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.     

Gazelle bone marrow yields and Epipalaeolithic carcass exploitation strategies in the southern Levant

We measure the bone marrow yields of mountain gazelle (Gazella gazella) carcasses to reconstruct gazelle exploitation strategies during the Epipalaeolithic periods in the southern Levant. We present experimentally derived data on the bone marrow content of seven fresh gazelle carcasses, determine the range of marrow yields among individuals of different sexes and seasons of death, and compare the new data to gazelle skeletal element abundances from five Epipalaeolithic assemblages from Israel. We found extensive variation in marrow fat content among individual gazelles. Animals with the highest marrow yields were killed in the spring while animals killed in the early autumn had lower fat contents. Nevertheless, our results suggest that gazelle marrow provided a reliable, albeit small fat resource for prehistoric foragers in all seasons. Strong relationships between bone fragmentation and marrow content demonstrate that Epipalaeolithic people preferentially processed bones with high marrow yields.     

Gazelle bone fat processing in the Levantine Epipalaeolithic

We investigate mountain gazelle (Gazella gazella) carcass processing to reconstruct resource intensification strategies during the Epipalaeolithic period of northern Israel. We adopt a multivariate taphonomic approach to identify the processes that most influenced bone survivorship in five gazelle assemblages. All of the assemblages are characterized by significant density-mediated biases, yet in situ attrition played a minimal role in assemblage formation. In contrast, the survivorship of hare (Lepus capensis) skeletons is not mediated by bone density indicating that different prey taxa experienced independent taphonomic histories. Both gazelle cortical and cancellous bone is highly fragmented and the degree of fragmentation and survivorship are strongly correlated with fat yields. Results of multiple tests point to intensive marrow and grease extraction as the primary determinant of gazelle bone survivorship. Although gazelle carcasses were intensively utilized throughout the Epipalaeolithic, the intensity of use is stable across the duration of the period.     

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.     

“After 20 Years”: A Taphonomic Re-evaluation of Nahal Hadera V,an Epipalaeolithic Site on the Israeli Coastal Plain

We carried out a detailed taphonomic study of the faunal remains from Nahal Hadera V (NHV), a large Kebaran open-air site on the northern coastal plain, Israel. Faunal remains were previously identified to bone element and species (Saxon, Martin & Bar-Yosef, 1978). However, NISP and MNI values were calculated based on teeth and epiphyses only. In this study we identified the maximum number of skeletal elements, including head fragments, vertebrae, ribs, carpals/tarsals, and shaft fragments. Further analyses included search for various bone surface modifications and the study of mode of bone fragmentation. We found minor loss of bones caused by various diagenetic processes, and no signs of carnivore activity. These results suggest that most of the bone destruction was caused during time of occupation, probably for marrow consumption, as indicated by mode of bone fragmentation. Absence of selective transport for gazelle (Gazella gazella) and fallow deer (Dama mesopotamica), the major prey species (with 70% and 20%, respectively), is supported by homogeneous representation of head-limb-toe elements. However, the carcasses may have been processed before transport to the site. Thus, a taphonomic analysis from NHV helps to illuminate the economy of NHV inhabitants and fine-tunes our understanding of the depositional history of the site. However, it does not overthrow results of earlier research using classical methods.     

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.     

Pinniped behavior, foraging theory, and the depression of metapopulations and nondepression of a local population on the southern Northwest Coast of North America

Models derived from foraging theory suggest that high-value prey will be depressed (encounter rates will decrease) relative to low-value prey as human predation intensifies. Numerous case studies in ethnographic and zooarchaeological settings indicate depression of prey is very common. Exceptions to depression are few and have been cited as evidence of conservation and resource management. Reanalysis of pinniped and ungulate data from sites on the Oregon coast indicates that the metapopulation (entire population in the region) of otariids was depressed but a single local population of Steller sea lions exploited over a span of 600 years reached an equilibrium with human predation and was not subsequently depressed. This local population was not historically documented, and thus must have been extirpated by commercial sealing in the late nineteenth century. This singular case of prehistoric epiphenomenal conservation was an unintentional result of human predation that took advantage of the lack of escape behaviors among breeding male Steller sea lions.     

