The context of Stw 573, an early hominid skull and skeleton from Sterkfontein Member 2: taphonomy and paleoenvironment more

Journal of Human Evolution 46 (2004) 279–297 The context of Stw 573, an early hominid skull and skeleton from Sterkfontein Member 2: taphonomy and paleoenvironment Travis Rayne Pickeringa,b,c*, Ron J. Clarkec, Jason L. Heatona,b,c b a Department of Anthropology, Indiana University, 130 Student Building, Bloomington, IN 47405, USA Center for Research into the Anthropological Foundations of Technology (CRAFT), Indiana University, Bloomington, IN 47405, USA c Sterkfontein Research Unit, University of the Witwatersrand, WITS 2050, Johannesburg, South Africa Received 25 July 2003; accepted 3 December 2003 Abstract The reconstructed taphonomic and paleoenvironmental contexts of a ca. 4 million-year-old partial hominid skeleton (Stw 573) from Sterkfontein Member 2 are described through presentation of the results of our analyses of the mammalian faunal assemblage associated stratigraphically with the hominid. The assemblage is dominated by cercopithecoids (Parapapio and Papio) and felids (Panthera pardus, P. leo, Felis caracal, and Felidae indet.), based on number of identified specimens, minimum number of elements and, minimum number of individuals. In addition, the assemblage is characterized by a number of partial skeletons and/or antimeric sets of bones across all taxonomic groups. There is scant indication of carnivore chewing in the assemblage. These observations, in addition to other taphonomic data, suggest that the remains of many animals recovered in Member 2 are from individuals that entered the cave on their own—whether accidentally by falling through avens connecting the cave to the ground surface above or by intentional entry—and were then unable to escape, rather than primarily through systematic collection by a biotic, bone-accumulating agent. The taphonomic conclusion that animals with climbing proclivities (i.e., primates and carnivores) are preferentially preserved over other taxa, ultimately because of those proclivities, urges caution in assessing the fidelity of the assemblage for reconstruction of the Member 2 paleoenvironment. With that caveat, we infer that the Member 2 paleoenvironment was typified by rolling, rock-littered and brush- and scrub-covered hills, indicated by the abundant F. caracal and cercopithecoid fossils recovered and the identified presence of the extinct Caprinae Makapania broomi. In addition, the valley bottom may have retained standing water year-round, perhaps supporting some tree cover—a setting suitable for the well-represented ambush predator P. pardus and suggested by the presence of Alcelaphini. Finally, the reconstructed taphonomic and paleoenvironmental settings of Sterkfontein Member 2 are compared to penecontemporaneous sites in South and East Africa.  2004 Elsevier Ltd. All rights reserved. Keywords: Australopithecus; Sterkfontein; Paleontology; Taphonomy; Paleoenvironment * Corresponding author. Tel.: +1-812-856-5260; fax: +1-812-855-4358 E-mail address: trpicker@indiana.edu (T.R. Pickering). 0047-2484/04/$ - see front matter  2004 Elsevier Ltd. All rights reserved. doi:10.1016/j.jhevol.2003.12.001 280 T.R. Pickering et al. / Journal of Human Evolution 46 (2004) 279–297 Introduction The skull and associated skeleton of a Pliocene hominid (Stw 573) from Member 2 of Sterkfontein Cave (South Africa) is remarkable for its completeness, for its estimated great age (ca. 4 million-years-old [myr]; Partridge et al., 2003), and for the unusual events leading to its discovery (Clarke, 1998, 1999, 2002a; Clarke and Tobias, 1995). In addition, although further study is required, Stw 573 may represent a species of Australopithecus hitherto unknown in southern Africa (Clarke, 1998; White, 2002). Here we describe the taphonomic and paleoenvironmental contexts of Stw 573, through presentation of the results of our analyses of the mammalian faunal assemblage associated stratigraphically with the hominid. We emphasize that our analyses are the first to be based strictly on fauna recovered from excavation of in situ Member 2 breccia. This excavation was initiated by our team in 1997 and conducted in the eastern end of the Silberberg Grotto, a deep underground cavern at Sterkfontein (Fig. 1). Member 2 contains multiple facies because a large stalagmite was forming in the cavern during the later stages of the accumulation of the deposit, effectively dividing that unit in places (Partridge, 2000; Clarke, 2002b). The flowstone sealed the Stw 573 skeleton, which was already buried on the western face of the Member 2 talus cone. Thus, the skeleton is separated spatially from the abundance of other fossils preserved in Member 2 at the eastern end of the cavern, but is still part of the same deposit. Except for that of Turner (1997), previous taxonomic studies of purported Member 