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Areas 1- Ern Africa Kroeber Anthropological Society Papers, Nos. 71-72, 1990 Diet, Species Diversity and Distribution of African Fossil Baboons Brenda R. Benefit and Monte L. McCrossin Based on measurements ofmolarfeatures shown to befunctionally correlated with the proportions of fruits and leaves in the diets ofextant monkeys, Plio-Pleistocenepapionin baboonsfrom southern Africa are shown to have included more herbaceous resources in their diets and to have exploited more open country habitats than did the highlyfrugivorousforest dwelling eastern African species. The diets ofall species offossil Theropithecus are reconstructed to have included morefruits than the diets ofextant Theropithecus gelada. Theropithecus brumpti, T. quadratirostris and T. darti have greater capacitiesfor shearing, thinner enamel and less emphases on the transverse component ofmastication than T. oswaldi, and are therefore interpreted to have consumed leaves rather than grass. Since these species are more ancient than the grass-eating, more open country dwelling T. oswaldi, the origin ofthe genus Thero- pithecus is attributed tofolivorous adaptations by largepapionins inforest environments rather than to savannah adapted grass-eaters. Reconstructions ofdiet and habitat are used to explain differences in the relative abundance and diversity offossil baboons in eastern andsouthern Africa. INTRODUCTION abundance between eastern and southern Africa is observed for members of the Papionina (Papio, Interpretations of the dietary habits of fossil Cercocebus, Parapapio, Gorgopithecus, and Old World monkeys have been based largely on Dinopithecus). [We follow Szalay and Delson analogies to extant mammals with lophodont teeth (1979) in recognizing two tribes of cercopithe- (Jolly 1970; Napier 1970; Delson 1975; Andrews cines, Cercopithecini and Papionini, and three 1981; Andrews and Aiello 1984; Temerin and subtribes of the Papionini: Theropithecina (gela- Cant 1983). With the exception of Jolly's (1972) das, fossil and modern), Macacina (macaques, study of Theropithecus oswaldi, such inter- fossil and modern) and Papionina (baboons, pretations have focused on the origins of drills, mandrills and mangabeys, fossil and mod- Cercopithecoidea with relatively litde attention ern)]. Eighty-four percent of the fossil monkeys given to more recent dietary diversification widtin in southern Africa are papioninans (Freedman the superfamily. 1976), while only 10% of those in eastern Africa In this study the dietary habits of Plio- are members of this subtribe. In southern Africa Pleistocene cercopithecine monkeys from fossil at least nine species ofpapioninan monkeys have sites in eastern and southern Africa are recon- been recovered, but only three species are known structed on the basis of dental features shown to to have occurred in eastern Africa (although the be functionally correlated to diet among extant actual number is presendy indeterminable due to cercopithecoids (Kay 1977, 1978, 1981, 1984; the fragmentary and incomplete nature ofthe ma- Kay and Covert 1984; Kay and Hylander 1978; terial). Reconstructions of the dietary habits of Benefit 1987). A major concern ofthe study is to these extinct animals, in combination with infor- better understand the differences in the patterns of mation about the habitats in which they lived and species diversity and the relative abundance of studies of food consumption and habitat use by cercopithecine monkeys which existed in the eas- living mammals, are used to describe a new sce- tern and southern regions of the African continent nario about the evolution, diversity, distribution during the Plio-Pleistocene. and relative abundance of fossil Theropithecus In cave deposits of southern Africa Thero- and other baboons during the Plio-Pleistocene. pithecus is rare, comprising 7% of the total cercopithecoid fauna collected prior to 1976 (Freedman 1976). In contrast, 84% of the mon- MATERIALS AND METHODS keys collected at the Omo and 85% of those collected at Koobi Fora, both in eastern Africa, Fossil cercopithecine monkeys were sampled prior to the same date belong to the genus Thero- from Plio-Pleistocene deposits in southern Africa pithecus (Eck 1976; Leakey 1976). In addition, (Sterkfontein Member 4, Kromdraai Members A Theropithecus is more diverse in deposits from and B, Taung, and Swardcrans Member 1) and eastem Africa, with five species occurring in the eastern Africa (Laetoli and Olduvai, Tanzania; fossil deposits, as opposed to only two in south- Olorgesailie and Koobi Fora, Kenya -- Areas 1- ern Africa. The opposite pattem ofdiversity and 203; Omo, Ethiopia -- Usno Formation, Shun- 78 gura Formation Members B- Table 1. Numbers of fossil cercopithecine molars