Meat-eating by early hominids at the FLK 22Zinjanthropussite, Olduvai Gorge (Tanzania): an experimental approach using cut-mark data

The meat-eating behavior of Plio-Pleistocene hominids, responsible for the bone accumulations at the earliest archaeological sites, is still a hotly-debated issue in paleoanthropology. In particular, meat-eating and bone marrow consumption are often presented as either complementary or opposing strategies of carcass exploitation. The presence of cut marks on fossil archeofauna is a potential source of information that has not been consistently used as evidence of carcass consumption by hominids. Some authors interpret cut marks as the result of hominids manipulating meat-bearing bones, while others argue that they can also be the result of hominids extracting marginal scraps of carcass flesh that have survived carnivores’ initial consumption. In this study, a referential framework concerning the interpretation of cut marks is presented, based on a set of experiments conducted by the author. It is suggested, according to these experiments and data drawn from the FLK “Zinj” site, that hominids processed meat-bearing bones (on which flesh was abundant) rather than defleshed carcasses from felid kills.     

Cutmarked bones from Pliocene archaeological sites at Gona, Afar, Ethiopia: implications for the function of the world's oldest stone tools

Newly recorded archaeological sites at Gona (Afar, Ethiopia) preserve both stone tools and faunal remains. These sites have also yielded the largest sample of cutmarked bones known from the time interval 2.58-2.1 million years ago (Ma). Most of the cutmarks on the Gona fauna possess obvious macroscopic (e.g., deep V-shaped cross-sections) and microscopic (e.g., internal microstriations, Herzian cones, shoulder effects) features that allow us to identify them confidently as instances of stone tool-imparted damage caused by hominid butchery. In addition, preliminary observations of the anatomical placement of cutmarks on several of the recovered bone specimens suggest that Gona hominids may have eviscerated carcasses and defleshed the fully muscled upper and intermediate limb bones of ungulates--activities that further suggest that Late Pliocene hominids may have gained early access to large mammal carcasses. These observations support the hypothesis that the earliest stone artifacts functioned primarily as butchery tools and also imply that hunting and/or aggressive scavenging of large ungulate carcasses may have been part of the behavioral repertoire of hominids by c. 2.5 Ma, although a larger sample of cutmarked bone specimens is necessary to support the latter inference.     

Hominid carnivory and foraging strategies, and the socio-economic function of early archaeological sites.

New evidence for the tissue types exploited by early hominids from carcasses possibly acquired through scavenging is derived from the larger mammal bone assemblages from FLK I, level 22 (Zinjanthropus floor), and FLKN levels 1 and 2 from Bed I, Olduvai Gorge, Tanzania. Published skeletal part profiles from the two archaeological sites are evaluated using (i) modern observations on the sequence by which carnivores consume carcass parts in order to assess the timing of hominid access to carcasses, and (ii) measurements of flesh and marrow yields to assess the tissue types sought and acquired. These results suggest that the maximization of marrow (fat) yields, not flesh (protein) yields, was the criterion shaping decisions about carcass processing. Because of evidence for density-dependent destruction of some flesh-bearing parts by scavengers of the hominid-butchered assemblages, however, it is uncertain whether carcass parts were transported and acquired by hominids in a largely defleshed condition. The results on tissue types acquired are combined with a discussion of predation risk, feeding competition and the equipment needs of carcass processing in an attempt to identify archaeological test implications of competing hypotheses for the socio-economic function of the earliest archaeological sites.     

New estimates of tooth mark and percussion mark frequencies at the FLK Zinj site: the carnivore-hominid-carnivore hypothesis falsified

Traditional interpretations of hominid carcass acquisition strategies revolve around the debate over whether early hominids hunted or scavenged. A popular version of the scavenging scenario is the carnivore-hominid-carnivore hypothesis, which argues that hominids acquired animal resources primarily through passive opportunistic scavenging from felid-defleshed carcasses. Its main empirical support comes from the analysis of tooth mark frequency and distribution at the FLK Zinj site reported by Blumenschine (Blumenschine, 1995, J. Hum. Evol. 29, 21-51), in which it was shown that long bone mid-shafts exhibited a high frequency of tooth marks, only explainable if felids had preceded hominids in carcass defleshing. The present work shows that previous estimates of tooth marks on the FLK Zinj assemblage were artificially high, since natural biochemical marks were mistaken for tooth marks. Revised estimates are similar to those obtained in experiments in which hyenas intervene after humans in bone modification. Furthermore, analyses of percussion marks, notches, and breakage patterns provide data which are best interpreted as the results of hominid activity (hammerstone percussion and marrow extraction), based on experimentally-derived referential frameworks. These multiple lines of evidence support previous analyses of cut marks and their anatomical distribution; all indicate that hominids had early access to fleshed carcasses that were transported, processed, and accumulated at the FLK Zinj site.     

