Pits and pitfalls: taxonomic variability and patterning in tooth mark dimensions

Archaeologists use experimentally derived tooth mark frequencies, locations, and size data to infer (a) the extent of carnivore involvement in the formation and modification of faunal assemblages, (b) the size classes of predators marking those assemblages, and (c) whether hominins accessed fleshy or defleshed carcasses (Blumenschine and Pobiner, 2007; Dominguez-Rodrigo et al., 2007). These inferences are often debated in part because frequency counts can vary widely among observers and because the carnivore taxa for which tooth mark dimensional data are available are limited. This study contributes to the body of actualistic/experimental tooth mark data and presents a methodology for collecting these data. We offer a greatly simplified method that may encourage others to collect and quantify tooth mark dimensions. We present dimensional data from feeding experiments with 16 omnivore and carnivore species of known age and mass, ranging in size from skunks to tigers, significantly expanding the taxonomic and size range of carnivores for which we have tooth pit data. Our results demonstrate considerable, but not complete, overlap in tooth pit dimensions among size, class, and taxon. Tooth mark dimensions on epiphyses and metaphyses were relatively strongly correlated with body mass, whereas diaphyseal tooth marks exhibited the weakest correlation. Human consumption of animal tissue produced tooth marks comparable in size to medium felids and small canids, suggesting the possibility that some tooth marks on Early Stone Age (ESA) faunal assemblages could, as suggested by Oliver (1994), result from small carnivore and/or hominin consumption.     

Ascertaining an agent: using tooth pit data to determine the carnivore/s responsible for predation in cases of suspected big cat kills in an upland area of Britain

The identification of the involvement of a particular carnivore in the modification of bone assemblages concerns a number of fields of research including archaeological and palaeontological enquiry. Taphonomy provides a methodology by which bone assemblages can be analysed and interpreted and this is more often undertaken with archaeological or palaeontological assemblages. A taphonomic analysis is undertaken here in order to determine the perpetrator of predation attacks on domestic stock from a modern-day setting. Recently reported techniques using tooth marks preserved on bone surfaces made by known carnivores are successful at determining some class sizes of predators and are used here to determine the perpetrator(s). Although a class size of carnivore is readily identified by this methodology, a particular carnivore taxon is not. Tooth morphology and dental configuration are reported here as better criteria for identifying a particular taphonomic agent. Tooth pit dimensions are used here to identify the class size of carnivores involved, and tooth morphology and cusp spacing to suggest a medium sized felid and fox as taphonomic agents. The identification of the medium-sized felid may support observations and reports of alleged “big” cat kills in the area. The study has important implications for the interpretation of fossil sites where felids may have been involved in the modification of animal carcasses but are archaeologically invisible in terms of their fossil remains.     

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.     

Evidence of butchery and hominid activities at Torralba and Ambrona; an evaluation using microscopic techniques

Torralba and Ambrona have been interpreted as butchery sites for many years, a contention recently challenged; natural death or carnivore activities are invoked as an alternative explanation. Scanning electron microscopy (SEM) of bone surface replicas distinguishes among hominid-produced cutmarks, carnivore tooth scratches, and other types of bone damage. A sample of 102 replicas, comprising the most likely cutmarks on a combined sample of roughly 3000 fossils from Torralba and Ambrona, were scanned to determine the major agent of damage. Microscopically verified cutmarks are present, but rare, occurring in less than 1% of the bones in the combined sample. Carnivore tooth scratches are comparably rare. In contrast, evidence of sedimentary abrasion, which obliterates the diagnostic features of cutmarks, is present on nearly every bone from Torralba and Ambrona. It remains unresolved whether cutmarks were initially more common on these bones and were subsequently obliterated by abrasion, or whether the incidence of cutmarks was always low. These data demonstrate clearly that hominids and carnivores each damaged some bones at Torralba and Ambrona, but the frequency of each type of mark is too low to confirm strongly the interpretation of these sites as either butchery or carnivore remains.     

