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.     

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.     

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. Am J Phys Anthropol, 2004. © 2004 Wiley‐Liss, Inc.     

Small mammal diversity of Mt. Kenya based oncarnivore fecal and surface bone remains

Ecological dynamics and faunal diversity documentation is normally conducted by direct observation and trapping of live animals. However,surveys of carnivore scat prey and surface bone remains, which are relatively inexpensive, can provide complementary data that expand carnivore diet breadth and may improve accuracy regarding inferences of the ecological dynamics of a given ecosystem. We used this inexpensive method to document species diversity variation with elevation on the leeward(Sirimon) and windward(Chogoria)areas of Mt. Kenya. Bone and fecal specimens were opportunistically collected by walking 2 km in opposite directions from transect points selected at 200-m intervals along the elevational gradient of the study areas. We collected a total of 220 carnivore fecal and owl pellet specimens from both study sites, which were mainly deposited by the spotted hyena(Crocuta crocuta), leopard(Panthera pardus),serval(Leptailurus serval), genet(Genetta sp.), and Mackinder's Cape owl(Bubo capensis mackinderi).Serval scats were the most common, followed by those of the spotted hyena. Scats and bones were found at the lowest density at the lowest elevations,peaked at mid-higher elevations, and then declined at the highest elevations. Based on skeletal analysis only, there were more species in Sirimon(19) than in Chogoria(12). Small fauna(rodents to duiker size bovids) formed the bulk of the identified remains,representing 87.9% of the Sirimon fauna and 90.9% of the Chogoria fauna. The genus Otomys was the dominant prey of the owl and serval in both sites. Three giraffe teeth were found at 3 500 m a.s.l. in Chogoria on the edge of Lake El is, suggesting that it is an occasional visitor to such high elevations. This study underscores the value of fecal and bone surveys in understanding the diet and diversity of mammals in ecological ecosystems,but such surveys should be complemented with analysis of hairs found in scats to obtain a more complete list of carnivore prey at Mt. Kenya.     

Hominid tool-making versus carnivore scavenging

The effects of hyena scavenging on the remains of 75 animals in the Awash National Park, Ethiopia, were examined to provide a modern analogy to the fossil assemblage from Makapan. Brain's recent work ('70, '76) suggests that the actions of carnivores may account for the characteristics of the Makapan assemblage, a theory which contrasts with Dart's ('49, '55, '57, '59) interpretation of the fossils as osteodontokeratic tools and weapons manufactured, collected, and used by australopithecines. The modern assemblage was like that from Makapan in terms of relative representation of skeletal elements and patterns of bone breakage. Observations of hyenas in the Awash also lend support to the hypothesis that the Makapan assemblage may represent the uneaten remains of carnivore kills. The representation of major vertebrate groups at Makapan was compared with their representation at Awash and with their representation in hyena diets in the Serengeti and Ngorongoro Crater (Kruuk, '72). This technique of faunal analysis was unable to detect differences between these groups, despite the fact that the three modern assemblages represent animals that died of different causes. Thus extreme caution must be used in deducing any specific information about the mode of death or taphonomic history of the animals represented in an assemblage on the basis of such faunal analyses.     

The TD6.3 faunal assemblage of the Gran Dolina site (Atapuerca,Spain): a late Early Pleistocene hyena den

Ethological studies have shown that besides human groups, large-medium carnivores have bone-collecting habits. The research developed since the last half of the twentieth century has attempted to characterise the carnivore’s accumulations and to identify them in the archaeo-paleontological record. At present, we have diagnostic criteria that define the accumulations produced by hyenids (mainly, Crocuta crocuta), thereby allowing us to differentiate them from the other accumulating agents. The faunal assemblage recovered at the Early Pleistocene TD6.3 layer of the Gran Dolina site is characterised by the presence of typical elements described in hyena dens: presence of small carnivores remains, high bone breakage, low epiphysis survival and a high frequency (>30%) of specimens with carnivore induced modifications, including large amounts of digested bones. However, attritional mortality profiles, hyena’s cubs remains, mid-shaft bone cylinders or differential anatomical composition among different weight-sizes, have not been observed or are ambiguous. In addition, anthropic evidences are scarce and concentrated in the uppermost section of the layer. TD6.3 is the result of an accumulation produced by hyenas using the cave as a den, in alternation with sporadic occupations by hominin groups. TD6.3 shows that hyena fossil accumulations may present great anatomical and taxonomic variability.     

