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.     

The biological and archaeological implications of the beetle Aglenus brunneus (Gyllenhal) in ancient faunas

The distribution and ecology, as far as they are known, of the Colydiid beetle Aglenus brunneus (Gyll.) are summarized and the significance of its records from archaeological sites considered. The supposition that A. brunneus originated in North America (Peyerimhoff, 1945) is questioned in the light of the ancient records. The species is used to illustrate some of the general principles and problems involved in the theoretical reconstruction of past environments from ancient faunas.     

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.     

The first fossil crow (Corvus sp. indet.) from the Early Pleistocene Nihewan Paleolithic sites in North China

We describe a three dimensional preserved right tarsometatarsus, representing the first fossil crow from the well-known Early Pleistocene Nihewan paleolithic sites in North China. The new specimen is distinguished from other known species of the genus Corvus by the combined morphological features, especially the peculiar hypotarsus. The phylogenetic significance of the arrangement of canals at the proximal hypotarsus in passerines is discussed. Although the fossil was too incomplete to warrant the erection of a new species of Corvus, it nonetheless increases the taxonomical diversity of the Nihewan fauna and improves our knowledge on Pleistocene birds in China. Finally the paleoenvironment of the Nihewan sites are discussed in terms of its faunal composition.     

Neolithic and Bronze Age ungulate footprint-tracks of the Severn Estuary: Species,age, identification and the interpretation of husbandry practices

Ungulate footprint-tracks provide information regarding the species and age of animals. Combined with other datasets, this contributes to interpretation of seasonal husbandry patterns in the Severn Estuary, focusing on Bronze Age intertidal footprint-tracks at Redwick and Goldcliff East and the Late Neolithic site of Oldbury. Metric dimensions and morphology of modern contemporary ungulate footprint-tracks are used as analogues to help understand the species and age of prehistoric ungulates. Findings indicate that Dexter cattle and Soay sheep are metrically similar to British prehistoric ungulates. The prehistoric sites have a concentration of neonatal and juvenile individuals. Along with evidence provided by environmental data, faunal skeletal assemblages and lipid and isotopic analysis, this leads to the conclusion that the presence of younger animals and evidence at Brean Down for dairying is consistent with saltmarsh grazing activity in spring and summer.     

Radiocarbon and U-series dating of the endemic deer Praemegaceros cazioti (Depéret) from “Grotta Juntu”, Sardinia

Radiocarbon and U-series methods (²³⁰Th/U and ²³¹Pa/²³⁵U) for absolute age determination have been applied to some fossil samples from Grotta Juntu, in North Eastern Sardinia (Italy). The remains belong to the endemic deer species Praemegaceros cazioti, which is here represented by an almost complete skeleton. The three dating methods lead to concordant ages of about 7500 years BP, indicating that the skeleton was maintained as closed system after burial. Taking into consideration these results, Praemegaceros cazioti from Grotta Juntu is now the youngest representative of this species in Sardinia.     

Domestic and wild ungulate dietary traits at Kouphovouno (Sparta,Greece): implications for livestock management and paleoenvironment in the Neolithic

The objective for this study is to explore interspecific variations in domestic and wild ungulate diets and management at the Neolithic site of Kouphovouno (Sparta, southern Greece). We tested four hypotheses related to environmental context and livestock management using, for the first time, a combination of mesowear and microwear analyses on a Neolithic site. We identified interspecific differences between ungulate taxa, especially between wild and domestic ungulates, however there is no significant difference between Ovis and Capra. There is no diachronic change in diet from the Middle Neolithic to the Late Neolithic. Changes in animal utilization are not reflected in their diet. Whatever the purpose of the husbandry, animals had access to the same food resources. The study of dental wear patterns gave some insights regarding the management practices for the domestic livestock as well as paleoenvironment through the study of wild ungulates.     

Disentangling long bones of foxes (Vulpes vulpes and Alopex lagopus) from arctic archaeological sites

To discern the presence of two anatomically close species within an archaeological assemblage is always a problem for the analyst. A particularly interesting case is the distinction between the red fox (Vulpes vulpes) and the arctic fox (Alopex lagopus), which is often determined using non-rigorous criteria and producing results that must be regarded with caution. Classical methods for separating these two species and determining sexual dimorphism, and eventually the sex ratio, such as analyses using uni- and bivariate plots, are based on arbitrarily fixed limits between sexes. In this text, a more robust statistical method termed mixture analysis (MA) is used to determine the species of limb bones from foxes. First, the MA is applied to a sample of each species using metric data taken from a collection of present-day fox skeletons. Afterwards, the MA is applied to archaeological samples dated to the Dorset period and retrieved from the Tayara site (near Salluit, South Hudson Strait, Nunavik). The results clearly demonstrate that the greatest length (GL) of long bones, especially the humerus and tibia, is the best measurement for distinguishing the species, followed by the distal breadth and the proximal breadth; and that GL is better suited for sexing foxes. The results of MA applied to the Tayara site collection lead to the conclusion that the red fox is present along with the arctic fox among the assemblage, and that there is no fox body size change during the last two millennia in the Eastern Arctic.     