Taphonomic perspectives on hominid site use and foraging strategies during Bed II times at Olduvai Gorge, Tanzania

The faunal assemblages excavated by Mary Leakey in Bed II of Olduvai Gorge, Tanzania, have, like the more well-known Bed I assemblages, traditionally been interpreted as the result of hominid butchering activities in the lake margin and riverine settings of the paleo-Olduvai Basin. A reexamination of all of Leakey's Bed I sites has shown that hominids played little or no role in the formation of all but one of those faunal assemblages, a finding that prompted the reanalysis of the Bed II sites presented here. We expand upon a previous taphonomic study that provided systematic data for HWK East Levels 1–2, MNK Main, and BK. In addition to these assemblages, we provide data on HWK East Levels 3–5, FC West, TK, and SHK. Our data contradict previous interpretations of MNK Main as a hominid accumulation but uphold the contention that BK represents a primarily hominid accumulation reflecting early access to carcasses. The small and poorly preserved assemblages from FC West and TK are difficult to link unambiguously to either hominids or carnivores. Site MNK Main and HWK East Levels 3–5 appear to be death arenas where carcasses accumulated via natural deaths and/or serial predation. Site SHK is severely biased by selective retention and therefore little can be said of its formational history. Nevertheless, no hominid modifications were documented in this assemblage. Comparisons with other Olduvai sites indicate a more conspicuous hyena taphonomic signal during Bed II times than Bed I times, which appears to mirror the changing configuration of the large carnivore guild. These findings also beg the question of what activities were being carried out by hominids with the stone tools discarded at these sites. Although it seems clear that hominids were utilizing stone tools to carry out subsistence activities unrelated to carcass butchery, more excavation and techniques such as phytolith analysis should be employed to explore alternative explanations.     

Modern hunting behavior in the early Middle Paleolithic: faunal remains from Misliya Cave, Mount Carmel, Israel

Understanding the behavioral adaptations and subsistence strategies of Middle Paleolithic humans is critical in the debate over the evolution and manifestations of modern human behavior. The study of faunal remains plays a central role in this context. Until now, the majority of Levantine archaeofaunal evidence was derived from late Middle Paleolithic sites. The discovery of faunal remains from Misliya Cave, Mount Carmel, Israel (>200 ka), allowed for detailed taphonomic and zooarchaeological analyses of these early Middle Paleolithic remains. The Misliya Cave faunal assemblage is overwhelmingly dominated by ungulate taxa. The most common prey species is the Mesopotamian fallow deer (Dama mesopotamica), followed closely by the mountain gazelle (Gazella gazella). Some aurochs (Bos primigenius) remains are also present. Small-game species are rare. The fallow deer mortality pattern is dominated by prime-aged individuals. A multivariate taphonomic analysis demonstrates (1) that the assemblage was created solely by humans occupying the cave and was primarily modified by their food-processing activities; and (2) that gazelle carcasses were transported complete to the site, while fallow deer carcasses underwent some field butchery. The new zooarchaeological data from Misliya Cave, particularly the abundance of meat-bearing limb bones displaying filleting cut marks and the acquisition of prime-age prey, demonstrate that early Middle Paleolithic people possessed developed hunting capabilities. Thus, modern large-game hunting, carcass transport, and meat-processing behaviors were already established in the Levant in the early Middle Paleolithic, more than 200 ka ago.     

Validation of bone surface modification models for inferring fossil hominin and carnivore feeding interactions, with reapplication to FLK 22, Olduvai Gorge, Tanzania

Resolving the issue of how Early Stone Age hominins acquired large mammal carcasses requires information on their feeding interactions with large carnivores. This ecological information and its behavioral and evolutionary implications are revealed most directly from the tooth, cut, and percussion marks on bone surfaces generated by hominin and carnivore feeding activities. This paper employs a bootstrap method, a form of random resampling with replacement, to refine published neotaphonomic models that use the assemblage-wide proportions of long bones bearing feeding traces to infer the sequences in which Plio-Pleistocene hominins and carnivores accessed flesh, marrow, and/or grease from carcasses. Results validate the sensitivity of the models for inferring hominin feeding ecology, which have been questioned on grounds shown here to be unfounded. The bootstrapped feeding trace models are applied to the late Pliocene larger mammal fossil assemblage from FLK 22 (Zinjanthropus site), Olduvai Gorge, Tanzania. High frequencies of tooth and percussion marking on long bone midshaft fragments from FLK 22 are most consistent with those feeding trace models that simulate hominin scavenging from carcasses defleshed by carnivores, while cut mark data indicate that hominins more often had access to upper forelimb flesh than upper hind limb flesh. Together, the bone surface modification data indicate that hominins typically gained secondary access to partially defleshed carnivore kills, but they also allow for the possibility of some carcasses being processed only by carnivores and only by hominins.     