Late Pleistocene hyena skeleton remains of a communal/prey depot cave den in the Bohemian Mountains (Czech Republic) – its osteology,taphonomy and palaeoecology

A total of 366 bones are arranged to six incomplete individual skeletons (=MNI) of different age classes and taphonomic preservation from the Late Pleistocene Crocuta crocuta spelaea (Goldfuss 1823) hyena den Srbsko Chlum–Komín Cave, Czech Republic, Bohemian Mountains (Central Europe). The hyena assemblage counts no siblings, 37% cub, 33% early adult/subadult and 30% adult/senile individuals. A senile scavenged individual demonstrates the cannibalism within the cave. About 10% of the 3569 megafauna bones are from hyenas which larger amounts are typical at communal/prey depot den sites. This site is different to three other larger natal/birth and communal or prey depot hyena den cave sites in Czech Republic. The Chlum–Komín Cave hyena prey bone assemblage (=NISP) consists, similar as at two other Czech Republic sites, mainly of Equus ferus przewalskii (50%), including pregnant females being hunted in spring/early summer. The fewer amounts of Coelodonta antiquitatis (4%), Bison priscus (2%), Rangifer tarandus (15%), and alpine Rupicapra rupicapra (2%) or Capra ibex (1%) and the absence of mammoth correlate to hyena den bone assemblages in middle high elevated mountain regions of northern/central Europe. The rareness of cave bears in the Bohemian Mountains, on which hyenas specialized to feed additionally in European mountain regions, explains predominant predation on horses.     

河南栾川孙家洞遗址出土的穿孔标本

孙家洞遗址初步测年为40万年左右,出土了少量石制品、古人类牙齿以及大量动物化石,其中一件骨骼标本存在非常规整的穿孔,其整体形态和大小与人工穿孔比较相似。通过系统的宏观观测与显微镜下分析,并与已明确性质的考古标本(自然穿孔和人工穿孔标本)和实证材料(现代鬣狗洞发掘出土材料)进行对比后发现,孙家洞遗址出土的穿孔标本与食肉类啃咬、甲虫类啮食以及人工造成的穿孔标本存在明显差异,而具备了被鬣狗胃酸腐蚀标本的所有典型特征。     

Faunal assemblage seriation of southern African Pliocene and Pleistocene fossil deposits

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 Faunal 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 Hrdli?ka deposits, Sterkfontein Member 5 (in part) and Kromdraai B, Kromdraai A and Swartkrans 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. © 1995 Wiley-Liss, Inc.     

Bone mineral density and survival of elements and element portions in the bones of the Crow Creek massacre victims

The interpretation of archaeologically-derived skeletal series is dependent on the elements and portions of elements preserved for examination. Bone and bone portion survival is affected by factors, both intrinsic and extrinsic to the elements themselves, that influence deterioration and preservation. Among the intrinsic variables, the density of the element and element portion are particularly important with respect to the degree of preservation. Recently reported bone mineral density values from a contemporary human sample are compared to the survival of prehistoric limb bones of the Crow Creek specimens, a fourteenth-century massacre skeletal series. The contemporary density values are positively correlated with Crow Creek element and element portion survival. Two calculations of bone mineral density, however, are more closely related to preservation than a third. Such density information has implications for assessing minimum number of elements and individuals and documenting taphonomic processes. Am J Phys Anthropol 104:513–528, 1997. © 1997 Wiley-Liss, Inc.     

