Reconsideration of criteria for differentiating faunal assemblages accumulated by hyenas and hominids

Zooarchaeologists have established several criteria for differentiating hominid‐ and hyena‐derived faunal assemblages. In some cases, however, the patterns of skeletal part representation and bone surface modification on which these criteria are ultimately based have been observed in fossil bone assemblages of unknown origin, rather than in modern assemblages of known origin. When the proposed criteria are evaluated within an actualistic framework, only three are able to differentiate between hominid‐ and hyena‐created faunal assemblages. I suggest that only these three criteria—proportions of carnivores to ungulates in the assemblage, the preserved condition of long bone specimens (either as whole cylinders or as splintered shaft fragments), and the types of bone surface modifications—should be retained as important factors in a diagnosis of the ancient bone‐collector. The remaining four criteria—the relative proportion of horn pieces in the assemblage, the relative representation of podial bones, the relative representation of small and large bovid skeletal parts, and bovid mortality profiles—are not relevant or applicable to the problem of differentiating hominid‐ from hyena‐derived faunal assemblages. Copyright © 2002 John Wiley & Sons, Ltd.     

Avian taphonomy: observations at two Magellanic penguin (Spheniscus magellanicus) breeding colonies and their implications for the fossil record

Actualistic studies on vertebrate taphonomy have centred on mammals, and to a much lesser extent on flying birds. Members of the Spheniscidae are anatomically specialized making it problematic to make direct inferences from previous taphonomic work. Modern bone assemblages from two Spheniscus magellanicus breeding colonies on the southern Atlantic coast of continental Patagonia, Argentina, were surveyed on foot along linear transects, and features were determined that are useful in interpreting the fossil record. Present day bone accumulations have specific properties resulting from penguin anatomy and osteology as well as from the composition of the vertebrate community and regional environmental conditions. Most salient are the high spatial density of bones, the dominance of Spheniscidae remains over those of other vertebrates, the high percentage of disarticulated bones, the predominance of limb bones, the low frequencies of carnivore bone modifications, and the abundance of fractured bones. Many of these features observed in the modern assemblages can be identified in fossil avian bone deposits.     

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.     

A taphonomic study of bone modification and of tooth‐mark patterns on long limb bone portions by suids

This work presents new taphonomic data on bone modification by suids, including domestic pig, wild and hybrid boars. The intense modification undergone by bones from animals smaller than 100 kg is shown, together with a more moderate modification on bones from larger animals. Both the ravaging pattern (with preferential deletion of cancellous tissue) and the tooth‐marking frequencies are similar to those documented among hyenas and dogs when having primary access to complete bones. A dual‐patterned experimental model consisting of the interaction of humans and suids was also considered. Here it is shown how suid modification of hammerstone‐broken bone assemblages is different from that documented among canids and hyaenids, as experimentally replicated. These results increase the number of non‐anthropogenic bone‐modifying agents and posit new issues on equifinality processes. Copyright © 2008 John Wiley & Sons, Ltd.     

A taphonomic study of the African wild dog (Lycaon pictus)

Carnivore taphonomy has been traditionally used for the interpretation of archaeological sites in order to discriminate human-generated or modified from non-anthropic bone assemblages. In most of this actualistic research, the focus has mainly been placed on hyenas and felids, neglecting other carnivores. This paper analyzes the taphonomic impact of the African wild dog (Lycaon pictus) on equid bones and compares it with the bone modification patterns produced by other canids, such as wolves (Canis lupus) in order to compare medium-/large-sized canid variability on bone modification patterns and elaborate a referential framework which could be feasibly applied to the zooarchaeological record to detect canid intervention on archaeological assemblages in the past.     

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.     

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.     

