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Patterns of Diet and Body Mass of Large Ungulates from the Pleistocene of Western Europe, and Their Relation to Vegetation

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Patterns of Diet and Body Mass of Large Ungulates from the Pleistocene of Western Europe, and Their Relation to Vegetation Palaeontologia Electronica palaeo-electronica.org Patterns of diet and body mass of large ungulates from the Pleistocene of Western Europe, and their relation to vegetation Juha Saarinen, Jussi Eronen, Mikael Fortelius, Heikki Seppä, and Adrian M. Lister ABSTRACT Ungulate diets may vary following differences in vegetation, and their body size is affected by a complex set of ecological and physiological variables. Here we analyse Middle and Late Pleistocene British and German ungulate palaeocommunities to test whether there are significant correlations of diet and body size of ungulate species with vegetation openness. We also evaluate the role of interspecific interactions on the diet and body mass of the ungulate species. We use mesowear for dietary analyses and regression equations for estimating body mass from skeletal measures. The results show a correlation between ungulate mesowear and non-arboreal pollen percentages of the localities, but there are marked differences between species. Body masses of rhinoceroses (Rhinocerotidae) and deer (Cervidae) are on average higher in open environments, whereas aurochs (Bos primigenius) does not show clear connection of body size with vegetational conditions, and bison (Bison spp.) and wild horses (Equus ferus) have on average smaller mean size in more open ecosystems, possibly because of high population densities and resulting resource limitations. It is evident that the correlation of body size and vegetation openness is not straightforward and is likely to reflect the varying effects of population density, ecological adaptations and environmental conditions on body size in different species. Juha Saarinen. Department of Geosciences and Geography, University of Helsinki, P.O. Box 64, Gustaf Hällströmin katu 2a, 00014 Helsinki, Finland. [email protected] and Department of Earth Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK. Jussi Eronen. Department of Geosciences and Geography, University of Helsinki, P.O. Box 64, Gustaf Hällströmin katu 2a, 00014 Helsinki, Finland. [email protected] Mikael Fortelius. Department of Geosciences and Geography, University of Helsinki, P.O. Box 64, Gustaf Hällströmin katu 2a, 00014 Helsinki, Finland. [email protected] Heikki Seppä. Department of Geosciences and Geography, University of Helsinki, P.O. Box 64, Gustaf Hällströmin katu 2a, 00014 Helsinki, Finland. [email protected] Adrian M. Lister. Department of Earth Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK. [email protected] Keywords: mesowear; body mass; ungulate communities; vegetation openness; Pleistocene Submission: 25 November 2013 Acceptance: 20 July 2016 Saarinen, Juha, Eronen, Jussi, Fortelius, Mikael, Seppä, Heikki, and Lister, Adrian M. 2016. Patterns of diet and body mass of large ungulates from the Pleistocene of Western Europe, and their relation to vegetation. Palaeontologia Electronica 19.3.32A: 1-58 palaeo-electronica.org/content/2016/1567-pleistocene-mammal-ecometrics Copyright: Palaeontological Association September 2016 SAARINEN ET AL.: PLEISTOCENE MAMMAL ECOMETRICS INTRODUCTION mesophytic forest, which is supported by the over- all composition of the mammal fauna, whereas The composition of living herbivorous mam- Eppelsheim would likely have included at least mal communities generally reflects vegetation pat- seasonally abundant grassy vegetation, which is terns: grazing and grass-dominated mixed-feeding supported by palaeobotanic finds (Kaiser, 2004). species are more abundant in grassland environ- Another example is the study by Rivals et al. ments than in forest environments, which are char- (2015a) showing intraspecific variation in the diets acterised by browsers and browse-dominated of proboscidean species Anancus arvernensis, mixed-feeders (Gordon and Prins, 2008). However, Mammuthus rumanus and M. meridionalis diets within individual ungulate species can vary between Early Pleistocene localities in Europe, considerably in open versus closed environments based on tooth microwear analysis. and in different ungulate assemblages following It can be hypothesised that differences in the the availability of resources, as well as resource diets of local fossil populations of a species reflect competition with, and resource facilitation by, other differences in local vegetation, and probably also ungulate species (e.g., Langvatn and Albon, 1986; the presence of competitors. We have included all Hobbs et al., 1996; Latham, 1999; Arsenault and the most common species of large ungulates from Owen-Smith, 2002; Stewart et al., 2002).Variation several British and German Middle