measured. H, and Kalam Area). Spe- L = Lower, U = Upper. cies sampled are listed by deposit in Table 1. [A EAST AFRICA complete list of specimens LM1 LM2 LM3 UMI UM2 UM3 sampled is given in Benefit Cercocebus sp., KOOBI FORA 5 T7 1T 5 (1987).] Taxonomic identi- fications for the southern Cercocebus ado, OLDUVAI 1 1 1 African fossils are largely Parapapio ado, LAETOLI 5 8 10 4 8 5 based on Freedman (1957, 1961a, 1961b, 1965, Parapapio jonesi, KANAPOI 1 1 1 1976), Freedman and Brain Parapapio sp., KOOBI FORA 2 3 6 2 2 2 (1972), Freedman and Stenhouse (1972), Maier Papionini indet, BARINGO 1 (1971a, 1971b) and Eisen- Papio sp., OLDUVAI 2 3 1 2 hart (1974). Identification Theropithecus oswaldi, of fossil monkeys from KOOBI FORA & ILERET (combined) 13 61 40 12 44 17 eastern Africa are based on AREA 1 (1.5-1.6 my) 3 1 descriptions by R.E. Leakey AREA 8 (1.5-1.6 my) 8 1 2 8 1 AREA 103 (1.6-1.7 my) 5 1 1 (1969), Jolly (1970, 1972), AREA 10 (1.7-1.9 my) 3 1 1 1 M.G. Leakey (1976, 1982), AREA 123 (1.7-1.9 my) 3 5 Leakey and Leakey (1973a, AREA 130 (1.8-1.9 my) 3 8 5 1 4 2 Eck AREA 104 (1.7-2.0 my) 2 4 3 1 1 1973b, 1976), (1977), AREA 106 (2.0 my) 1 Eck and Howell (1982), AREA 116 (2.0 my) 1 2 1 1 1 Eck and Jablonski (1984) and Delson OMO SHUNGURA FORMATION and Leakey Member G 4 (1987). Member H 1 Biostratigraphic dating of Kalam area (Members J-L) 1 the southern African cave deposits indicates that Ma- OLDUVAI 2 11 12 4 8 8 kapansgat is the oldest site OLORGESAILIE 39 35 23 35 25 14 at approximately 3.0 million Theropithecus brumpti years (my), followed by KOOBI FORA/TULU BOR Sterkfontein (3.2-2.5 my AREA 117 &204 (3.3 my) 4 2 6 1 2 3 and 1.75-1.4 my), Krom- AREA 203 (3.3 my) 2 6 6 1 draai (2.7-1.8 my), Taung OMO, SHUNGURA FORMATION (2.3-1.0 my) and Swart- Member C krans (1.8-1.6 my and Member D 2 1.25-0.9 my) (Vrba 1982). Theropithecus quadratirostris Eastern African deposits are OMO USNO FORMATION more securely dated on ra- Member 11 1 diometric grounds. Laetoli is considered to be 3.8-3.5 my (Leakey et al. 1976; SOUTHERN AFRICA Leakey and Hay 1982), the Ml LM2 LM3 UMi UM2 UM3 Usno and Shungura Forma- Dinopithecus ingens, SWARTKRANS 3 5 7 4 7 5 tions at the Omo range in age from 2.9-1.3 my Gorgopithecus major, KROMDRAAI 1 3 4 3 4 (Shuey et al. 1974; Brown Papio angusticeps, KROMDRAAI 2 2 2 1 2 2 and Nash 1976; Brown et TAUNG 3 1 2 2 al. 1985), Koobi Fora and Papio robinsoni, SWARTKRANS 6 10 10 5 15 10 Ileret Formations range in Parapapio ionesi, SWARTKRANS 2 1 2 1 2 2 age from 3.3-1.5 my (Feibel 1 et al. 1989), Olduvai from STERKFONTEIN 3 7 3 4 5 2.2-0.6 my (Leakey and Parapapio whitei, STERKFONTEIN 2 4 4 2 2 2 Hay 1982) and Olorgesailie from 1.3-0.5 my (Potts Parapapio broomi, STERKFONTEIN 3 4 3 2 5 3 1989). Because of the long Theropithecus darti, SWART[RANS 2 4 2 3 79 time ranges represented at mostt of the deposits, grinding takes place during the second phase of samples of monkeys were examiiined according to occlusion as the jaw moves lingually and mesi- their stratigraphic unit of proveinience whenever ally, with the protoconid and hypoconid coming possible. into direct contact with the protocone and hypo- Dietary estimates for the fossil species are cone as a result (Crompton and Hiiemae 1970; based on the relationship betwreen molar mor- Kay and Hiiemae 1974; Kay 1975, 1978). The phology and diet in nine extarit species which tendency for monkeys that include large amounts have been studied extensively in the wild and for of leaves in their diets to have longer shear crests which the proportions of leaves and fruits in the than monkeys which eat greater amounts of fruits annual diet are known (Table 2). The species and seeds has been demonstrated by Kay (1975, considered were Colobus baditus (n=33), Colo- 1978, 1981; Kay and Covert 1984). bus guereza (n=22), Colobus satanas (n=2), In this study eight shear crest lengths for Presbytis melalophos (Kay anId Covert 1984), each upper and lower molar were measured using Presbytis obscura (Kay and Cc vert 1984), Cer- an ocular micrometer mounted in a stereoscopic cocebus galeritus (n=20), Cercocebus albigena microscope (Figure IA). Shearing crest develop- (n=25), Macaca nemestrina (n,=6) and Macaca ment was then appraised in four ways: 1) from fascicularis (n=34). Additional comparative data calculation of Kay's (1984) shear quotient (SQ) were taken from Kay (1978, 1981; Kay and for lower second molars; 2) from the sum of Covert 1984) and Benefit (198' 7). Dental termi- shear crests gauged against molar length (abbre- nology and measurements usted in this paper viated PERS); 3) from the sum of lingual (for follow Jolly (1972), Delson (1')73), Kay (1978, lower molars) or buccal (for upper molars) shear 1981) and Benefit (1987).
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