Recognizing hominoid-modified bones: the taphonomy of colobus bones partially digested by free-ranging chimpanzees in the Kibale Forest, Uganda

We present a taphonomic study of bones that have passed though the digestive tracts of free-ranging chimpanzees from the Kibale Forest in Uganda. The bone assemblage can be characterized as having a very low species diversity; low number of identifiable specimens (NISP) per scat; bones extremely broken up (very small size range); skeletal part frequencies similar in some ways to those resulting from carnivore partial digestion; and sometimes articulated specimens. Modifications to the bones include corrosion, tiny tooth scores and pits, cracking, and fraying of bone edges. Together, these characteristics suggest that hominoid bone digestion may be recognizable, despite some similarities with leopard-, canid-, and eagle-modified bone. Chimpanzees are well-documented hunters of medium-sized vertebrates such as monkeys. This is significant in the study of human evolution if, as it seems, the last common ancestor of chimpanzees and humans was chimpanzee-like. It suggests there was a pre-stone-tool-using hunting phase in human evolution, perhaps by australopiths or the last common ancestor. Taphonomically, pre-stone tool meat eating has been very difficult to detect in the fossil record. However, if chimpanzees leave a recognizable taphonomic signature on the bones of their prey, we will be able to look for analogous signatures in fossil bones associated with fossil hominoids and hominids.     

An ecological reassessment of the southern African carnivore guild: a case study from Member 4, Sterkfontein, South Africa

The southern African late Pliocene to early Pleistocene carnivore guild was much larger than that of the present day. Understanding how this guild may have functioned is important for the reconstruction of carnivore-hominin interactions and to assess the potential for hominin scavenging in southern Africa. In modern ecosystems, the coexistence of larger carnivore species is constrained by several factors, which include high levels of interspecific competition. Here, the composition of the fossil carnivore guild is examined using Sterkfontein Member 4 (Cradle of Humankind, South Africa) as a case study. Sterkfontein Member 4 contains 10 larger carnivore taxa (body mass > 21.5 kg) and may also contain two Australopithecus species. Two possible causes of higher numbers of carnivore species in the South African fossil record are initially considered. First, that there is a bias introduced through comparing assemblages of differing sizes; second, carnivore biodiversity may have been artificially inflated due to previous taxonomic splitting of carnivore species, such as Crocuta. These possibilities are rejected and modern ecological data are used to construct a simple spatial model to determine how many carnivores could have co-existed. Although the resulting model indicates that the carnivore taxa present in Member 4 could have co-occurred, modern ecological studies indicate that it is highly unlikely that they would have co-existed simultaneously. Considering the complex depositional processes that operate in the southern African cave sites, it is proposed that the larger carnivore guild observed in the Sterkfontein Member 4 fossil assemblage is a palimpsest created by time-averaging. In light of this, we suggest that sites which have a large number of carnivore taxa should be examined for time-averaging, while those sites which have relatively few species may be a better reflection of carnivore communities.     

Mammalian collagenase predisposes bone surfaces to osteoclastic resorption

Summary The cell-free endocranial surface of young adult rat parietal bones was used as a substrate for bone cell-derived mammalian collagenase. Incubation of parietal bones in a concentration of enzyme comparable to that secreted by osteoblastic cells in vitro caused destruction of surface osteoid, and resulted in exposure of mineral onto the bone surface. Bones so pre-treated were considerably more susceptible to osteoclastic resorption than bones preincubated in the absence of collagenase. These results are consistent with the view that the osteoid layer which covers bone surfaces acts as a barrier to osteoclastic contact with underlying, resorption — stimulating bone mineral; and that cells of the osteoblastic lineage induce osteoclastic resorption through collagenase secretion which, by digestion of the surface osteoid, exposes bone mineral to osteoclastic contact.     

