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.     

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.     

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.     

The Taphonomy of Resource Intensification: Zooarchaeological Implications of Resource Scarcity Among Bofi and Aka Forest Foragers

Zooarchaeological analyses often rely on bone fragmentation, cut marks, and other taphonomic indicators to bolster interpretations of resource intensification that are based on observed changes in prey types and frequencies. While these taphonomic indicators are assumed to be good proxy measures of processing effort, this assumption is based on inadequate actualistic data and analysts often conflate one or more taphonomic indicators as manifestations of the same process. In this paper, we present zooarchaeological data from two villages occupied by Central African forest foragers with very different foraging efficiencies. These data provide the first case where known disparities in diet breadth and foraging efficiency are matched with prey assemblages and taphonomic attributes. Observational and quantitative data show differences between the villages in diet breadth and access to high-ranked prey, but specific taphonomic indicators such as cut mark distribution and intensity do not match predictions generated from models of resource intensification. We propose that linking different taphonomic processes to resource scarcity and intensification can provide powerful adjunctive information. However, because different processing outcomes may be associated with different kinds of resource intensification in response to different kinds of scarcity, we need to strengthen the validity of purported taphonomic indicators with more rigorous independent studies.     

Assessing the variability in taphonomic studies of modern leporid remains from Eagle Owl (Bubo bubo) nest assemblages: the importance of age of prey

The Eagle Owl (Bubo bubo) is an important leporid remains accumulator at archaeological sites. Taphonomical studies of modern leporid remains from nest assemblages of this nocturnal raptor aim to establish the main characteristics that define the taphonomic signature of Eagle Owls in order to distinguish them from that of other leporid predators, namely diurnal raptors, terrestrial carnivores and humans. However, analyses carried out so far have shown that the identification of diagnostic features of Eagle Owls leporid nest accumulations is not a straightforward issue – variability has been observed. Age and size of prey are pointed among others as accountable factors but little research has been done on the study of these variables. Here we present the analysis of two samples of modern leporid remains recovered from Eagle Owls nests with the aim of contributing new data to this subject. The high presence of juvenile individuals in both samples enables us to assess anatomical representation, breakage and digestion patterns of immature leporid remains and also to compare them with other assemblages richer in adult prey. Results show that the main characteristic features noted in previous works which recognise Eagle Owls as agents of leporid bones accumulations in the fossil record, are not significantly affected by the age of prey. Less variability than originally thought occurs which means other variables need to be further investigated.     

Bone damage patterns found in the avian prey remains of crested caracara Caracara plancus (Aves,Falconiformes)

The following paper presents the results of the analysis of the avian prey bones found in uneaten remains of crested caracara (Caracara plancus, Aves, Falconiformes) from La Pampa province, Argentina. Anatomical parts representation and taphonomic modifications were evaluated and compared to results of the evaluation of bone remains recovered from crested caracara’s pellets and to previous studies of other diurnal birds of prey. The results suggest a preferential consumption of some body parts of avian prey, as evidenced in the high frequency of wing elements in the uneaten prey remains. This analysis helps to support interpretative data concerning the origins of avian remains in the zooarchaeological and paleontological record, and contributes to the knowledge of a common predator found throughout diverse environments in South America.     

Who brought in the rabbits? Taphonomical analysis of Mousterian and Solutrean leporid accumulations from Gruta do Caldeirão (Tomar,Portugal)

In the Iberian Peninsula, leporids, and specifically rabbits, play a key role in the understanding of hunter-gatherer economies. They appear to have been especially important in the Tardiglacial, when large numbers of small prey animals and of the European rabbit (Oryctolagus cuniculus) in particular, are a ubiquitous feature of faunal assemblages from archaeological sites. Since a large number of non-human predators can also contribute to the formation of such assemblages, the ability to discriminate between bones accumulated by humans and by other kinds of predators is a key prerequisite to their interpretation. On the basis of systematic actualistic studies carried out on modern leporid remains produced by mammalian terrestrial carnivores, nocturnal and diurnal raptors, and humans, we identified diagnostic taphonomic indicators of the different predators. In this paper, the patterns observed on the modern material are applied to the taphonomical analysis of two archaeological samples of rabbit and hare remains from Mousterian and Solutrean layers of Gruta do Caldeirão, a cave site located in Central Portugal. Our results suggest that Eagle Owl (Bubo bubo) were mainly responsible for the Mousterian accumulations, whilst the Solutrean ones were most likely the result of human activity. These data support the notion that, in Iberia, significant reliance on rabbits does not become a feature of subsistence strategies until later Upper Palaeolithic times.     

