnfd “I $(96961) Weg “AO uLpTy DU ulvig “SIO oumogeM *D “A ANY AI ‘NOILVOIIENd JLvVuvdas GNYV YNNVA ONIAALLNAGI NOsudd (8261) 930A 29 yuownvog “19z1nNg ‘(8/6]) 72 72 JuOUNog (8L6T) 930A yuouInesg “1ozjng ‘ (8/61) 72 72 yuounvog (9L61) uooeed “f “H (SL61) Sole (@ecisd core) *slod) yuounvag ‘(TL6]) SIeIe 2 [e30A (‘wu0d ‘sisd) Joyoolg “WH (€961) 24909 “IO (0 ‘G1ISd Z) eed Jopi0og Cd‘ 000 $9-000 SZ *2) ¢ a8vIS' adojos] (VITSAZ—-ATSAT) PARD Jopsog Cd‘ 000 ZE-000 #9 *9) € adds adojosy (q [9A9]) Woog noYy[a AL (uolyedn550 JeMO]) TUaMSUOYS ({ JoAe]) sueTysoussuruneyy (d“ 000 ZI-000 ZE *?) Z asvig adojosy ojepuoiy (S[SAQ] UOIIAA PUe UPMSUOWOd) sMsuOWIOg (d'@ 0-000 ZI °2) paiwiuasaffipun ‘7 asvjg' adojosy (961) wove ‘f “H (CL61) Uoseved “ft (‘Wwu0d ‘siod) JUOWNeEg (‘WIUI0D *siod) yuounveg ‘(TL6]) SlereW] 2 [930A (qndun) 3njq (6961) HOI * (6961) MnOT (9261) uooeed ‘f “H (CL61) UOoRe ‘f (LL61) }4NOSTAIEq (‘WIIOD “sied) MeYs}19qoOxY (SL61) seed (WIUIOS ‘sied) JUOWUNeEg (‘wiuw0d ‘sied) yUoWNvog aOUNOS TvaaNAS (AU-AIN S[PAe]) Woogynoy ya (P-DE SIPA2]) WOITAA Pinel (Z “€ Siohv]) SUBIYSOUSSUTUNSFT Jayays yoy uewysng (d’@ 000 S-000 ZI *?) 4]4va ‘T asvig adojosy (GINO “AVO S[PA9]) Woog noyy[s] (AE-T S[SA9]) WOT (LV) Meys uuy MOIAIIC-T (uonednsso Jaddn) tuasmMsuoys Tuaqequig yuu Cd 0-000 § °2) ain] ‘[ asvIS' adojosy aLis “QU0ZODq ULI[PRASULI] SY) Ul S[EWUIEUT ode] JO SUIeUIOI poprAoId oALY Jey} So}Is QUSDO[OH{ pue oUed0}sIE[q Jedd p) 240 € a1avL 241 ~”a a < = = < = w 16) % < 4 a ° wn Zi ie) A < e =| A 2 4 < oO _ le) [e} = e) oO a Qa v4 <_< c Z a = (4 ° 4 > Zz a euou VSW VSW auou 9suou VSI VSW VSN VSI VSN SNOILVIOOSSV TVOISOTOAUVHOUVY (POL6T) SIP “H “I (2961) 94909 ‘§ “dH (TL61) SUINOgPA *D “A ‘(Z961) 24909 'S “‘@ “H uoyysney *§ 7 A21|P9Z “A “AV (€961) 24009 *§ ‘A °H :(1S61) SIM ‘H “1 24909 ‘S ‘dH (pL61) uosdwieg ul pojiodol se ulelg “yy ‘O (LL61) USI “DU (LL61) UPI “DU (LL61) UPI “DU 8nd “I (96961) WeIg “I'D ANV dI ‘NOILVOIIdNd ALVUVdAS GNY VNOVa ONIAALLNAGI Nosaddd (6961) sseD (7961) uoseW (8561) 72 12 UoSseI *(L96T 7961) UOSRIN (9161) Ae[e9Z (IS61) Slowing 2 puog (€961) 24909 “I'D sooly SUJIPOFT JO OARS yueqg yey SYIOMIOVeAA OAPMEING JowyjeyO (SIOAP] WSJ) OMSUOWOg Cd’ 000 2-000 8ZI *2) paroijuasaffipun “¢ so/puv “p ‘¢ sasnjg adojosy (8261) [930A 29 yuowmnrog “1ozing ‘(8/6]) 72 72 JUOUINvOg (8L61) 1930, 29 quOUINvog “IozIng °(8/6]) 72 42 yuOUNLSg (8L61) 980A 29 quounvog ‘1ozjng ‘(8L61) 72 42 yuouNvog (‘doid ul) Jezjng -(qndun) 3njd *(696T) HOI (6961) AnoT (aq ‘dngg5]) sAvD Jopi0g PG asvjS adojosy (VME “ITSAE) PAD Jepiog q¢ aspjs adojosyT (dng@eé “VA\Z) 2A2D Jopiog DG asvjs adojosy (S[SA9T WS 197k]) YOoY, UewYysng Cd“ 000 ZE-000 SZ *2) paioijuadaffipun “p puv ¢ sasoig adojosy qaOUNOS TyyudNao aLis 242 ANNALS OF THE SOUTH AFRICAN MUSEUM which establishes a general correspondence between apparently colder episodes, as reflected in sediments, and “bushier’ fauna. Unfortunately, the faunal samples | from various levels are too small for a truly detailed study of the relationship between changes in sediment parameters and species frequencies. Yet further north, environments different at one or more times during the Upper Pleistocene from modern ones are probably implied by the presence of common springbok and of blesbok/bontebok at the Cave of Hearths and especially at Kalkbank. Both sites are located north of the areas in which these species were distributed in historic times (Kettlitz 1962). In the vicinity of both sites, the impala apparently fills the niche occupied by the springbok elsewhere, yet in the fossil fauna from Kalkbank, for which species frequency estimates are available, springbok is actually more common than impala. The occurrence of springbok and of bontebok/blesbok in earlier Upper Pleistocene contexts at the Cave of Hearths and Kalkbank was perhaps to be expected in view of their presence in broadly contemporaneous deposits at Redcliff yet further north, and probably reflects the same vegetational change (less bush, more grass) as at Redcliff. THE KALAHARIAN ECOZONE This ecozone corresponds broadly to that part of the southern African interior plateau that is often known as the Kalahari desert, though it is not really a desert in either climatic or vegetational terms. With regard to modern political units, the Kalaharian Zone covers eastern South West Africa, the western two-thirds of Botswana, and a large portion of the adjacent (northern) Cape Province of South Africa. Rainfall in the Kalaharian Zone is erratic and comes almost entirely in summer. The average decreases from roughly 500 mm/a in the north-west to as little as 200 mm in the south-west. Highly porous, sandy soils soak up rainfall rapidly, so that surface water is relatively rare, even in areas where the average rainfall is fairly high. Vegetation cover is highly variable, from relatively luxuriant acacia savanna with an important grass component in the better watered parts (particularly in the north) to sparse shrub acacia savanna and bushveld in the more arid parts (particularly in the south). At least historically, the grass cover over much of the Kalaharian region was sufficient to support fair numbers of gregarious, migratory grazing ungulates, particularly ones which are capable of obtaining their moisture requirements largely from plants. Springbok and gemsbok were especially common, and blue wildebeest, hartebeest, Burchell’s zebra, and warthog were at least locally abundant. Browsing animals, including especially giraffe and greater kudu, were generally less common. Among mixed feeders, eland and steenbok were wide- spread. Bushbuck (a browser), Cape buffalo and roan antelope (grazers), and impala (a mixed feeder) occurred in some areas of denser bush. Among non- ungulate herbivores, hare(s), springhare, porcupine, baboon, and rock hyrax were (and in some cases still are) widespread and abundant. The principal ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS 243 carnivores preying on these creatures or scavenging on their carcasses were lion, leopard, cheetah, brown hyena, spotted hyena, Cape hunting dog, and jackals. Upper Pleistocene and Holocene environmental change within the Kalaharian Zone has been best documented by geomorphic research at its south-eastern margin, particularly along the Gaap Escarpment in the northern Cape Province (Butzer, Stuckenrath et al. 1978). Alternation of subhumid and semi-arid climatic phases is apparent in the Gaap sequence, with the earliest radiocarbon-dated subhumid phase fixed between >21 000 and 14000 years ago. Subsequent subhumid phases are dated between 9 700 and 6 500 B.P. and between 4 500 and 400 B.P. Long-term fluctuations in precipitation appear to be broadly in phase with those recorded in the Basutolian Zone to the east, but not necessarily with those in the Transvaalian Zone to the north (see above and Heine 1978). Beds of cryoclastic rubble document several past episodes of relatively intense cold along the Gaap Escarpment and presumably throughout the Kalaharian Zone. The most recent very cold interval clearly coincided with deep-sea stage 2. The greatest cold appears to have preceded and followed the marked subhumid phase between >21 000 and 14 000 B.P., indicating that the relationship between past temperature and precipitation change was a complex one. The Upper Pleistocene and Holocene sites in the Kalaharian Zone which have provided remains of large mammals are listed in Table 4, along with the probable correlation of the sites (or of levels within them) with various oxygen- isotope stages. Most of the sites are located on the south-eastern margin and date from the late Holocene. The late Holocene faunal samples are dominated by hare(s), springhare, rock hyrax, Burchell’s zebra or quagga, warthog, black wildebeest and/or Cape hartebeest, springbok, and steenbok. The mountain reedbuck is also relatively common, reflecting the relatively rugged topography surrounding many of the sites. In no case is there any clear suggestion of an