Reappraisal of ‘chronospecies’ and the use of Arvicola (Rodentia,Mammalia) for biochronology

The water vole, genus Arvicola, is characterised by a broad geographic distribution throughout Europe and is widespread during the late Middle and Upper Pleistocene. This genus is used as a major biostratigraphic tool within the Quaternary. Specific determinations using the Schmelzband-Differenzierung-Quotient or SDQ have identified many chronospecies within the fossil species Arvicola cantiana (Hinton, 1910). As SDQ calculation remains limited, this study reappraises the Arvicola genus in terms of morphodiversity and morphospace using outline analysis which takes into account the tooth as a whole. Outline analysis suggests that one single species of Arvicola, A. cantiana, was present during the Pleistocene. This species shows great variability with no trends or patterns in morphospace. Thus, these results call into question the reliability of SDQ for specific determinations and throw doubt on the biochronological framework based on Arvicola.     

New results on deer from swanscombe,and the stratigraphical significance of deer in the middle and upper pleistocene of Europe

The value and limitations of mammalian fossils as biostratigraphic indicators in the Pleistocene are discussed. Aspects of a species' history which are of potential biostratigraphic value are its overall stratigraphic range, shifts in geographical distribution, evolutionary transitions, and changing abundance in the fauna. These form a complex pattern which must be established in some detail before a species can be used in dating a “new” locality. The major features of stratigraphic range, distributional history, and evolution of each of the nine species of cervid from Britain and mainland Europe during the Middle and Upper Pleistocene are outlined, with notes on their possible biostratigraphic value. Some new results on the deer remains from Swanscombe are presented. Fallow deer can be referred to subspecies Dama dama clactoniana only in the Basal and Lower Gravels and Lower Loam. There is no evidence of change in its body size within the sequence. An idiosyncrasy in the lower dentition of Swanscombe fallow is described. The red deer was the “coronate” form, and was of small body size. The giant deer was of small body size, and may have shared the broad brow tines of the Steinheim population. Roe is rare and elk is absent. The Swanscombe deer suggest a post-Cromerian, interglacial dating of all faunal levels at the site. They are consistent with a wholly Hoxnian age, but precise contemporaneity with Hoxne Bed 1 is unlikely, and the possibility of additional stages being represented cannot be excluded.     

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.     

Stable isotopes and diet at Ancient Kerma,Upper Nubia (Sudan)

Carbon and nitrogen stable isotope values were measured from bone collagen extracted from archaeological Nubian human (n = 54) and faunal (n = 61) populations from the site of Kerma, Sudan. Collagen suitable for isotopic analysis was extracted from 22 faunal and 48 human samples from the Eastern cemetery site, dated to the Middle Kerma (c. 2050–1750 BC) and Classic Kerma (c. 1750–1500 BC) periods respectively. The isotopic data indicate that the human dietary regimen included a mix of C₃ and C₄ plant-derived components, with a larger C₄ component than previously reported in archaeological Egyptian Nile Valley populations. Elevated δ¹⁵N values are attributed to consumption of dietary resources from a ¹⁵N-enriched terrestrial ecosystem. The faunal isotope data also indicate the consumption of C₃ and C₄ plants. The large range of δ¹³C values measured in both the human and faunal samples supports previous work suggesting that a significant portion of the populations buried at Kerma may have originated elsewhere, further confirming the Nile Valley as a major corridor for population movement in the region.     

Stone age predation on large African bovids

Crown height measurements are used to establish age distributions for several species of larger bovids represented in faunal samples from the Middle Stone Age (earlier Upper Pleistocene) deposits of the Klasies River Mouth Caves and the Later Stone Age (later Upper Pleistocene/Holocene) deposits of Nelson Bay Cave, South Africa. There are no obvious differences between the sites in the age distributions of the species they share, but there are significant differences in age distributions among species. Two basic patterns are apparent. In the first, characterizing the blue antelope (Hippotragus leucophaeus), roan antelope (Hippotragus equinus), Cape buffalo (Syncerus caffer), and giant buffalo (Pelorovis antiquus), the archaeological samples contain numerous very young animals and relatively few prime-age adults. At least in the buffalo samples, there is also a fair representation of old adults. In the second pattern, characterizing the bastard hartebeest (Damaliscus dorcas or D. niro) and especially of the eland (Taurotragus oryx), prime adults are far more prominent relative to younger and older age groups. The first pattern is similar to the natural pattern of attritional mortality that probably characterizes all healthy, stable populations of free-ranging large ungulates, while the second is more reminiscent of the age structure of live herds. The first pattern may reflect hunting focused on individual animals, particularly those whose age made them most vulnerable, while the second may reflect the susceptibility of certain species to driving, so that whole groups could be killed in traps in which differences in age had no meaning.     