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.     

A study of carnivore competition in riparian and open habitats of modern savannas and its implications for hominid behavioral modelling

Some of the models proposed to explain Plio-Pleistocene hominid behavior and the formation of early East African archaeological sites are based on the assumption that the riparian habitats in which most of them occur were places of low interspecific competition. Competition is expressed here in terms of carnivore and hominid interactions. In this paper, a study of carnivore interaction in open and closed habitats is presented. The results indicate that riparian woodland shows the lowest degree of competition in savanna ecosystems. This suggests that if Plio-Pleistocene carnivores were adapted like their modern counterparts, the paleoecological settings of early sites could have provided hominids with enough safety to process carcasses and behave as shown in "central-place", "near-kill location" and "refuge" foraging models.     

Cranial base morphology and temporal bone pneumatization in Asian Homo erectus

The external morphological features of the temporal bone are used frequently to determine taxonomic affinities of fossils of the genus Homo. Temporal bone pneumatization has been widely studied in great apes and in early hominids. However, this feature is rarely examined in the later hominids, particularly in Asian Homo erectus. We provide a comparative morphological and quantitative analysis of Asian Homo erectus from the sites of Ngandong, Sambungmacan, and Zhoukoudian, and of Neandertals and anatomically modern Homo sapiens in order to discuss causes and modalities of temporal bone pneumatization during hominid evolution. The evolution of temporal bone pneumatization in the genus Homo is more complex than previously described. Indeed, the Zhoukoudian fossils have a unique pattern of temporal bone pneumatization, whereas Ngandong and Sambungmacan fossils, as well as the Neandertals, more closely resemble the modern human pattern. Moreover, these Chinese fossils are characterized by a wide midvault and a relatively narrow occipital bone. Our results support the point of view that cell development does not play an active role in determining cranial base morphology. Instead, pneumatization is related to available space and to temporal bone morphology, and its development is related to correlated morphology and the relative disposition of the bones and cerebral lobes. Because variation in pneumatization is extensive within the same species, the phyletic implications of pneumatization are limited in the taxa considered here.     

Adults only. Reindeer hunting at the middle palaeolithic site salzgitter lebenstedt, northern Germany

The Middle Palaeolithic site Salzgitter Lebenstedt (northern Germany), excavated in 1952, is well known because of its well-preserved faunal remains, dominated by adult reindeer (Rangifer tarandus). The archaeological assemblage accumulated in an arctic setting in an earlier part of the last (Weichsel) glacial (OIS5-3). The site is remarkable because of the presence of unique Middle Palaeolithic bone tools and the occurrence of the northernmost Neanderthal remains, but this paper focuses on an analysis of its reindeer assemblage. The results indicate autumn hunting of reindeer by Middle Palaeolithic hominids. After the hunt, carcasses were butchered and in subsequent marrow processing of the bones a selection against young and sub-adult animals occurred. Adults were clearly preferred, and from their bones, again, poorer marrow bones were neglected. This focus on primeness of resources has been documented in other domains of Neanderthal behaviour, but Salzgitter Lebenstedt is the best example yet known in terms of systematic and routinized processing of game. The Salzgitter Lebenstedt assemblage displays some remarkable similarities to the Late Glacial reindeer assemblages from the Ahrensburg tunnel valley sites. The subsequent review of the evidence on subsistence strategies from earlier periods of the European Palaeolithic shows that hunting of large mammals may have been a part of the behavioural repertoire of the Middle Pleistocene occupants of Europe from the earliest occupation onwards. At the same time, it is suggested that these early hunting strategies were incorporated in ways of moving through landscapes ("settlement systems") which were different from what we know from the middle parts of the Upper Palaeolithic onwards.     