2 fauna (e.g., Broom, 1945a,b; Broom and Schepers, 1946; McKee, 1996) were based on fossils recovered from dumps created by lime miners during their operations in the Silberberg Grotto. However, it is important to note that the Silberberg Grotto also contains the fossil-bearing breccia of Member 3, in addition to that of Member 2 (Partridge, 1978; Partridge and Watt, 1991). Thus, earlier taxonomic lists for Member 2 (and arguments based on those lists) must be regarded with caution. Taxonomy An updated taxonomic list for Sterkfontein Member 2 is provided in Table 1. In this section, we preliminarily describe better preserved examples of newly recorded Member 2 species from the Silberberg Grotto (excluding the Australopithecus skeleton, Stw 573, and the partial Chasmaporthetes nitidula skull, S94-13225, which have been initially described elsewhere; Clarke and Tobias, 1995; Turner, 1997; Clarke 1998, 1999). Systematic paleontology of new records Family Cercopithecidae Cercopithecoides williamsi Mollett, 1947 Extinct colobus monkey Material Un-numbered mandible (Fig. 2). Description The specimen includes both sides of the mandible. The left horizontal ramus is still partially embedded in the hard breccia matrix so that only the lingual surfaces of the right M2 and M3 are currently visible (the left incisors are also observable); the ascending ramus of the left side is missing. The right dentition and right side of the mandible are complete except for the missing ascending ramus. The specimen is crushed medially so that the opposing horizontal rami are in contact and the right side is levered ventrocaudally relative to the left side. However, we are confident in the taxonomic assignment of the specimen because the corpus of the mandible is relatively shallow, with a marked lateral ridge, and the teeth are relatively small, with V-shaped buccal clefts, hypsodont cusps, and large central foveae on the molars (non-metric characteristics of C. williamsi are reviewed in Jablonski, 2002). Parapapio jonesi Broom, 1940 Extinct papionini Material SWP-2914, left maxilla fragment with M2–M3; SWP-2931, partial basicranium and lower splanchnocranium; SWP-2935, right mandible fragment ¯ with C–M2; SWP-2936, right mandible fragment T.R. Pickering et al. / Journal of Human Evolution 46 (2004) 279–297 281 Fig. 1. South-north section through the Sterkfontein Formation showing the relative position of Member 2 within the Silberberg Grotto (adapted from Partridge and Watt, 1991; Partridge, 2000). with P3–M3; SWP-2938, mandible fragment with right M2–M3 and left P3–M3; SWP-2947, nearly complete cranium, with partially exposed endoI cranial cast and right C–M3 and left I1, C–M3. I Description All specimens are illustrated in Fig. 3. We diagnosed Parapapio cranial specimens based on the presence of straight nasal profiles and short 282 T.R. Pickering et al. / Journal of Human Evolution 46 (2004) 279–297 Table 1 Taxonomy of the Silberberg Grotto fossil fauna Taxon Primates Hominidae Australopithecus sp. Cercopithecidae Papio izodi Parapapio jonesi Parapapio broomi Cercopithecoides williamsi Carnivora Hyaenidae Chasmaporthetes silberbergi Chasmaporthetes nitidula Hyaenidae indet. Felidae Dinofelis barlowi Megantereon cultridens Felis caracal Panthera pardus cf. Panthera leo Acinonyx jubatus Artiodactyla Bovidae Alcelaphini indet. Makapania broomi Non-Alcelaphini Size Class 2d a Member 2a Silberberg Grottob from Pp. broomi based on the smaller overall size and relatively larger canines and premolars of the former taxon. + + +c + + + + Family Felidae cf. Panthera leo Linnaeus, 1758 Lion Material SG 97-68, left distal humerus (Fig. 4). Description The specimen is broken into two major pieces just superior to the metaphysis and displays many pre- and post-fossilization cracks, chips and breaks over much of it. In relation to humeri of both extinct and extant large African felids, it compares most favorably to lions in the following features used by Turner (1987: 338) to distinguish P. leo from Megantereon: overall size, the angle of penetration of the epicondylar foramen, the depth of the radial fossa, the relative development of the ridge that is a distal extension of the deltoid ridge, and the size of the distal articular surface relative to the whole epicondylar region. + + + + + + + + + + + + + + Taxa recovered confidently from Member 2 during the 1997–current excavation. Each new taxonomic record for the Silberberg Grotto fauna, based on the current study and that of Turner (1997), is indicated with a bold +. b Taxa recovered from Member 2 and possibly Member 3 deposits during previous excavations and examination of dump materials. Annotated taxonomic lists and relevant references for these earlier described materials appear in Broom (1945a,b), Broom and Schepers (1946), McKee (1996), and Turner (1997). c Berger et al. (2002: 194) state incorrectly in Table 