environmental setting that differed significantly from the historic one. With the important exception of the fauna from Equus Cave, which has been sorted and identified, but not yet analysed, the faunas from Kalaharian sites of Pleistocene age are either small, poorly excavated, incompletely reported, or all three. Most of them are also very imprecisely dated. All this makes it difficult to assess their palaeoenvironmental significance. However, the occur- rence of blesbok/bontebok at Black Earth Cave, Witkrans Cave, and especially at Gobabis far to the north-west of the species’ historic range, suggests a vegetation cover in which perennial grasses may have played a greater role than they did historically in the Kalaharian region. The occurrence of vaalribbok at Black Earth Cave, Ochre Cave, and Boetsap may have broadly similar implica- tions, depending upon how common it was in the deposits (the samples presently available for study are all highly selected and therefore not suitable for estab- lishing species frequencies). The vaalribbok occurred near all the sites in historic times, but was not especially common. It has so far not been recorded in local VST VSI VST VST VSI VST VSTI VST VSI VST VST ANNALS OF THE SOUTH AFRICAN MUSEUM SNOLLVIOOSSY TVOIDOTOAVHOUV 244 suINOGPM “D W (TL6T) 2UINOG|EM “D “A Lic)p: arom: t *(qndun) 24005 ‘§ “‘€ "“H Lic)p: aro lest -(qndun) 93005 ‘§ ‘€ “H UP Da up[xa “Da (-qndun) 24000 ‘s ‘a “H (-qndun) 24005 ‘s§ ‘a ‘H SIISM “H “TI 9400) °§ “a “H suINOg PM “D “aX peyuepran suryoy “£ (6L61) USM “DU (6L61) USI “Da (6L61) UPI DU (P6L61) USI “DU Ue DO U:(PU)2ex000 Sa H ¢ (PU) 9400) ‘§ “dH ANY Al ‘NOILVOIIaNd ALVUuvdas GNV VNNVA ONIAAILNACI NOSadd (6L61) UTA 2 SyOOIg ‘(8/61) UoIs[OH =(1L61) USTIPA ([2A2] VSIA) 184 (6961) uose, (Ajl[eoo] Jeunes jiupsueurodd¢) ar1g s,osunyy (F561) Apoqeod desjaog (ps61) Apoqeed SAR) Ye YOR (d'@ 000 ZE-000 82I *?) parwyuasaffipun *¢ 10/puv p “¢ sadvjg' adojosy (‘WIUIOD ‘sied) Jozing ‘(0g/6]) UOsqooer sIquqod (8L61) 72 12 YyeIUEyonIG “Iezjng ‘(wos ‘sied) Aeppoeysg 2 JUOUINvEg aAeD snnby (8261) 72 42 qyeIUSyoNIg ‘Iozing =(TL61) ALBIO :-S6t) Apoqeod (S[SA°] VSIA) PAD sUBIIITAN Cd“ 000 ££-000 8ZI *?) ¢ asv1g adojosy (pS61) Apoqeod Sina NU Cd’ 0-000 ZI °2) paiviuasaffipun ‘7 a8vj5' adojosy (‘uiur09 ‘sied) Jezing ‘(‘wiuI09 ‘sied) jUOUINeeg ‘(¢F61) SIPM 8% URTV «(TH6T) OOD 7% uryey (.PISgurUg Jorres,) SARD YoOMIOpuo AA Cd‘ 000 $-000 ZI *9) Aj4va ‘7 asvig adojosy (6L61) UPT19A 7 Syooig (PA2] VST) IBF (‘wu05 *sied) Jozing ‘(‘WwUuIOS “sied) yuouNeeg (PJoeyUIMUWIS 10}e],) SAVD YIOMIOpPUO MA (PL6]) F981YSsog 3 JUOWINeogG = Ulo}UOJUIOOG (S161) skeryduinyy Jay[9Y§ s.[[eyoing (deid ur pure ‘p/61) skeryduinyy 7 “I yosoqgyig (‘deid ur) skeryduinyy Z ‘T yoor9swry (8261) Skosrydunyy {(ps61) Apoqrog SARD SSNOyIEMOg WIUIOD “sIod) JuOUINvEg *(7S61) Apoqred SUPDIUM 31T (ps61) Apoqeod ([PA9] ,UOIIAA,) SARD SUBIAITAA Cd‘@ 0-000 £ *2) ain] ‘T asvig' adojosy aOUNOS TyuwaNad ALIS *QUOZOOW ULIILYL[ey 9} Ul S[eUIWIUT oSIe] JO SUIPUIOI PepIAOId SAY 3eU} SO}IS SUDDO;OF{ pue ouUad0}sSI0;q IeddqQ p A1EvL — ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS 245 late Holocene faunas (unless the fauna from Ochre Cave dates from this interval). It is a hillside grazer which is most at home in the Basutolian and Cape Zones, where its frequency in both live communities and archaeological sites tends to be highly correlated with and subequal to that of mountain reedbuck. Moister conditions may well be implied if it were as frequent as mountain reedbuck in a Kalaharian fossil fauna. THE BASUTOLIAN ECOZONE This zone comprises the Drakensberg Mountains and the high plateau country adjacent to them. In modern political terms, it covers the southern Transvaal, the western quarter of Swaziland, the highlands of western Natal, all of Lesotho, most of the Orange Free State, and a portion of the adjacent eastern Cape Province. The area is characterized by warm, relatively moist summers (average precipitation generally between 620 and 750 mm) and cold, dry winters. East of the Drakensberg, the historic vegetation of the Basutolian Zone was primarily open grassland with patches of temperate forest at the heads of river valleys and areas of acacia savanna at lower altitudes. West of the Drakens- berg, in the area known in South Africa as the ‘highveld’, the vegetation was nearly pure grassveld with trees largely confined to the river valleys. Historically, the fauna of the Basutolian region was dominated over- whelmingly by large, migratory, gregarious grazers, especially Burchell’s zebra, the recently extinct quagga, black wildebeest, blesbok, and springbok. Their numbers may even have exceeded those of their counterparts in the east African grasslands, and it is probable that they were interdependent in a grazing succession similar to that recently observed in east Africa. They probably migrated with the seasons in search of good pasture, and it is likely the migrations took in the eastern part of the Karoo-Namaqualian Zone into which the Basutolian one merges imperceptibly. In keeping with the open nature of the vegetation, browsers (greater kudu, bushbuck, etc.) and grazers favouring wooded country (roan, sable, Cape buffalo) were rare or absent in the Basutolian Zone. The impala was completely displaced by the springbok, but eland and steenbok, also mixed feeders, were widespread. Warthogs were numerous, but bushpigs were generally absent. Mountain reedbuck, vaalribbok, and to a lesser extent klipspringer were common in areas of high relief. Among smaller mammals, hares, springhare, and rock hyrax are still abundant in many places. Pollen-analytical studies undertaken at Florisbad by Van Zinderen Bakker (1957) and at Aliwal North by Coetzee (1967), both located near the western margin of the Basutolian Zone, reveal that the local grassveld was replaced at various times in the later Pleistocene by semi-desert shrub of the Karoo- Namaqualian Zone. At Aliwal North, where the sequence is reliably dated between approximately 13 200 and 9 600 B.P., a replacement of grassveld by Karoo shrub (and the reverse) occurred three times, reflecting relatively rapid fluctuations between cooler/moister and warmer/drier conditions similar to the ——— 24" FO™M 246 ANNALS OF THE SOUTH AFRICAN MUSEUM kind of relatively rapid climatic fluctuations that are known to have characterized the contemporaneous terminal Pleistocene/Holocene transition in Europe. At Florisbad, the pollen spectra are all much older and precise dating is a problem, though a grassveld phase indicating relatively moist conditions occurs in deposits that probably correlate with deep-sea isotope stage 2. Studies of alluvial cut-and-fill sequences in the Upper Orange drainage by Butzer (1971b) also indicate that stage 2 times were generally wet in the Basutolian Zone, as they were in the neighbouring Kalaharian one. Yet earlier wetter and drier phases are difficult to date, and it is clear that the ones reflected in pollen spectra at Florisbad are beyond the range of radiocarbon. The earliest part of the Florisbad sequence, in fact, probably dates from the later mid-Pleistocene. The Upper Pleistocene and Holocene sites in the Basutolian Zone which have provided remains of large mammals are listed in Table 5, along with the probable correlation of the sites (or of levels within them) with various oxygen- isotope stages. Most of the sites are located near the western margin of the zone and either date very clearly from the late Holocene or are difficult to date precisely. The late Holocene faunas are dominated by hare(s), rock hyrax, Burchell’s zebra and/or quagga, black wildebeest, springbok, and steenbok, suggesting an environment broadly similar to the historic one. Faunas coming from sites located in more rugged topography are clearly marked by a higher frequency of mountain reedbuck and vaalribbok, as would be expected. The pre-Holocene faunas are also dominated by large gregarious grazers, suggesting general limits