Evolution of femur and tibia in higher primates: Adaptive morphological patterns and phylogenetic diversity

Seventy six metrical traits measured on the femur and tibia of three higher primate groups —Ceboidea, Cercopithecoidea, Hominoidea have been processed by various univariate and multivariate statistical methods to survey the process of evolution of the morphology of the femur and tibia in higher primates. Intragroup and intergroup variability, similarity and differences as well as various aspects of scaling and sexual dimorphism have been analyzed to study adaptive trends and phylogenetic diversity in higher primates, in individual superfamilies and to explore the adaptive morphological pattern of early hominids and basic differences between hominids and pongids. Two basic morphotypes of the femur and tibia in higher primates have been determined. They are (1) advanced hominoid morphotype (hominids and pongids) and (2) ancestral higher primate morphotype (platyrrhine and cattarrhine monkeys, early hominoids, and hylobatids). Cebid lower limb bones are adapted to arboreal quadrupedalism with antipronograde features while femur and tibia of cercopithecid monkeys are basically adapted to the semi-arboreal locomotion. Early hominoids (Proconsul) and hylobatids are morphologically different from pongids; some features are close toAteles or other monkey species. Pongids and hominids are taken as one major morphological group with different scaling and some functional and morphological similarities. Numerous analogous features were described on the lower limb skeleton ofPan andPongo showing analogous ecological parameters in their evolution. Major morphological and biomechanical trends are analyzed. It is argued that early advanced hominoid morphology is ancestral both to the pongids and to early hominids. The progressive morphological trend in early hominids has been found fromA. afarensis with ancestral hominid morphology, toH. habilis with an elongated femur and structural features similar to advanced hominids. A detailed phylogenetic analysis of higher primate femur and tibia is also presented.     

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.     

Who was the first? An experimental application of carnivore and hominid overlapping marks at the Pleistocene archaeological sites

There are many types of evidence that provide information about methods for obtaining animal nutrients. Several researchers suggest that the main element to be considered is the skeletal representation of the different species identified in the faunal assemblage. This element must be associated to the animals’ age at death and the localisation of processing marks of the carcasses (both those of anthropic origin and those produced by carnivores). Occasionally, these marks coincide on the same point of the bone, giving cause for overlapping marks. These marks can be considered an aid more to identify the anthropic manner for obtaining animal recourses. However, these cases are very unusual at archaeological sites, and it is not always easy to identify which of the two predators has obtained the prey first. Through the experimental process presented in this article, we have observed diagnostic elements on overlapping marks that show the action sequence of the predators (hominids and carnivores) on carcasses. These experimental criteria were applied to different archaeological sites of the Lower and Middle Pleistocene in the Iberian Peninsula: Bolomor Cave (Valencia, Spain) and level TD10-1 and TD6-2 of Gran Dolina (Atapuerca, Burgos). In these assemblages, we were able to distinguish hunting and scavenging events through overlapping marks, providing a new element to the general interpretation of these sites.     

泥河湾盆地板井子晚更新世遗址古人类对动物资源的消费策略

本文以泥河湾盆地板井子遗址（约90 kaBP）2015年出土的动物骨骼为材料,通过骨骼表面痕迹分析、骨骼单元出现频率研究及长骨破碎方式的统计,辨识自然作用与人类行为对动物骨骼造成的不同影响,讨论深海氧同位素5阶段(MIS 5)中国北方古人类获取、搬运及处理动物资源的综合性策略。研究结果显示：1)遗址出土的动物遗存为古人类活动所遗留,自然营力对动物骨骼有改造作用,但并未影响我们对古人类行为的辨识;2)古人类在捕获马科动物之后,将其完整地带回了遗址并肢解;3)在处理猎物遗骸时,肌肉和骨髓富集的部位是古人类利用的重点。结合河南许昌灵井遗址的动物遗存研究结果,我们认为,在MIS 5阶段前后,中国北方已经出现了可以高效利用动物资源的人类群体;稳定、充沛的能量来源,或是他们应对北半球中高纬度干旱草原环境气候波动的重要生存策略。     

What happens to dead birds left in a cave and its vicinity for a quarter of a century? A taphonomic experiment

Abstract

This paper describes a bird carcass-monitoring experiment carried out in a limestone cave and its immediate vicinity in southern Poland for almost a quarter of a century. Some specimens deposited outside the cave were preserved almost intact, others were only a little weathered. Soft tissues and remnants of feathers were preserved in some of the samples inside the cave. Stages of bone modifications that can occur in a cave have been distinguished and illustrated for the first time. Also, bird bones from owl pellets have been used in a carcass-monitoring experiment for the first time. Bone modifications in the form of wrinkling, bending and/or collapsing of the bone surface were recorded only on the bones of birds that decomposed inside the cave, and not on bones that had been isolated from owl pellets prior to their deposition in the cave. Therefore, it is postulated that avian – and possibly also small mammal – bones that have been defleshed, either by people or animals, have a greater chance to survive and fossilize than bones deposited as complete carcasses in a cave. This agrees with the generally accepted notion that larger assemblages of small vertebrates in caves are usually the result of predator accumulation.     

