An exclusively hyena-collected bone assemblage in the Late Pleistocene of Sicily: taphonomy and stratigraphic context of the large mammal remains from San Teodoro Cave (North-Eastern Sicily,Italy)

A detailed taphonomic analysis of the large mammal assemblage from 1998 to 2006 excavations at San Teodoro Cave is presented, taking into account the stratigraphic context of the deposits. Three not strictly contemporary fossiliferous levels having different lithological features have been detected, here named B-I, B-II, and B-III. Fossil remains are prevalently accumulated in B-I and B-II. The three levels are characterized by evidence of Crocuta crocuta spelaea occupation, represented by their skeletal remains, coprolites, and distinctive damages on the bones, similar to fossil and modern spotted hyena dens from Europe and Africa. A differential distribution of coprolites and small digested bones, probably due to different humidity conditions, has been recognized in B-I and B-II, and can be related to different topographic locations within the cave or to different climate conditions during the sedimentation phases. The very low density of fossil remains in B-III, which is the oldest level, could indicate an area that was less inhabited by hyenas, probably due to geomorphological conditions. Taphonomic comparison of the three fossiliferous levels of the San Teodoro Cave deposits points to a long-term, perhaps cyclic, occupation of the cave by hyenas and confirms the cave as one of the most important Pleistocene hyena dens in Europe.     

Late Pleistocene Crocuta crocuta spelaea (Goldfuss, 1823) clans as prezewalski horse hunters and woolly rhinoceros scavengers at the open air commuting den and contemporary Neanderthal camp site Westeregeln (central Germany)

Late Pleistocene Ice Age Crocuta crocuta spelaea ( Goldfuss, 1823) hyenas from the open-air gypsum karst site Westeregeln (Saxony-Anhalt, central Germany) is dated into the early to middle Late Pleistocene. Hyena clans apparently used the karst for food storage and as “commuting den”, where typical high amounts (15% of the NISP) of hyena remains appear, also faecal pellets in concentrations for den marking purposes. Additionally small carnivores Meles, Vulpes and Mustela appear to have used some cavities as dens. Several hundreds of lowland “mammoth steppe fauna” bones (NISP = 572) must have been accumulated primarily by hyenas, and not by Neanderthals at the contemporary hyena/human camp site. Abundant caballoid horse remains of “E. germanicus Nehring, (1884)” are revised by the holotype and original material to the small E. c. przewalskii horse. Woolly rhinoceros Coelodonta antiquitatis remains are also abundant, and were left in several cases with typical hyena scavenging damages. Rangifer tarandus (11%) is mainly represented by numerous fragments of shed female antlers that were apparently gathered by humans, and antler bases from male animals that were collected and chewed in few cases (only large male antlers) by hyenas. The large quantities of small reindeer antlers must have been the result of collection by humans; their stratigraphic context is unclear but such large quantities most probably resulted from schamanic activities. The hyena site overlaps with a Middle Palaeolithic Neanderthal camp, as well as possibly with a later human Magdalénian site.     

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.     

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.     

Late Pleistocene Eemian hyena and steppe lion feeding strategies on their largest prey—Palaeoloxodon antiquus Falconer and Cautley 1845 at the straight-tusked elephant graveyard and Neanderthal site Neumark-Nord Lake 1, Central Germany

The scavenging activities of hyenas and lions at the Neumark-Nord Lake 1 (Central Germany) site on straight tusk elephant carcasses is commonly documented by slight bone damages and initial skeleton partitioning. Canine bite marks are found at several joints of elephant long bones, as well as on the vertebrae and skulls. Deep bite and scratch marks from 5 to 7 mm in width made by large carnivore canine teeth (Panthera leo spelaea, Crocuta crocuta spelaea) on the ventral vertebral columns and pelvises of two Palaeoloxodon antiquus straight-tusked elephant skeletons suggest that the intestines and inner organs were consumed by both large lion/hyena predators as is commonly observed when modern African lions and spotted hyenas feed on contemporary elephant carcasses. A scavenging strategy can be demonstrated using 24 partly preserved Palaeoloxodon skeletons, where Ice Age spotted hyenas appear to be the main scavenger (95 %) and sole skeleton decompositor such as bone destructor. While Crocuta may feed anywhere on an elephant carcass, they are known to specialize to eat the complete feet, whereas scavenging from the anus into the body cavity (intestine/internal organ feeding) is more typical of lions. Shallow water at Neumark-Nord Lake 1 prevented hyenas from completely scavenging elephants; thus, bone damage at this site was constrained to initial stages of destruction and final carcass partitioning by high water stands and floods.     