A new element of trampling: an experimental application on the Level XII faunal record of Bolomor Cave (Valencia,Spain)

In Level XII of Bolomor Cave (Valencia, Spain) modifications have been observed on some of the bone fragments that make up the faunal record. These alterations consist of small notches located on the oblique fracture angles of some bones. The frequency at which these modifications appear in the assemblage has obliged to plan several experimental series to know the process that causes them. To be more exact, this study aims to verify whether these small chips are the result of trampling. For this purpose, two experimental series have been developed. The first of which has the objective of differentiating between the variables related to the original state of the bone (green, semi-dry and dry) and the second is aimed at reproducing the modifications observed in the archaeological set, taking into account the data obtained in the previous series. Following these experiments, the appearance of small notches is observed on the oblique fracture angles of some bones in semi-dry state. These resulting chips are similar to those documented in Bolomor Cave, Level XII. Therefore, it is deduced that trampling is the principle cause of these modifications in the bone record. This fact has important archaeological implications, as the spatial distribution of this modification can provide information about the intrasite movements of the human groups.     

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.     

Breaking Bones to Obtain Marrow: A Comparative Study between Percussion by Batting Bone on an Anvil and Hammerstone Percussion

The fat‐ and nutrient‐rich marrow of animal bones can be extracted using different techniques. Passive hammerstone percussion has been the primary focus of experimental bone breaking and the main analogy to understand archaeological bone breakage. Here, the term ‘passive’ is applied because the bone to be broken passively receives the impact from a hammerstone. In addition to this technique, there is another bone‐breaking method that also requires direct percussion, but in an active way. This method is percussion by ‘batting’, in which the bone is actively hit against an anvil until the bone breaks. This technique has rarely been considered at an experimental level and, therefore, has been omitted in the majority of the archaeological interpretations of faunal assemblages with pre‐use of fire technologies. In this study, we attempt to analytically characterize this type of bone‐breaking technique through a systematic comparison with hammerstone percussion. The applied statistical tests will allow us to distinguish some diagnostic modifications, such as the outlines of the fracture planes and the type of notches or their location with respect to the longitudinal axis of the bone. These features and their proportions allow the consideration of the use of this technique in Pleistocene anthropogenic faunal assemblages.     

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.     

Food for thought: using game cameras to better understand the movement of bones by scavenging in archaeological faunal assemblages

This paper questions our understanding of the movement of bones by animal scavengers in the archaeological record. Since assumptions regarding the effects of animal scavenging shape final interpretations of skeletal element frequencies in archaeological faunal assemblages, they are important for our understanding and reconstruction of ancient human behaviour. The results of a 4-year actualistic kangaroo scavenging study from Australia are used to question our understanding of the movement of the bone by contrasting visual data captured by motion-activated digital game cameras with traditional taphonomic studies using skeletal element frequencies. Game cameras are commonly used by ecologists to capture the behaviour of living species but have not yet been used in experimental archaeology where visually documenting animal scavenging behaviour can be used to understand the movement of carcasses and individual bones. Results suggest that traditional zooarchaeological analyses may not be accurate indicators of hunted versus scavenged prey in archaeological faunal assemblages. Moreover, they most certainly fail to document the entire suite of animals scavenging a carcass. These implications are discussed with particular reference to the ability to definitively ascertain the role of humans in the megafaunal extinction debate in Australia.     

Bone Modifications Resulting from Captive Chimpanzee Mastication: Implications for the Interpretation of Pliocene Archaeological Faunas

In order to obtain chimpanzee (Pan troglodytes) mastication damage on bones, cleaned, disarticulated ribs and long bones of bovids and cervids were coated with food substances found palatable by captive chimpanzees. The bones were then presented to four groups of mixed sex and mixed age chimpanzees from the Tulsa Zoo (Oklahoma) and the Southwest Foundation for Biomedical Research (San Antonio, Texas). In the process of feeding on the coated bones, the chimpanzees inflicted discernible mastication damage on 73·61% of the total recovered bone specimens. Analysis of these chimpanzee-induced bone modifications reveals that non-technological hominoids of the same approximate size and with comparable dentition and bite force to the gracile australopithecines and earliestHomoare very capable of inflicting the same range and degree of damage to bones as are feeding carnivores. This finding implies that zooarchaeologists must take a contextual (configurational) approach when analysing mastication damage on arch-aeological bones, rather than automatically attributing all such damage to carnivores.     

