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

¹ Paul Palmqvist. Departamento de Geología y Ecología (Área de Paleontología), Facultad de Ciencias, Campus Universitario de Teatinos, 29071 Málaga, Spain. Paul.Palmqvist{at}uma.es
² Darren R. Gröcke. Department of Geology, Royal Holloway, University of London. Egham Hill, Egham, Surrey TW20 0EX, United Kingdom. d.grocke{at}gl.rhul.ac.uk
³ Alfonso Arribas. Museo Geominero, Instituto Geológico y Minero de España (IGME), Ríos Rosas, 23, 28003 Madrid, Spain. a.arribas{at}igme.es
⁴ Richard A. Fariña. Departamento de Paleontología, Facultad de Ciencias, Universidad de la República, Iguá, 4225, 11400 Montevideo, Uruguay. fari~a{at}fcien.edu.uy

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.

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