Hominin vertebrae and upper limb bone fossils from Sterkfontein Caves, South Africa (1998-2003 excavations).

OBJECTIVES: We provide descriptions and functional interpretations of 11 >2.0 Ma hominin vertebral and upper limb fossils from Sterkfontein. MATERIALS AND METHODS: We employed taphonomic methods to describe postmortem damage observed on the fossils. We used osteometric tools and measurements to generate quantitative descriptions, which were added to qualitative descriptions of the fossils. These observations were then interpreted using published data on the same skeletal elements from extant and extinct hominoid taxa. RESULTS: Six of the fossils carry carnivore tooth marks. Two vertebrae show morphologies that are consistent with fully developed lordosis of the lumbar spine, but which are not completely consistent with bipedal loading of the same intensity and/or frequency as reflected in the lumbars of modern humans. A clavicle shows a combination of humanlike and apelike features, the latter of which would have endowed its hominin with good climbing abilities. When combined, analyses of fragmentary radius and ulna fossils yield more ambiguous results. DISCUSSION: The new fossil collection presents a mix of bipedal and climbing features. It is unclear whether this mix indicates that all Sterkfontein hominins of >2.0 Ma were terrestrial bipeds who retained adaptations for climbing or whether the collection samples two differently adapted, coeval hominins, Australopithecus africanus and Australopithecus prometheus, both of which are represented at Sterkfontein by skull remains. Regardless, the significant frequency of tooth-marked fossils in the sample might indicate that predation was a selection pressure that maintained climbing adaptations in at least some Sterkfontein hominins of this period.

Ages-at-death distribution of the early Pleistocene hominin fossil assemblage from Drimolen (South Africa).

OBJECTIVES: A prevailing hypothesis in paleoanthropology is that early Pleistocene hominin bones were accumulated in South African caves by carnivores, which used those shelters, and the trees surrounding them, as refuge and feeding sites. We tested this hypothesis at the site of Drimolen, by comparing its hominin age-at-death distribution to that of the nearby and roughly contemporaneous site of Swartkrans. MATERIALS AND METHODS: We employed standard dental aging systems in order to categorize the Drimolen hominin teeth into age classes of 5 years each. We then compared the age-at-death distribution for Drimolen with the published data available for the Swartkrans hominins. RESULTS: Age-at-death distributions indicate that the age category "young adults" is the best represented age category at Swartkrans and the most poorly represented one at Drimolen. Moreover, Drimolen has a preponderance of infant specimens. Both sites have a low frequency of old adult specimens. CONCLUSIONS: Differences observed in frequencies of the age-at-death categories suggest different mechanisms of hominin skeletal accumulation at Drimolen and Swartkrans. Swartkrans' frequency curve reflects mortality in a population subjected to predation and is thus consistent with the carnivore-accumulating hypothesis. In contrast, the Drimolen curve is similar to that of wild populations of living apes. Living primates have been observed exploiting caves as sleeping shelters, for nutritional, security, drinking, and thermoregulatory purposes. We suggest that similar cave use by Pleistocene hominins can explain, in large part, the accumulation of hominin bones at Drimolen. Such a conclusion is another illustration of the growing awareness that a "one-size-fits-all" taphonomic model for South African early Pleistocene hominin sites is probably insufficient.

New early Pleistocene hominin teeth from the Swartkrans Formation, South Africa.

We describe 14 hominin teeth and tooth fragments excavated recently from Swartkrans Cave (South Africa). The fossils derive from Members 1 (Lower Bank) and 3, from the Member 2/3 interface and from two deposits not yet assigned to member (the "Talus Cone Deposit" and the "Underground North Excavation" [UNE]) of the Swartkrans Formation, and include the first hominin fossil from the UNE, the two smallest Paranthropus robustus deciduous maxillary second molars in the entire hominin fossil record, and one of the smallest P. robustus permanent maxillary second molars from Swartkrans. The small permanent molar is accompanied by another tooth from a different individual but from the same stratigraphic level of the Swartkrans Formation; this second tooth is among, if not, the largest P. robustus permanent maxillary first molars known from anywhere-lending credence to assertions that degrees of body size sexual dimorphism previously ascribed to this species may be underestimated. It is more equivocal whether this evidence also supports hypotheses proposing that P. robustus assemblages from Swartkrans (as well as those from other South African cave sites) formed through the taphonomically biasing actions of large carnivores.