Faunal assemblage composition and paleoenvironment of Plovers Lake, a Middle Stone Age locality in Gauteng Province, South Africa

Plovers Lake is a dolomitic cave infill located approximately 45 km northwest of Johannesburg in the Bloubank Valley, Gauteng Province, South Africa. Excavations between 2002–2004 revealed a rich and diverse fauna, a moderate-sized stone tool assemblage of Middle Stone Age (MSA) character, and human skeletal remains. Two principal depositional units are recognized: 1) a disturbed ex situ component that was likely displaced from 2) an otherwise relatively undisturbed in situ component from which the human skeletal material was recovered. The in situ depositional unit is bracketed by 2 flowstone layers, with U-series dates of 62.9 (± 1.3) ka for the capping flowstone and 88.7 (±1.6) ka for the underlying flowstone. A single isochron ESR date of 75.6 (± 5.6) ka corroborates the U-series dates. This paper presents an analysis of the mammalian, bird, and reptile faunas recovered from these two units. The two faunal assemblages show close correspondence in taphonomic, taxonomic, and ecological composition, supporting a common origin for both the ex situ and in situ components. Although human skeletal material, cut-marked bone, and stone tools have been recovered, these indications are too rare to consider Plovers Lake a human occupation site. Instead, a high abundance of carnivores, coprolites, and carnivore damaged bones point to brown hyenas as the principal, though not exclusive, bone accumulating agent. In the absence of a significant taphonomic bias relating to accumulating agent, Plovers Lake allows us to document an environment occupied by MSA humans, even if the humans were not resident in the cave itself. We reconstruct the paleoenvironment of Plovers Lake as predominantly grassland, though it was colder, moister, and more wooded than at present. Paleoclimatic conditions appear to have been as different from historic norms as those seen in several fossil localities in the Western Cape, pointing to greater environmental heterogeneity than has previously been documented for the region in the later Pleistocene.     

New evidence for hominin carcass processing strategies at 1.5 Ma, Koobi Fora, Kenya

Reconstruction of early Pleistocene hominin carcass acquisition and processing behaviors are necessarily based at least in part on butchered fossil bones. This paper provides zooarchaeological and taphonomic analyses and behavioral interpretations of three approximately 1.5 million-year-old archaeofaunas from areas 1A and 103 in the Okote Member of the Koobi Fora Formation, northern Kenya: FwJj14A, FwJj14B, and GaJi14. These sites are all located in similar paleoenvironmental contexts, near shallow water with swampy, seasonally flooded areas, and some evidence for more wooded or gallery forest settings. Both individual specimen--and assemblage-level analyses of butchery-marked bones indicate that the hominins appear to have practiced similar butchery strategies at all of these sites, with butchery (defleshing, disarticulation, and marrow extraction) of both high- and low-ranked skeletal elements with no apparent preference for prey size, skeletal region, limb class, or limb portion. Only four tooth-marked specimens, including one likely crocodile-tooth-marked bone, are preserved in all three archaeofaunas. A paucity of limb epiphyses suggests that bone-crunching hyenids may have deleted these portions subsequent to hominin butchery. Strangely, there are no stone tools preserved with the 292 cut-marked and 27 percussion-marked faunal specimens (out of a total of 6,039 specimens), suggesting that raw material availability may have conditioned hominin lithic discard patterns at these locales. These assemblages increase our knowledge of the dietary behavior and ecology of Homo erectus, and provide support for variability in early Pleistocene hominin carcass foraging patterns.     

Taphonomic analysis of the Middle Stone Age faunal assemblage from Pinnacle Point Cave 13B, Western Cape, South Africa