to the extent of later Pleistocene environmental change. However, the presence of lechwe or waterbuck at Vlakkraal and Koffiefontein, of Cape buffalo at Koffiefontein and Driefontein, of impala at Koffiefontein, and of roan at Driefontein, may reflect moister conditions at one or more times during the earlier Upper Pleistocene. The lechwe is also present at Florisbad, but its stratigraphic provenience within the site is unknown. It may have come either from Upper Pleistocene levels, from levels that probably date to the late mid-Pleistocene, or from both. The same problems of provenience, reflecting relatively uncontrolled excavations, make it impossible to relate various elements in the Florisbad fauna to the palynological and geomorphic observations that have been made at the site, but the fauna is further interesting for the presence of hippopotamus, suggesting a time(s) when the pan next to the site may have contained a lake. The occurrence of water mongoose and clawless otter may reflect the same moist interval(s). Perhaps even more intriguing is the occurrence of giraffe, which must indicate that trees once grew near by, though the area was treeless historically, and Van Zinderen Bakker found virtually no arboreal pollen in any of the samples he examined from the site. THE KAROO—-NAMAQUALIAN ECOZONE This zone has two major components: (1) the Namib Desert, a narrow strip up to 160 km wide along the Atlantic coast, extending from the mouth of the Orange River through South West Africa to beyond Mossamedes in Angola; — ———— Sw 247 ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS VSWE VST VST VST VST VST VSI VSI VST VSI VST VST VST VST VST SNOILVIOOSSV TYOISOTOAVHOUV (PL6T) USI “D “A ‘SPM “HT JONIeD “Td Joe) “Td 2400) 'S ‘a “H (€961) 93909 ‘§ ‘€ “H *SIIPM 2 29D *§ “€ “H (€961) 94909 '§ ‘€@ “H ‘SIIPAA 7% 2400D ‘§ ‘€ 'H (€961) 91909 "S “@ "H ‘SIEM 9Y00D 'S ‘A “H Joep “I'd UPI Da é (06161) UTA “DU (P6161) UI “DU (76161) WIM "DU SUIYuY “f peynueprun (P6L61) UIA “DU peynueprun Joe) “I'd JOM’ “Td Joe “I ‘d Joe) “Td UP “OD “a ANV dI ‘NOILVOITaNd ALVUVdaS GNV VNOVd ONIASILNACI NOsddd (G0L61) SIIPAA (PLET) [P80A 2 J0}IBD *(LL61) 199IeD (LL61) 19348) Cd" 000 ZE-000 8ZI (‘WIUIOS ‘sod puv “91L61) 10ZIN ‘(8P61) 40D ‘S ‘A ‘H (‘WIWIOD *sied pure 1.) eee tek oe wen 260 ANNALS OF THE SOUTH AFRICAN MUSEUM for which detailed numerical data are available. Some authorities believe that it is not really possible to use fossil faunas to document environmental change | because it is rarely possible to know the relationship between relative species abundance in a fossil fauna and relative abundance in the live fauna from which the fossils were derived. However, the Cape data show that this is not an insuper- able problem, and the contrast between the Cape Zone and others indicates that widespread deficiencies in dating, sample size, and sample description are far more serious obstacles to the palaeoenvironmental interpretation of faunal data. MAMMALIAN EVIDENCE FOR CULTURAL CHANGE In archaeological sites, changes in environment and changes in culture are the principal causes of shifts in relative species abundance through time. In situations where the intervals between shifts represent thousands of years, where the shifts are repetitive or cyclical, where they appear to be correlated with changes in environment suggested by other lines of evidence, and where they occur in the absence of evidence for any significant cultural change or innovation, the writer feels it is most economical to ascribe the species fluctuations to environmental change. Most of the fluctuations in large mammal frequencies that can be documented through Upper Pleistocene and Holocene time in southern Africa seem to the writer to occur in circumstances such as the ones that have just been listed, and they have therefore been discussed in the previous section on ‘Mammalian Evidence for Environmental Change’. There are, however, some instances of mammalian frequency changes which are not clearly related to environmental change, which are not repetitive or cyclical, and which occur in contexts where there is evidence for major cultural change or innovation. The most obvious instance of a culturally determined species frequency change is the introduction of domestic stock to southern Africa by Iron Age mixed farmers, beginning about 2 000 years ago (Phillipson 1977). The stock were diffused far beyond the areas ever occupied by Iron Age farmers themselves (H. J. Deacon et al. 1978, with references), and some of the late Holocene faunas listed in the last section may be used to establish the route(s) of diffusion (Klein 1979a) or the impact the stock may have had on the abundance of some indigenous wild species (Klein 19745). For the open grasslands of the Basutolian region, Maggs (1975) has documented faunal differences that probably reflect major social and technological differences between Iron Age people and broadly contemporaneous Stone Age ones in the same area. The Iron Age faunal samples are richer in large gregarious grazing ungulates, probably because Iron Age peoples could mobilize more manpower for surrounds and drives and could more easily dig game pits that, combined with drives, would constitute the most effective means of obtaining large ungulates in the Basutolian Zone. The principal concern in this section, however, is not with changes in species frequencies that reflect the introduction of domestic stock or of Iron Age technology and social organization, but rather with a much earlier, more subtle | ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS 261 shift in mammal species frequencies that the writer believes may reflect important differences between Middle Stone Age and Later Stone Age peoples in their ability to hunt. In southern Africa, the term Middle Stone Age (MSA) is currently applied to artefact assemblages dominated by large stone flakes and blades, sometimes altered by retouch into side-scrapers, end-scrapers, points, denticulates, notches, backed pieces, and other tool types conventionally recognized by archaeologists. Handaxes are absent, and microlithic tools are rare. Bone artefacts are also rare, and items of personal adornment or art objects are unknown. Typological and technical variability through space and time is relatively limited, and many differences in typology or flaking technique among MSA assemblages in different regions or at different times may reflect differences in raw material availability more than anything else. The earliest Middle Stone Age assemblages in southern Africa may be as much as 200 000 years old (Butzer, Beaumont & Vogel 1978), while the latest are probably all older than 30 000 years, on the basis of a large series of recently obtained radiocarbon dates (see especially Vogel & Beaumont 1972; Beaumont ef al. 1978; H. J. Deacon 1979; Klein 1974a). It is presently impossible to characterize the Later Stone Age (LSA) succinctly, in part because the artefact assemblages involved are more variable in time and space than Middle Stone Age ones and in part because only Later Stone Age assemblages post-dating 20000 B.P. are reasonably well known. LSA assemblages ante-dating 20 000 years have been found at only a handful of sites in southern Africa (Kalemba, Leopard’s Hill, Redcliff, Heuningsneskrans, Border Cave, Apollo 11, Elands Bay, and Boomplaas; with references in Tables 2-7), where the samples are either small or remain incompletely described or both. In most LSA assemblages post-dating 20 000 B.P., microlithic tools are a prominent component, but this is not universally true, and, in the Cape Ecozone, there is clear evidence for a ‘macrolithic’ industry sandwiched between two ‘microlithic’ ones (J. Deacon 1978). Generally speaking, LSA peoples appear to have produced ‘macrolithic’ flakes and blades with less care than many MSA peoples. At least the better known LSA assemblages younger than 20 000 B.P. also regularly include standardized bone