New hominid fossils from Member 1 of the Swartkrans formation, South Africa.

Member 1 of the Swartkrans Formation is comprised of two sedimentary infills, the Lower Bank (LB) and the Hanging Remnant (HR). Together, the LB and HR preserve fossils of early Homo and Paranthropus robustus, Earlier Stone Age lithic artifacts, purported bone digging tools and butchered animal bones. Collectively, this evidence was the first to establish the co-existence of two early Pleistocene hominid species and also led to inferences of plant root harvesting and meat-eating by one or both of those species. P. robustus is the more abundant of the two hominids at Swartrkrans, represented in Member 1 by hundreds of fossils that derive from at least 99 individuals. Thus, Swartkrans Member 1 stands as the world's single largest repository of that extinct species. Here we add to the Member 1 sample of hominid fossils with descriptions of 14 newly discovered specimens.

Savanna chimpanzees use tools to harvest the underground storage organs of plants.

It has been hypothesized that plant underground storage organs (USOs) played key roles in the initial hominin colonization of savanna habitats, the development of the distinctive skull and tooth morphology of the genus Australopithecus, and the evolution of the genus Homo by serving as "fallback foods" exploited during periods of food shortage. These hypotheses have been tested mostly by morphological, isotopic, and microwear analyses of hominin bones and teeth. Archaeological evidence of USO digging technology is equivocal. Until now relevant data from studies of chimpanzees, useful in behavioral models of early hominins because of their phylogenetic proximity and anatomical similarities, have been lacking. Here we report on the first evidence of chimpanzees using tools to dig for USOs, suggesting that exploitation of such resources was within the cognitive and technological reach of the earliest hominins. Consistent with scenarios of hominin adaptation to savannas, these data come from Ugalla (Tanzania), one of the driest, most open and seasonal chimpanzee habitats. USOs are, however, exploited during the rainy season, well after the period of most likely food shortage, contradicting the specific prediction of fallback food hypotheses. The discovery that savanna chimpanzees use tools to obtain USOs contradicts yet another claim of human uniqueness and provides a model for the study of variables influencing USO use among early hominins.

Newly discovered fossil- and artifact-bearing deposits, uranium-series ages, and Plio-Pleistocene hominids at Swartkrans cave, South Africa.

We report on new research at Swartkrans Cave, South Africa, that provides evidence of two previously unrealized artifact- and fossil-bearing deposits. These deposits underlie a speleothem dated by the uranium-thorium disequilibrium technique to 110,000+/-1,980 years old, the first tightly constrained, geochronological date available for the site. Recovered fauna from the two underlying deposits-including, prominently, the dental remains of Paranthropus (Australopithecus) robustus from the uppermost layer (Talus Cone Deposit)-indicate a significantly older, late Pliocene or early Pleistocene age for these units. The lowest unit (LB East Extension) is inferred to be an eastward extension of the well-known Lower Bank of Member 1, the earliest surviving infill represented at the site. The date acquired from the speleothem also sets the maximum age of a rich Middle Stone Age lithic assemblage.

The "bear" essentials: actualistic research on Ursus arctos arctos in the Spanish Pyrenees and its implications for paleontology and archaeology.