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.     

Sources of variation in carnivore tooth-mark frequencies in a modern spotted hyena (Crocuta crocuta) den assemblage,Amboseli Park,Kenya

Distributions and frequencies of carnivore tooth-marks on large mammal long-bone fragments are commonly used to infer the timing of hominin and carnivore access to prey resources in archaeofaunal assemblages. The strength of these inferences, however, is limited by a broad and currently inexplicable range of tooth-mark frequencies across experimental and archaeological assemblages. Controlling for this variation first requires that the sources be identified. Several sources of variation are examined here in an analysis of tooth-marked bone recovered from a modern spotted hyena (Crocuta crocuta) den assemblage in Amboseli Park, Kenya. Results indicate that tooth-mark frequencies: (1) depend on fragment size, (2) vary across mammals of different size classes, (3) are highly variable across equivalent portions of different long-bone elements, and (4) on certain long-bone portions are correlated with bone density and can be depressed in archaeological assemblages subjected to density-mediated attrition. Stronger inferences based on tooth-mark frequencies will require that such variation be taken into consideration, and methods for doing so are suggested.     

Evolving ecosystems: ecological data from an Iron Age small mammal accumulation at Alorda Park (Catalonia,Spain)

Small mammals are considered to be good indicators of ancient environments. One particular layer in a fortification tower of a Mediterranean Iberian village, Alorda Park (4th century BC), provided a large amount of well-preserved small mammal bones (about 19,200 remains) probably accumulated by barn owls (Tyto alba). Such an accumulation is rarely available from the Iron Age. It provides not only palaeoenvironmental data, but also further information about other studies such as morphometrics, ancient DNA and zoogeography. This article focuses on the ecological data, and attempts to develop an accurate taphonomic study to assess the reliability and statistical significance of this archaeological sample.     

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 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.     

The archaeology and paleoenvironment of an Upper Pleistocene hyena den: An integrated approach

Data of high resolution for reconstructions of archaeological site formation processes can be obtained only by the use of precise excavation methods and comprehensive recovery techniques using fine-mesh water screening, followed by meticulous sorting and quantitative studies of all the small organic components from the screens. These methods are generally used at human occupation sites, but are not often employed at paleontological or mixed human-carnivore sites. At the site of Bois Roche, an Upper Pleistocene hyena den, we have used these meticulous recovery techniques in combination with absolute dating, geoarchaeological observations, taphonomic analyses, studies of microvertebrates and pollen analysis. Correlation of the site environmental data from pollen and microvertebrate assemblages with the vegetation record of a deep-sea core taken in the Bay of Biscay have allowed us to constrain the ages provided by ESR dating of bovid teeth and to reconstruct the physical environment of the site with a degree of confidence rarely achieved by the use of fossil remains from archaeological sites alone.     

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.     

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.     

Carnivore competition, bone destruction, and bone density

In carnivore-modified archaeofaunal assemblages it is important to evaluate the degree to which carnivores have overprinted hominin behavioral signals. To examine the signals of increased competition for discarded bone, we present controlled experimental data on 33 simulated archaeological sites subjected to secondary consumption by spotted hyenas. We examine the relationship between competition, as measured by controlled numbers of hyenas and limb bones, and resultant levels of destruction and correlations between long-bone portion survivorship and bone density. Our results indicate that levels of destruction are equivalent regardless of the numbers of hyenas or long-bones included in the experimental assemblages. Correlations between long-bone epiphyseal and near-epiphyseal portions and bone density, however, do provide an indication of the level of competition. Results from the experimental study are used to highlight divergent levels of carnivore competition for hominin-discarded bone at the Plio-Pleistocene localities FLKN-Zinjanthropus and FLKN levels 1–2 from Olduvai Gorge, Tanzania.     

Applying dynamics and comparing life tables for Pleistocene Equidae in anthropic (Bau de l'Aubesier,Combe-Grenal) and carnivore (Fouvent) contexts with modern feral horse populations (Akagera,Pryor Mountain)

In this study, we propose a method based on life tables currently used in population dynamics and ecology to compare the mortality curves of Equidae from anthropic and carnivore fossil assemblages with those from modern populations. The combined analysis of frequency (f_x) with various ecological parameters such as mortality rate (q_x), survival rate (l_x), fecundity rate (m_x), and killing factor (k_x), established for each age class, allows us to characterize the stability of fossil populations. These ecological parameters, developed in demography, correspond to many curves and constitute a complementary approach to interpret the two main mortality models, attritional and catastrophic, often evoked in zooarchaeology. The age structures of fossil horses from the anthropic sites of Bau de l'Aubesier (Vaucluse, France) and Combe-Grenal (Dordogne, France) and the hyena den of Fouvent (Haute-Saône, France) are compared to typical catastrophic modern populations of Equidae from the National Park of Akagera (Rwanda). We also used data collected by the Bureau of Land Management for present day feral horses from Pryor Mountain (Montana, USA) which constitutes a reliable referential of a stable living population. The characterization of age structures in those different contexts is discussed as well as the impact of Neanderthals and hyenas on fossil assemblages.     

