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.     

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.     

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.     

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.     

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.     

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.     

An experimental study of large mammal bone modification by crocodiles and its bearing on the interpretation of crocodile predation at FLK Zinj and FLK NN3

The taphonomic signature of crocodiles as agents of bone modification has been previously identified by specific tooth mark types (e.g., bisected pits) and by a conspicuous presence of these marks: more than 75% of bones modified by crocodiles bear at least one of these distinctive marks. Therefore, crocodile tooth-marking would be notably prevalent in bone assemblages resulting from crocodile predation and active scavenging. The present study contributes to refine this diagnosis by showing greater variability of these types of marks, a different degree of tooth-marking intensity, and a somewhat different distribution of tooth-marked elements according to skeletal parts from previous experiments with crocodiles. Some of these differences are due to different experimental variables and conditions and this highlights the need to understand behavioral variability in crocodile ecological settings. This variability in crocodile tooth-marking probably results from several as-yet-inadequately measured behavioral and ecological factors, such as intensity of feeding competition and differences between male and female crocodile feeding behaviors, among others. Furthermore, this study also contributes to a better definition of the microscopic criteria that can be used to distinguish crocodile-inflicted marks from other types of bone surface modifications. In light of these and previous experimental frameworks, we reevaluate the application of these analogs to modifications documented in hominin fossils from Olduvai Gorge (OH8 and OH35) and the resulting inferences about the hazard posed by crocodiles on the paleolandscape where FLK North North and FLK Zinj (Bed I) were formed. The taphonomic analysis also shows that Olduvai OH8 and OH35 were probably not preyed upon by crocodiles. It is concluded that no tangible evidence can be used to support the interpretation that OH35 was modified by crocodiles and that the overall presence of crocodiles in FLK North North and FLK Zinj was rather marginal, based on the virtual absence of crocodile-modified bones in both archaeofaunal assemblages.     

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.     

Dietary ecology of extant guanaco (Lama guanicoe) from Southern Patagonia: seasonal leaf browsing and its archaeological implications

The analysis of dietary traits of ungulates through tooth microwear and mesowear has been applied to archaeological sites to investigate seasonal changes in settlements by hunter–gatherers. In this paper we propose to test the hypothesis that tooth microwear (combined to mesowear) is able to indicate seasonality in the diet of extant ungulates in arid habitats (semi-deserts or steppe). The material analyzed comes from six faunal monospecific assemblages of guanaco (Lama guanicoe) resulting from a mass mortality event in winter 2000 near the Cardiel Lake in Southern Patagonia (Province of Santa Cruz, Argentina). Mesowear results indicate that the guanacos from the Cardiel Lake area are mixed feeders, and thus, have a diet that shifts seasonally. Moreover, microwear analysis supports the hypothesis that tooth microwear is able to indicate seasonality in the diet of extant guanaco in arid habitats. The pattern is clear for the winter sample and needs to be confirmed for a summer sample. Consequently, tooth microwear is proposed as a new potential proxy for detecting seasonal occupation in archaeological sites in Patagonia and other arid environments.     

Changing patterns of carnivore modification in a landscape bone assemblage, Amboseli Park, Kenya

This study compares the landscape-scale taphonomic signal of carnivore modification to the surficial bone assemblage in Amboseli Park, Kenya as it was in 1975 and 2002–2004. Change in predator abundances over time provides a means of assessing the taphonomic signal of carnivore-mediated bone consumption and destruction under differing ecological conditions and varying levels of conspecific competition for resources. The landscape assemblage indicates taxonomic variation in the patterning of carnivore modification to ungulates of different size classes as well as within equivalent size classes. Analyses of long bone elements indicate that the differential destruction of limb ends and the strength of the correlation between limb end abundance and bone mineral density provide an indication of the intensity of carnivore modification to a faunal assemblage. The ability to infer levels of carnivore modification based on limb elements can provide faunal analysts with the tools to determine whether the taphonomic signals in the fossil record relate to carnivore modification, hominin transport of appendicular elements, or both.     

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.     