Deciphering bone depositional sequences in caves through the study of manganese coatings

The presence of black-coloured bones in caves has been sometimes associated to the use of fire by humans, but more frequently the agent responsible for black colour is the natural deposition of manganese oxides. Manganese coatings were not usually studied once human action was discarded. However, they can provide valuable information about the origin and the sequence of formation of a palaeontological or archaeological deposit. In this paper, we present a quantitative and qualitative study of the manganese coatings that affect most of the bones recovered in the palaeontological site of Liñares cave (NW of Iberian Peninsula) together with other taphonomic marks. The manganese coatings are superimposed on other features previously existing on the surface of the bones. In turn, the coatings can be affected by other agents that modified them totally or partially. The combined interpretation of the different taphonomic marks provides us with valuable information on the sequence where the different taphonomic processes were produced, as for example the intervention of carnivores in the disarticulation of some elements before the burial, the preservation of anatomic connections until after the burial of the carcasses, and the existence of reworking processes in some parts of the infill.     

Carcass damage and digested bone from mountain lions (Felis concolor): implications for carcass persistence on landscapes as a function of prey age

This actualistic study examines the taphonomic signatures of modern free-ranging mountain lions (Felis concolor) on prey skeletal remains left at kill sites and digested bone fragments in the lions’ scats. Conducted in western Texas and southeastern New Mexico (U.S.A.), the study and its outcomes are relevant to models of carcass persistence and scavenging opportunities on ancient landscapes. Mountain lions in the study area ingested disproportionate quantities of bone from very young prey. This tendency holds true irrespective of prey body size. The results meanwhile confirm a relatively mild pattern of damage to adult deer carcasses. Digestive erosion of the surfaces of bones that passed through the gut was relatively severe, but many of these bone and tooth specimens retained identifiable features. It is clear that the mountain lions quickly remove the carcasses of very young prey from the pool of potentially scavenge-able resources. The non-linear relation between bone destruction from feeding by the cats and the skeletal maturity of prey also has consequences for prey mortality patterns, specifically a bias against the representation of very young individuals. This effect is not sufficient, however, to produce a global bias to prime-adult prey because older juveniles are much less affected.     

Neotaphonomic measures of carnivore serial predation at Ngamo Pan as an analog for interpreting open-air faunal assemblages

Here I present a neotaphonomic account of natural bone accumulations that have resulted from carnivore serial predation at Ngamo Pan, a vast complex of seasonal water holes located in Hwange National Park, Zimbabwe. Reconstructing the taphonomic histories of faunal assemblages from open-air archaeological sites is burdened with a host of interpretive complications, and this study is offered as a referential means for evaluating instances where stone tools and bone accumulations are associated in apparent archaeological contexts. While the presence of stone tools implies some involvement on the part of humans, open-air sites near water would also have served as prime locations for serial predation by large carnivores to ambush prey—a situation that, over time, can mimic archaeological bone accumulations. The taphonomic and zooarchaeological signatures of carnivore serial predation at Ngamo Pan show marked similarities with the open-air faunal accumulation from Kalkbank, a late Pleistocene site in Limpopo Province, South Africa, located along the margins of a relict pan. Many potential archaeological sites within the interior of southern Africa dating to the Middle Stone Age are known from open-air settings near permanent or ephemeral bodies of water, and the ability to decipher between hominin and non-hominin carnivore involvement with bone accumulations is paramount in determining the hunting and scavenging behaviors of our early ancestors. As much of our understanding of hominin subsistence during the Middle Stone Age is drawn from coastal cave locations, this study is intended to encourage a broadening of our perspective on the taphonomic histories of faunal accumulations dating to the Middle Stone Age by incorporating supplementary evidence provided by these open-air sites.     