and Late Pleis- of body size in herbivorous mammal species is tocene localities (see Table 1) to test the correla- also probably influenced, at least in part, by open tion of their dietary signals with environmental versus closed habitats and by the quality and proxy data obtained from pollen records. Non-arbo- abundance of vegetation, in ways that may vary real pollen percentages (NAP %) can be used as with species’ adaptations (Jarman, 1974; Langvatn rough estimates of vegetation openness (Sven- and Albon, 1986; Guthrie, 1990; Bro-Jørgensen, ning, 2002), although they do not allow precise 2008). In this study we analyse local patterns of quantitative estimates of the extent of open land- diet of some of the most abundant ungulate spe- scape, or productivity (Sugita et al., 1999). It can cies from Middle and Late Pleistocene mammal be predicted that the broad dietary composition of localities of Western and Central Europe in order to the ungulate community is correlated with the explore their relationship with vegetation recon- degree of openness of vegetation, although it is structions. We also examine whether body size is likely that some species are so specialized in utiliz- correlated with vegetation type and diet. Moreover, ing certain kinds of plant material that their diet we compare the patterns of diet and body size does not vary greatly across localities with different across ungulate species to evaluate the role of diff- plant communities, and they would be absent if ent ecological strategies and potential resource their preferred food were missing. competition between the species. Correlations of ungulate body mass with envi- Several recent studies have demonstrated ronmental characteristics can also be predicted but dietary variation in fossil ungulate species from they are likely to be complex. Body size is affected localities with different environmental conditions. by a variety of physiological and ecological varia- For example, Kahlke and Kaiser (2011) used bles, and it may be difficult to separate their effects mesowear analysis on the Middle Pleistocene rhi- on the body size of individual species (e.g., Jar- noceros Stephanorhinus hundsheimensis in Ger- man, 1974; Peters, 1983; West et al., 1997; Gillo- many. The population from the cool-climate of oly et al., 2002; Haskell et al., 2002; Damuth, Süssenborn has, on average, a mesowear signal 2007). Body size has been postulated to relate to indicating a more abrasive diet than that of the temperature, via its effects on heat conservation interglacial of Voigtstedt. Kaiser (2004) did a similar (e.g., Bergmann, 1847; Peters, 1983), but as a analysis for the Late Miocene populations of the general explanation of mammalian size variation equid Hippotherium primigenium from Höwenegg this idea has been strongly criticized (Geist, 1987; and Eppelsheim, Germany. He noted that the pop- Lister, 1992). For herbivorous mammals, more ulation from Höwenegg had a mesowear signal likely determinants include habitat structure, social indicating browsing diet roughly similar to that of structure (e.g., Jarman, 1974), and the availability the modern Sumatran rhinoceros (Dicerorhinus and abundance of their optimal food (e.g., Lang- sumatrensis), whereas the population from vatn and Albon, 1986). In particular, large body Eppelsheim had a dietary signal resembling that of size could be associated with open environments the common waterbuck (Kobus ellipsiprymnus), for a number of reasons. First, because of mostly which is a grazer. Palaeoenvironmental data indi- seasonally harsh climates but long sunshine hours cate that Höwenegg was probably a subtropical 2 PALAEO-ELECTRONICA.ORG and fertile soils created by glacial erosion, the open Torquay Museum (TM), the Sedgwick Museum of environments of glacial Europe comprised season- Geology, Cambridge (SMC), University Museum of ally highly productive plant communities with low Zoology, Cambridge (UMZC), Ipswich Museum chemical defences, thus providing abundant and (IM), British Geological Survey at Keyworth (BGS), good-quality sources of food for herbivores Staatliches Museum für Naturkunde, Stuttgart (Guthrie, 1990; Zimov et al., 1995; Geist, 1998). (SMNS), Staatliches Museum für Naturkunde, Karl- Second, large body size enables the processing of sruhe (KMNS) and the Quaternary Palaeontology large quantities of this often digestively challenging Research Station of the Senckenberg institute, plant food, and also increases resistance to sea- Weimar (IQP). sonal shortages of food and water (Peters, 1983; To ensure adequate sample sizes, only locali- Lindstedt and Boyce, 1985; Clauss et al., 2003) ties rich in Middle or Late Pleistocene large mam- during the harsh winters of glacial episodes. Third, mal remains were chosen. Even so, not all species large size may help in predator avoidance because are represented by statistically significant
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