Taphonomic analysis of rodent bone accumulations produced by Geoffroy's cat (Leopardus geoffroyi, Carnivora,Felidae) in Central Argentina

Rodent prey remains recovered from Geoffroy's cat (Leopardus geoffroyi) scats were analyzed in order to identify taphonomic features produced by this predator. The analysis includes a year-long sampling discriminated by the seasons. Modifications produced by digestion are heavy. Taphonomic variables in samples discriminated by the seasons did not show major differences with respect to the total sample; thus, scats collected in any season clearly show the modifications on the bones of prey rodents made by Geoffroy's cats. The results presented here and its interpretation could be extrapolated to an analysis of zooarchaeological or paleontological assemblages.     

Micromammal taphonomy of el-Wad Terrace,Mount Carmel,Israel: distinguishing cultural from natural depositional agents in the Late Natufian

A taphonomic analysis of micromammal remains from the Late Natufian deposits of el-Wad Terrace, Mount Carmel, Israel, was conducted in order to test the long-standing premise of owl deposition as the primary accumulating agent. The inferred taphonomic sequence was modeled within an actualistic (recent) comparative framework incorporating a locally derived barn owl pellet collection and an off-site control assemblage of micromammal remains from the cliff overhanging the terrace. The sequence was reconstructed based on multiple types of recorded taphonomic data, comprising skeletal modifications (breakage, digestion, weathering, gnawing, and charring) and age structure. Evidence for post-depositional processes including fluvial transport, trampling, and weathering was isolated and consequently the typical owl imprints were traced through the two actualistic and the archaeological assemblages. Based on the extent of breakage and patterning in skeletal element frequencies it was also possible to scale the preservation potential of primary data in el-Wad Terrace as a discrete site type, intermediate between true cave and open-air depositional environments. Verifying the role of owls as principal agents of accumulation of the el-Wad Terrace micromammal remains enabled the detection of two minor superimposed cultural patterns: consumption of mole rats (Spalax sp.) by the Natufians and commensalism of mice (Mus spp.).     

When wildcats feed on rabbits: an experimental study to understand the taphonomic signature of European wildcats (<Emphasis Type="Italic">Felis silvestris silvestris</Emphasis>)

Studies of the feeding ecology of the European wildcat (Felis silvestris silvestris) demonstrate that leporids, mostly European rabbit (Oryctolagus cuniculus), dominate their diet in regions where they are present. The remains of wildcats have been found at Pleistocene and Holocene archaeological sites, raising the possibility that they actively accumulated leporid bones in caves and shelters shared with other terrestrial carnivores, raptors and humans. We present the first taphonomic study of rabbit remains consumed by this terrestrial carnivore, with the ultimate aim of understanding their role in bone accumulations at archaeological sites. An experimental study was carried out with a wildcat female, who was fed with nine complete rabbit carcasses. Non-ingested remains and scats were recovered for the analysis of anatomical representation, breakage and bone surface modification. This revealed that non-ingested remains and scats of the European wildcat can be discriminated from most other agents of accumulation. The referential framework provided will permit the discrimination of hominids and wildcats as agents of fossil accumulations of rabbits.     

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.     

The recovery and interpretation of microvertebrate bone assemblages from archaeological contexts

The analysis of microvertebrate remains recovered from buried contexts is frequently problematic for archaeologists. Diminutive size is a major obstacle to recovering, identifying, and interpreting microfaunal bone materials successfully. Their taphonomic history is often obscure, thus the significance of microvertebrate accumulations for settlement, subsistence, and ecological interpretation is difficult to evaluate. Recognizable and reliable archaeological signatures are sought for assessing the effect of taphonomic history on microvertebrate assemblages. Relevant signatures of bone damage, skeletal survivorship, taxonomic composition, and context are examined in assemblages produced and modified through accidental non-predator-related death, predator accumulation, and postmortem alteration.