artefact types (such as ‘awls’, ‘points’, ‘needles’, ‘hide-burnishers’, and ‘fish-gorges’), as well as easily recognizable items of personal adornment or art objects (ostrich egg-shell beads, incised or engraved pieces of ostrich egg-shell or bone, shell pendants, etc.). On the evidence from Border Cave, it seems likely that the manufacture of beads and standardized bone artefacts was practised from the very beginning of the Later Stone Age, more than 30 000 years ago. People making Later Stone Age artefacts of various kinds were still living in much of southern Africa at time of historic contact. The available evidence suggests strong parallels between the Middle Stone Age and Later Stone Age of southern Africa, as outlined here, and the Middle Palaeolithic (Mousterian) and Upper Palaeolithic of Europe, as they have long — — se ivarr se _ 2 a E ae SAA CS ee TT An _ a Fa 262 ANNALS OF THE SOUTH AFRICAN MUSEUM been known. Although the Middle Stone Age may have begun somewhat earlier than the Middle Palaeolithic, throughout much of their existence the two’ were clearly contemporaneous and their terminal dates are very similar. Both exhibit less temporal and spatial variability than the culture-stratigraphic units that succeed them, and both are characterized by the absence of art objects and standardized bone artefact types. One major point of non-comparability may be in the kinds of people associated with them. While the makers of Middle Palaeolithic artefacts were Neanderthals (Homo sapiens neanderthalensis), clearly distinct from modern people, the makers of MSA tools may have been anatomically modern (Homo sapiens sapiens) (Rightmire 1979), though a pattern of well-documented associations between MSA artefacts and diagnostic human remains will be necessary to show this with reasonable certainty. The makers of Upper Palaeolithic artefacts were certainly anatomically modern, as were at least those Later Stone Age people who lived after 20 000-18 000 B.P. In Europe, there is substantial evidence to argue that the Upper Palaeolithic represents a quantum advance over what preceded it (Klein 1973), and at least for the moment, the writer feels it is reasonable to hypothesize that the Later Stone Age represents basically the same phenomenon in southern Africa. Under these circumstances, differences between MSA and LSA faunas from sites occupied under broadly similar environmental conditions could be interpreted to reflect cultural evolution. In orde1 to establish such faunal differences, it is of course necessary to have large MSA and LSA faunal samples whose palaeo- environmental context has been reasonably well established on independent grounds, and these conditions are so far met only in the Cape Ecozone, especially by the Middle Stone Age faunas from Klasies River Mouth and Die Kelders 1 and the Later Stone Age ones from Nelson Bay Cave and Byneskranskop 1. At all four sites, in levels that were formed when the coast was at or near its present position, bones of seals and penguins are common, but only in the two Later Stone Age sites are they accompanied by large numbers of bones from fish and from flying marine birds. The writer believes this indicates that Later Stone Age people were capable of active fishing and fowling, while Middle Stone Age people were not. With regard to the terrestrial mammal remains that are the principal focus of this paper, there is also an interesting contrast. Comparing layers at Nelson Bay formed during the Holocene to ones that appear to have formed during broadly similar portions of the Last Interglacial at Klasies, the Nelson Bay deposits are significantly richer in remains of pigs and poorer in remains of eland. The LSA levels of Byneskranskop 1 contrast with the MSA ones at Die Kelders in essentially the same way (the comparisons here are restricted to pairs of sites which are located in very similar environments today), though the extent of past environmental comparability is less certain than in the Klasies/Nelson Bay case. In any event, the writer has suggested that the higher frequency of wild pig and lower frequency of eland in the LSA sites reflects the enhanced ability of LSA people to deal with prey that are likely to mount an effective counter-attack on the hunter. Using data on the ages of animals ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS 263 represented in the Nelson Bay and Klasies faunas in addition to the contrasts in relative species frequencies, the writer has further suggested that even when MSA people hunted basically the same species as their LSA successors, they were less effective, that is, they took a smaller proportion of the available animals (Klein 19796). Clearly, the hypothesis that LSA hunters were more effective than MSA ones would be more secure if it could be demonstrated at additional sites in the Cape Zone and also at sites outside it. The writer has examined the available faunal data from all the other ecozones considered in the previous section of this paper, and only in the Zambesian Zone are there sufficient numerical data for even a preliminary test. Pigs (especially warthog) are more common in both MSA and LSA sites in the Zambesian Zone than in contemporaneous Cape sites, probably reflecting the fact that pigs have always been more abundant in the Zambesian Zone, as they were historically. At the same time, in those Zambesian sites for which numerical data are available, pigs are relatively more frequent in LSA levels (Kalemba, Redcliff, Gwisho, Leopard’s Hill, and Makwe) than in MSA ones (Kalemba and Redcliff). However, at Redcliff and Kalemba, where MSA and LSA pig frequencies may be compared within the same sites, the relative increase in pigs is not statistically significant, and it remains possible that the observed differences in pig frequencies between the MSA sites and various LSA ones reflect differences in local environment rather than cultural evolution. Sorting out the alternatives will be possible only with better palaeo- environmental controls and larger faunal samples. More generally, as in the case of environmental change, the principal obstacle to documenting cultural change from large mammal remains is the shortage of appropriately large, well- described, and well-dated samples. THE ECOLOGY AND DEMISE OF EXTINCT SPECIES Not very long ago, it was widely believed that Acheulean cultures had survived into the Upper Pleistocene and that faunas associated with later Acheulean artefacts could even be as recent as 40000 B.P. The richest such fauna in southern Africa is the one from Elandsfontein (Hopefield) in the south- western Cape Province (Hendey 1974 with references), which is characterized by at least 19 extinct large mammal species out of approximately 50 that have been identified. The extinct species include a giant gelada baboon (Theropithecus (Simopithecus) sp.), a sabre-toothed cat (Megantereon sp.), an elephant (Loxodonta atlantica), a large horse or zebra (Equus capensis), two different kinds of pigs (Mesochoerus sp. and Metridiochoerus sp.), a sivathere (Sivatherium maurusium), a small kudu (Tragelaphus (Strepsiceros) sp.), a giant buffalo (Pelorovis sp.), a giant hippotragine antelope (Hippotragus gigas), the ancestor (Rabataceras arambourgi) of the modern hartebeests, an extinct bastard harte- beest (Damaliscus niro), a giant alcelaphine antelope (Megalotragus sp.), a large grysbok (Raphicerus sp.), a gazelle (Gazella sp.), springboks (Antidorcas recki GANA oo. YY CS ty i 4g Tet r 7.3 1 om ams Se Le 7” Ferran Sl 34 NoN 264 ANNALS OF THE SOUTH AFRICAN MUSEUM and A. australis), and some peculiar antelopes whose affinities will perhaps be clarified following E. S. Vrba’s current detailed study of the Elandsfontein bovids. It is now very clear that the Acheulean did not survive into the Upper Pleistocene, and it seems increasingly probable that it was replaced by the Middle Stone Age within the later part of the Middle Pleistocene, more than 130 000 years