Factors affecting Early Stone Age cut mark cross-sectional size: implications from actualistic butchery trials

Early Stone Age cut marks are byproducts of hominins' tool-assisted animal carcass consumption and provide a potential avenue of inference into the paleoecology of hominin carnivory. If diagnostic cut mark characteristics can be linked to flake and core tool use or the completion of distinct butchery actions, it may be possible to infer ancient tool preferences, reconstruct the consumption of specific muscular tissues, and illuminate landscape-scale stone resource use. Recently, diagnostic morphological criteria including cut mark width and depth have been used to identify marks made by different classes of experimental and archaeological stone tools (Bello, S.M., Parfitt, S.A., Stringer, C., 2009. Quantitative micromorphological analyses of cut marks produced by ancient and modern handaxes. Journal of Archaeological Science 36: 1869–1880; de Juana, S., Galan, A.B., Dominguez-Rodrigo, M., 2010. Taphonomic identification of cut marks made with lithic handaxes: an experimental study. Journal of Archaeological Science 37: 1841–1850; Dominguez-Rodrigo, M., de Juana, S., Galan, A. B., Rodriguez, M., 2009. A new protocol to differentiate trampling marks from butchery cut marks. Journal of Archaeological Science 36: 2643–2654). The work presented here adds to this experimental butchery database by using measurements of cut mark cross-section taken from bone surface molds to investigate how stone tool characteristics including flake versus core tool type, edge angle, and tool weight, influence cut mark width and depth, ultimately testing whether cut mark size is a useful indicator of tool identity. Additionally, these experiments investigate the influence of contextual factors, including butchery action, carcass size, and bone density on cut mark size. An experienced butcher used replicated Oldowan flakes and bifacial core tools in experimental trials that isolated skinning, bulk and scrap muscle defleshing, and element disarticulation cut marks on goat and cow skeletons. This sample explores cut mark traces generated under realistic butchery scenarios and suggests the following results: 1) Core and flake tools were equally efficient at completing all butchery tasks in size 1 and 3 bovid carcasses. 2) Samples of cut mark width and depth produced by core and flake tools were similar and cut marks could not be accurately classified to a known tool type. 3) Skinning and disarticulation activities produced significantly wider and deeper marks than defleshing activities. 4) Cut marks on cows tended to be wider and deeper than those on goats. 5) Cut mark width is negatively correlated with bone density when carcass size and bone portion are taken into consideration. These results suggest that a general quantitative model for inferring tool type or edge characteristics from archaeological cut mark size is not warranted.     

Bone surface modifications in zooarchaeology

Cutmarks made by stone tools, conchoidal flake scars from hammerstone percussion, carnivore tooth marks, striations from sedimentary abrasion, and other surface modifications on bones from archaeological sites constitute a crucial body of evidence for investigating the role of human behaviors and of nonhuman taphonomic processes in site formation. This paper describes the various kinds of bone surface modifications produced by humans and by nonhuman processes and assesses the current status of bone surface modification studies with regard to such issues as the need for greater analytical standardization, the selection of instruments for examining bone specimens, tactics for identifying the origins of marks on bones, and strategies for inferring human behaviors.     

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.     

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.     

Cut Mark Cluster Geometry and Equifinality in Replicated Early Stone Age Butchery

Early Stone Age cut marks created during tool‐assisted carnivory potentially offer inferences into hominin butchery behaviour and access to complete or defleshed carcasses. Actualistic butchery trials of 16 goat and cow half‐carcasses were completed by an experienced butcher with replicated Oldowan tools to investigate how the geometric organisation of cut mark clusters reflects flake versus core tool use and bulk muscle versus scrap defleshing. A cluster of cut marks is defined as a series of adjacent cut mark striations that occur at an anatomical location and are bounded by unmarked cortical surface. Tool type and butchery action were predicted to differentially mark certain long bone portions and influence cluster attributes. Moulds of 613 cut mark clusters were photographed and measured using ImageJ software (National Institutes of Health, Bethesda, Maryland, USA) for cluster area, cut mark count, median cut mark length and standard deviation of cut mark length and angle. Analysis suggests the following results: (i) cluster attributes are correlated;(ii) changes in cluster geometry are related to increasing cut mark count and length but not tool type or defleshed muscle amount; (iii) large clusters occur on large animals; and (iv) long bone midshaft portions are cut‐marked during both bulk and scrap muscle defleshing. Analysis of 179 cut mark clusters on long bone shafts of sizes 1–4 mammals from three Okote member assemblages at Koobi Fora, Kenya, shows that archaeological clusters have a similar number of marks when compared with experimental clusters but are significantly smaller, have shorter median marks and include less deviation in mark length and angle. Archaeological clusters corroborate that increasing area is positively correlated with cut mark count, median mark length and standard deviation of mark length and angle. A quantitative inferential model that links cut mark cluster geometry to tool type or the amount of muscle defleshed is not supported by these data. Copyright © 2015 John Wiley & Sons, Ltd.