Examining criteria for identifying and differentiating fossil faunal assemblages accumulated by hyenas and hominins using extant hyenid accumulations

Numerous authors have put forth criteria for distinguishing between assemblages collected by hyenas and hominins. Of the seven most recognised criteria used to distinguish hyenid from hominin assemblages, it has recently been suggested that four be rejected and three retained. The four rejected criteria are: an excessive proportion of horns and horn cores in hyena accumulated assemblages; the absence of small, hard, compact bones; mortality profiles; and the ratio of cranial bones to postcranial bones. The three criteria previous researchers suggested be retained are: a carnivore MNI ratio of ≥20%; an abundance of cylinder fragments; and hyena‐inflicted damage upon the bones. In this examination of over 27,000 faunal remains associated with all three species of extant bone‐collecting hyenids from four countries and two continents, six of the seven previously established criteria and reconsiderations of criteria have been evaluated. The results of the present study indicate that of the six criteria examined, none, as written, are indicative of hyenid activity on bone assemblages of unknown origin. Copyright © 2008 John Wiley & Sons, Ltd.     

Spotted hyena (Crocuta crocuta) den use and taphonomy in the Masai Mara National Reserve,Kenya

Hyena taphonomy is of great importance to studies of hominid evolution, since these carnivore taxa have the highest potential both to produce large osseous assemblages and to modify existing hominid-accumulated assemblages throughout the Old World. The three extant hyena species (brown: Parahyaena brunnea; striped: Hyaena hyaena; and spotted: Crocuta crocuta) are all significant bone collectors and modifiers. Spotted hyenas generally have the lowest potential to accumulate osseous remains, and the rate of accumulation varies based upon the type of den. The present research examines the remains accumulated by spotted hyenas in Masai Mara Cave, Masai Mara National Reserve, Kenya. The contents of this den were collected twice by the authors, with an 11-year span interceding. The taphonomic signatures of spotted hyena interaction with bone are presented, including species and skeletal element representation, breakage patterns, tooth marks, tooth puncture, edge polish, and gastric corrosion. Other taphonomic factors examined include rodent gnawing and weathering stage. The cave den assemblage was accumulated at a rate of 30.4 identified specimens and a minimum of 4.1 prey individuals per year. In addition, the osseous remains accumulated by spotted hyenas at multiple burrow dens within the Reserve were examined for taxonomic representation and multiple taphonomic parameters. Rates of accumulation at this type of den tend to be very low, due to differential usage by spotted hyenas and the more ephemeral nature of earthen dens.     

Baboon Bone Mineral Densities: Implications for the Taphonomy of Primate Skeletons in South African Cave Sites

C. K. Brain (e.g., 1981) documented an interesting difference in relative skeletal part representation between primates and bovids of similar live body size recovered from the fossil cave site of Swartkrans (South Africa). Hominids and baboons are represented primarily by skull parts and a paucity of postcranial bones, while small sized bovids (Size Classes 1 & 2) are represented by more equivalent abundances of all skeletal elements. Brain argued that the Swartkrans bones were primarily carnivore collected, and postulated that the difference in element frequencies between primates and bovids is the manifestation of less durable primate postcranial skeletons relative to those of bovids when either are subjected to a destructive process such as carnivore feeding. Experiments by Brain (1981) in which baboon and bovid carcasses were fed to large, African carnivores lend support to this hypothesis. Bovid postcranial elements survived carnivore feeding more frequently and more completely than those of primates. Assuming that bulk bone mineral density (bulk BMD) is a measure of durability, we supplement Brain's observational data by presenting the first systematic, element-by-element comparison of baboon and bovid postcranial bulk BMD measurements obtained by dual energy X-ray absorptiometry (DXA). While the bovid in our sample exhibits greater absolute bulk BMD values in 57·8% of all measured bone areas, when grouping areas by body region, a statistically significant difference in absolute bulk BMD values between the baboons and bovids is documented only in the hindlimb region (i.e., femur, tibia and patella). Density differences in other body regions are not statistically significant. Further, in only one case (i.e., small bovids in the Member 2 [1948–1953/1965–1975 excavation] assemblage) is there a significant and positive correlation between skeletal part frequencies of primates or bovids and bulk BMD. This suggests that, for the most part, primate and bovid skeletal part representation at Swartkrans is not the result of density-mediated processes. Differences in skeletal part representation between primates and bovids at Swartkrans may thus be attributable to factors other than bone density—such as bone size, length, shape, and/or the relative palatability of surrounding soft tissues on bones.     

