Generalism as a subsistence strategy: advantages and limitations of the highly flexible feeding traits of Pleistocene Stephanorhinus hundsheimensis (Rhinocerotidae,Mammalia)

The so-called Hundsheim rhinoceros, Stephanorhinus hundsheimensis, was a very common faunal element of the Early to early Middle Pleistocene period in the western Palaearctic. In this study, individuals from two different central European populations of the Hundsheim rhinoceros were analysed in order to determine whether their local dietary signals could reflect differing food availability between the two populations, and whether such information could provide a better understanding of the ecological role of S. hundsheimensis within corresponding faunal assemblages, and of its principal subsistence strategy in the western Palaearctic. The mesowear traits observed in the studied S. hundsheimensis populations have been interpreted as representing biome-specific signals, indicating grassland vegetation at the site of Süßenborn, and dense to open forests at Voigtstedt (both localities in Germany). The analyses performed on the fossil rhino material demonstrate the most pronounced dietary variability ever established for a single herbivorous ungulate species by mesowear studies. This variability ranges from an attrition dominated grazing regime, to a one of predominantly browsing, and characterises S. hundsheimensis as the most ecologically tolerant rhinoceros of the Palaearctic Plio-Pleistocene. Although such dietary flexibility proved an effective enough subsistence strategy over a period of 600–900 ka (1.4/1.2–0.6/0.5 Myr) in the western Palaearctic, the situation changed dramatically after 0.6 Myr BP, when the new species of rhinoceroses, Stephanorhinus hemitoechus and Stephanorhinus kirchbergensis, appeared and started to compete for both the grass and the browse. For the generalist S. hundsheimensis, this bilateral interference was detrimental to its success in all of its habitats. The successful competition of specialised forms of rhinoceroses, which might have originated as a result of the development of 100 ka periodicity in the global climatic record, is proposed as the main reason for the extinction of S. hundsheimensis during the early Middle Pleistocene.     

Western Palaearctic palaeoenvironmental conditions during the Early and early Middle Pleistocene inferred from large mammal communities,and implications for hominin dispersal in Europe

Large-scale fluctuations in global climate and resulting changes in ecology had a profound effect on human evolution and dispersal. Though hominin remains are scarce, studies focussing on the more abundant records of fossil land mammal communities can contribute greatly to our knowledge of the palaeoenvironmental circumstances that influenced and directed the global spread of hominins. To produce a comprehensive and accurate account of the evolution of western Palaearctic habitat diversity between 2.6 and 0.4 Ma BP, information generated from large mammal communities from 221 key sites has been included in this study.The palaeoecological conditions of the western Palaearctic during the Early and early Middle Pleistocene were principally controlled by the following key factors: (1) a widespread trend of temperature decrease, (2) the periodicity of the global temperature record, (3) the intensity of single climatic stages, (4) the temporal pattern of climatic variation, (5) geographical position, and (6) the distribution of continental water resources. A general picture of the evolution of western Palaearctic habitat diversity saw the replacement of extensive forested terrain by an alternating sequence of varied savannah-like and forested habitats during the 2.6–1.8 Ma span, as well as an alternation between different types of predominantly open habitats between 1.8 and 1.2 Ma. Both of these processes were governed by 41 ka temperature periodicity. During the 1.2–0.9 Ma time span, irregular climatic fluctuations were more common and habitat variability increased. The subsequent 0.9–0.4 Ma interval, a period controlled by 100 ka periodicity, was by comparison more stable, with longer climatic cycles alternating between open and forested landscapes. During the entire Early and early Middle Pleistocene, assemblages of large mammal communities reveal a distinct trend of decreasing continentality between Eastern and South-Eastern Europe on the one hand, and South-Western and North-Western Europe on the other. This trend was due to the effect of the Atlantic Ocean, while in Southern Europe the relatively low continentality was balanced by influences from the Mediterranean Sea.When plotted against evidence of hominin occurrence, the data on western Palaearctic habitat diversity inferred from large mammal communities indicate clear environmental stimuli for the earliest human dispersal in Europe. These are: (1) a wide range of habitats, implying a high diversity of resources; (2) mild climates with low seasonality, implying a lack of strong environmental fluctuations. Around 1.8 Ma at the latest, hominins of African origin entered the western Palaearctic for the first time, taking advantage of the diversity of habitats and resources, particularly along large river systems. Their subsequent westward spread between 1.7 and 1.3 Ma was restricted to Mediterranean-influenced areas, which offered a high variability of habitats and relatively low seasonality. The increase in environmental diversity, which occurred from 1.2 Ma onwards, opened up South-Eastern and Eastern Europe for hominin occupation. According to the available records, North-Western and Central Europe were initially colonized during late Early to early Middle Pleistocene interglacials, when these regions experienced periods of low seasonality and considerable habitat diversity.