1 of their publication that Pp. jonesi has been previously recorded from Member 2. Our identification of this species in the Member 2 assemblage is actually the first record of the taxon from the Silberberg Grotto. d Based on SG 97-119, a fragment of a Size Class 2 bovid hemimandible (Brain, 1974; Brain, 1981), with the single cusp of a lower molar. The occlusal surface of the molar is absent, but the lingual lobe is clearly pointed—a feature uncharacteristic of the Alcelaphini, but characteristic of the Tragelaphini and Antilopini. Family Bovidae Alcelaphini indet. Medium-size alcelaphine Material SG 97-100, fragmentary left maxilla, with M1–M2 (Fig. 5). Description The maxillary bone portion of the specimen is encased in hard breccia, with the exposed (but badly damaged and displaced) molars displaying the very rounded lingual lobes and central cement cavity characteristic of the Alcelaphini (reviewed in Gentry, 1978). Makapania broomi Wells and Cooke, 1956 Extinct Caprinae muzzles (reviewed in Jablonski, 2002). In addition, the Parapapio supraorbital torus is slight, if present at all. Finally, Pp. jonesi was distinguished T.R. Pickering et al. / Journal of Human Evolution 46 (2004) 279–297 283 Fig. 2. Cercopithecoides williamsi mandible. Right horizontal ramus (with I1–M3 visible) is seen in the foreground compressed medially against the left side, with its M2 and M3 apparent. The specimen is still cemented in its breccia matrix, and the large tooth partially visible in the background above the Cercopithecoides right dentition is a Chasmaporthetes carnassial. Material SG 97-106, articulated foot bones, including distal end of proximal phalanx, complete intermediate phalanx, both distal phalanges and two sesamoids (Fig. 6). Description M. broomi belongs to the subfamily Caprinae, and diagnostic caprine morphology is evident in the distal phalanx of SG 97-106, including: an underdeveloped processus extensorius, a dorsal rim that is convex in outline and mediolateral compression of the bone distal to the articular surface (Brink, 1999). Taphonomy Skeletal part representation No taxon from the Member 2 assemblage (except possibly Australopithecus) is represented by an anatomically complete skeleton (Tables 2 and 3). However, primates and felids display the widest range of different skeletal parts; primates are, in part, represented by articulating bone specimens and the felid subassemblage preserves antimeric specimens (Fig. 7). Interestingly, there are additional articulating specimen-sets in the bovid subassemblage as well. 284 T.R. Pickering et al. / Journal of Human Evolution 46 (2004) 279–297 Fig. 3. Parapapio jonesi cranial and mandibular specimens: (a) SWP-2947; (b) SWP-2931; (c) SWP-2914; (d) SWP-2938; (e) SWP-2936; (f) SWP-2935. Several other Plio-Pleistocene fossil assemblages from the Sterkfontein Valley that also preserve high numbers of primates and felids have been interpreted, at least in part, as carnivore-collected (e.g., Sterkfontein Member 4: Brain, 1981; Pickering et al., in press; Swartkrans Hanging Remnant, Lower Bank and Member 2: Brain, 1981, 1993; Pickering, 2001a; Carlson and Pickering, 2003; Carlson and Pickering, in press; Pickering and Carlson, 2002; Kromdraai B: Vrba, 1981; Vrba and Panagos, 1982). In terms of skeletal part representation, the primate subassemblage from Sterkfontein Member 2 most closely resembles a carnivore refuse assemblage (i.e., an assemblage composed of those bone specimens not ingested by feeding carnivores), rather than a carnivore scat assemblage (i.e., an assemblage composed of those bone specimens ingested and voided by feeding carnivores) (Pickering, 2001a,b; Carlson and Pickering, 2003). Fig. 8 illustrates the close fit in pattern (although not in absolute frequencies) between Member 2 and a modern refuse assemblage, composed of bones from ten complete baboon carcasses fed to leopards and spotted hyenas; there is agreement between the assemblages in high skull representation, low axial element survivorship, peaks in limb bone representation, and a paucity or absence of the bones of the hand and feet. T.R. Pickering et al. / Journal of Human Evolution 46 (2004) 279–297 285 Fig. 4. cf. Panthera leo left humerus (SG 97-68): anterior (left); posterior (right). Bone surface modifications Bone surface damage is an additional line of evidence with which to assess the hypothesis that the Member 2 primate subassemblage was collected primarily by carnivores. Identification of bone surface modifications was undertaken by inspecting each fossil specimen under a strong oblique light source, with the aid of at least 10 magnification, as recommended by several analysts (e.g., Bunn, 1981, 1991; Bunn and Kroll, 1986; Blumenschine and Selvaggio, 1988, 1991; Blumenschine and Marean, 1993; Blumenschine, 1995; Blumenschine et al., 1996). Carnivore tooth marks are the major types of biotic