Carnivores and their prey in the Wezmeh Cave (Kermanshah,Iran): a Late Pleistocene refuge in the Zagros

Wezmeh Cave is located on the northeastern edge of the Islamabad plain, a high intermontane valley in the western‐central Zagros. In 1999 a disturbed but large faunal assemblage was recovered from this site. The abundant and extremely diverse faunal spectra present at Wezmeh Cave has highlighted the importance of this assemblage. Carnivore remains constitute the bulk of the assemblage; red fox (Vulpes vulpes) has the highest number of identified specimens followed by spotted hyena (Crocuta crocuta), brown bear (Ursus arctos), wolf (Canis lupus), felids (lion, leopard, lynx/caracal and wildcat), mustelids (badger, polecat, marten) and viverrids (mongoose). Artiodactyls (bovid, cervid, suid), equids, rhinoceros (Dicerorhinus sp.) and small animals (Cape hare, porcupine, tortoise, snake, birds) are also present. According to U‐series dating, the site was occupied from around 70 ka BP through to sub‐recent periods by carnivores. Amongst this rich assemblage, a human fossil tooth was also found and dated by non‐invasive spectrometry gamma dating to 20–25 ka BP. A preliminary zooarchaeological and taphonomic study shows that Wezmeh Cave was used by multiple carnivore species, a unique phenomenon in the Zagros Mountains in particular and southwest Asia in general. Copyright © 2008 John Wiley & Sons, Ltd.     

FOCUS: on the use of the petrous bone for estimating cranial abundance in fossil assemblages

Low representation of braincase bones in zooarchaeological assemblages suggests that skulls have been intensively processed by Levantine Epipalaeolithic foragers; most cranial elements are often unidentifiable and are considered poor candidates for quantifying crania. In contrast, the petrous bone is usually found complete, and was found to be easily identifiable to body size category. Use of the petrous bone in fossil assemblages analyses leads to better estimation of the occurrence of cranial elements, and thus of skeletal part representation. We therefore suggest use of the petrous bone for detecting bone destruction and selective transport in faunal assemblages.     

Taphonomy and zooarchaeology of the Upper Palaeolithic cave of Dzudzuana,Republic of Georgia

We present the results of a detailed taphonomic and zooarchaeological study of the faunal remains from the Upper Palaeolithic layers of Dzudzuana Cave, Republic of Georgia. This study presents the first carefully analysed Upper Palaeolithic faunal assemblage from the southern Caucasus and thus serves as a significant point of reference for inter‐regional studies of Upper Palaeolithic subsistence in Eurasia. A series of intra‐site taphonomic comparisons are employed to reconstruct the depositional history of the bone assemblages within the different occupational phases at the site and to investigate subsistence, meat procurement and bone‐processing strategies. Caucasian tur (Capra caucasica), aurochs (Bos primigenius) and steppe bison (Bison priscus) were the major prey species throughout the Upper Palaeolithic. Their frequencies do not change significantly over time, and nor does bone preservation vary by layer. The assemblage is characterised by significant density‐mediated biases, caused by both human bone‐processing behaviours and in situ post‐burial bone attrition. Bone marrow extraction produced large numbers of unidentified bone fragments, many exhibiting green bone fractures. The density and size of bone assemblages and the extent of fragmentation indicate that Dzudzuana Cave was repeatedly occupied by Upper Palaeolithic foragers over many years. Skeletal part representation and butchery marks from all stages of carcass processing suggest that prey occasionally underwent field butchery. Intra‐site taphonomic comparisons highlight uniform patterns of cultural and economic behaviours related to food procurement and processing strategies. Copyright © 2007 John Wiley & Sons, Ltd.     

