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Ecometrics of Large Herbivorous Land Mammals in Relation to Climatic and Environmental Changes During the Pleistocene

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Ecometrics of Large Herbivorous Land Mammals in Relation to Climatic and Environmental Changes During the Pleistocene View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Helsingin yliopiston digitaalinen arkisto Ecometrics of large herbivorous land mammals in relation to climatic and environmental changes during the Pleistocene JUHA SAARINEN ACADEMIC DISSERTATION To be presented, with the permission of the Faculty of Sciences of the University of Helsinki, for public examination in auditorium XII, University main building, on 16th December 2014, at 12 noon. DEPARTMENT OF GEOSCIENCES AND GEOGRAPHY A28 / HELSINKI 2014 © Juha Saarinen (synopsis and Paper III) © AAAS Science (Paper I) © Royal Society Publishing (Paper II) © BioOne (Paper IV) Cover illustration: Juha Saarinen Author´s address: Juha Saarinen Department of Geosciences and Geography P.O.Box 64, 00014 University of Helsinki, Finland [email protected] Supervised by: Professor Mikael Fortelius Department of Geosciences and Geography University of Helsinki, Finland Professor Heikki Seppä Department of Geosciences and Geography University of Helsinki, Finland Docent Jussi Eronen Department of Geosciences and Geography University of Helsinki, Finland Reviewed by: Professor P. David Polly Department of Geological Sciences Indiana University, USA Docent Suvi Viranta-Kovanen Institute of Biomedicine University of Helsinki, Finland Opponent: Professor Danielle Schreve Department of Geography Royal Holloway University of London, Great Britain Department of Geosciences and Geography A ISSN 1798-7911 ISBN 978-952-10-9467-5 (paperback) ISBN 978-952-10-9468-2 (PDF) http://ethesis.helsinki.fi Unigrafia Helsinki 2014 Saarinen J., 2014. Ecometrics of large herbivorous land mammals in relation to climatic and environmental changes during the Pleistocene. Unigrafia. Helsinki. 42 pages and 4 figures. Abstract The climatic cooling during the Cenozoic (65 dry conditions and often rapid climatic changes) Ma – present) culminated in the Pleistocene Ice could have favoured large body sizes in large Ages (ca. 2.6 Ma – 10 000 BP) during which terrestrial mammals through mechanisms which the global climate oscillated between relatively are more complicated than the often cited benefit warm climatic phases and very cold and dry of large size for heat conservation (Bergmann’s glacial phases when extensive continental rule). Large size increases the ability to survive glaciers formed in the Northern hemisphere. over seasonal shortages of resources such as food The oscillation between the cold and warm and water and enables long-distance migrations climatic stages caused dramatic cyclic changes to areas of better resource availability. On the in the structure of vegetation varying at its other hand, strong erosional processes caused by extreme between relatively humid forests and glaciers produced fertile soils and harsh climates very dry and cold mammoth steppes in Europe. reduced the chemical defences of plants, which These constantly changing and harsh climatic resulted in seasonally high primary production and environmental conditions caused strong and plant quality, which would have enabled extinction and evolution pressures on mammalian herbivorous mammals to grow into large sizes species. In this thesis I will discuss how two during seasons of high productivity. major ecometric variables, body size and diet, The main factor driving fine-scale body size of large herbivorous land mammals have varied variations in ungulate populations has been shown during the Pleistocene and how these patterns by several studies to be resource availability, are connected with climate, environmental which is regulated by primary productivity, conditions and competing mammal species. plant quality, population densities of the ungulate Mammals diversified and started to occupy species (intraspecific resource competition) the niches of large vertebrates after the Late and interspecific resource competition. The Cretaceous mass extinction which caused the comparisons of ungulate body sizes from Middle extinction of large non-avian dinosaurs. The and Late Pleistocene of Britain and Germany frequency of maximum body size in archaic with vegetation