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We report quantitative paleoecologic data on the large mammal assemblage preserved in lower Pleistocene deposits at Venta Micena (Orce, Granada, southeastern Spain). Taphonomic studies show that bones were collected mainly by hyaenids, which transported and deposited them near shallow dens. Differential fragmentation of major long bones was produced by hyaenas as a function of their density and marrow content. Strong selection of prey by carnivores—which preferentially killed juveniles, females, and individuals with diminished locomotor capabilities among ungulate prey species of larger body size—is indicated by (1) the abundance of remains of juvenile ungulates in relation to the average weight of adult individuals in each species, (2) attritional mortality profiles for ungulate species deduced from crown height measurements, (3) the presence of many metapodials with different osteopathologies in their epiphyses, such as arthrosis, and (4) a biased intersexual ratio of large bovids. Comparison of the frequencies with which modern African carnivores kill and scavenge ungulates from various size classes with the abundance of these size categories in the assemblage suggests that the Venta Micena hyaena (Pachycrocuta brevirostris) was a bone-cracking scavenger that fed largely on carcasses of ungulates preyed upon and partially consumed by fresh meat-eating carnivores such us saber-toothed felids (Homotherium latidens and Megantereon whitei) and wild dogs (Canis falconeri).

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Ecomorphological and biogeochemical (trace element, and carbon, nitrogen, and oxygen isotope ratios) analyses have been used for determining the dietary niches and habitat preferences of large mammals from lower Pleistocene deposits at Venta Micena (Guadix-Baza Basin, Spain). The combination of these two approaches takes advantage of the strengths and overcome the weakness of both approaches. The range of δ13Ccollagen values for ungulate species indicates that C3 plants were dominant in the diet of these mammals. δ13Ccollagen values vary among ungulates: perissodactyls have the lowest values and bovids the highest ones, with cervids showing intermediate values. The hypsodonty index measured in lower molar teeth and the relative length of the lower premolar tooth row indicate that the horse, Equus altidens, was a grazing species, whereas the rhino, Stephanorhinus etruscus, was a mixed feeder in open habitats. The similar δ13Ccollagen values shown in both perissodactyls does not reflect differences in feeding behavior with other ungulates, but rather a lower isotope enrichment factor in these monogastric herbivores than in ruminants, owing to their lower metabolic efficiency. The cervids Eucladoceros giulii and Dama sp. show low hypsodonty values, indicating that they were mixed feeders or browsers from forested habitats, an ecomorphologically based conclusion corroborated in the former by its low δ15Ncollagen content (canopy effect). Bovid species (Bovini aff. Leptobos, Soergelia minor, and Hemitragus albus) presumably inhabited open environments, according to their comparatively high hypsodonty and δ15Ncollagen values. Carnivore species (Homotherium latidens, Megantereon whitei, Pachycrocuta brevirostris, Canis falconeri, and Canis etruscus) exhibit higher δ15Ncollagen values than ungulates. These results record the isotopic enrichment expected with an increase in trophic level and are also supported by low bone Sr.Zn ratios. The elevated δ15Ncollagen value for a sample of Mammuthus meridionalis, which came from an individual with unfused epiphyses, confirms that it was a suckling animal. The δ15Ncollagen value of the scimitar-cat H. latidens is well above that obtained for the young individual of Mammuthus, which indicates that juvenile elephants were an important part of its diet. The hippo, Hippopotamus antiquus, yielded unexpectedly high δ15Ncollagen values, which suggest feeding on aquatic, non-N2-fixing plants. The high δ18Ohydroxyl values of bovids Hemitragus and Soergelia and of cervid Dama indicate that these ungulates obtained most of their water requirements from the vegetation. The megaherbivores and Eucladoceros exhibit the lowest δ18Ohydroxyl values, which suggest increased water dependence for them. Paleosynecological analysis was based on the relative abundance of species of large mammals from different ecological categories, determined by feeding behavior and locomotion types. The comparison of the frequencies of such categories in Venta Micena with those found in modern African communities indicates that the composition of the paleocommunity closely resembles those of savannas with tall grass and shrubs. The net above-ground primary productivity estimated for the on-crop biomass of the mammalian species preserved in the fossil assemblage also yields a figure congruent with that expected for an open environment.

An ecomorphological analysis of the skeletal remains of large canids, Canis (Xenocyon) falconeri and Canis etruscus (Mammalia, Carnivora, Canidae), preserved in an assemblage of large mammals from the lower Pleistocene site at Venta Micena (Guadix-Baza Basin, Orce, Granada, southeastern Spain) is reported. Mean body mass of adult individuals was estimated to be around 10 kg for C. etruscus and approximately 28 kg for C. falconeri using multiple regression.A comparative study of tooth measurements in modern canids, using principal components and discriminant function analysis, infers quite different ecomorphological adaptations and feeding behavior for both fossil species. The craniodental morphology of C. falconeri is similar to that shown by extant hypercarnivorous canids whose diet include more than 70% of vertebrate meat, whereas C. etruscus shows a cranial morphology similar to those of modern omnivorous species, thus indicating a dietary niche in which vertebrate meat represented less than 70% of its diet, with other feeding resources making up the balance. These results suggest that there was a marked ecological segregation between both sympatric species of large canids. The find of a complete skull of C. falconeri showing bilateral asymmetry and marked dental anomalies could suggest high levels of genetic homozygosis in the population which inhabited this region during early Pleistocene times, possibly as a consequence of isolation and the low number of individuals. This may have subsequently led to the extinction of C. falconeri in the Western fringe of Europe. The survival of this pathological individual to adulthood indicates that this species may have developed cooperative behavior similar to that of modern African wild dogs. OCANIS (XENOCYON) FALCONER], CANIS ETRUSCUS, ecomorphology, principal component analysis, discriminant analysis, cranial fluctuating asymmetry, Venta Micena, lower Pleistocene. Paul Palmqvist [Paul.Palmqvist@uma.es],