“Measure for measure”: a taphonomic reconsideration of the Kebaran site of Ein Gev I,Israel

We present the results of a detailed taphonomic study of the faunal remains from Ein Gev I, a Kebaran open-air site near the Sea of Galilee, Israel. In an earlier study faunal remains were assigned to bone element and species using a rapid method technique based on the identification of teeth and epiphyses only (Davis, 1974). In this study we identified the maximum number of skeletal elements, including fragments of long bone shafts. We also employed a series of taphonomic analyses to reconstruct the depositional history of the bone assemblage and to investigate subsistence, meat procurement and bone processing strategies. We compared the results obtained by a complete bone sampling method with those derived using a rapid method. The bone assemblage is dominated by mountain gazelle (Gazella gazelle), Mesopotamian fallow deer (Dama mesopotamica) and bezoar goat (Capra aegagrus). The assemblage is characterized by significant density-mediated biases, caused by both human bone processing behaviours and in situ post-burial bone attrition. Our attempts to compare species abundance derived from the two different identification protocols revealed no significant difference between the two sampling methods, both in terms of species diversity (richness and evenness) and skeletal element abundance (SEA). These comparisons contrast previous studies that expected to find ubiquitous head-and-foot dominated SEA profiles in zooarchaeological studies that were based on the incomplete identification of faunal remains.     

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.     

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.     

Eagle owl (Bubo bubo) pellets from Roman Sagalassos (SW Turkey): distinguishing the prey remains from nest and roost sites

Two concentrations of animal bones, almost exclusively from small mammals and wild birds, were found within the destruction debris of a Roman bath complex in Sagalassos (SW Turkey). The overall species spectrum, skeletal element representation, fragmentation and preservation condition of the bones indicate that they represent the prey remains of a large nocturnal avian predator, more precisely the eagle owl (Bubo bubo). Differences in skeletal element representation and in prey species' spectrum show that the two bone clusters derive from pellets deposited near a nest site and a roost site, respectively. Radiocarbon dates obtained from the bones indicate that eagle owls lived in the collapsing bath complex during the second half of the 6th to the beginning of the 7th century AD, before the final abandonment of the town. The MNI of the prey animals found at the nest site, confronted with the daily dietary needs of a female eagle owl and its young, indicates repetitive use of the same place during several years. Copyright © 2008 John Wiley & Sons, Ltd.     

Zooarchaeological perspectives on the Chinese Early and Late Paleolithic from the Ma’anshan site (Guizhou,South China)

Ma’anshan is a Paleolithic cave site in the Guizhou province of southern China. The total area of the excavations is ca. 48 m², and the cultural deposit is divided into two layers representing the Chinese Late Paleolithic and the later part of the Early Paleolithic. The upper layer dates between 19,295  BP and 31,155 BP by AMS technique, and the lower layer is dated to around 53,000 BP by U-Series technique. Thousands of bone fragments were recovered from the two layers. Species abundance, bone surface modifications, skeletal element representation, and mortality patterns were studied in an investigation of assemblage formation history and patterns of prey selection and meat consumption by hominins. Interpretations of the faunal data are strengthened by reference to experimental studies and ethnoarchaeological data. The zooarchaeological data indicate that the later occupants of Ma’anshan Cave hunted mainly medium and small game animals, while the earlier occupants tended to prey upon larger animals. In the later period, hominins also made fuller use of the carcasses. The breadth of the meat diet increased with time, due to the inclusion of quick small animals (bamboo rats and birds) in the later period. The differences between the two faunal assemblages are consistent with the chronologic boundary currently drawn between the Chinese Early and Late Paleolithic ca. 30–27 ka BP, based on technological and other evidence.     

Taphonomy of the bones of rodents consumed by Andean hog-nosed skunks (Conepatus chinga, Carnivora,Mephitidae) in central Argentina

The Andean hog-nosed skunks (Conepatus chinga) are generalized consumers. Rodents are a substantial part of their diet. In order to learn about the effects of this predator on the bones of the prey they consume, the bone remains from scats collected in La Pampa, Argentina, were analyzed from a taphonomic point of view. Analysis confirms that C. chinga produces severe destruction of bone remains, mainly during the chewing process. This allows inclusion of this predator in the category of extreme modifier of the bones of its prey. Data presented here and their interpretation provide criteria to identify the intervention of this predator in archaeological and paleontological sites.     