New evidence for the establishment and management of the European fallow deer (Dama dama dama) in Roman Britain

The semi-domestic status of the European fallow deer (Dama dama dama) renders its ancient biogeography a reflection of human activity with the potential to provide important insights into the movement, trade patterns and ideology of past societies. Given this potential, fallow deer have received surprisingly little attention from scientists within the fields of archaeology, biology and zoology. Here we present new AMS radiocarbon dates, stable carbon and nitrogen isotope data and genetic evidence (the first ancient DNA sequences for the species) resulting from the analysis of a set of remains recovered from the Roman settlement at Monkton on the Isle of Thanet, Kent, UK. By viewing our results against the very limited available comparative data, this paper provides new information for the establishment and management of fallow deer in Britain. We argue that much more could be achieved with even a slight increase in sample sizes and a plea is made for greater research into this culturally significant species.     

Pacific cod (Gadus macrocephalus) as a paleothermometer: otolith oxygen isotope reconstruction

Stable isotope studies are increasingly important for understanding past environmental and cultural developments along the North Pacific Rim. In this paper, we present methods for using Pacific cod (Gadus macrocephalus) otoliths as a paleothermometer using a case study from Kodiak Island, Alaska. The results of this study indicate that Pacific cod otoliths record variable paleoenvironmental conditions during the Little Ice Age. The broad distribution of Pacific cod and success in using the otoliths as a paleothermometer makes this method widely applicable to researchers working throughout the northern Pacific Rim.     

Molecular markers for the discrimination of Triticum turgidum L. subsp. dicoccum (Schrank ex Schübl.) Thell. and Triticum timopheevii (Zhuk.) Zhuk. subsp. timopheevii

The persistent uncertainty on the classification of the “new” glume wheat found in Neolithic and Bronze Age sites from Greece and other European settlements might be resolved only through analysis of its ancient DNA. Tools able to discriminate among different Triticum species on the basis of scarce, very damaged DNA, are therefore essential. While current attempts concentrate on DNA fragments sequencing and comparison, in some instances PCR-based selective amplification techniques might offer a cheaper and quicker alternative. The purpose of this research was therefore the identification of species-specific primers, able to distinguish caryopses of Triticum timopheevii subsp. timopheevii from those of Triticum turgidum subsp. dicoccum. Primers and their working conditions were defined and optimized using DNA from modern accessions. The ribosomal primers ITS1 tim and ITS2 tim, and the nuclear primer acetyl-coenzyme A tim clearly discriminated the sequences of Triticum timopheevii from other species. Finally, Neolithic charred wheat grains found in the sites of Sammardenchia (Pozzuolo del Friuli, Udine) and La Marmotta (Lago di Bracciano, Roma), belonging to the “new” wheat type or to emmer, were tested with the three selected primers. However, the results were not conclusive, because the samples analysed were apparently too degraded to yield useful DNA.     

Comparative human and deer (Odocoileus virginianus) taphonomy at the Richards site,Ohio

The Richards site is attributed to the Philo phase of the Fort Ancient tradition of the Ohio Valley area. Human skeletal material from the site shows evidence of peri‐ and post‐mortem taphonomic changes, including cut marks, burning and fracturing. Previous analyses have discussed explanations for these changes, including secondary burial, ritual destruction and cannibalism. Researchers have theorised that, allowing for differences in anatomy among species, humans and animals butchered for the same purpose (consumption) will show similar patterns of taphonomic changes associated with butchery. The human remains at the Richards Site were disposed in general midden pits containing mixed cultural debris and faunal remains. White‐tailed deer (Odocoileus virginianus) constitutes approximately 60% of all the faunal bone, indicating that it was a major food resource. To test a cannibalism explanation, a comparative analysis of human‐induced taphonomy in human and deer skeletal remains was performed, using chi‐square and odds ratio tests. If humans were being used as a food resource, the pattern of butchery seen would mirror that of the deer. The analysis described here compares the patterns of treatment and disposal of human and deer skeletal elements at the Richards site, to test whether both species were used as food resources. Similar types of evidence for human‐induced taphonomic changes, including cutting, chopping, burning and breakage, can be seen in both species. However, results indicate that, in general, human remains show much more evidence of perimortem treatment than do deer remains. In fact, the common odds ratio for perimortem treatment in all bones is 3.25, indicating that a human bone is 3.25 times as likely as a deer bone to be affected by burning, cutting or chopping. This probably indicates that perimortem treatment of humans was greater than that necessary simply for butchering for consumption. Copyright © 2005 John Wiley & Sons, Ltd.     