Neotaphonomic studies of large carnivores are used to create models in order to explain the formation of terrestrial vertebrate fossil faunas. The research reported here adds to the growing body of knowledge on the taphonomic consequences of large carnivore behavior in temperate habitats and has important implications for paleontology and archaeology. Using photo- and videotrap data, we were able to describe the consumption of 17 ungulate carcasses by wild brown bears (Ursus arctos arctos) ranging the Spanish Pyrenees. Further, we analyzed the taphonomic impact of these feeding bouts on the bones recovered from those carcasses. The general sequence of consumption that we charted starts with separation of a carcass's trunk; viscera are generally eaten first, followed by musculature of the humerus and femur. Long limb bones are not broken open for marrow extraction. Bears did not transport carcasses or carcass parts from points of feeding and did not disperse bones appreciably (if at all) from their anatomical positions. The general pattern of damage that resulted from bear feeding includes fracturing, peeling, crenulation, tooth pitting and scoring of axial and girdle elements and furrowing of the upper long limb bones. As predicted from observational data, the taphonomic consequences of bear feeding resemble those of other non-durophagus carnivores, such as felids, and are distinct from those of durophagus carnivores, such as hyenids. Our results have paleontological and archaeological relevance. Specifically, they may prove useful in building analogical models for interpreting the formation of fossil faunas for which bears are suspected bone accumulators and/or modifiers. More generally, our comparative statistical analyses draw precise quantitative distinctions between bone damage patterns imparted respectively by durophagus (modelled here primarily by spotted hyenas [Crocuta crocuta] and wolves [Canis lupus]) and non-durophagus (modelled here by brown bears and lions [Panthera leo]) carnivorans.

First records of talon cusps on baboon maxillary incisors argue for standardizing terminology and prompt a hypothesis of their formation.

Dental characters can provide vital clues for understanding intra- and intertaxonomic morphological variation and its underlying genetic and environmental components. However, the unambiguous identification of particular traits and their comparative study is often confounded by lack of consistent terminology in the relevant literature. This difficulty is exacerbated when the etiologies are not completely understood, as is the case with talon cusps. To date, research on talon cusps has focused on modern humans. In many instances, descriptions of talon cusps appear in clinical case studies focusing on their treatment and removal. What is lacking in those discussions, though, is a comparative framework, in which the occurrence of talon cusps in nonhuman primates, and possibly other mammals, is established and understood. Here, we report on a taloned upper central incisor of a wild baboon (Papio hamadryas ursinus) from South Africa. The anomalous incisor of this individual includes an exaggerated accessory cusp diagnosed as a Type II talon. Microcomputed tomographic and radiographic analyses show that the taloned cusp possesses enamel, dentin, and pulp. In addition, we identified an unclassifiable talon cusp on a central maxillary incisor of a baboon skull housed in the Smithsonian Institution's Natural History Museum collection. Our observations of talon cusps on baboon incisors demonstrate that, with regard to this phenomenon, systematic study of nonhuman primates is much needed, along with a consistent use of terminology in the anatomical and anthropological literature. Finally, we present a hypothesis of the formation of talon cusps on mammalian incisors.

First partial skeleton of a 1.34-million-year-old Paranthropus boisei from Bed II, Olduvai Gorge, Tanzania.

Recent excavations in Level 4 at BK (Bed II, Olduvai Gorge, Tanzania) have yielded nine hominin teeth, a distal humerus fragment, a proximal radius with much of its shaft, a femur shaft, and a tibia shaft fragment (cataloged collectively as OH 80). Those elements identified more specifically than to simply Hominidae gen. et sp. indet are attributed to Paranthropus boisei. Before this study, incontrovertible P. boisei partial skeletons, for which postcranial remains occurred in association with taxonomically diagnostic craniodental remains, were unknown. Thus, OH 80 stands as the first unambiguous, dentally associated Paranthropus partial skeleton from East Africa. The morphology and size of its constituent parts suggest that the fossils derived from an extremely robust individual who, at 1.338±0.024 Ma (1 sigma), represents one of the most recent occurrences of Paranthropus before its extinction in East Africa.

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

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.

Role of carnivores in the accumulation of the Sterkfontein Member 4 hominid assemblage: a taphonomic reassessment of the complete hominid fossil sample (1936-1999).