openness (percentages of non- mammal orders shows a significant global arboreal pollen from associated pollen records) peak in the Middle Eocene (ca. 40 Ma) as a show that species with different ecological result of the diversification and niche filling after strategies have different body size patterns the Late Cretaceous mass extinction, but after in relation to the vegetation structure. The that maximum size frequency in mammalian connection between body size patterns and orders was low until it peaked significantly ecological strategies could explain the different again the Pleistocene Ice Ages. This indicates responses of body size to vegetation openness. that the Pleistocene climatic and environmental Species which tend to have relatively small conditions favoured particularly large body group sizes (e.g. deer) show on average larger sizes in mammals. The overall “harshness” of body sizes in environments where the vegetation the Ice Age climate (seasonal, mostly cold and structure is open, whereas gregarious, open DEPARTMENT OF GEOSCIENCES AND GEOGRAPHY A adapted species (e.g. horses) tend to have smaller In order to extend the palaeodietary and average body sizes in open habitats. I suggest this palaeoecological analyses based on mesowear is because open habitats favour large body size signals of herbivorous mammas, a new tooth wear in ecologically flexible species with small group -based dietary analysis method was developed sizes due to high resource availability and quality for elephants and other lamellar toothed per an individual (relatively low population proboscideans, based on measuring occlusal densities), less size-restricted manoeuvrability relief of their molar teeth as angles. The benefits and enhanced capability to escape predators, of that approach compared with other available whereas resource limitations for each individual methods are that it is easy-to-do, fast and robust, caused by high population densities can become and it gives consistent and comparable results for a limiting factor for individual body size in open- species with different dental morphologies. The adapted, gregarious species which are efficient preliminary results from that study indicate that open-vegetation feeders and form large groups in the angle measurement method is a powerful open habitats. In closed environments, the body tool for reconstructing proboscidean diets from size of the open-adapted, gregarious species is the fossil record. 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 locality- level. 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 browse- dominated 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. 4 Tiivistelmä (in Finnish) Sama ilmiö on nähtävissä maksimikoon esiintymisfrekvenssissä hyvin erilaisissa nisäkäslahkoissa ja maailmanlaajuisesti. Erityisen Kenotsooisella ajalla (65 miljoonaa vuotta voimakas maksimiruumiinkoon esiintymisen sitten – nykyaika) maapallon ilmasto on lämpimän huippuvaihe on pleistoseenin jääkausiajalla, alkuvaiheensa jälkeen muuttunut enimmäkseen mikä viittaa siihen että ilmaston kylmeneminen kylmemmäksi, keskileveyksillä kuivemmaksi ja siihen liittyvät ympäristönmuutokset suosivat ja kausittaisemmaksi. Tämä ilmaston erityisen suuren ruumiin koon kehittymistä pitkäaikainen kehitys huipentui pleistoseeniin eli useimmissa nisäkäslahkoissa. Tämän ilmiön jääkausiaikaan, joka vallitsi noin 2,6 miljoonaa aiheuttivat todennäköisesti monet ilmaston – 10 000 vuotta sitten. Pleistoseeniajalla ilmasto ja ympäristöjen muutoksiin liittyvät tekijät, vaihteli voimakkaasti ja sen seurauksena jotka ovat monimutkaisempia kuin usein pohjoisella pallonpuoliskolla esiintyi vuorotellen esitetty suuren koon hyöty ruumiinlämmön melko lämpimiä ilmastovaiheita joiden aikana ylläpitämiseksi kylmässä ilmastossa (niin metsät olivat hallitsevia elinympäristöjä ja kutsuttu Bergmannin sääntö). Suuri koko auttaa kylmiä glasiaalivaiheita, jolloin kylmän ja nisäkkäitä selviämään kausittaisesta ravinnon kuivan ilmaston seurauksena syntynyt aro- ja niukkuudesta ja mahdollistaa tehokkaan tundrakasvillisuuden vyöhyke, niin kutsuttu vaeltamisen ravintokohteita seuraten. Toisaalta mammuttiaro, levittäytyi laajalle alueelle jäätiköiden aiheuttama voimakas eroosio tuotti pohjoisessa Euraasiassa. Eläimistön piti sopeutua ravintorikkaita maannoksia mahdollistaen näihin voimakkaisiin ilmaston ja elinympäristöjen
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