ago (Klein 19766; Butzer, Beaumont & Vogel 1978; Szabo & Butzer 1979). Furthermore, the overwhelming majority of extinct species repre- sented at Elandsfontein and other Acheulean sites are not represented in Middle Stone Age faunas and in non-archaeological faunas that are contemporaneous with them (see the lists of sites in Tables 2—7 above). Particularly striking in this context is the absence of most of the extinct Elandsfontein species in large earlier Upper Pleistocene faunas in the (same) Cape Ecozone, especially in the very large samples from Klasies River Mouth, where dating of the MSA artefacts and associated fauna to the early Upper Pleistocene is secure. In fact, it appears increasingly likely that most of the extinct species present at Elandsfontein and other Acheulean sites disappeared well before the early Upper Pleistocene, since they are not represented in the large faunal sample from Florisbad, most of which probably comes from a later mid-Pleistocene horizon at the site (Butzer pers. comm.). The Florisbad fauna contains five extinct species of large mammals—the ‘giant Cape horse’ (Equus capensis), a large warthog (Metridiochoerus sp.), a giant buffalo (Pelorovis antiquus), a giant alcelaphine (Megalotragus priscus), and Bond’s springbok (Antidorcas bondi). With the addition of the southern springbok (Antidorcas australis), these are also the only extinct species which have been found in Upper Pleistocene contexts in southern Africa. Some dentitions assigned to the extant Damaliscus dorcas (bontebok/blesbok) in Upper Pleistocene faunas may actually belong to the extinct form Damaliscus niro, but horn-cores to prove the presence of D. niro are lacking. Additionally, in contrast to Gentry (1978), the writer believes that D. niro may be ancestral to Upper Pleistocene and recent D. dorcas, in which case it is not extinct in the same sense as the other species considered here. Extinct forms of wildebeest and reed- buck are common in Upper Pleistocene sites in the Cape Ecozone, but were probably just local variants (subspecies) of the black wildebeest and southern reedbuck that survived elsewhere. The purpose in this section is to present the information that is available on the distribution and ecology of the extinct species followed by a brief considera- tion of the causes of extinction. Teeth of the various extinct forms are illustrated in Figures 3-5. EQUUS CAPENSIS The writer has followed Churcher & Richardson (1978; also Churcher 1970) in assigning all large later Pleistocene horses in southern African sites to the species Equus capensis (including ‘E. helmei’, ‘E. plicatus’, et al.) Upper Pleisto- cene specimens assignable to Equus capensis have been found in all six ecozones ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS 265 Metridiochoerus sp. Equus capensis (RC VII EE 1) (RC VI S) Fig. 3. A fragmentary third molar of Metridiochoerus sp. and a lower third molar of Equus capensis. Both specimens come from Redcliff Cave, Zimbabwe Rhodesia, and are reproduced natural size. (Drawings by K. Scott.) considered earlier: at Redcliff in the Zambesian Zone; at Pomongwe, Chelmer, Kalkbank, the Cave of Hearths, Bushman Rock Shelter, Border Cave, and Aloes in the Transvaalian Zone; at Equus Cave, Black Earth Cave, and gi in the Kalaharian Zone; at Florisbad, Vlakkraal, and Koffiefontein in the Basutolian Zone; at Apollo 11 and Elands Bay Caves in the Karoo-Namaqualian Zone; and at Sea Harvest, Hoedjies Punt, Swartklip, Bloembos, Duinefontein, Byneskranskop 1, and Boomplaas in the Cape Zone. On the evidence from Apollo 11, Elands Bay, Byneskranskop 1, and Boomplaas, Equus capensis probably made its last appearance in the Karoo— Namaqualian and Cape Zones between 12000 and 10000 years ago. In the Zambesian Zone, it is known from the “Tshangula’ horizons at Redcliff, where it may be as young as 20 000 B.P. In other zones, it cannot be shown to have survived 40 000-30 000 B.P., but this probably reflects the very small number and small size of faunal samples that date between 40 000-30 000 and 10 000 B.P. ee ‘> la . — +. eo _— ~ 24 - Ctr, Rass a — KS fo a ~ [-j een (9) ey, 2 ¥om™N 266 ANNALS OF THE SOUTH AFRICAN MUSEUM in these zones. In essence, the time when E£. capensis made its last appearance outside the Karoo—Namaqualian and Cape Zones remains to be established. In most Upper Pleistocene sites in southern Africa and in all those which have provided large samples, Equus capensis is accompanied by one or more of its smaller, historic relatives—Burchell’s zebra, quagga, or mountain zebra. In most sites, it is much less common than the smaller form(s), but in Karoo- Namadqualian sites, located in what is and perhaps always has been the driest of the southern African ecozones, and in sites in the drier, western portion of the Cape Ecozone, it is relatively more common (versus smaller equids). Additionally, in the Zambesian Zone, at Redcliff, where E. capensis is not particularly frequent overall, it is most frequent in the ‘later Bambata’ horizons, which sedimentological evidence suggests were formed under comparatively arid conditions. The possibility that E. capensis preferred or was relatively well . adapted to arid environments is especially interesting, since some authorities believe it was ancestral to the living Grevy’s zebra of northern Kenya, southern Ethiopia, and Somalia (Churcher & Richardson 1978). Among the living zebras, Megalotragus priscus (RC ND) Pelorovis antiquus (RC 15'B) Fig. 4. Lower molars of Megalotragus priscus and Pelorovis antiquus. Both specimens come from Redcliff Cave, Zimbabwe Rhodesia, and are reproduced natural size. (Drawings by K. Scott.) f~ een ew 267 ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS _ — SA As LS ee eee lee eee — a aetlUm6RE (‘nosg “yy Aq s8UIMeI) “9ZIS [eInyeu poonpoider ore suounioads yJog “eIsepoyy eMqequulZ “oAvD Pyopoy Wor [ dippeMg Woy syossnv sposopyup JO e{QIpueUl VW °¢ 1puog SvI4OpYUp JO Te[OUI IaMO] & pue “eOTIFY YINOG “soUlACI adea U10}seM-43N0s (a SZ Ou) ipuog soo2s0pijuy/ « iG ‘31d (28 MZ) syo4snD spo40pnuy 268 ANNALS OF THE SOUTH AFRICAN MUSEUM Grevy’s appears to be the least dependent upon water. It inhabits some of the most arid country in east Africa, overlapping Burchell’s zebra on the margins of its range, but occurring alone in the arid core. METRIDIOCHOERUS SP. The writer has followed*White and Harris (1977) in assigning all large warthog-like pigs of the later Pleistocene to the genus Metridiochoerus (including “Stylochoerus’, ‘Tapinochoerus’, et al.), though not all specialists agree with their taxonomy (Cooke 1978). In any case, the large pigs involved are so far very rare in Upper Pleistocene contexts, having been recorded only at Koffiefontein, Vlakkraal, and Florisbad in the Basutolian Ecozone (the Florisbad specimens may be largely or wholly of mid-Pleistocene age), and at Redcliff Cave in the Zambesian Zone. Among these occurrences, the latest is perhaps at Redcliff, where a large warthog-like pig is still present in the “Tshangula’ horizons, — tentatively dated to between 30 000 and 20 000 B.P. in discussion above. Estimates of Metridiochoerus abundance are available only from Redcliff where the creature is very rare, too rare to search for frequency covariation with extant species that might indicate shared habitat preferences. Its highly hypsodont molars, similar in structure to those of the warthog, plainly indicate it was a grazer, which is in keeping with the predominance of grazers at Redcliff and in the Basutolian faunas in which Metridiochoerus has been identified. PELOROVIS ANTIQUUS The writer has followed Gentry (1978) in assigning all the ‘giant’, long- horned Upper Pleistocene buffaloes of southern Africa to this species (including especially material formerly assigned to ‘Homoioceras baini’). Fossils of Pelorovis antiquus have been found at Redcliff in the Zambesian Zone; at Kalkbank and the Cave of Hearths in the Transvaalian Zone; at #gi in the Kalaharian