damage observed on the Member 2 fossils, and include both conspicuous and inconspicuous tooth scores (Blumenschine et al., 1996), pits, punctures, and crenulation. No tooth notches or chipped back edges were observed (damage types reviewed in Pickering and Wallis, 1997). There are only two isolated primate dental specimens in the assemblage. Unlike isolated teeth, the remaining fossil specimens represent skeletal portions (bone-only specimens or specimens that are composed of both teeth and tooth-bearing bony structures—i.e., partial crania and mandibles) that are consistently tooth-marked in high frequencies in modern assemblages of carnivoreravaged primate bones (e.g., Simons, 1966; Brain, 1981; Haglund et al., 1988, 1989; Haglund, 1997; Pickering, 2001a,b). A scoring system of the 286 T.R. Pickering et al. / Journal of Human Evolution 46 (2004) 279–297 Fig. 5. Alcelaphini indet. left M2 (SG 97-100). T.R. Pickering et al. / Journal of Human Evolution 46 (2004) 279–297 287 Fig. 6. Makapania broomi partial foot skeleton (SG 97-106): sesamoids (top); distal phalanges (bottom left); intermediate phalanx (middle); proximal phalanx (right). relative integrity of non-dental bone surfaces (Pickering, 1999; Pickering et al., 2000) was applied to the Member 2 fossils. This system classifies bone specimens with minor cortical exfoliation (comparable to Stage 0 through Stage 2 in Behrensmeyer’s [1978] surface weathering scheme) as fair to high integrity. Results indicate that 82% of the primate total number of identified specimens (NISP=121) have fair to high integrity bone surfaces. Thus, a majority of the primate fossils from Member 2 possess bone surfaces which are suitably preserved to maintain discernable modifications had any modifications been inflicted on those specimens in the biostratinomic phase. However, only two specimens display certain carnivore chewing damage, while five others display probable chewing damage. Thus, at best, only 6% (n=7) of the total primate NISP is damaged by carnivore teeth. This low percentage of toothmarked primate specimens is likely reflective of a low frequency of chewing damage in the original deposited assemblage. In support of this assertion is the observation that the remainder (non-primate portion) of the identifiable (n=137) and nonidentifiable (n=68) Member 2 assemblage preserves only six other possibly tooth-marked specimens (76% of this portion of the faunal assemblage is scored as fair to high integrity). 288 T.R. Pickering et al. / Journal of Human Evolution 46 (2004) 279–297 Table 2 Skeletal part frequencies (NISP/MNE/MNI) for non-hominid primates, carnivores, and bovids in the Member 2 fossil assemblagea Skeletal part Cranium Mandible Vertebrae Sacrum Rib Clavicle Humerus Radius Ulna Radioulna Carpals Metacarpal Os Coxae Femur Tibia Tarsals Metatarsal Metapodial Proximal Phalanx Intermediate Phalanx Distal Phalanx Total a Cercopithecidae b b Hyaenidae 1/1/1 1/1/1 Small Felidae 2/2/2 3/2/2 Medium Felidae 2/2/2 Large Felidae 2/1/1 2/1/1 6/6/– Small Bovidae 1/1/1 Large Bovidae 2/1/1 5/5/1 2/2/2 3/3/1 1/1/1 8/7/6 1/1/1 1/1/1 5/4/4 2/2/1 2/1/1 5/3/2 2/2/1 3/2/1 1/1/– 6/6/3 1/1/1 3/2/2 1/1/1 5/5/– 6/6/3 8/8/– 13/13/– 5/5/– 6/6/– 75/69/3 (7%) 3/3/3 2/2/1 1/1/1 5/3/2 1/1/1 1/1/1 1/1/1 8/7/4 9/6/4 7/4/3 5/5/5 3/3/1 4/4/– 1/1/1 3/3/– 1/1/1 3/2/2 2/2/2 2/2/1 3/2/2 1/1/1 1/1/1 3/3/2 4/4/1 4/4/2 4/4/1 (2%) 24/21/3 (7%) 58/50/28 (66%)b 6/6/1 (2%) 21/18/4 (10%) 7/5/2 (5%) NISP=number of identified specimens; MNE=minimum number of elements; MNI=minimum number of individuals. Highest MNI for each taxon is indicated in the Total row, and that group’s proportion of the total mammal MNI (=42) is indicated parenthetically. Cercopithecidae=craniodentally-identified Papio izodi, Parapapio jonesi, Pp. broomi, Cercopithecoides williamsi; Hyaenidae=craniodentally-identified Chasmaporthetes nitidula; Small Felidae=craniodentally-identified Felis caracal; Medium Felidae=craniodentally-identified Panthera pardus; Large Felidae=craniodentally-identified Panthera leo; Small Bovidae=craniodentally-identified Size Class 2 non-alcelaphine; Large Bovidae=craniodentally-identified Alcelaphini indet. and Makapania broomi. b For details on Cercopithecidae skull bone counts, see Table 3. Other indications of assemblage formation There are additional taphonomic indications that the Member 2 faunal assemblage does not represent primarily a carnivore-collected accumulation. While there is a high percentage of carnivore individuals in the assemblage, none of dependent juvenile age is represented—the presence of which can be suggestive of a prehistoric den site (e.g., Cruz-Uribe, 1991; Klein et al., 1991; Stiner, 1991; Brugal et al., 1997; Pickering, 2002). Moreover, no carnivore coprolites or digested bone specimens have been recovered in the assemblage, which would be expected had the cave served as a feeding/den site during