A new experimental study on percussion marks and notches and their bearing on the interpretation of hammerstone-broken faunal assemblages

Preliminary experiments on bone breakage have shown the potential utility of quantifying some of its diagnostic features (percussion marks, percussion notches) for taphonomic inferences about human and carnivore involvement in bone breakage in faunal assemblages. The present study increases the range of experiments undertaken to understand the identification of hammerstone percussion (dynamic loading) and its differences from carnivore bone breakage through tooth pressure (static loading). This study contributes to a better understanding of frequencies of percussion marks, uncovers and quantifies those percussion marks that lack key diagnostic features to be identifiable and that could be mistaken with carnivore tooth marks, quantifies notch types, shows different size ranges for notches on bones from small and large fauna than previously reported, quantifies the proportion of notches bearing percussion marks, and introduces new size data for percussion (impact) flakes and percussion marks. Furthermore, all these variables have been applied to a dual experimental set: one experiment using non-modified hammerstones and another based on the use of modified hammerstones. Results vary considerably according to hammerstone type. Some of these taphonomic variables increase the range of equifinality when identifying marks and notches created by different human and non-human agents. This calls for further caution when using isolated variables and features rather than a holistic approach to make taphonomic inferences.     

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.     

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.     

A cautionary note on the use of captive carnivores to model wild predator behavior: a comparison of bone modification patterns on long bones by captive and wild lions

A study with wild lions in Tarangire National Park (Tanzania) and with captive lions in Cabárceno Reserve (Spain) has yielded two different bone modification patterns, probably as a result of the differences in environmental contexts. Captive lions have modified bones more intensively, both in the form of total number of tooth-marked bones and number of tooth marks per tooth-marked bone, probably because of stereotypic behaviors. This emphasizes the importance of environmental contexts to understand carnivore behavior and their resulting bone modification patterns. It also shows that analogical models based on experiments carried out with captive carnivores may be biased and inadequate as proxies for wild carnivore bone modification behaviors.     

The Middle Pleistocene argali (Ovis ammon antiqua) assemblages at the Caune de l'Arago (Tautavel,Pyrénées‐Orientales,France): were prehistoric hunters or carnivores responsible for their accumulation?

The argali (Ovis ammon antiqua) assemblages from the Middle Pleistocene site of the Caune de l'Arago (Tautavel, southern France) were studied in terms of zooarchaeology and taphonomy. It is possible to discern palaeobiological information lost during fossilisation, as well as the palaeoethology of the bone collector, by the observation of taphonomic details preserved on the bone assemblages. The observations leave no doubt that both humans and carnivores were involved in the accumulation of argali carcasses in the cave. In some assemblages, the type of bones found in articulation and the gnawing marks observed are characteristic of carnivores. In other levels, the intense fracturing of the major limb bones in relation to their marrow content and mineral density, and butchering marks found on specimens in the earlier levels, are in favour of human accumulation, the modalities of which are discussed. The results suggest that the degree of carnivore activity seems to have been higher in levels M, N and O than in level F. Copyright © 2006 John Wiley & Sons, Ltd.     

Hiding to eat: the role of carnivores in the early Middle Pleistocene from the TD8 level of Gran Dolina (Sierra de Atapuerca,Burgos, Spain)

Damage generated by large and small carnivores is common in many Middle Pleistocene sites. However, identifying the predator that produces the faunal accumulations is often a difficult task. In order to recognize the main type of carnivore that acts on a faunal assemblage, a combination of several characteristics should be taken into account: taxonomic and skeletal element representation, age profiles, carnivore damage (location, frequencies and dimensions of tooth-marks, bone breakage and digested bones), degree of fragmentation and frequencies of coprolites. But, adding environmental characteristics and the ethology of non-human predators/scavengers is also important. All these aspects are applied to the faunal assemblage from the TD8 level of the Gran Dolina site (Sierra de Atapuerca, Burgos, Spain). Paleomagnetic data combined with ESR and U-series place the TD8 level at the beginning of the Middle Pleistocene, specifically circa 700 kyr ago. The TD8 level contains a large faunal accumulation primarily composed of ungulate skeletal elements, and to a lesser extent carnivore remains. This assemblage is characterized by an overrepresentation of fallow deer (Dama vallonetensis), a skeletal profile biased towards cranial remains and limb bones, diversity of ages at death, a high proportion of carnivore damage and tooth-marks of large size, and an absence of human activity. According to these data, the accumulation seems to have been produced primarily by large carnivores, possibly hyenas. This observation does not rule out the possible occasional activity by other carnivores. Nevertheless, the characteristics of the TD8 assemblage do not correlate entirely with those traditionally used to define carnivore dens. In TD8, there are (1) no immature carnivore remains (remains of just one young Mosbach wolf); (2) scarce traces related to the end stages of consumption and some anatomical connexions; (3) few coprolites; (4) high proportion of adult ungulates and; (5) high quantities of whole bones and epiphyses. From this perspective, the TD8 faunal assemblage seems to correspond to a succession of carnivore occupations that allows the development of a suite of features to identify the activities of several species of predators that may have used the cave in different ways and durations. This study aims to emphasize the importance of these analyzes in order to know the behaviour of different non-human predators/scavengers in the European Middle Pleistocene sites.     