A detailed taphonomic analysis is provided for the mammalian and tortoise faunal assemblages from Pinnacle Point Cave 13B (PP13B). It is the first of several reports on the fauna from this site, and must necessarily precede analyses focused on higher level interpretations of Middle Stone Age (MSA) butchery, transport, and hunting behavior. The taphonomic work shows that the faunal assemblage is well preserved and there are discernable differences in the taphonomic pathways to which the fauna was subjected at PP13B between the Middle and Late Pleistocene, between the front and back of the cave, and between body size classes. The largest mammals (size classes 2-5, body weight >24?kg) were mainly accumulated by MSA hominins. Size class 1 ungulates also exhibit a degree of hominin modification consistent with some hominin accumulation of fresh carcasses, but this is more variable through time and includes an observable degree of independent carnivore contribution. Basic taxonomic comparisons reveal a low representation of small mammals, tortoises, and marine mammals at PP13B relative to larger (>4.5?kg) terrestrial mammals. This is a different pattern from other MSA sites along the southwestern coast of South Africa, where small mammals and tortoises are abundant. A microscopic study of the bone surfaces confirms that MSA hominins exploited these small faunal components opportunistically, while focusing most heavily on large terrestrial ungulates. All faunal components show evidence of carnivore scavenging of hominin food debris and a high degree of density mediated destruction. Raptors are at no point implicated as major accumulators of any fauna. The study demonstrates that the full spectrum of MSA faunal exploitation can only be understood when the large mammal, small mammal, and tortoise components of fossil assemblages have all been subjected to comprehensive taphonomic analyses.     

Leftover prey remains: a new taphonomic mode from the Late Miocene Cerro Azul Formation of Central Argentina

The Cerro Azul Formation (La Pampa Province, Argentina) comprises a rich vertebrate fauna of small mammals dominated by notoungulates and rodents. The fauna pertains to the Late Miocene specifically to the Huayquerian Stage/Age. Taphonomic analysis of micro‐mammals from Estancia Ré locality evidenced that the faunal assemblage was accumulated by the activity of a predator. This assemblage was compared with others from the Cerro Azul Formation in Telén and Caleufú localities (La Pampa Province), previously interpreted as products of predator activities. These microfossil accumulations differ from assemblages attributed to pellets and faeces produced by modern predators (nocturnal and diurnal bird raptors and carnivore mammals). However, due to their anatomical representation, degree of bone breakage and presence of tooth marks, they are interpreted as accumulations of uneaten prey remains discarded by the predator. The predator involved could not be determined with certainty, although the presence of tooth marks in some skeletal remains and the presence of coprolites in Telén and Caleufú suggest that it could be a carnivore mammal. Similarities in the accumulation mechanism, patterns of preservation and sedimentary contexts in the three assemblages support the recognition of a new taphonomic mode, termed ‘leftover prey remains’.     

Are local patterns of anthropoid primate diversity related to patterns of diversity at a larger scale?

Aims (1) To determine the relationship between local and regional anthropoid primate species richness. (2) To establish the spatial and temporal scale at which the ultimate processes influencing patterns of primate species coexistence operate. Location Continental landmasses of Africa, South America and Asia (India to China, and all islands as far south as New Guinea). Methods The local–regional species richness relationship for anthropoid primates is estimated by regressing local richness against regional richness (independent variable). Local richness is estimated in small, replicate local assemblages sampled in regions that vary in total species richness. A strong linear relationship is taken as evidence that local assemblages are unsaturated and local richness results from proportional sampling of the regional pool. An asymptotic curvilinear relationship is interpreted to reflect saturated communities, where strong biotic interactions limit local richness and local processes structure the species assemblage. As a further test of the assumption of local assemblage saturation, we looked for density compensation in high‐density local primate assemblages. Results The local–regional species richness relationship was linear for Africa and South America, and the slope of the relationship did not differ between the two continents. For Asia, curvilinearity best described the relationship between local and regional richness. Asian primate assemblages appear to be saturated and this is confirmed by density compensation among Asian primates. However, density compensation was also observed among African primates. The apparent assemblage saturation in Asia is not a species–area phenomenon related to the small size of the isolated islands and their forest blocks, since similar low local species richness occurs in large forests on mainland and/or peninsular Asia. Main conclusions In Africa and South America local primate assemblage composition appears to reflect the influence of biogeographic processes operating on regional spatial scales and historical time scales. In Asia the composition of primate assemblages are by‐and‐large subject to ecological constraint operating over a relatively small spatial and temporal scale. The possible local influence of the El Niño Southern Oscillations on the evolution and selection of life‐history characteristics among Asian primates, and in determining local patterns of primate species coexistence, warrants closer inspection.