New analytical methods for comparing bone fracture angles: A controlled study of hammerstone and hyena (Crocuta crocuta) long bone breakage

Accurate interpretation of the cause and timing of bone breakage is essential for understanding the archaeological record. However, many variables potentially influencing break morphology have yet to be systematically explored. Focusing primarily on hammerstone breakage, we introduce new analytical methods for comparing fracture angles using the absolute values of the angle from 90°. We systematically control for intrinsic variables such as taxon, skeletal element, limb portion and skeletal age. We also compare experimental assemblages of femora broken by hammerstone and spotted hyena (Crocuta crocuta). We show that fracture angles are influenced by breakage plane, skeletal element and limb portion. While the latter two have been suggested before, this is the first time the differences have been quantified. We suggest that researchers stratify their assemblages by these variables if they are using fracture angles in analyses. At the assemblage level, hyenas created more oblique fracture angles on oblique breaks than did hammerstones.     

Skeletal element abundances in archaeofaunal assemblages: economic utility,sample size,and assessment of carcass transport strategies

Economic utility indices provide a means of interpreting butchery and transport decisions reflected in the relative abundance of skeletal elements. Because of destructive taphonomic processes, interpreting skeletal element abundances in terms of carcass transport strategies requires that faunal analysts consider only those elements which accurately reflect their original abundances following human discard. In this study we use resampling techniques to examine the impact of sample size on correlations between high-survival skeletal element frequencies and economic utility in four simulated population assemblages reflecting distinct carcass transport strategies. Correlations alone do not accurately reflect the true relationship between bone abundance and economic utility as particular transport strategies have a tendency to generate high frequencies of Type II errors as sample size decreases. We show that the Shannon evenness index can be used as a quantitative means of distinguishing between bone assemblages characterized by subtle variations in skeletal element abundances. The evenness index can also be used to evaluate whether observed correlations reflect sampling error. Results from our simulations are applied to three published faunal assemblages to evaluate likely carcass transport strategies.     

Why are cut mark frequencies in archaeofaunal assemblages so variable? A multivariate analysis

Cut mark frequencies in archaeological faunal assemblages are so variable that their use has recently created some skepticism. The present study analyses this variability using multivariate statistics on a set of 14 variables that involve differential skeletal element representation, fragmentation processes, carnivore ravaging impact, carcass size and tool type. All these variables affect the resulting cut mark frequencies reported in archaeological sites. A large sample of archaeofaunal assemblages has been used for this study. It was concluded that the best estimator of cut mark frequency in any given assemblage is the percentage of cut-marked long bone specimens (probably due to its better preservation than other anatomical areas), which is determined by fragmentation and carnivore ravaging. Carcass size and tool type also play a major role in differences in cut mark frequencies. Fragmentation is also a key variable determining the abundance of cut-marked specimens. It is argued that general cut mark percentages are of limited value, given the number of variables that determine them, and that a more heuristic approach involves quantifying cut marks in a qualitative manner.     

TESTING ANALOGICAL TAPHONOMIC SIGNATURES IN BONE BREAKING: A COMPARISON BETWEEN HAMMERSTONE‐BROKEN EQUID AND BOVID BONES

Current analogical data used to infer prehistoric human bone breakage rely on a plethora of experimental hammerstone‐broken bovid bone sets. Several criteria have been argued to be diagnostic of bone breakage by humans, among which the most important are: a specific range of broken specimens bearing percussion marks, a specific distribution of different percussion mark types, metric properties of notches, differential notch type distribution, and the angle of oblique breakage planes. The present work shows that those properties derived from the breakage of bovid bones cannot be universally applied to all types of animals. As an example, here it is experimentally demonstrated that hammerstone‐broken equid bones (with different thickness and structural properties compared to bovid bones) show different values in all these variables and some of them overlap with criteria documented in bones broken by static loading. This suggests that the agents of equid bone breakage are more difficult to identify, and that the number of variables that can be successfully used to that end is smaller than in bovid bones.     