New taphonomic data on the Sterkfontein Member 4 (South Africa) fossil hominid assemblage are presented. The previous estimate of hominid individuals represented in the deposit (45) is increased to 87. New minimum numbers of hominid skeletal elements are provided, and incidences of bone surface damage inflicted by prehistoric biological agents are summarized. The hominid sample from Member 4 is composed predominately of gnathic remains and has a paucity of postcrania. This dearth of postcrania limits, to some extent, inferences about the formation of the Sterkfontein assemblage. However, carnivore tooth marks on some fossil specimens and an overall broad similarity in patterns of skeletal part representation between Sterkfontein and primate bone assemblages created by extant carnivores suggest that carnivores did have some involvement in the accumulation of the fossil hominid assemblage. Thus, this study provides support for the "carnivore-collecting hypothesis" of Brain (Brain [1981] The Hunters or the Hunted? Chicago: University of Chicago Press), which implicates large carnivores as prominent collecting agents of hominid body parts in Sterkfontein Member 4. Evidence of bone surface damage is, however, too scant to make confident inferences about specific carnivore taxon/taxa involved in hominid bone collection at the site.

Disentangling Early Stone Age palimpsests: determining the functional independence of hominid- and carnivore-derived portions of archaeofaunas.

Determining the extent to which hominid- and carnivore-derived components of fossil bone palimpsests formed independently of each other can provide valuable information to paleoanthropologists interested in reconstructing the foraging adaptations of hominids. Because stone tool cutmarks, hammerstone percussion marks, and carnivore tooth marks are usually only imparted on bone during nutrient extraction from a carcass, these bone surface modifications are particularly amenable to the types of analyses that might meet this goal. This study compares the percentage of limb bone specimens that preserve evidence of both hominid- and carnivore-imparted bone damage from actualistic control samples and several Plio-Pleistocene archaeofaunas, including new data from Swartkrans Member 3 (South Africa). We argue that this procedure, which elucidates the degree of hominid-carnivore independence in assemblage formation, will allow researchers to extract for focused analyses high integrity components (hominid and carnivore) from presumably low integrity sites. Comparisons suggest that the hominid- and carnivore-derived components from sites in Olduvai Gorge Bed II (Tanzania), the ST Site Complex at Peninj (Tanzania), and Swartkrans Member 3 formed largely independent of each other, while data from the FLK 22 Zinjanthropus (FLK Zinj) site (Olduvai Gorge Bed I) indicate significant interdependence in assemblage formation. This contrast suggests that some Early Stone Age assemblages (e.g., the Olduvai Gorge Bed II sites, the Peninj ST Site Complex, and Swartkrans Member 3) are probably more useful than others (e.g., FLK Zinj) for assessing the maximal carcass-acquiring abilities of early hominids; in such assemblages as those in the former set, sole hominid-contribution is more confidently discerned and isolated for analysis than in assemblages such as FLK Zinj.

Beyond leopards: tooth marks and the contribution of multiple carnivore taxa to the accumulation of the Swartkrans Member 3 fossil assemblage.

The ca. 1.0 myr old fauna from Swartkrans Member 3 (South Africa) preserves abundant indication of carnivore activity in the form of tooth marks (including pits) on many bone surfaces. This direct paleontological evidence is used to test a recent suggestion that leopards, regardless of prey body size, may have been almost solely responsible for the accumulation of the majority of bones in multiple deposits (including Swartkrans Member 3) from various Sterkfontein Valley cave sites. Our results falsify that hypothesis and corroborate an earlier hypothesis that, while the carcasses of smaller animals may have been deposited in Swartkrans by leopards, other kinds of carnivores (and hominids) were mostly responsible for the deposition of large animal remains. These results demonstrate the importance of choosing appropriate classes of actualistic data for constructing taphonomic inferences of assemblage formation. In addition, they stress that an all-encompassing model of assemblage formation for the hominid-bearing deposits of the Sterkfontein Valley is inadequate and that each must be evaluated individually using not just analogical reasoning but also incorporating empirical data generated in the preserved fossil samples.