Member 2 times. Finally, we note again the presence of several articulated specimen sets. Especially impressive in illustrating the distinctly “non-ravaged” condition of the assemblage is the hominid skeleton Stw 573. Although Stw 573 is not yet completely excavated, it appears undamaged by biotic taphonomic agents. We have observed no tooth marks on its exposed bone surfaces, its upper and lower teeth are perfectly occluded, its zygomatic arch is intact, and its uncovered postcranial bones mostly adhere to living natural positions relative to one another (e.g., Clarke, 1998, 1999, 2002a). These observations are largely inconsistent with the conditions seen in dozens of modern baboon carcasses fed-on by large carnivores (TRP, personal observations T.R. Pickering et al. / Journal of Human Evolution 46 (2004) 279–297 Table 3 Cercopithecidae skull part representation (NISP/MNE/MNI) by taxon and age group in the Member 2 fossil assemblagea Skull partb Frontal Maxilla Mandible Total MNI a 289 Adult Papio izodi 2/2/2 1/1/1 2 (7%) Adult Parapapio jonesi 1/1/1 3/3/3 3/3/3 3 (11%) Adult Parapapio broomi 3/3/3 7/7/7 10/8/8 8 (29%) Adult Papionini indet. 7/5/5 7/4/4 1/1/1 5 (18%) Juvenile Papionini indet. 9/9/9 5/5/5 3/3/3 9 (32%) Adult Cercopithecoides williamsi 1/1/1 1 (4%) NISP=number of identified specimens; MNE=minimum number of elements; MNI=minimum number of individuals. Highest MNI for each taxon/age is indicated in the Total row, and that group’s proportion of the total Cercopithecidae MNI (=28) is indicated parenthetically. b The most commonly occurring skull portions in the primate subassemblage from Member 2 are frontal, maxillary, and mandibular specimens. Some single cranial specimens include both frontal and maxilla portions and are thus counted in each category. Two maxillary specimens are isolated teeth. 1995–2001; see, e.g., Simons, 1966; Brain, 1981; Pickering, 2001a,b). In contrast, the condition of Stw 573 closely resembles non-ravaged, mummified baboon remains observed in modern cave sites. These types of primate remains are often found in difficult-toaccess portions of cave systems, e.g., a juvenile baboon mummy discovered by TRP at the bottom of a precipitous drop in a cave on the John Nash Nature Reserve, South Africa, and four dehydrated baboon carcasses found at the bottom of a 70 foot deep caldera blowhole in Mount Suswa Cave, Kenya, reported on and illustrated by Simons (1966). At least partial mummification of the hominid skeleton seems to be indicated by the calcium carbonate-lined spaces observed around the left hand and forearm bones, which suggest that the encasing matrix had calcified before the soft tissues had decayed completely (Clarke, 1999). Alternatively, perhaps sediment calcification was remarkably rapid and soft tissue decay proceeded at a “normal” rate. In any case, this type of extraordinary preservation suggests that the cave was not conducive to carnivores carrying out normal activities such as denning, carcass-ravaging and bone-collecting. Rather, the depositional situation at Sterkfontein during Member 2 times may have been characterized largely as a death-trap. We propose a deathtrap scenario in its broadest sense: meaning not only the accidental ensnarement of some animals that tumbled into the cave via avens, but also the “capture” of others that may have wandered into the cave intentionally and were then unable to escape. The fact that the assemblage is dominated taxonomically by primates and carnivores—taxa that are more agile than are ungulates—is suggestive that the cave was accessible, but not easily accessible or escapable, to motivated, proficient climbers. Cooke (1991) made much the same suggestion to explain the predominance of primates and carnivores in Peabody’s Pit 23 (UCMP Locality V-4888) at Bolt’s Farm (South Africa). Support for this idea is that 32% of the total primate subassemblage is composed of subadults (Table 3), individuals that might have been less savvy than were adults in finding their ways back out of the cave. Finally, we note that “intentional entry” need not always mean an animal scampering down a near-vertical avens. For example, there is a horizontal tunnel that still leads into the Silberberg Grotto, ending in a precipitous 10 m drop (unapparent in the cavern’s darkness) onto the talus cone below. This is also a plausible route of entry into the cavern for some animals during Member 2 times. Paleoenvironment Following the discussion above, it is apparent that the Member 2 faunal assemblage is likely biased taxonomically. Animals that are relatively good climbers (i.e., primates and carnivores) are 290 T.R. Pickering et al. / Journal of Human Evolution 46 (2004) 279–297 Fig. 7. (a) SWP-2922, a cercopithecoid foot skeleton, and (b) SG 97-83 and SG 97-84, the right and left radii of a young, large-bodied felid (a felid phalanx is visible cemented with breccia