Zone; at Florisbad, Vlakkraal, Koffiefontein, and Driefontein in the Basutolian Zone; at Elands Bay Cave in the Karoo-Namaqualian Zone; and at Sea Harvest, Hoedyjies Punt, Ysterfontein, Bloembos, Swartklip, Die Kelders, Nelson Bay, Boomplaas, and Klasies River Mouth in the Cape Zone. The species was thus very widespread, but it is generally not common in fossil faunas, except in the Basutolian Zone, where it has also been recorded as an apparently isolated fossil at various localities, for example, in the alluvium of the Modder River, which provided the first giant buffalo fossil to be scientifically recorded in southern Africa (Cooke 1955). Finds at Nelson Bay and Elands Bay Caves suggest P. antiquus made its last appearance in the Cape and adjacent parts of the Karoo-Namaqualian Zone between 12 000 and 10000 years ago. Elsewhere, it cannot be shown to have survived beyond 40 000-30 000 B.P., but, as in the case of Equus capensis, this probably reflects the absence of large faunal samples from the interval between 40 000-30 000 and 10 000 B.P. The date of its last appearance outside the Cape and Karoo—Namaqualian Zones thus remains to be established. The hypsodont teeth and exceptionally long horns of P. antiquus (spanning ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS 269 2-3 m) both point to a preference for relatively open, grassy environments, also perhaps suggested by the relative abundance of the species in Basutolian sites. It seems likely that P. antiquus was more at home in open settings than its closest living relative, the Cape buffalo. Ecological distinction from the Cape buffalo is implied at Klasies River Mouth, where the frequencies of the two species vary independently of one another from level to level within the deposits. At Klasies, the frequency of giant buffalo is correlated most closely with that of eland, which may suggest a common environmental preference. Eland, as studied by Hillman (1974), appear to favour open country with large islands and galleries of tree and shrub growth. The Basutolian sites at which giant buffalo are common are all located near streams or springs where such islands or galleries probably occurred. The giant buffalo may have gravitated to them for shade rather than for food. At Klasies, there are enough giant buffalo dentitions for an analysis of the ages at which individual animals died or were killed by the Middle Stone Age occupants of the site (Klein 1978d, 19796). Most of the individuals present were within the first 10 per cent of potential lifespan (many were newborn), while among older animals, prime adults (between 20% and 50% of lifespan) were especially rare. The Klasies giant buffalo age distribution is very similar to the natural (attritional) mortality pattern in all free-ranging large ungulates, to the age distribution in recently observed Cape buffalo killed by lions, and to the age distribution of Cape buffalo in the same Klasies MSA levels. Recent observations indicate that it is large size and membership in large herds that make Cape buffalo prime adults largely immune to lion predation. The age distribution of Cape buffalo at Klasies suggests that the MSA occupants were constrained by the same features as lions in dealing with Cape buffalo, and the similarity between the Cape buffalo and giant buffalo distributions suggests that large size and herd membership may also be the reasons that giant buffalo prime adults are rare at Klasies. Given the resemblances between the giant buffalo age profile at Klasies and the probable natural (attritional) mortality pattern in giant buffalo, it is even conceivable that the Klasies people did not prey on giant buffalo at all, but scavenged individuals which had died naturally or been killed by other predators. MEGALOTRAGUS PRISCUS The writer has followed Gentry (1978) in assigning all the later Pleistocene ‘giant alcelaphines’ of southern Africa to the single species Megalotragus priscus (including ‘Pelerocerus’ spp, ‘Lunatoceras mirum’, “Alcelaphus helmei’, et al.). Upper Pleistocene fossils of M. priscus have been found at Leopard’s Hill and Redcliff in the Zambesian Zone; at Chelmer and Kalkbank in the Transvaalian Zone; at Equus Cave, Black Earth Cave, Gobabis, and “gi in the Kalaharian Zone; at Florisbad, Viakkraal, K offiefontein, and Driefontein in the Basutolian Zone; and at Hoedjies Punt, Boomplaas, and Nelson Bay Cave in the Cape Zone. Outside the Basutolian Zone, M. priscus is generally not a common a rr tay... ya. BAAN Sea, = 123. —_ - ae a 270 ANNALS OF THE SOUTH AFRICAN MUSEUM element in fossil faunas; within this zone, it is not only well represented at the sites listed above but has been found as an isolated fossil or with presently | unclear associations at a variety of other sites, including especially the alluvium of the Modder River, which provided the holotype specimen. Its hypsodont teeth, the dietary preferences of all its closest living relatives (hartebeests, wildebeests, and bastard hartebeests), its former distribution, and its clear tendency to be most common in fossil faunas heavily dominated by grazers, indicate beyond all doubt that M. priscus was a grazer. Its latest known occurrence is in deposits at Nelson Bay Cave radiocarbon-dated to approxi- mately 16 000 B.P., but it is a rare element in the Nelson Bay fauna, and the possibility is good that it persisted, like Equus capensis and Pelorovis antiquus, in the Cape Zone somewhat later, perhaps to between 12 000 and 10 000 B.P. At Leopard’s Hill in the Zambesian Zone, its provenience within the site is uncertain, but it is almost certainly younger than 24 000 radiocarbon years (the approximate age of the oldest fossiliferous deposits at the site). Elsewhere, it cannot be shown to have survived beyond 40 000-30 000 B.P., but again, as in the case of Equus capensis and Pelorovis antiquus, this may well reflect the rarity of large faunal samples from the interval 40 000-30 000 to 10 000 B.P., rather than the absence of the species. ANTIDORCAS BONDI Cooke and Wells (1951) initially described this hyperhypsodont antilopine antelope as Gazella bondi. However, Vrba (1973) has demonstrated that the frontals of this species were inflated below the horn-cores, a feature that is characteristic of Antidorcas and not of Gazella. For this and other reasons, the species is better placed in Antidorcas (Gentry 1978). Upper Pleistocene fossils of A. bondi are known from Redcliff in the Zambesian Zone; from Chelmer, the Cave of Hearths, and Border Cave in the Transvaalian Zone; from Gobabis, Witkrans, Equus Cave, and Black Earth Cave in the Kalaharian Zone; and from Florisbad, Vlakkraal, and Driefontein in the Basutolian Zone. Its absence so far in the Karoo-Namaqualian Zone may be a result of inadequate sampling, but its failure to occur in Cape faunas may reflect true absence, since the Cape samples in which it might be expected are relatively large. The latest record of A. bondi is at Border Cave in a level radiocarbon-dated to approximately 38 000 B.P. However, the date of its last appearance remains unestablished, since large samples dating to between 38 000 and 10 000 B.P. are unknown in those ecozones where the species was most common. Its hyperhypsodonty, geographic distribution, and the species with which it occurs all indicate clearly that A. bondi was primarily a grazer. This is shown particularly well at Border Cave, where there is alternation between levels dominated by grassland species and ones dominated by species that prefer bushier settings. A. bondi is plainly most common in those levels where grassland animals are most abundant. — ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS 271 ANTIDORCAS AUSTRALIS Hendey and Hendey (1968) originally described this species as a subspecies of the common springbok, Antidorcas marsupialis. However, Hendey (1974) concluded it was more likely to be a separate species, since, at the mid- Pleistocene locality of Elandsfontein, it