The Bearded Vulture (Gypaetus barbatus ) as an Accumulator of Archaeological Bones. Late Glacial Assemblages and Present-day Reference Data in Corsica (Western Mediterranean)

Assemblages of 751 bone remains of an endemic Corsican deer (Megaloceros cazioti) from Late Glacial layers at Luri- Grı̀tulu cave (Northern Corsica) show an uncommon taphonomic pattern, mainly characterized by high dominance of third phalanxes and high rate of digestion marks. The authors demonstrate that this pattern cannot result from accumulation by human beings or by large mammal carnivores. The bearded vulture, which is able to accumulate large mammal bones in its nest and the presence of which is attested in the faunal remains at Grı́tulu, is the principal putative accumulator. In order to test this hypothesis, bone contents from 11 nests of modern Corsican bearded vultures (871 bone remains, i.e., more than 105 individual carcasses) are analysed from a taphonomic point of view. They are mainly characterized by overwhelming quantities of third phalanxes from middle-size ruminants, abundance and location of digestion marks around proximal articulations, and few bone flakes. A comparison of the Grı́tulu cave assemblages with these new present-day reference data leads to the conclusion that they have actually been accumulated by Late Glacial bearded vultures. Small discrepancies can, however, be explained by differences in food availability between the Late Glacial and modern times.     

A study of dimensional differences of tooth marks (pits and scores) on bones modified by small and large carnivores

The use of tooth mark sizes to infer carnivore types when analyzing the modification of faunal assemblages has been criticized on the base of intense overlap in tooth mark size among differently sized carnivores. The present study analyzes this overlap and presents some critical explanations for it. This work is based on the largest collection of tooth pit dimensional data collected to date for some of the most relevant carnivore types. The study empirically shows that small and large carnivores can be clearly differentiated when using tooth pit size, with a higher discrimination when using tooth marks on dense shafts than on cancellous ends. It is argued that most previous studies of tooth mark sizes have reproduced a higher overlap probably because sample sizes were small, and experiments were carried out using small carcasses (which require a smaller bite force) or for a combination of factors.     

The Red Fox (Vulpes vulpes) as an Accumulator of Bones in Cave‐like Environments

Identifying the behavioural patterns of bone collecting animals is a crucial aspect of taphonomic studies. Although many studies have established criteria for identifying animal‐collected or animal‐modified bones, very few papers describe the distinguishing features of fox‐made bone assemblages. The bone assemblage collected in an inactive underground stone mine in Potok‐Senderki (Poland) is diagnostic of a red fox (Vulpes vulpes) den. This site provides an ideal opportunity to develop an understanding of the bone collecting behaviour of red foxes in cave‐like environments. This study showed that bones collected by red foxes are concentrated in clusters. The bones represent a broad spectrum of local fox prey species, with most bones showing the marks of gnawing. Each cluster may contain from <10 to >100 bones. Furthermore, the long axes of the bones in clusters frequently show specific orientation. The analysis of bones at this site might make an important contribution towards the establishment of baseline criteria for the identification and evaluation of fox‐accumulated bone assemblages. Copyright © 2012 John Wiley & Sons, Ltd.     

Was it the deer or the fox?

Herbivores, as taphonomic agents, can modify and consume bones and antlers for no nutritive purpose. This unusual behavior is due to a nutritional dysfunction (osteophagia) that allows them to supplement a lack of minerals in their diet through ingestion of minerals contained in bones. When chewing, herbivores change skeletal element morphology and produce a characteristic forked shape. At an incipient stage of modification, herbivore chewing may mimic that of carnivores. In this paper, we provide diagnostic criteria to distinguish bone modification made by herbivores from that produced by other taphonomic agents, mainly carnivores.