to the distal end of the left radius SG 97-84), illustrating the partially complete condition of the remains of many animals that contributed to the Sterkfontein Member 2 faunal assemblage. over-represented in the accumulation relative to less adept climbers—even though primates and carnivores may not have been relatively abundant on the ground above the cave during Member 2 times, and might thus be less informative to paleontologists for habitat reconstruction. In addition, the Member 2 fauna is clearly timeaveraged, which means that the assemblage likely samples an evolving paleo-community over a substantial time span. Thus, we emphasize that the following paleoenvironmental reconstruction is provisional and subject to revision with additional faunal recovery and possibly isotopic work. Cercopithecoids predominate in terms of their proportional representation in the Member 2 assemblage, based on number of identified specimens (NISP), minimum number elements (MNE) and minimum number of individuals (MNI) (Tables 2 and 3). While this abundant and varied primate fauna (MNI=28, 66% of the total MNI) indicates some tree cover, many of the extinct primate species represented have been reconstructed as substantially terrestrial. For instance, a variety of studies on the postcranium suggest that Cercopithecoides williamsi was much more terrestrial than many modern colobines (e.g., Birchette, 1982; Ciochon, 1993; Elton, 2000; see also, Szalay and Delson, 1979; Delson, 1984), and was adapted to open and mixed habitats (Elton, 2000, 2001). In addition, studies of the dentition of Cercopithecoides from East Africa, which identified abundant grit microwear on nonocclusal surfaces (Ungar and Teaford, 1996), agree with an inference of a terrestrial adaptation for this T.R. Pickering et al. / Journal of Human Evolution 46 (2004) 279–297 291 Fig. 8. Histogram of skeletal element representation as measured by %MAU for primate remains in the Sterkfontein Member 2 fossil assemblage and in two modern, experimental assemblages composed of specimens from an original number of ten complete baboon carcasses. The refuse assemblage consists of specimens not ingested by leopards and spotted hyenas; the scat assemblage consists of those specimens ingested and subsequently voided (see experimental details in Pickering, 2001a,b). Note that there is close agreement between Member 2 and the refuse assemblage in high skull representation, low axial element survivorship, peaks in limb bone representation, and a paucity or absence of the bones of the hand and feet. genus. Similarly, recent interpretations of postcranial and dental adaptations in Parapapio (the most abundant identified genus in the Member 2 faunal assemblage, MNI=11) point to a terrestrial component in the behavioral suite of that genus. Ciochon (1993) suggests that, like modern baboons (Papio spp.), Parapapio was terrestrial, with good tree-climbing abilities. Felids are the next most abundant taxon in the Member 2 faunal assemblage (MNI=9, 21% of the total MNI). Two of those felid individuals represented are leopards (Panthera pardus), a species that, because of its remarkable adaptability across wide geographic and habitat-type ranges in both historic and prehistoric times (Brain, 1981), is a very poor indicator of paleoenvironment. Likewise, lions (P. leo), of which there is at least one represented in the Member 2 faunal assemblage, are not truly habitat specific; today, they occur in open (i.e., <20% canopy cover) and mixed (i.e., approximately 20% canopy cover) habitats (Lewis, 1997). The most abundant cat in 292 T.R. Pickering et al. / Journal of Human Evolution 46 (2004) 279–297 the Member 2 fauna, the caracal (Felis caracal) (MNI=4), is also the most habitat-specific of the three positively identified felid species. Today, this small cat prefers “plains and rocky hills . Acacia and Commiphora thickets and Karoo scrub . The Arabic name ‘rock cat’ reflects the animal’s preference for . stony outcroppings.” (Kingdon, 1997: 280–281). The final carnivore species thus far identified in the Member 2 faunal assemblage is the extinct, long-legged hyena Chasmaporthetes nitidula. The carnassial blades of Chasmaporthetes are long and set parallel to the long axis of the cheektooth rows. This morphology and arrangement is catlike and indicates that Chasmaporthetes teeth were well adapted for slicing meat away from bones, and were not particularly well adapted for bonecracking, as are those of most extant hyena species (e.g., Kurten and Werdelin, 1988; Werdelin and Turner, 1996). This, in turn, suggests that Chasmaporthetes would have been at a relative disadvantage if forced to scavenge largely defleshed carcasses, appropriated from and/or abandoned by other predators. Based on this functional interpretation of the dental anatomy of Chasmaporthetes, it is likely that it was primarily a hunter, using