Taphonomy of ungulate ribs and the consumption of meat and bone by 1.2-million-year-old hominins at Olduvai Gorge,Tanzania

The phenomenon of equifinality complicates behavioral interpretations of faunal assemblages from contexts in which Pleistocene hominins are suspected bone accumulators. Stone tool butchery marks on ungulate fossils are diagnostic of hominin activities, but debate continues over the higher-order implications of butchered bones for the foraging capabilities of hominins. Additionally, tooth marks imparted on bones by hominins overlap in morphology and dimensions with those created by some non-hominin carnivores, further confounding our view of early hominins as meat-eating hunters, scavengers or both. We report on the manual/oral peeling of cortical layers of ungulate ribs as taphonomically diagnostic of hominoid/hominin meat- and bone-eating behavior that indicates access to large herbivore carcasses by hominins at the site of BK, Olduvai. Supporting these inferences, we show that certain types of rib peeling damage are very rare or completely unknown in faunas created by modern carnivores and African porcupines, but common in faunas modified by the butchery and/or consumption activities of modern humans and chimpanzees, during which these hominoids often grasp ribs with their hands, and then used their teeth to peel strips of cortex from raggedly chewed ends of the ribs. Carnivores consume ungulate ribcage tissues soon after kills, so diagnostic traces of hominin butchery/consumption on ribs (i.e., peeling and butchery marks) indicate early access to ungulate carcasses by BK hominins. Tooth marks associated with the peeling and butchery marks are probably hominin-derived, and may indicate that it was not uncommon for our ancestors to use their teeth to strip meat from and to consume portions of ribs. Recognition of rib peeling as a diagnostic signature of hominoid/hominin behavior may also aid the search for pre-archaeological traces of hominin meat-eating.     

Fluvial transport of bovid long bones fragmented by the feeding activities of hominins and carnivores

This study explores the hydraulic transportability of bovid long bone fragments created through hominin and carnivore carcass consumption in order to determine the effect of fluvial transport on the incidences of hominin- and carnivore-induced bone surface modifications. Transportability was determined using an oval race track flume and 311 long bone fragments from modern control collections of hominin- and carnivore-modified bone. Results show that the fluvial transport of long bone fragments is predicted by animal size class and bone specimen size, as measured by maximum cortical thickness, maximum length, and maximum width. All of these variables can be measured on fossil specimens. Long bone portion (midshaft, near-epiphysis, and epiphysis) does not affect transport and hydraulic transport does not substantially modify the incidences of tooth, percussion, and cut marking in transported or lag assemblages in low energy fluvial environments. Implications of this study are: 1) animal size classes, and cortical thickness, length, and width of long bone fragments can be used to identify fluvial winnowing in fossil assemblages; 2) analyses concerning the relative timing of hominin and carnivore carcass consumption based on the proportions of long bone fragments bearing tooth, percussion, or cut marks can be meaningfully applied to fossil assemblages deposited in low-energy fluvial environments.     

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.     

Hunted or Scavenged Neanderthals? Taphonomic Approach to Hominin Fossils with Carnivore Damage

Carnivore damage on Neanderthal fossils is a much more common taphonomic modification than previously thought. Its presence could have different explanations, including predatory attacks or scavenging scenarios, which are both situations with important implications concerning Neanderthal behaviour. In the present paper, we analyse several Neanderthal hominin fossils from a taphonomic and forensic perspective in order to infer the nature of the modifications observed on the bone surfaces. Fossils displaying carnivore modifications from Spain, Germany, Belgium and Greece are evaluated from a taphonomic perspective for the first time in a significant sample of hominin specimens. Our results show that the materials analysed have been modified by small to large carnivores and that both attacks and strictly carnivore scavenging events can be inferred. This study also points out the importance of developing taphonomic approaches to the analysis of hominin bone surfaces, which can contribute significantly to knowledge of several aspects of Neanderthal behaviour. Copyright © 2017 John Wiley & Sons, Ltd.     