Feeding the scavengers. Actualistic taphonomy in the Jordan Valley,Israel

This paper describes the results of a 3‐year project carried out in the Jordan Valley, Israel, in which the influence of specific local environmental conditions, such as scavenging, weathering and scattering of carcasses, were related to the rate of their bone survival. During the 3 year period, the taphonomy of 16 whole animal carcasses, deposited in a nature reserve and agricultural cultivated area were compared. The exact age, date of death and weight of all animals was known. Results of animal scavenging activities on the carcasses, such as disappearance of bones and damage, such as tooth marks, breakage, gnawing to the leftover bones, were recorded at the end of the initial, intensive scavenging period (Phase I), and at later dates. Bone scattering maps were drawn at the end of Phase I and 2.5 and 3 years later. Weathering of surviving bones at the sites is described. Results show that in the Jordan Valley Nature Reserve, scavenging is the most important taphonomic agent causing total disappearance of all bones of young dairy calves, and reducing amount of bones remaining of more mature animals to a minimum. These small number of leftover bones, after only 2–3 years are the only candidates for eventual burial and fossilization. The loss of most of the bones through scavenging in the nature reserve might cause serious bias to conclusions about subsistence behaviour of prehistoric man. This is a taphonomic actualistic analogue to eventual conditions in prehistoric times. Copyright © 2000 John Wiley & Sons, Ltd.     

Prehistoric seal carcass exploitation at the Shag Mouth site,New Zealand

Seal populations in New Zealand declined dramatically during the prehistoric period. The loss of this important resource significantly affected the foraging practices at the Shag River Mouth site. Previous research documented substantial changes to the diet with the decline of seals and the corresponding decline in foraging efficiency. In this study, I examine how New Zealand foragers altered their use of seal carcasses as the availability of these marine mammals declined. Otariid seal data from the Shag River Mouth site in southern New Zealand are analyzed to test changes in butchery/transport and skeletal element breakage patterns expected with resource depression and declining foraging efficiency. This research shows that at Shag Mouth, seal carcasses were used more intensively over time. However, bone breakage patterns showed little change in the exploitation of within-bone nutrients.     

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.     

Gazelle exploitation in the early Neolithic site of Motza,Israel: the last of the gazelle hunters in the southern Levant

We studied the faunal remains from the Early and Middle PPNB site of Motza, Judean Mts., Israel, in order to gain insight into the economic basis prior to livestock husbandry, with a focus on gazelle hunting. Taphonomic analysis showed that bone preservation at the site was excellent. The subsistence economy in Motza was based on a broad spectrum of hunted species, with mountain gazelle (Gazella gazella) as the dominant prey, similar to many Epipalaeolithic and Pre-Pottery Neolithic sites in the southern Levant. We studied gazelle exploitation patterns, in order to learn about the interaction of humans with this species prior to ungulate domestication. Analysis of the demography of the gazelle herd, which included aging and sexing, revealed no age preferences and no selective culling. Moreover, the PPNB gazelle population of Motza does not exhibit allometric changes in morphology, that are allegedly correlated to increased hunting pressure on gazelle populations prior to livestock domestication.     

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.     

Bone Mineral Density of Wild Turkey (Meleagris gallopavo) Skeletal Elements and its Effect on Differential Survivorship

Zooarchaeologists recognize that density-mediated attrition is a bias that demands consideration in the analysis of archaeofaunal samples. This paper presents bone mineral density values (aereal and volumetric) for the wild turkey (Meleagris gallopavo) and represents the first application of Dual-energy X-ray Absorptiometry (DEXA) to birds. The relationship between density and the survivorship of avian skeletal elements and portions is demonstrated by examining 20 assemblages from the archaeological record of the northeastern United States. The results suggest that density may account for the differential survivorship of least 35% of the samples tested. Further analysis of a single site demonstrates the importance of examining bone mineral density when interpreting past bird hunting and use.     

Cut marks on the Middle Pleistocene elephant carcass of Áridos 2 (Madrid,Spain)

Áridos 1 and Áridos 2 (Madrid, Spain) are two Middle Pleistocene sites belonging to the isotopic stages 9–11. Both places contain partial carcasses of Elephas (Paleoxodon) antiquus associated to Acheulian stone tools. In this work, the taphonomic study of the elephant remains of Áridos 2 is presented. This study has documented several cut marks on different bones, which indicate bulk flesh and viscerae extraction by Middle Pleistocene hominins. Several arguments are provided to support that at least some of the cut marks were made with handaxes, further suggesting that some of these artifacts were butchering tools in this stage of human evolution. Although cut marks on elephant carcasses have been documented at some Middle Pleistocene sites, very few have been published in detail to allow consideration of their status as hominin-imparted marks. By doing so, the present study provides more evidence of large carcass exploitation by hominins during this period.