Analysis of a bone assemblage made by chimpanzees at Gombe National Park, Tanzania

Chimpanzee hunting provides information on prey characteristics and constraints acting on a large-bodied primate lacking a hunting technology, and has important implications for modeling hunting by fossil hominids. Analysis of the remains of five red colobus monkeys captured and consumed by Gombe chimpanzees in a single hunting bout provides one of the first opportunities to investigate the characteristics of prey bones surviving chimpanzee consumption. Four of the five individuals (an older infant, two juveniles and one subadult) were preserved in the bone assemblage; a neonate was entirely consumed. Cranial and mandibular fragments had the highest survivorships, followed by the scapulae and long bones. Post-cranial axial elements had the lowest survivorships. A high percentage (80%) of the long bones and ribs surviving consumption were damaged, most commonly through crenulation and step fracturing of bone ends. One of two partially reconstructed crania preserves a canine puncture through its left parietal. Proposed characteristics of faunal assemblages formed through chimpanzee-like hunting include small modal prey size, limited taxonomic diversity, a high proportion of immature individuals and a high frequency of skull bones. These characteristics would not uniquely identify hunting by fossil primates in the geological record, necessitating a contextual approach to diagnose hunting by hominids not forming an archeological record.Hominid utilization of vertebrate tissue is first unambiguously documented at 2.5 m.y.a. Rather than representing a strict "scavenging phase" in the evolution of hominid-prey interactions, Oldowan hominid carnivory may represent the overlay of large mammal scavenging on a tradition of small mammal hunting having a low archeological visibility.     

The parietal bone of indonesian homo erectus

A comparative study of Indonesian parietal bones from Sangiran, Sambungmachan 1 and Ngandong has been undertaken. This study comprises a morphological and metrical analysis of the individual parietal bones, followed by consideration of the biparietal vault. The results are compared with other hominids from earlier and later periods. These hominids were found in China (Sinanthropus II, III, X, XI and XII), in Africa (ER 3733, OH 9, Ternifine, Broken Hill and Saldanha) and in Europe (Arago XLVII, Petralona, Swanscombe, Steinheim, Le Lazaret, La Chaise (Abri Suard) and Cova Negra). These European Middle Pleistocene hominids are attributed toHomo erectus by various authors (Lumley 1973;Hemmer 1972;Spitery 1982;Lumley andFournier 1982) and to an early Neanderthal group, pre-Neanderthal orHomo sapiens sensu lato (Neanderthals+modern humans) by others (Stringer 1980, 1981, 1983, 1984,Wolpoff 1980,Holloway 1982). The discussion about the classification of those hominids is not closed, but it is not the subject of this paper and not our intention to solve it here. So we have chosen to call this fossil material ‘Anteneandertals’ (Lumley 1973). It appears that some morphological metrical features allow us to separate the Sangiran and Ngandong samples. Sambungmachan 1, whose chronological age is not well established, appears to be closer to Ngandong men.     

Multivariate analysis of the sexual dimorphism of the hip bone in a modern human population and in early hominids

A large sample of hip bones of known sex coming from one modern population is studied morphologically and by multivariate analysis to investigate sexual dimorphism patterns. A principal component analysis of raw data shows that a large amount of the hip bone sexual dimorphism is accounted for by size differences, but that sex-linked shape variation is also very conspicuous and cannot be considered an allometric consequence of differences in body size between the sexes. The PCA of transformed (“shape”) variables indicates that the female hip bones are different in those traits associated with a relatively larger pelvic inlet (longer pubic bones, a greater degree of curvature of the iliopectineal line, and a more posterior position of the auricular surface), as well as a broader sciatic notch. The analysis of nonmetric traits also shows marked sexual dimorphism in the position of the sacroiliac joint in the iliac bone, in the shape of the sciatic notch, in pubic morphology, and in the presence of the pre-auricular sulcus in females. When the australopithecine AL 288-1 and Sts 14 hip bones are included in the multivariate analysis, they appear as “ultra-females.” In particular these early hominids exhibit extraordinarily long pubic bones and iliopectineal lines, which cannot be explained by allometry. © 1994 Wiley-Liss, Inc.