was apparently sympatric with A. recki, the probable ancestor of the common springbok. Upper Pleistocene fossils of A. australis have been found only in the Cape Ecozone where it was widespread. Its last recorded occurrence is in deposits at Nelson Bay Cave dated to between 12 000 and 10 000 B.P. A. australis had horn-cores that were more mediolaterally compressed than those of A. marsupialis and that did not bend sharply backwards and outwards. It was also somewhat smaller on average than A. marsupialis, but in most other important respects, including the morphology of the dentition, it was very similar to A. marsupialis. A. marsupialis is not known to have occurred in the Cape Zone in Upper Pleistocene (or recent) times, except on the peripheries, and it seems likely that A. australis filled the niche for a small selective grazer also capable of some browsing that A. marsupialis filled and still fills in neighbouring ecozones. Like the frequency of A. marsupialis, that of A. australis in fossil sites is closely linked to the frequency of ‘plains’ species, such as wildebeest, bastard hartebeest, and zebra. THE CAUSES OF EXTINCTION As in other parts of the world, the causes of late Pleistocene mammalian extinctions in southern Africa are a subject of considerable interest, particularly given the very real possibility that early people were involved. The other major ‘cause’ which has been postulated is environmental change. The writer believes that environmental change played a role in late Pleisto- cene extinctions, but it can nowhere be the sole reason, since the species that became extinct repeatedly survived the same kinds of environmental change earlier on in the Pleistocene. This can be shown especially clearly in the Cape Ecozone where the extinction of the giant Cape horse, giant buffalo, southern springbok, local subspecies of the black wildebeest and southern reedbuck, and probably also the giant alcelaphine occurred roughly 12 000—10 000 years ago, at a time when it is clear that fynbos, bush, and forest were replacing much of the grassland that had existed previously. The species that disappeared were all primarily grazers, which probably did suffer a shrinkage in both numbers and range as a result of the vegetational change. However, the same kind of vege- tational change characterized similar climatic shifts earlier on in the Cape Zone (that is, earlier transitions from ‘glacial’ to ‘interglacial’ conditions). During these earlier shifts, at least some of the extinct species appear to have become less numerous, but they all survived. Environmental change is thus not a sufficient explanation for extinctions in the Cape Zone, and the principal factor which differentiated the period of extinction, 12000-10000 years ago, from earlier periods of comparable —_ wool = “he ee \cowe | a Ay Be 0 | — \"' we. RAS _| Os..vwsi = \w,..\ws zo Die ANNALS OF THE SOUTH AFRICAN MUSEUM environmental change was perhaps the presence of more proficient hunters, as discussed in the previous section of this paper. The writer believes that it is entirely possible that Stone Age people in the Cape, faced with a decline in the ‘plains’ game on which they had depended for generations, intensified their pursuit of those that were left, perhaps through technological innovation driving the numbers of some species below a critical threshold and impairing their reproductive capacity. Extinction would follow, even if people were not responsible for killing the very last animal. Outside the Cape Ecozone, the hypothesis that people were responsible for late Pleistocene extinctions is much more tenuous. It seems likely that the extinctions occurred after Later Stone Age peoples had replaced Middle Stone Age ones in other ecozones, but evidence that these Later Stone Age peoples were more proficient hunters is so far lacking. Additionally, the timing of extinctions outside the Cape Zone remains unclear, mainly because there are no large faunal assemblages elsewhere dating from the critical period between 40 000-30 000 and 10 000 B.P. Finally, even if it is assumed that the extinctions outside the Cape Zone took place 12 000-10 000 B.P., as they did inside it, the nature of environmental change in this interval is not as well established as in the Cape, and it is far from certain that it would have adversely affected the extinct species in the way it probably did in the Cape. This is important, because the hypothesis offered above requires an environmental stimulus for any human role in extinction. Clearly, further elucidation of the causes of late Pleistocene extinctions in southern Africa will require the recovery of large late Pleistocene faunal samples from various ecozones. These are necessary not only for more precise estimates of the timing of extinctions and of the nature of any environmental change that may have accompanied them, but also for determining if the extinct species experienced long periods of decline or if they disappeared rather abruptly, as appears to be the case in the Cape Zone. Additionally, large samples may allow the construction of age/sex profiles to determine if the extinct species underwent demographic changes through time that might indicate impaired reproductive capacity or changing patterns of human predation. In sum, as in the case of demonstrating environmental and cultural change from faunal remains, elucidating the causes of extinction is not so much a theoretical problem as it is a practical one of obtaining large, well-excavated, and well-described faunal assemblages from contexts where good data on artefacts, sediments, pollen, and so forth provide controls for comparisons designed to separate the cultural and environmental variables responsible for assemblage composition. CONCLUSIONS In the overwhelming majority of cases, it is impossible to assume a one-to- one relationship between the relative abundance of species in a fossil mammal assemblage and their relative abundance in the live community from which they were drawn. Almost always, it is far safer to assume that the agency of accumu- ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS 273 lation has altered the original frequency relationships, and it is usually impossible to say how much. Thus, for example, the fact that a particular archaeological faunal assemblage consists three-quarters of grazers and one-quarter of browsers does not mean that the grazer/browser ratio in the ancient environment was 3 : 1 or even that grazers were more numerous than browsers. It may be that the people responsible for the bones found it easier to obtain grazers, or that grazers, since they are often larger, provided a higher return for time and effort in the hunt. Whatever the case, the extent to which the people may have altered or ‘piased’ the original grazer/browser ratio is probably unknowable. Superficially then, detailed interpretation of fossil mammal assemblages may appear impossible from the outset. However, this is only the case if the goal is to make precise statements about the live abundance of various species or about the subsistence behaviour of people at single instants in past time. It is not true if the focus is on changes in live abundance or in cultural practices through time, since, in this case, it is possible to introduce controls for the effects of environment or culture. As an example, take two faunas of different ages from neighbouring archaeological sites or from different levels within the same site. If there is evidence from pollen, sediments, oxygen-isotope ratios in associated marine shells, etc., that the faunas were accumulated under very similar environ- mental circumstances, it seems reasonable to suppose that any differences in species frequencies between them reflect differences in human behaviour. Similarly, if there is sound associated evidence for differences in past environment and no artefactual evidence for significant differences in culture, then it is probably most