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Faunal Assemblage Seriation of Southern African Pliocene and Pleistocene Fossil Deposits

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Faunal Assemblage Seriation of Southern African Pliocene and Pleistocene Fossil Deposits AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 96235-250 (1995) Faunal Assemblage Seriation of Southern African Pliocene and Pleistocene Fossil Deposits JEFFREY K. McKEE, J. FRANCIS THACKERAY, AND LEE R. BERGER Department of Anatomy and Human Biology, University of the Witwatersrand, Johannesburg (J.K.M, L.R.B.) and Department of Palaeontology, Transvaal Museum, Pretoria (J.F.T.),South Africa KEY WORDS Faunal seriation, Southern African biochronology ABSTRACT Fossil assemblages from the Pliocene and Pleistocene of southern Africa were seriated in order to give a better idea of their relative chronology. Time-sensitive mammals were selected for calculation of the Fau- nal Resemblance Index among 17 site units. On the basis of a logistical seriation and subsequent site analysis, the following sequence of sites was deemed most probable: Makapansgat Member 3, Makapansgat Member 4, Taung Dart deposits, Sterkfontein Member 4 and Taung HrdliEka deposits, Sterkfontein Member 5 (in part) and Kromdraai B, Kromdraai A and Swart- krans Member 1, Swartkrans Member 2, Swartkrans Member 3, Plovers Lake, Cornelia, Elandsfontein Main Site, Cave of Hearths Acheulian levels, Florisbad and Equus Cave and Klasies River Mouth. o 1995 Wiley-Liss, Inc. Associations of the early hominid fossils and Partridge, 1983; Vogel, 1985) led to the with a wealth of Pliocene and Pleistocene suggestion that the Taung child may have fauna from southern Africa provide vital been close to the hypodigm of A. robustus clues to the past environments and evolu- (Tobias, 1973,1975,1978; Wolpoff, 1974) or tionary processes that shaped human evolu- that it represented a derived form of A. afri- tion. Due to the nature of the cave deposits canus (Tobias, 1991a,b). Recent faunal dat- in which these ancient assemblages are typ- ing of Taung (Delson, 1988; Cooke, 1990; ically found, where absolute geochronologi- McKee, 1993a), placing the site in time cal dating techniques are inhibited, the ab- closer to Sterkfontein and Makapansgat, solute and even relative temporal positions opens the possibility that the Taung homi- of these sites have proved difficult to ascer- nid is a more typical or even primitive repre- tain. Nevertheless, it is of pivotal impor- sentative ofA. africanus. tance in studies of mammalian evolution, Turner (1983, 1990) and Vrba (1985a, and particularly in paleoanthropological re- 1988,1992) have argued that changes in glo- search, to establish a chronology with which bal climate during the Pliocene caused we can assess the evolutionary roles of the mammalian evolutionary changes in Africa. varied fauna found in southern African cave Recently Hill et al. (1992) have suggested deposits. that the origin of Homo corresponded to Understanding the timing of deposition in these putative climatic changes on the basis southern African hominid fossil sites is of of the age of a specimen they feel represents continuing importance, as illustrated by the the genus. However, in southern Africa it is influence dates have on interpretations of hominid phylogeny and theories of the causes of evolution. For example, the pro- Received August 21,1993; accepted August 17,1994. posed late date for the type specimen ofdus- Address reprint requests to Dr. J.K. McKee, Department of Anatomy and Human Biology, University of the Witwatersrand tralopithecus africanus from Taung, at circa Medical School, 7 York Road, Parktown, 2193 Republic of South 1 mya (Partridge, 1973, 1982, 1985; Vogel Mrica. 0 1995 WILEY-LISS, INC. 236 J. K. MCKEE ET AL. not clear whether the apparent faunal turn- referred to as Sterkfontein Member 5), overs seen in the fossil record predate or Swartkrans, and Kromdraai. The Vaal-Cor- postdate any climatic change, or whether or nelia Faunal Span included the Vaal not some of the faunal turnovers are indeed Younger gravels, Elandsfontein (formerly real (McKee, 1991, in press). Thus an under- known as Hopefield or Saldanha), and Cor- standing of cause and effect in mammalian nelia, followed by the Florisbad-Vlakkraal evolution requires at least an understand- Faunal Span encompassing the important ing of the succession of sites if not the actual Homo-bearing sites of Florisbad and Broken age of deposition. Hill. Opportunities to determine the ages of As progress is made in both excavation sites by geological dating have been limited and taxonomic identification of southern Af- by the nature of cave deposits. Sedimento- rican fossil fauna, sufficient samples are logical correlations of sites (Butzer, 1974), emerging for the refinement of the early nick-point analysis of cave openings (Par- methods that simply recognized faunal tridge, 19731, sedimentological rates (Par- spans. Toward this end, biochronologies tridge, 1985), and isotopic techniques (Vogel within taxonomic families have been estab- and Partridge, 1983; Vogel, 1985) have pro- lished for bovids (Vrba, 1975, 1982,1985b1, vided unsatisfactory results (De