its long legs and feet effectively as a cursorial predator (e.g., Berta, 1981). Thus, on balance, this indirect evidence suggests that Chasmaporthetes may have preferred relatively open country and/or broken woodland, in which its cursorial adaptations could be used to best advantage. The relatively meager bovid subassemblage from Member 2 is also fairly informative from a paleoenvironmental perspective. Makapania broomi (MNI=1) is typically classified as an ovibovine (Gentry, 1970, 1996), although Brink (1999) has argued recently that this large extinct bovid seems to have stronger morphological affinities to the Caprini than to the Ovibovini. Regardless, both tribes are contained within the subfamily Caprinae, whose extant members are very “goatlike” in their general agility and ability to traverse hilly and rocky terrain efficiently (e.g., Gentry, 1996; Kingdon, 1997). Morphology consistent with these locomotor behaviors is expressed strongly in the phalanges of these animals, and also in those of M. broomi (Brink, 1999). Finally, the presence of at least one Alcelaphini individual in the Member 2 faunal assemblage might indicate the proximity of permanent water. Most modern alcelaphines are obligate drinkers and inhabit open woodlands and grasslands, preferring to graze in wetter parts of the habitat (Reed, 1996; Kingdon, 1997). In summary, the mammalian fauna from Member 2 indicates a paleohabitat that was probably typified by rolling, rock-littered and brushand scrub-covered hills (suitable for caracals and Makapania, and also commonly exploited by papionins). The valley bottom might have retained standing water year-round, and perhaps supported a tree line or restricted riverine forest, fringed by open woodland or grassland—a setting appropriate for Alcelaphini, the abundant monkeys, and ambush predators, such as leopards. Discussion and conclusions The inferred paleoenvironment of Sterkfontein Member 2 conforms broadly to that of the peneconteporaneous1 4.2–3.8 myr old Australopithecus anamensis localities at Kanapoi and Allia Bay (Leakey et al., 1995, 1998; Wynn, 2000; Schoeninger et al., in press; C.S. Feibel, pers. comm., cited in Heinrich et al., 1993 and Ward et al., 1999a). Recovered fauna from these Kenyan sites and isotopic studies indicate the presence of riverine gallery forest, with surrounding bushland and open country (Leakey et al., 1995; Ward et al., 1999a; Wynn, 2000; Schoeninger et al., in press). In addition, the ca. 4 myr old fauna from Jacovec Cavern at Sterkfontein (Partridge et al., 2003), including Australopithecus, multiple species of Parapapio, a colobine, and Chasmaporthetes, is largely consistent with that recovered from Member 2, and led Kibii (2000; in prep) to 1 Contrary to claims of a more recent age for Sterkfontein Member 2 (Berger et al., 2002), we reassert our current estimate of 4.0 Ma for the unit, based on cosmogenic 26Al and 10Be burial dates (Partridge et al., 2003). It is not our purpose here to issue a rejoinder to Berger et al. (2002), as that was done previously by Clarke (2002b). T.R. Pickering et al. / Journal of Human Evolution 46 (2004) 279–297 293 reconstruct a mosaic environment similar to that proposed for Member 2, Kanapoi, and Allia Bay. Based on taxonomic assessment of the recovered microfauna, the Waypoint 160 deposit at Bolt’s Farm (South Africa) may also be at least 4 myr old (and possibly 5 myr old) (Senegas and Avery, ´ ´ 1998), but no paleoenvironmental interpretations have yet been offered for this important deposit. Similarly, little has yet been published on the inferred paleoenvironments of the hominid localities of Belohdelie (3.89–3.86 myr old, Clark et al., 1984; White et al., 1993) and FJ-4 Fejej (4.18–4.00 myr old, Asfaw et al., 1991; Fleagle et al., 1991; Kappleman et al., 1996), both in Ethiopia. However, it is interesting that researchers infer substantial open components in the paleoenvironments of many mid-Pliocene hominid sites2 (i.e., ca. 3.6–3.0 myr old), such as is reconstructed for the earlier-occurring sites of Sterkfontein Member 2, Kanapoi, and Allia Bay. Taken together then, what is notable about most early hominid sites spanning ca. 4.0–3.0 Ma is that they included open components as a larger proportion of the total habitat compared to paleoenvironments at hominid-bearing sites that predate them. For example, although the exact paleohabitat of the TM 266 Sahelanthropus tchadensis hominid locality is still under investigation (Vignaud et al., 2002), the other known late Miocene hominids, Ardipithecus ramidus (e.g., WoldeGabriel et al., 1994; WoldeGabriel et al., 2001) and Orrorin tugenensis (Pickford and Senut, 2001), have been recovered in association with faunas indicating predominately closed woodlands. East African sites of relevant age include: (1) all A. afarensis sites in Ethiopia, Tanzania, and Kenya, dated radioisotopically and biochronologically to ca. 3.6–2.9 Ma (reviewed in White, 1995, 2002); (2) localities in the Unso Formation, Member U-10 and the Shungura Formation, Member B, Omo, Ethiopia, dated radioisotopically and biochronologically to ca. 3.4–3.2 Ma (de Heinzelin, 1983; Feibel et al., 1989); (3) localities from South Turkwel, Kenya, dated stratigraphically andbiochronologically to ca. 3.5 Ma (Ward et al., 1999b); and (3) sites from Lomekwi, Kenya, dated radioisotopically and biochronologically ca. 3.5 Ma (Leakey et al., 2001). The sole north central African site of relevance is KT 12, Bahr el Gahzal, Chad (dated biochronologically to ca. 3.5–3.0 Ma, Brunet et al., 1995). 