Alfonso Arribas

Paleoecological reconstruction of a lower Pleistocene large mammal community using biogeochemical (δ13C, δ15N, δ18O, Sr:Zn) and ecomorphological approaches

Abstract 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.

Taphonomy and palaeoecology of an assemblage of large mammals : hyaenid activity in the lower Pleistocene site at Venta Micena (Orce, Guadix-Baza Basin, Granada, Spain)

Abstract We report taphonomic and palaeoecologic data on the rich, diverse and well preserved assemblage of large mammals from lower Pleistocene deposits at Venta Micena (Orce, Granada, south-east Spain). The biostratinomic and diagenetic characteristics of the assemblage are congruous with the sedimentary context deduced from the study of the site, and both confirm that: (i) the assemblage represents an accumulated taphonomic stage, (ii) it was formed by demic, autochtonous palaeobiologic entities, which were preserved and recorded in situ, and (iii) it is the result of biological processes and agents. Interspecific analysis of size/abundance patterns in ungulates shows that the main taphonomic bias affecting the bones was produced by biological destruction before burial, and that the loss of information was greater for species of smaller body size. Factor correspondence analysis was used to compare the frequencies at which some groups of postcranial elements are represented in several recent and archaeological bone assemblages accumulated by carnivores, rodents and hominids. The results obtained strongly suggest that the bones from Venta Micena were collected mainly by hyaenids, which deposited them near shallow dens excavated around the ponds that surrounded the Pleistocene lake of Orce. An analysis of the abundance of major long bones has shown that differential fragmentation was produced by hyaenas as a function of their structural density and mean marrow content. All these data allow to formulate a descriptive-quantitative model for the characterization of bone assemblages generated from hyaenid activity, in which Venta Micena is an example of bone concentration and modification activities by Pachycrocuta brevirostris . Strong selection of prey by carnivores (which killed preferably juveniles, females and individuals with diminished locomotive capabilities among ungulate prey species of larger body size) is indicated by (i) the abundance of juvenile individuals with deciduous teeth in relation to the average weight estimated for adults in each ungulate species, by (ii) the U-shaped attritional mortality profiles deduced from crown heigth measurements, by (iii) the presence of many metapodials with different osteopathologies, and by (iv) a biased sexual ratio deduced from the metacarpals of large bovids. Comparison between the frequencies in which modern African carnivores kill and scavenge ungulates from different size classes and the abundance of these size categories in the assemblage suggests that the Venta Micena hyaena was a bone-cracking scavenger which fed largely on carcasses of ungulates preyed upon and partially consumed by flesh-eating carnivores such as saber-toothed felids and wild dogs.

Prey selection by terrestrial carnivores in a lower Pleistocene paleocommunity

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 ).

Taphonomic decoding of the paleobiological information locked in a lower Pleistocene assemblage of large mammals

Abstract The processes of fossilization have usually been perceived by paleontologists as destructive ones, leading to consecutive (and in most cases irretrievable) losses of paleobiological information. However, recent developments of conceptual issues and methodological approaches have revealed that the decrease in paleobiological information runs parallel to the gain of taphonomic information. This taphonomic imprinting often makes it possible to decode the fraction of paleobiological information that was lost during fossilization, and may also contribute new data for deciphering paleobiological information that was not originally preserved in the assemblage, such as paleoethology. A good example is the study of the macrovertebrate assemblage from the lower Pleistocene site at Venta Micena (Orce, southeastern Spain). Taphonomic analysis showed that the giant, short-faced hyenas (Pachycrocuta brevirostris) selectively transported ungulate carcasses and body parts to their maternity dens as a function of the mass of the ungulates scavenged. The fracturing of major limb bones in the dens was also highly selective, correlating with marrow content and mineral density. Important differences in bone-cracking intensity were related to which species the bones came from, which in turn biased the composition of the bone assemblage. The analysis of mortality patterns deduced for ungulate species from juvenile/adult proportions revealed that most skeletal remains were scavenged by the hyenas from carcasses of animals hunted by hypercarnivores, such as saber-tooths and wild dogs. Analytical study of the Venta Micena assemblage has unlocked paleobiological information that was lost during its taphonomic history, and has even provided paleobiological information that was not preserved in the original bone assemblage, such as the paleoethology of P. brevirostris, which differed substantially from modern hyenas in being a strict scavenger of the prey hunted by other carnivores.

A Mammalian Lost World in Southwest Europe during the Late Pliocene

Background Over the last decades, there has been an increasing interest on the chronology, distribution and mammal taxonomy (including hominins) related with the faunal turnovers that took place around the Pliocene-Pleistocene transition [ca. 1.8 mega-annum (Ma)] in Europe. However, these turnovers are not fully understood due to: the precarious nature of the periods fossil record; the “non-coexistence” in this record of many of the species involved; and the enormous geographical area encompassed. This palaeontological information gap can now be in part bridged with data from the Fonelas P-1 site (Granada, Spain), whose faunal composition and late Upper Pliocene date shed light on some of the problems concerning the timing and geography of the dispersals. Methodology/Principal Findings This rich fossil site yielded 32 species of mammals, among which autochthonous species of the European Upper Villafranchian coexist with canids (Canis), ovibovines (Praeovibos) and giraffids (Mitilanotherium) from Asia. Typical African species, such as the brown hyena (Hyaena brunnea) and the bush pig (Potamochoerus) are also present. Conclusions/Significance This assemblage is taxonomically and palaeobiogeographically unique, and suggests that fewer dispersal events than was previously thought (possibly only one close to 2.0 Ma) are responsible for the changes seen around 1.9–1.7 Ma ago in the fauna of the two continents.

The last Iberian gomphothere (Mammalia, Proboscidea): Anancus arvernensis mencalensis nov. ssp. from the earliest Pleistocene of the Guadix Basin (Granada, Spain)

This work describes a new finding of Anancus arvernensis ̶ a maxilla fragment that preserves M2 and M3 ̶ from the earliest Pleistocene (c.a. 2.5-2.4 Ma) at the Fonelas SCC-3 site (Cuenca de Guadix, Granada, Spain). This fossil is attributed to a new chronosubspecies based on the combination of anatomical features shown by M3: a primitive anatomical pattern plus derived features. The primitive features include the hexalophodont condition, a massive, rectangular distal outline, inconspicuous enamel folding, and indiscernible anancoidy. The derived features ̶ tooth-valleys covered by cement and the small overall size of the tooth ̶ are typical of the last representatives of the lineage. This mosaic of features allows a new chronosubspecies to be proposed: Anancus arvernensis mencalensis nov. ssp. This would be the youngest representative of the genus Anancus known for the Iberian Peninsula (MNQ 17a), and represents an intermediate evolutionary stage between Anancus arvernensis arvernensis and Anancus arvernensis chilhiacensis, the last known representative of the European lineage. A. arvernensis mencalensis therefore forms part of a temporal cline in the configuration of M3 over the Plio-Pleistocene transition. These anatomical changes could have occurred as a response to the aridification that began around 2.5 Ma, which led to changes in the composition of plant communities. Guiomar Garrido. Estación Paleontológica Valle del Río Fardes, Instituto Geológico y Minero de España (IGME), Ríos Rosas 23, 28003 Madrid, Spain g.garrido@igme.es Alfonso Arribas. Estación Paleontológica Valle del Río Fardes, Instituto Geológico y Minero de España (IGME), Ríos Rosas 23, 28003 Madrid, Spain a.arribas@igme.es