Paleodemographic analysis of a fossil porcupine (Hystrix refossa Gervais, 1852) population from the Upper Pleistocene site of Geula Cave (Mount Carmel,Israel)

For decades zooarchaeological studies mainly focused on teeth age profiles to derive central hypotheses concerning human subsistence during the Pleistocene. Behavior was often related to hunting and/or scavenging strategies according to two models: attritional versus catastrophic. However, few studies have estimated basic demographic parameters using standard methods of population ecology, which are necessary for a more confident interpretation of animal paleopopulation structures. Thanks to a remarkably rich fossil bone assemblage from the Upper Pleistocene cave of Geula (Mount Carmel, Israel), in this paper we perform a paleodemographic analysis of Hystrix refossa, Gervais 1852 (Mammalia, Rodentia). We first consider phylogenetic and morphometric dental criteria for the comparison of Pleistocene porcupine species, followed by an examination of taphonomic conditions of the bone assemblage. We focus strongly on the age structure of the fossil population, presenting a new methodological approach that combines life tables and Leslie matrix models – one of the best-known methods in population ecology for assessing population growth and age distribution. We detail demographic parameters derived from cross-sectional life tables and past trends in the porcupine population at Geula Cave using simple matrix projection model. Finally, we were able to perform an elasticity analysis to identify which demographic component was potentially the most critical for influencing growth. We show that the population of H. refossa did not decline or shrink but was stable. Our results permit us to characterize Geula Cave as a natural shelter for porcupines, with limited evidence to scavenging and hunting by either hyenas or humans.     

Starch grain characterization of Prosopis chilensis (Mol.) Stuntz and P. flexuosa DC,and the analysis of their archaeological remains in Andean South America

The fruit (pods) of Prosopis (Fabaceae) are frequently recovered from pre-Hispanic Argentinian archaeological sites, suggesting that this genus was of importance in ancient economies in this region. Yet it is only recently that archaeobotanists have begun to carry out systematic research into this genus. Therefore many questions remain to be addressed concerning the food value of Prosopis fruit, and its potential contribution. This paper examines starch from the pods of two species, Prosopis flexuosa and Prosopis chilensis, for the purposes of describing and classifying their morphological features and biometrical parameters. Pods of both species were gathered from two extremes (northern and southern) of the Hualfín Valley, Catamarca, Argentina. Starch abundance is estimated and compared with that of Zea mays, an economically important plant with high concentrations of starch. This paper reveals that Prosopis pods contain very low amounts of starch compared to starch-rich edible species. Despite this, the recovery of Prosopis starch grains can be successfully used to assess archaeological tool uses. Prosopis starch grains were found to be highly variable in shape. Grain size ranged between 10 and 20 μm. Granule irregularity and the high birefringence under polarized light are two of the most important diagnostic features. No significant statistical differences were found in the structure (morphology, size and hilum) of the starch of the same species from both localities. Finally, the implications for the role of this plant in past societies are discussed.     

Endangered species,archaeology, and stable isotopes: huemul (Hippocamelus bisulcus) isotopic ecology in central-western Patagonia (South America)

The management and conservation of animal species should be based upon a long-term analysis that considers its geographical distribution and feeding patterns. The study of carbon and nitrogen stable isotopes on skeletal remains provides a quantitative approach to the paleodietary reconstructions and constitutes a potent tool for comparing behavioral aspects of the fauna. In this paper we present the first set of isotopic values for modern and archaeological samples of Huemul (Hippocamelus bisulcus) from the forests of Central-western Patagonia, Chile. Contrary to initial expectations, our analysis indicates that there is no evidence of an important incidence of the canopy effect on the δ¹³Ccollagen values, which we suggest is due to an ecological selection that drives the huemul to focus its predation on open sectors within the forest. On the other hand, a wide range of δ¹⁵Ncollagen values for huemul was verified. This could reflect the N impoverishment of the forest soils and provide an interesting ecological indicator. The long-term information offered by the archaeological record provides the necessary context for decision-making conducive towards the preservation of Hippocamelus bisulcus in Patagonia.