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New Species of Cercopithecoides from Haasgat, North West Province, South Africa

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New Species of Cercopithecoides from Haasgat, North West Province, South Africa Journal of Human Evolution 60 (2011) 83e93 Contents lists available at ScienceDirect Journal of Human Evolution journal homepage: www.elsevier.com/locate/jhevol New species of Cercopithecoides from Haasgat, North West Province, South Africa Jeffrey K. McKee a,b,*, Acacia von Mayer c, Kevin L. Kuykendall d a Department of Anthropology, The Ohio State University, 4034 Smith Laboratory, 174W. 18th Avenue, Columbus, OH 43210-1106, USA b Department of Evolution, Ecology, and Organismal Biology, The Ohio State University, USA c School of Anatomical Sciences, University of the Witwatersrand, 7 York Road, Parktown 2193, South Africa d Department of Archaeology, University of Sheffield, Northgate House, West street, S1 4ET, Sheffield, UK article info abstract Article history: Analyses of new cercopithecid fossil specimens from the South African site of Haasgat point to cranio- Received 27 August 2008 facial affinities with the genus Cercopithecoides. Detailed metric and non-metric comparisons with South Accepted 28 August 2010 African Cercopithecoides williamsi, and other East African Cercopithecoides species, Cercopithecoides kimeui, Cercopithecoides meaveae, Cercopithecoides kerioensis, and Cercopithecoides alemyehui demon- Keywords: strate that the Haasgat fossils have distinct craniofacial morphology and dental metrics. Specifically, Cercopithecoides material from Haasgat probably represents one of the smaller Cercopithecoides, differing from the others Haasgat in its particular suite of features that vary within the genus. It is unique in its more vertical ramus, South Africa Pleistocene associated with a relatively lengthened mandibular body. Haasgat Cercopithecoides has a particularly narrow interorbital region between relatively larger ovoid orbits, with articulation of the maxillary bones at a suture above the triangular nasal bones. Furthermore, the maxillary arcade is more rounded than other Cercopithecoides, converging at the M2 and M3. The conclusion drawn from this analysis is that the Pleistocene Haasgat fossils are colobines representing a distinct taxon of Cercopithecoides, Cercopithe- coides haasgati, thus adding a second species of the genus to southern Africa. Ó 2010 Elsevier Ltd. All rights reserved. Introduction Martini, 1990). Haasgat was mined for calcite flowstone in the first quarter of the past century (Keyser and Martini, 1990), with In 1988, excavation of the South African fossil site Haasgat fossiliferous breccia having been dumped down the steep slope yielded a sample of craniofacial fossils with characteristics of the below the cave (Plugh and Keyser, 1994). The material used in this extinct genus Cercopithecoides (Keyser and Martini, 1990; Von study came from these dumps. The breccia is thought to have come Mayer, 1998). These fossils appeared to be different from the from the siltstone beds immediately above the floor flowstone, but representatives of this genus previously found at other South has no accurate in situ context (Keyser and Martini, 1990). African sites, all of which are currently referred to a single species, Keyser and Martini (1990) initially suggested that Haasgat was Cercopithecoides williamsi, and from other members of the genus older than the lower members of Sterkfontein and Makapansgat, from East Africa. Here we present background on the context of the because of the high elevation of the cave and the fact that most of fossils and species of Cercopithecoides. its proximal deposits appeared to have been weathered away. Subsequent faunal analyses have suggested a more recent age. Identified mammalian species from Haasgat show an ambiguous Haasgat cave site, geological age, and paleoenvironment mix of mammals (Table 1). Whereas two of the species are known from sites dating to the Pliocene/Pleistocene border, two others The Haasgat cave is located in the Brits District of the North only appear in the South African fossil record in the late Pleistocene. 