Ecometrics of large herbivorous land mammals in relation to climatic and environmental changes during the Pleistocene

The climatic cooling during the Cenozoic (65 Ma – present) culminated in the Pleistocene Ice Ages (ca. 2.6 Ma – 10 000 BP) during which the global climate oscillated between relatively warm climatic phases and very cold and dry glacial phases when extensive continental glaciers formed in the Northern hemisphere. The oscillation between the cold and warm climatic stages caused dramatic cyclic changes in the structure of vegetation varying at its extreme between relatively humid forests and very dry and cold mammoth steppes in Europe. These constantly changing and harsh climatic and environmental conditions caused strong extinction and evolution pressures on mammalian species. In this thesis I will discuss how two major ecometric variables, body size and diet, of large herbivorous land mammals have varied during the Pleistocene and how these patterns are connected with climate, environmental conditions and competing mammal species. Mammals diversified and started to occupy the niches of large vertebrates after the Late Cretaceous mass extinction which caused the extinction of large non-avian dinosaurs. The frequency of maximum body size in archaic mammal orders shows a significant global peak in the Middle Eocene (ca. 40 Ma) as a result of the diversification and niche filling after the Late Cretaceous mass extinction, but after that maximum size frequency in mammalian orders was low until it peaked significantly again the Pleistocene Ice Ages. This indicates that the Pleistocene climatic and environmental conditions favoured particularly large body sizes in mammals. The overall “harshness” of the Ice Age climate (seasonal, mostly cold and dry conditions and often rapid climatic changes) could have favoured large body sizes in large terrestrial mammals through mechanisms which are more complicated than the often cited benefit of large size for heat conservation (Bergmann’s rule). Large size increases the ability to survive over seasonal shortages of resources such as food and water and enables long-distance migrations to areas of better resource availability. On the other hand, strong erosional processes caused by glaciers produced fertile soils and harsh climates reduced the chemical defences of plants, which resulted in seasonally high primary production and plant quality, which would have enabled herbivorous mammals to grow into large sizes during seasons of high productivity. The main factor driving fine-scale body size variations in ungulate populations has been shown by several studies to be resource availability, which is regulated by primary productivity, plant quality, population densities of the ungulate species (intraspecific resource competition) and interspecific resource competition. The comparisons of ungulate body sizes from Middle and Late Pleistocene of Britain and Germany with vegetation openness (percentages of nonarboreal pollen from associated pollen records) show that species with different ecological strategies have different body size patterns in relation to the vegetation structure. The connection between body size patterns and ecological strategies could explain the different responses of body size to vegetation openness. Species which tend to have relatively small group sizes (e.g. deer) show on average larger body sizes in environments where the vegetation structure is open, whereas gregarious, open 4 Department of Geosciences anD GeoGraphy a adapted species (e.g. horses) tend to have smaller average body sizes in open habitats. I suggest this is because open habitats favour large body size in ecologically flexible species with small group sizes due to high resource availability and quality per an individual (relatively low population densities), less size-restricted manoeuvrability and enhanced capability to escape predators, whereas resource limitations for each individual caused by high population densities can become a limiting factor for individual body size in openadapted, gregarious species which are efficient open-vegetation feeders and form large groups in open habitats. In closed environments, the body size of the open-adapted, gregarious species is not limited by high population density which enables them to attain larger individual sizes. Dietary signals of the key ungulate species in Middle and Late Pleistocene Europe based on mesowear analyses are on average significantly positively correlated with vegetation openness (non-arboreal pollen percentages) at localitylevel. However, there are significant interspecific differences. While most of the species show positive correlations between their mesowear signal and non-arboreal vegetation, others, especially the red deer (Cervus elaphus), do not show any correlation. Instead, the mesowear signal of the red deer is significantly more abrasive dominated when other browsedominated feeders, especially the roe deer (Capreolus capreolus) are present. This indicates that interspecific competition can obscure the effect of available plant material in the diet of ecologically flexible species. This should be taken into account when interpreting the feeding ecology of the key species in palaeocommunities, and especially when attempting to reconstruct palaeoenvironmental conditions from dietary proxies of mammals. Such attempts should ideally be based on as complete dietary analyses of fossil herbivore faunas as possible. In order to extend the palaeodietary and palaeoecological analyses based on mesowear signals of herbivorous mammas, a new tooth wear -based dietary analysis method was developed for elephants and other lamellar toothed proboscideans, based on measuring occlusal relief of their molar teeth as angles. The benefits of that approach compared with other available methods are that it is easy-to-do, fast and robust, and it gives consistent and comparable results for species with different dental morphologies. The preliminary results from that study indicate that the angle measurement method is a powerful tool for reconstructing proboscidean diets from the fossil record.