2 Taphonomically, Sterkfontein Member 2 and the other Sterkfontein Valley deposits of similar age (i.e., Jacovec Cavern and Bolt’s Farm), with their low number of hominid fossils (in terms of NISP, MNE and MNI), stand in contrast to the hominid-rich assemblages from more recent units at Sterkfontein, Swartkrans, Drimolen, and Kromdraai (e.g., Broom and Schepers, 1946; Broom and Robinson, 1952; Tobias et al., 1977; Brain, 1981; Brain et al., 1988; Keyser et al., 2000; Thackeray et al., 2001; Pickering et al., in press). This may suggest the real rarity of hominids in the Sterkfontein Valley prior to 3.0–2.0 Ma, or instead may be a sampling artifact. Additional work at known sites and survey for new localities of relevant age may elucidate this issue more fully. Few relevant taphonomic comparisons can be made between Sterkfontein Member 2 and penecontemporaneous sites in East Africa, because Member 2 is a cave deposit and the other sites are open-air. For example, many of the Kanapoi and Allia Bay hominid fossils are tooth-marked and inferred to have been ravaged by carnivores (e.g., Leakey et al., 1998; Ward et al., 1999a). In contrast, there seems to be little indication of carnivore involvement in the collection of the Sterkfontein Member 2 fossils. We interpret the assemblage as primarily autopod in origin (i.e., accumulation of animal bones that arrive at a site as the result of self-transport, Vrba, 1982) and explain the predominance of primates and carnivores as due to the greater likelihood of primates and carnivores intentionally entering (and dying in) caves than is the case with ungulates (cf. Cooke, 1991). Furthermore, in the absence of other taphonomic evidence to the contrary (e.g., abundant carnivore tooth marks, coprolites, etc.), this simple observation may also be sufficient to explain the lower representation of ungulates compared to primates and carnivores in some other South African cave assemblages. In other words, while we agree that in some cases carnivores did den in sites during certain depositional phases (or portions thereof) and perhaps also focused their predatory activities on primates (e.g., Brain, 1981, 1993; White, 1988), other cases in which there are elevated counts of primates and carnivores simply reflect the curiosity and climbing aptitude of these animals relative to 294 T.R. Pickering et al. / Journal of Human Evolution 46 (2004) 279–297 most ungulates. In those cases it is unnecessary to appeal to a complex accumulation process(es) or to recalculate estimates of taxonomic abundance using new methods (e.g., de Ruiter, 2001) in order to bring relative numbers of different taxa in line with expectations based on the trophic pyramid. In addition, cave accumulations do not necessarily indicate accurate taxonomic proportions of the local prehistoric biota— but rather simply reflect the consequences of varied behavioral potentials and predilections of a selection of the larger mammalian paleo-community. References Asfaw, B., Beyene, Y., Semaw, S., Suwa, G., White, T., WoldeGabriel, G., 1991. Fejej: a new paleoanthropological research area in Ethiopia. J. Hum. Evol. 21, 137–143. Behrensmeyer, A.K., 1978. Taphonomic and ecologic information from bone weathering. Paleobiology 4, 150–162. Berger, L.R., Lacruz, R., de Ruiter, D.J., 2002. 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TRP’s work on the Member 2 fauna was funded by a Faculty Summer Research Grant from Indiana University and his work on the baboon feeding experiments was funded by the National Science Foundation (SBR 9614930), the Wenner-Gren Foundation for Anthropological Research (6109), and the University of Wisconsin Natural History Museums Council. TRP thanks his family and Nick Toth and Kathy Schick for their steadfast encouragement, support, and enthusiasm. JLH’s work on the Member 2 fauna was funded by a Grant-In-Aid of Research from Sigma Xi and a Graduate and Professional Student Organization Grant from Indiana University. JLH thanks his family for support during his prolonged absence in the completion of this study. Blasting of the vertical breccia face of Member 2 was carried out by John Cruise and Dusty van Rooyen. 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