reasonable to conclude that any differences in species frequencies between faunas reflect differences in past environments. As a more concrete example, take two faunas from successive levels in the same site containing broadly similar artefacts, but in sedimentary contexts suggesting important differences in palaeoenvironment. If the fauna of one level contains relatively more grazers than the fauna of the other, it seems most reasonable to conclude that grazers were more common on the hoof at the time the first level formed, even though the live abundance of grazers in the ancient environment of either level remains unknown or unknowable. Basically then, faunal samples are far more useful for establishing changes in environment or culture through time than for reconstructing environments or cultural practices at particular points in time. However, it is obvious that changes may be determined only through comparisons among samples, with controls provided by accompanying pollen, sediments, artefacts, and so forth, as well as by detailed characteristics of the bone assemblages themselves. From this, it follows, first, that faunal analysis is inextricably dependent upon other fields — conventional archaeology, palynology, sedimentology, isotope geo-chemistry, etc.—if interpretations are to be secure. Second, it is obvious that no single sample is ever likely to be very informative. Rather, it is necessary to have many samples to permit as wide a range of controlled comparisons as possible. It is, of course, also important that the samples be large, well described, and well en _ eel ae 444... “He \ Wor, > {7 oe — \OW, | Css, \oW, w2/ =a wy, ws] = 282 ANNALS OF THE SOUTH AFRICAN MUSEUM APPENDIX VERNACULAR AND SCIENTIFIC NAMES OF THE EXTANT MAMMALIAN SPECIES MENTIONED IN THE TEXT VERNACULAR NAME SCIENTIFIC NAME hedgehog Erinaceus frontalis Cape hare Lepus capensis scrub hare Lepus saxatilis red rock hare Pronolagus crassicaudatus sptinghare Pedetes capensis cane rat Thryonomys swinderianus porcupine Hystrix africae-australis Cape mole-rat Bathyergus suillus chacma baboon Papio ursinus yellow baboon Papio cynocephalus vervet monkey Cercopithecus aethiops side-striped jackal Canis adustus black-backed jackal Canis mesomelas Cape hunting dog Lycaon pictus ale Sisece'S -™ A awl iite™ aN News spotted hyena brown hyena lion leopard cheetah Cape fur seal gazelle seal elephant seal -crab-eater seal rock hyrax tree hyrax elephant Grevy’s zebra Burchell’s zebra quagga mountain zebra black rhinoceros white rhinoceros hippopotamus warthog bushpig giraffe okapi eland Crocuta crocuta Hyaena brunnea Panthera leo Panthera pardus Acinonyx jubatus Arctocephalus pusillus Arctocephalus gazella Mirounga leonina Lobodon carcinophagus Procavia capensis Dendrohyrax arboreus Loxodonta africana Equus grevyi Equus burchelli Equus quagga Equus zebra Diceros bicornis Ceratotherium simum Hippopotamus amphibius Phacochoerus aethiopicus Potamochoerus porcus Giraffa camelopardalis Okapia johnstoni Taurotragus oryx ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS 283 VERNACULAR NAME bongo greater kudu nyala sitatunga bushbuck gemsbok roan antelope sable antelope waterbuck puku lechwe southern reedbuck mountain reedbuck Cape (= red) hartebeest Lichtenstein’s hartebeest tsessebe bontebok and blesbok bastard hartebeest blue wildebeest black wildebeest impala Grant’s gazelle Thomson’s gazelle springbok blue duiker common (= grey) duiker oribi klipspringer steenbok Sharpe’s grysbok Cape grysbok vaalribbok domestic sheep domestic goat Cape buffalo domestic cattle SCIENTIFIC NAME Boocercus euryceros Tragelaphus strepsiceros Tragelaphus angasi Tragelaphus spekei Tragelaphus scriptus Oryx gazella Hippotragus equinus Hippotragus niger Kobus ellipsiprymnus Kobus vardoni Kobus leche Redunca arundinum Redunca fulvorufula Alcelaphus buselaphus Alcelaphus lichtensteini Damaliscus lunatus Damaliscus dorcas Damailiscus spp. Connochaetes taurinus Connochaetes gnou Aepyceros melampus Gazella granti Gazella thomsoni Antidorcas marsupialis Cephalophus monticola Sylvicapra grimmia Ourebia ourebi Oreotragus oreotragus Raphicerus campestris Raphicerus sharpei Raphicerus melanotis Pelea capreolus Ovis aries Capra hircus Syncerus caffer Bos taurus See a [eae es A a > <7 ee en ee cS Om, . Qa, \OW, .ws/ ao w,...ws/ = eaecnnnie et pen 6. SYSTEMATIC papers must conform to the International code of zoological nomenclature (particularly Articles 22 and 51). Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. noy., comb. nov., syn. nov., etc. An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is transferred from its original genus. The name of a subsequent user of a scientific name must be separated from the scientific name by a colon. Synonymy arrangement should be according to chronology of names, i.e. all published scientific names by which the species previously has been designated are listed in chronological order, with all references to that name following in chronological order, e.g.: Family Nuculanidae Nuculana (Lembulus) bicuspidata (Gould, 1845) Figs 14-15A Nucula (Leda) bicuspidata Gould, 1845: 37. Leda plicifera A. Adams, 1856: 50. Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b). Nucula largillierti Philippi, 1861: 87. Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9. Note punctuation in the above example: comma separates author’s name and year “semicolon separates more than one reference by the same author full stop separates references by different authors figures of plates are enclosed in parentheses to distinguish them from text-figures dash, not comma, separates consecutive numbers Synonymy arrangement according to chronology of bibliographic references, whereby the year is placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable. In describing new species, one specimen must be designated as the holotype; other speci- mens mentioned in the original description are to be designated paratypes; additional material not regarded as paratypes should be listed separately. The complete data (registration number, depository, description of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.: Holotype SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973. Note standard form of writing South African Museum registration numbers and date. 7. SPECIAL HOUSE RULES Capital initial letters (a) The Figures, Maps and Tables of the paper when referred to in the text __ e.g. ‘.. . the Figure depicting C. namacolus...’; *. . . in C. namacolus (Fig. 10)...’ ' (b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by initials or full names e.g. DuToit but A.L.du Toit; Von Huene but F. von Huene (c) Scientific names, but not their vernacular derivatives e.g. Therocephalia, but therocephalian Punctuation should be loose, omitting all not strictly necessary Reference to the author should be expressed in the third person Roman’ numerals should be conyerted to arabic, except when forming part of the title of a book or article, such as ‘Revision of the Crustacea. Part VIII. The Amphipoda.’ ees Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial capital letter, provided the same generic name is used consecutively. ; Name of new genus or species is not to be included in the title: it should be included in the abstract, counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts. ~~ ARAA Se coo lan?) ae _—— 7 2 Sh Ty ‘> wer. ewe] = %, w,...\.w2/ =m \ te. \o Ww. .wes > WSOC %,.,..ws/ =— etltind RICHARD G. KLEIN ENVIRONMENTAL AND ECOLOGICAL IMPLICATIONS OF LARGE MAMMALS FROM UPPER PLEISTOCENE AND HOLOCENE SITES IN SOUTHERN AFRICA SF on ed _. 5) lees SOA SFR AON Ish Al CAE? Lv. tiles “AT "2 f2z—S “AU et - SAS = FR aN a Ee - z —- ove ~ ~- a“ = re an ape — ~S” GQ m Soy ~ = SUS faa ) FHSON, é ° ral ILNL! NS wo s\ az js] > = », we et. = tH \9 :) ANotiTy ie . #£O PS i = \s' wy = = rs nti ANSTITL = Py > i > “ ee) e) = 2 = — = & Z o Z i 2 = = 77) ae NOILNLILSNI LIBRARIES. 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