“Measure for measure”: a taphonomic reconsideration of the Kebaran site of Ein Gev I,Israel

We present the results of a detailed taphonomic study of the faunal remains from Ein Gev I, a Kebaran open-air site near the Sea of Galilee, Israel. In an earlier study faunal remains were assigned to bone element and species using a rapid method technique based on the identification of teeth and epiphyses only (Davis, 1974). In this study we identified the maximum number of skeletal elements, including fragments of long bone shafts. We also employed a series of taphonomic analyses to reconstruct the depositional history of the bone assemblage and to investigate subsistence, meat procurement and bone processing strategies. We compared the results obtained by a complete bone sampling method with those derived using a rapid method. The bone assemblage is dominated by mountain gazelle (Gazella gazelle), Mesopotamian fallow deer (Dama mesopotamica) and bezoar goat (Capra aegagrus). The assemblage is characterized by significant density-mediated biases, caused by both human bone processing behaviours and in situ post-burial bone attrition. Our attempts to compare species abundance derived from the two different identification protocols revealed no significant difference between the two sampling methods, both in terms of species diversity (richness and evenness) and skeletal element abundance (SEA). These comparisons contrast previous studies that expected to find ubiquitous head-and-foot dominated SEA profiles in zooarchaeological studies that were based on the incomplete identification of faunal remains.     

Variation in tooth mark frequencies on long bones from the assemblages of all three extant bone-collecting hyaenids

Tooth mark frequencies on long bones are examined from the assemblages of all three extant bone-collecting hyaenids. Comparisons are made with a recent study examining tooth mark frequencies and possible sources of variation from a single spotted hyaena (Crocuta crocuta) assemblage (Faith, J.T., 2007. Sources of variation in carnivore tooth-mark frequencies in a modern spotted hyena (Crocuta crocuta) den assemblage, Amboseli Park, Kenya. Journal of Archaeological Science 34 (10), 1601–1609). The factors that may influence tooth mark frequencies are fragment size, fragments from different sized animals, region of skeletal element and bone density. All four factors are examined in the present study and compared across species and with previous results. The results indicate that there is a great deal of variation in tooth mark frequencies not only between the species but also from the same species.     

Hominid–Carnivore Coevolution and Invasion of the Predatory Guild

Coevolution is defined as reciprocal selective pressures that operate to make the evolution of one taxon partially dependent on the evolution another. This process often involves multiple species exploiting shared limiting resources. In classic coevolutionary models, populations of sympatric species are seen to diverge in one or more morphological, ecological, or behavioral traits to effect more even partitioning of resources and reduce levels of interspecific competition. Character displacement and resource partitioning are thought to be central not only to how species coexist on limited resources, but also to how species invade new resource niches. Hominid invasion of the predatory guild at least 2 my ago would have brought them into contact with a range of new selective pressures including competition with a number of large-bodied predators. This study explores resource partitioning between hominids and other Plio-Pleistocene large-bodied predators through analyses of predator food transport strategies. The anatomical content (head/limb MNE) of hominid transported faunal assemblages at Bed I Olduvai and FxJj 50 (Koobi Fora), when compared to modern predator control cases, suggests that Plio-Pleistocene hominids practiced food transport strategies intermediate between those of top predators such as wolves and those of confrontational scavengers such as spotted hyaenas. Plio-Pleistocene hominid food transport strategies do not resemble those of nonconfrontational scavengers such as brown and striped hyenas. The highly regular patterns differentiating bone assemblages accumulated by top predators, hominids, and confrontational and nonconfrontational scavengers suggest that hominid invasion of the predatory guild involved resource partitioning potentially based on some form of character displacement.     