Isotopic analysis of the ecology of herbivores and carnivores from the Middle Pleistocene deposits of the Sierra De Atapuerca,northern Spain

Carbon and oxygen isotope values reveal resource partitioning among the large mammal fauna from three contemporaneous Middle Pleistocene hominid-bearing localities within the Sierra de Atapuerca (northern Spain). Carbon isotope values sampled from the tooth enamel of fauna present during Atapuerca Faunal Unit 6 show that a C₃-dominated ecosystem surrounded the area where fossils were preserved during this time. For the herbivores, Fallow deer isotope values are significantly different from Red deer and horses and show that this species did not forage in open environments at this locality. Red deer and horses show similar feeding strategies with less negative carbon values implying use of more open environments for these taxa. For the carnivores, carbon isotope values for Ursus deningeri are significantly different from either lions (Panthera leo) or foxes (Vulpes vulpes) and support the contention that this species is herbivorous. Special metabolic mechanisms involved in hibernation in U. deningeri might also have influenced its isotope values. The carbon isotope values of remaining carnivores were similar and suggest that each was typically a generalist carnivore, eating a wide variety of prey items. While the isotopic results generally correspond to ecology indicated by previous techniques, this study shows that isotope analyses can provide further insights that alternate techniques do not provide. Isotope analyses can help elucidate the ancient ecology of taxa present in the Sierra de Atapuerca during the Middle Pleistocene allowing for an accurate portrayal of the setting in which humans lived.     

South African micromammals and predators: some comparative results

This paper presents the results of a taphonomic, comparative study on some South African species of small carnivores, and the effects they have on micromammalian prey skeletons in terms of breakage patterns, body part representation and etching from digestion. The results from a study of modern serval scats suggest that the taphonomic pattern found on prey remains from this felid may share more in common with some of the owls, diurnal birds of prey and other small carnivores than previously realized.     

Large carnivores as taphonomic agents of space modification: an experimental approach with archaeological implications

At most Pleistocene archaeological sites it is difficult to observe structured complex spatial behaviour. This common phenomenon could be a taphonomic issue. Problems in the preservation of the original spatial intra-site distribution could be related to hominid–carnivore alternation in the use of space. In the present paper we analyse the results of our experimentation with large extant carnivores (bears, wolves, hyenas and lions) and propose these animals acted as hearth and hearth-related assemblage modifiers. In this sense, the role of carnivores in the modification of these elements can cause problems in the interpretation and visibility of modern and complex behaviour in the conception of space in the archaeological record.     

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.     

Early hominid hunting,butchering, and carcass-processing behaviors: Approaches to the fossil record

A major aim of paleoanthropology is to learn what ancient behaviors were related to the acquisition, processing, and consumption of meat and when these behaviors arose. For this reason, studies focusing on purported early hominid hunting and butchery sites are important if rigorous criteria for recognizing such sites are used. Different criteria currently used as evidence of hominid involvement with ancient bones are reviewed and it is concluded that the presence of cutmarks, verified by scanning electron microscope (SEM) inspection, is the most reliable. Successful application of this criterion depends upon a thorough knowledge of the normal variations in microscopic morphology of different types of marks that are found on bones. Therefore, variations in microscopic and gross morphology within and among a large sample of known stone tool cutmarks, carnivore tooth scratches, and rodent gnawing marks are documented. The effects of sedimentary abrasion, as caused by fluvial transport of bones, are also presented. Guidelines are presented for using microscopic criteria to identify unknown marks on fossils and possible applications of this approach are discussed. Further, it is suggested that evidence of hominid carcass-processing activities can be placed into one of three ranked categories of certainty according to the type of data used. Explicitly stating the category and type of evidence used to deduce hominid activities, and by extension to define site types (i.e., butchery, kill, base camp), may improve the clarity of hypotheses about and interpretations of early hominid behaviors.