0 0 West Province, at 25 51 31"E and 27 50 9"S. It lies on the steep Thus the Haasgat assemblage, if it comes from a single temporal western slope of the Witwatersrand Spruit Valley (Keyser and horizon, likely dates to the Pleistocene, but further resolution of its age is not possible at this time. Certainly there is the potential that the cercopithecid fossils are coeval with South African C. williamsi. * Corresponding author. Department of Anthropology, The Ohio State University, The most abundant species found at Haasgat is Oreotragus major 4034 Smith Laboratory, 174W. 18th Ave. Columbus, OH 43210-11106, USA. Tel.: þ1 (klipspringer [Plugh and Keyser, 1994]). The presence of other 614 292 2745. E-mail addresses: [email protected] (J.K. McKee), k.l.kuykendall@sheffield.ac. browsers such as giraffe and kudu suggests a savannah environment uk (K.L. Kuykendall). in which trees featured prominently (Plugh and Keyser, 1994). The 0047-2484/$ e see front matter Ó 2010 Elsevier Ltd. All rights reserved. doi:10.1016/j.jhevol.2010.08.002 84 J.K. McKee et al. / Journal of Human Evolution 60 (2011) 83e93 Table 1 Table 2 Identified mammalian species from Haasgat. Key published features of Cercopithecoides williamsi. Family Species Temporal frame Craniofacial features Bovidae Oreotragus oreotragus major Pliocene-Pleistocene Calvarium is well rounded, more so in the female Kobus leche Pleistocene-Present Calvarium shorter and higher in female, in the male it is slightly more ovoid Kobus ellipsiprymnus Late Pleistocene-present Temporal lines form a thickened ridge from the supraorbital Alcelaphus buselaphus Late Pleistocene-present torus to the calvarial roof Temporal lines converge but do not meet, and do not Cercopithecidae Papio angusticeps Pliocene/Pleistocene border reach the nuchal crest Post-orbital constriction is marked Procaviidae Procavia transvaalensis Pliocene/Pleistocene border Orbits are relatively large and rounded Based on: Plugh and Keyser (1994); McKee and Keyser (1994). Supraorbital tori are only moderately developed in males, and lie slightly below bregma Supraorbital tori are well developed in females and raised above the calvarium roof presence of waterbuck (Kobus ellipsiprymus) suggests that there was Muzzle is short and narrow relative to that of cercopithecines water in close proximity and Kobus leche, in particular, suggests Muzzle a square outline anteriorly Maxillary dental arcade maximum width at M2 swampy conditions or shallowly inundated flood plains (Smithers, Mandibular corpus is shallow and of constant depth 1983). Thus the proposed palaeoenvironment surrounding Haas- Mandibular fossa absent gat at the time of deposition included montane forest, a nearby river, Foramen symphyseosum presence variable and open woodland or savannah all in close proximity. Dental features Teeth are small and show sexual dimorphism only in the CeP3 complex 3 Review of the genus Cercopithecoides Reduced protocone on P M1 < M2 < M3 in size and breadth Mollett (1947) erected the genus and species C. williamsi for the Based on: Mollett (1947); Freedman (1957); Verheyen (1962); Maier (1970); specimen AD 1326-3 from Makapansgat, a male cranium and Eisenhart (1974); Szalay and Delson (1979); Kuykendall and Rae (2008); Jablonski et al. (2008). mandible. He named the genus Cercopithecoides because of its resemblance to Cercopithecus, and the species C. williamsi in honor of the technician, E.W. Williams, who found the skull. C. williamsi has now been described from the South African sites of Maka- which developed into two cuspules on M3; most notably, all the pansgat, Sterkfontein (Member 4 and the Graveyard site), Swartk- upper molars had a prominent intercusp cleft on their buccal rans, Cooper’s A, Bolt’s Farm, and Kromdraai B, as well as from surface, each with a cingular ridge across its alveolar margin; and Koobi Fora in Kenya (Jablonski et al., 2008), and Leba in Angola larger molars than those of other South African Cercopithecoides, 3 (Szalay and Delson, 1979; Delson, 1984). Other species of Cercopi- particularly in terms of the mesio-distal diameters of M2 and M ; thecoides have been found at Lothagam (Leakey et al., 2003), Koobi After recovery of new material from Makapansgat, Freedman Fora (Jablonski et al., 2008), Hadar (Frost and Delson, 2002), Old- (1960) argued that this new material bridged the gap between uvai Gorge (Leakey, 1982), Rawi (Frost et al., 2003), and Daka C. williamsi and C. molletti both in terms of the size differences and (Gilbert and Frost, 2008). the slight morphological differences. Thus, he synonomized the The colobine affinity of the genus Cercopithecoides was first species C. molletti with C. williamsi. Freedman (1960) also noted proposed by Robinson (1952). Based upon this proposal, Maier a dental size difference between the C. williamsi of, in order from (1970) initially assumed that Cercopithecoides would therefore smallest to largest, specimens from Sterkfontein, Makapansgat and have been adapted for an arboreal lifestyle, akin to modern colo- Swartkrans. These size differences were especially clear for M2 size. bines. However, it is now clear from the post-cranial evidence that However he concluded that the overall size variation of all these East African Cercopithecoides shows strong morphological adapta- lumped together is still less than that within the Papio ursinus tions for terrestrial existence (Birchette, 1981; Leakey, 1982; Frost species and thus not indicative of separate species status. and Delson, 2002; Jablonski et al., 2008). Carbon isotope studies Delson (1983) noted that the Kromdraai C. williamsi
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