Paleoecological reconstruction of a lower Pleistocene large mammal community using biogeochemical ({delta}13C, {delta}15N, {delta}18O, Sr:Zn) and ecomorphological approaches

摘要

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 13C_collagen values for ungulate species indicates that C₃ plants were dominant in the diet of these mammals. 13C_collagen 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 13C_collagen 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 15N_collagen content (canopy effect). Bovid species (Bovini aff. Leptobos, Soergelia minor, and Hemitragus albus) presumably inhabited open environments, according to their comparatively high hypsodonty and 15N_collagen values. Carnivore species (Homotherium latidens, Megantereon whitei, Pachycrocuta brevirostris, Canis falconeri, and Canis etruscus) exhibit higher 15N_collagen 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 15N_collagen value for a sample of Mammuthus meridionalis, which came from an individual with unfused epiphyses, confirms that it was a suckling animal. The 15N_collagen 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 15N_collagen values, which suggest feeding on aquatic, non-N₂-fixing plants. The high 18O_hydroxyl 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 18O_hydroxyl 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.