Identifying the Involvement of Multiple Carnivore Taxa with Archaeological Bone Assemblages

Information on the number of carnivore taxa that were involved with archaeological bone assemblages is pertinent to questions of site formation, hominid and carnivore competition for carcasses and the sequence of hominid and carnivore activity at sites. A majority of early archaeological bone assemblages bear evidence that both hominids and carnivores removed flesh and/or marrow from the bones. Whether flesh specialists (felids) or bone-crunchers (hyaenas), or both, fed upon the carcasses is crucial for deciphering the timing of hominid involvement with the assemblages. Here we present an initial attempt to differentiate the tooth mark signature inflicted on bones by a single carnivore species versus multiple carnivore taxa. Quantitative data on carnivore tooth pits, those resembling a tooth crown or a cusp, are presented for two characteristics: the area of the marks in millimetres, and the shape as determined by the ratio of the major axis to the minor axis of the mark. Tooth pits from bones modified by extant East African carnivores and latex impressions of tooth pits from extinct carnivore species are compared to those in the FLK Zinjanthropus bone assemblage. Data on tooth mark shape indicate greater variability in theZinj sample than is exhibited by any individual extant or extinct carnivore species in the comparative sample. Data on tooth mark area demonstrate that bone density is related to the size of marks. Taken together, these data support the inference that felids defleshed bones in the Zinj assemblage and that hyaenas had final access to any grease or tissues that remained.     

Do faunal remains reflect socioeconomic status? An ethnoarchaeological study among Central African farmers in the northern Congo Basin

Zooarchaeological analyses often draw inferences on socioeconomic status from the composition of bone assemblages associated with houses and other structures in residential sites. In this paper, we test how well faunal assemblages reflect socioeconomic differences among contemporary farmer households in two rural villages in the Central African Republic. Independent measures of wealth are tallied and ranked for six households in each village, including complete inventories of the types and numbers of material goods and the sizes of residential structures and agricultural fields. These data are compared against the associated food bones collected from household trash middens and activity areas, including skeletal abundances, large mammal body part representation, and taxonomic diversity. In most instances larger and more taxonomically diverse faunal assemblages are associated with houses of means and the faunas do, in fact, reflect differences in socioeconomic status. However, faunal “wealth” may be linked to factors unrelated to social or economic inequalities, notably the presence of active hunters. Our analyses suggest that small animals provide useful and important data in assessing socioeconomic means, and comparative studies of wealth in archaeological contexts should not be based on bones alone.     

Body Part Representation,Fragmentation and Patterns of Ertebølle Deer Exploitation in Northwest Zealand,Denmark

The faunal assemblages from Asnæs Havnemark, Fårevejle and Trustrup, located in northwest Zealand, Denmark, yielded extensive yet substantially fragmented bone material dating predominantly to the late Mesolithic Ertebølle culture, ca. 5400–3950 cal BC . This paper presents a combined assessment of skeletal part representation of red deer (Cervus elaphus) and roe deer (Capreolus capreolus) from these three sites in conjunction with quantification of overall fragmentation data for the assemblages. These data indicate that the observed pattern of skeletal part abundance is most consistent with a pattern of density‐mediated skeletal attrition, and therefore the whole carcasses were initially deposited. Deer were not part of a logistic exploitation strategy at these sites. This indicates variability during the Ertebølle period not only in which resources were exploited, but in how these resources were processed. Copyright © 2013 John Wiley & Sons, Ltd.     

Zooarchaeological perspectives on the Chinese Early and Late Paleolithic from the Ma’anshan site (Guizhou,South China)