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.     

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.     

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.     

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.     

A rallying point for different predators: the avian record from a Late Pleistocene sequence of Grotte des Barasses II (Balazuc,Ardèche,France)

The presence of processed birds in the archeological faunal record is considered key to assessing human dietary evolution. Taphonomic studies on birds from sites older than Marine Isotope Stage (MIS) 2 have become relevant in the last few years, leading to the proposal of more complex scenarios of human subsistence. Several works have demonstrated direct evidence of bird consumption by Homo prior to anatomically modern humans in Europe; however, others support the hypothesis of non-anthropogenic bird accumulations. This has led to the necessity of determining what elements or factors cause the human exploitation of birds in some archeological sites before the end of the Pleistocene. The Grotte des Barasses II site is located within this framework. Short-term human occupations have been attested by the presence of lithic tools and processed macrofaunal remains. Additionally, a small assemblage of bird bones has also been recovered. Here, we present a detailed taphonomic study with the aim of exploring possible relationships between these avian taxa and human occupations. Despite the fact that Neanderthals inhabited the cave, avian specimens show damage pointing to different causative agents. Direct evidence (digestion, gnawing) indicates that mammalian carnivores and nocturnal raptors were mainly involved in the accumulation of bird bones. We propose some factors that might determine whether or not small game was exploited in this specific locality and emphasize the importance of such analytical approaches in the general interpretations of the Pleistocene sites.     

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.     

A taphonomic study of a carcass consumed by griffon vultures (<Emphasis Type="Italic">Gyps fulvus</Emphasis>) and its relevance for the interpretation of bone surface modifications

Vultures are ubiquitous taphonomic agents in most biomes. However, taphonomic studies of vultures are scarce and very little is known of the damage they cause to bone surfaces when consuming carcasses. This study presents the results of a taphonomic analysis of bones modified by griffon vultures. Although some marks created by these agents are diagnostically unique, others are extremely similar to modifications caused by other biostratinomic processes. This renders interpretations of marks on bones very problematic when these modifications are considered in isolation rather than at the assemblage and contextual levels.     

Can Archaeozoology and Taphonomy contribute to knowledge of the feeding habits of the Iberian wolf?

The feeding behaviour of the Iberian wolf (Canis lupus signatus) has usually been studied using traditional methods, such as the analysis of scats or stomach contents of dead individuals. Such studies usually analyze the hair: mainly feathers and occasionally a few bones. In the present work, a study of skeletal remains found in current wolf scats from 7 natural areas on the Iberian Peninsula is presented. Methods of archaeozoological and taphonomic research are applied in the study of this sample. Data concerning the type of prey-species, the body size, MNI, age groups and skeletal representation of prey consumed and defecated are provided. In addition, some data relating to the physical and biochemical modifications produced by consumption and digestion are provided. This type of study is suitable for the analysis of the current faecal remains in order to complement other studies on the feeding habits of the Iberian wolf where traditional methods are used. However, some considerations must be taken into account in future studies.     

Probable evidences of bone accumulation by Pleistocene bearded vulture at the archaeological site of El Mirón Cave (Spain)

The bearded vulture (Gypaetus barbatus), a bird or prey that mainly feeds on bones of large mammal carcasses, has been recently evidenced as an accumulating agent with significant contributions to palaeontological assemblages in caves and shelters, but extremely rare at archaeological sites. The results of the taphonomic analyses carried out in the Upper Palaeolithic layers of El Mirón Cave (Cantabrian Spain), which are presented here, evidence the existence of special digestive marks and a typical skeletal pattern in a noticeable amount of small and medium-sized ungulates bones. This fact suggest that a representative part of the bone assemblage has been caused by bearded vultures, and strengthens the idea that this bird has to be definitely included among other potential non-human accumulators in archaeological sites.     

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.