Swardt, cercopithecids (Delson, 1984, 1988), carni- 1974; Butzer et al., 1978; Tobias et al., 1993; vores (Hendey, 1974; Turner, 1990), and McKee, 1993a). Palaeomagnetic chronolo- suids (Harris and White, 1979). A combined gies have given mixed results, with some picture of all the known mammalian fauna success at Makapansgat (Brock et al., 1977; may then be useful in the seriation of indi- McFadden et al., 1979; McFadden, 1980) vidual site units within and between the and Kromdraai (Jones et al., 19861, but few faunal spans. clues at other sites due to the dynamics of There are numerous factors that may lead cave sediments. Electron spin resonance to similarities or differences of faunal repre- (ESR) has provided age estimates for vari- sentation in site assemblages that must be ous sites, including Border Cave (Grun et considered when attempting to establish al., 1990a), Klasies River Mouth (Grun et biochronologies across taxonomic bound- al., 1990b), and Sterkfontein (Schwarcz et aries. Temporal determinants, those most al., 1993), but these are dependent on as- useful for faunal dating, interplay with sumptions regarding rates of uptake of ura- other sampling biases so that only a fraction nium (Griin and Stringer, 1991). Neverthe- of the total faunal community living at any less, there is considerable potential in the one time would be represented in a cave de- application of ESR dating with necessary ca- posit. A summary of the key factors that veats and corroborating evidence from other may lead to similarities or differences techniques. among faunal assemblages in southern Afri- Chronologies based on faunal correlations can cave deposits can be found in Table 1. have thus been the primary means of dating Although faunal dating may have its limi- southern African fossil sites. Many attempts tations due to the complex network of fac- have been made (reviewed by McKee, tors that determine site assemblages, it re- 1993a), but the most relevant work has its mains as the most reliable tool for cornerstone in the faunal spans delineated determining the sequence of fossil sites in by Ewer and Cooke (1964) and Cooke (1967). southern Africa. Our objective here is to es- The oldest of these temporal associations tablish a chronological framework for un- was the Sterkfontein Faunal Span, compris- derstanding mammalian evolution in the ing the Australopithecus-bearingdeposits of Pliocene and Pleistocene of southern Africa Sterkfontein, Makapansgat, and Taung, as through the seriation of site units. In order well as the Older Gravels of the Vaal River to achieve this with a total mammalian per- Basin. The subsequent Swartkrans Faunal spective, issues arising from the complicat- Span encompassed deposits from the Sterk- ing factors discussed above must be resolved fontein extension site (most of which is now with an appropriate method. SOUTHERN AFRICAN FAUNAL ASSEMBLAGE SERIATION 237 TABLE 1. Factors causing similarities or differences in sampled or faunally diverse site is more faunal representation at southern African caue sites likely to yield species identified from else- Environmental factors where than is a site with a paucity of fauna. Faunal community Floral community Simpson’s (1960) Faunal Resemblance In- Demography dex (FRI) provides a partial solution to the Climate sampling problem and is employed here. For Altitude Terrain each pair of sites, the FRI is calculated by Geographic proximity of sites taking the number of species shared (C) di- Availability of water Cave size and depth vided by the total number of species found in Taphonomic factors the smaller sample (N1),or simply FRI = C/ Taphonomic agents Nl x 100. These values are displayed in Ta- Bone preservation Mixing of deposits ble 3B. The advantage of the method is that Interpretive factors a small sample can exhibit an FRI value Selective excavation and preparation comparable to that of a much larger sample, Taxonomic biases Temporal factors thus standardizing the figures in the ma- Origins trix. It should be noted, however, that a site Radiations Extinctions with a large sample still has a greater prob- ability of sharing species with other sites and the use of an FRI does not address this issue. METHODS AND ASSUMPTIONS Complications from nontemporal factors Lists of identified mammalian species and of site similarities can cause spurious corre- genera were compiled from published litera- lations. For example, a long-lived but ecolog- ture for 23 Neogene site units (Fig. l and ically limited species, Hippopotamus am- Table 2). This amounted to 246 species of phibius, appears at the oldest and youngest 159 genera and 44 families. sites, but not at sites that span the time in It was our primary goal to seriate the sites between. Thus the analysis must be limited based on similarities of identified species be- to the species that are most sensitive to time tween site unit assemblages. The initial step and not restricted to highly specialized envi- is to create a matrix exhibiting the number ronments. of identified species shared between each Evaluation of temporally sensitive species pair of site units (Table 3A); the diagonal can be a somewhat subjective task.
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