Ma’anshan is a Paleolithic cave site in the Guizhou province of southern China. The total area of the excavations is ca. 48 m², and the cultural deposit is divided into two layers representing the Chinese Late Paleolithic and the later part of the Early Paleolithic. The upper layer dates between 19,295  BP and 31,155 BP by AMS technique, and the lower layer is dated to around 53,000 BP by U-Series technique. Thousands of bone fragments were recovered from the two layers. Species abundance, bone surface modifications, skeletal element representation, and mortality patterns were studied in an investigation of assemblage formation history and patterns of prey selection and meat consumption by hominins. Interpretations of the faunal data are strengthened by reference to experimental studies and ethnoarchaeological data. The zooarchaeological data indicate that the later occupants of Ma’anshan Cave hunted mainly medium and small game animals, while the earlier occupants tended to prey upon larger animals. In the later period, hominins also made fuller use of the carcasses. The breadth of the meat diet increased with time, due to the inclusion of quick small animals (bamboo rats and birds) in the later period. The differences between the two faunal assemblages are consistent with the chronologic boundary currently drawn between the Chinese Early and Late Paleolithic ca. 30–27 ka BP, based on technological and other evidence.     

The hunted hunter: the capture of a lion (Panthera leo fossilis) at the Gran Dolina site,Sierra de Atapuerca,Spain

Many Pleistocene caves and rock shelters contain evidence of carnivore and human activities. For this reason, it is common to recover at these sites faunal remains left by both biological agents. In order to explain the role that carnivores play at the archaeological sites it is necessary to analyse several elements, such as the taxonomical and skeletal representation, the age profiles, the ratio of NISP to MNI, the anthropogenic processing marks on the carcasses (location and purpose of cutmarks and burning and bone breakage patterns), carnivore damage (digested bones, location and frequencies of toothmarks and bone breakage), length of the long bones, frequencies of coprolites and vertical distribution of the faunal remains, inter alia. From this, the documentation of carnivores in a faunal assemblage with a clear anthropogenic component can be understood from three main phenomena: (1) the carnivores as accumulators and the use of the site as a den; (2) carnivores as scavengers of hominid refuse and; (3) carnivores as hominids’ prey. Of these three phenomena, the last one is the least documented at the Middle Pleistocene sites. From this perspective, here we present the case of the anthropogenic use of a lion (Panthera leo fossilis) from level TD10-1 of Gran Dolina (MIS 9, Sierra de Atapuerca, Burgos, Spain). The lion bone remains show signs of direct interaction between this big cat and human groups that occupied Gran Dolina in these chronologies. From this perspective, the aim of this paper is to contribute to the knowledge of the role developed by large carnivores in the anthropogenic contexts and to provide data on human use of these predators at the European Middle Pleistocene sites.     

Urban small vertebrate taphonomy: a case study from Anglo–Scandinavian York

Sampling of deposits at the Queens Hotel site, York, produced a substantial number of small terrestrial vertebrate remains from the Anglo–Scandinavian features. By studying bone surface modification, fragmentation and skeletal completeness as taphonomic indicators, it was possible to demonstrate that the assemblage had resulted from two very different modes of accumulation and deposition. Refuse pits situated within the boundaries of the tenements had acted as accumulators of the fragmented and abraded small mammal and amphibian bones that existed as a sub‐surface death assemblage within the local environment. In contrast, the excellent preservation and skeletal completeness of numerous frogs recovered from the basal fill of a wooden well could be accounted for by their direct entry into the burial environment as a result of pit‐fall trapping. This paper also discusses the implications that the temporal and spatial variation in deposition demonstrated by the micro‐faunal remains has for the reconstruction of local ecological and environmental conditions within this site, and for other such sites. Copyright © 2001 John Wiley & Sons, Ltd.     

Taphonomic evidence for site formation processes at fort Christanna

Fort Christanna was an early eighteenth century Indian trading post on the western frontier of Virginia. This paper describes and interprets the faunal remains recovered during three seasons of excavation at the site. Taxonomic and body part representation are examined to gain insight into diet on the frontier, food provisioning at the fort, and the relationship between the fort's occupants and the neighbouring Indian settlement. These interpretations are limited by the poor preservation apparent in the faunal assemblage. The patterning of a variety of different surface modifications is explored to understand the taphonomic history of the assemblage. These characteristics of the assemblage seem to be related largely to the bone refuse disposal practices of the fort's occupants, and thus give important insight into a crucial aspect of the site formation processes.