Stable Isotope Ecology of Miocene Large Mammals from Sandelzhausen, Southern Germany
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View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Serveur académique lausannois Pala¨ontol Z (2009) 83:207–226 DOI 10.1007/s12542-009-0011-y RESEARCH PAPER Stable isotope ecology of Miocene large mammals from Sandelzhausen, southern Germany Thomas Tu¨tken Æ Torsten Vennemann Received: 28 February 2008 / Accepted: 17 July 2008 / Published online: 10 February 2009 Ó Springer-Verlag 2009 Abstract The carbon, oxygen, and strontium isotope Hyotherium soemmeringi has the highest d13C and lowest composition of enamel from teeth of large Miocene her- d18O value of all analyzed taxa, possibly related to a fru- bivorous mammals from Sandelzhausen (MN5, late Early/ givorous diet. Most taxa were water-dependent browsers early Middle Miocene) in the North Alpine foreland basin, that record meteoric water d18O values of about were analyzed to infer diet and habitat. The mean enamel -5.6 ± 0.7% Vienna Standard Mean Ocean Water d13C value of -11.4 ± 1.0% (n = 53) for the nine taxa (VSMOW). Using a modern-day mean annual air temper- 18 analyzed (including proboscideans, cervids, suids, chali- ature (MAT)–d OH2O relation a MAT of 19.3 ± 1.5°C can cotheres, equids, rhinocerotids) indicates a pure C3 plant be reconstructed for Sandelzhausen. A Gomphotherium diet for all mammals. 87Sr/86Sr ratios of *0.710 higher subtapiroideum tusk serially sampled for d18O values does than those from teeth of the western Molasse Basin (0.708– not record a clear pattern of seasonality. Thus most taxa 0.709) seem to indicate preferential feeding of the mam- were C3 browsers in a forested and humid floodplain mals in the northeastern Molasse Basin. The sympatric environment in the Molasse Basin, which experienced a herbivores have different mean d13C and d18O values warm-temperate to subtropical climate and possibly low which support diet partitioning and/or use of different seasonality. habitats within a C3 plant ecosystem. Especially the three sympatric rhinoceroses Plesiaceratherium fahlbuschi, Keywords Carbon isotopes Á Oxygen isotopes Á Lartetotherium sansaniense, and Prosantorhinus germani- Strontium isotopes Á Mean annual air temperature Á cus show clear partitioning of plants and/or habitats. The Enamel Á Diet Á Drinking water Á Molasse Basin palaeomerycid Germanomeryx fahlbuschi was a canopy folivore in moderately closed environments whereas Kurzfassung Die Kohlenstoff-, Sauerstoff- und Stron- Metaschizotherium bavaricum (Chalicotheriidae) and tium-Isotopenzusammensetzung von Zahnschmelzproben P. germanicus (Rhinocerotidae) were browsers in more mioza¨ner herbivorer Großsa¨uger aus Sandelzhausen (MN5, closed forest environments. The horse Anchitherium aur- Unter-/Mittelmioza¨n) im no¨rdlichen Alpenvorlandbecken elianense was probably a more generalized feeder than wurde analysiert, um die Erna¨hrungsweise und den assumed from its dental morphology. The forest hog Lebensraum zu rekonstruieren. Der mittlere Zahnschmelz d13C-Wert von -11,4 ± 1,0% (n = 53) fu¨r die 9 analy- sierten Taxa (u.a. Ru¨sseltiere, Hirsche, Schweine, T. Tu¨tken (&) Klauentiere, Pferde, Nasho¨rner) zeigt, dass alle Sa¨ugetiere Steinmann Institut fu¨r Geologie, Mineralogie und Pala¨ontologie, eine ausschließlich C3-Pflanzen basierte Erna¨hrungsweise Poppelsdorfer Schloss, University of Bonn, 53115 Bonn, 87 86 Germany hatten. Die Sr/ Sr-Verha¨ltnisse von *0,710, die ho¨her e-mail: [email protected] sind als von Za¨hne aus dem westlichen Molasse Becken (0,707–0,709), deuten auf eine bevorzugte Nah- T. Tu¨tken Á T. Vennemann rungsaufnahme im Bereich des nordo¨stlichen Molasse Institut de Ge´ochimie, Universite´ de Lausanne, 1015 Lausanne, Switzerland Beckens hin. Die zeitgleich vorkommenden Herbivoren 13 18 e-mail: [email protected] haben unterschiedliche d C- und d O-Werte, welche 123 208 T. Tu¨tken, T. Vennemann auf die Nutzung unterschiedlicher Nahrungsressourcen vegetation and habitat in which the particular taxa lived und/oder Lebensra¨ume innerhalb eines C3-O¨ kosystems (e.g., van der Merwe and Medina 1991; Cerling et al. 2004; hindeuten. Insbesondere die drei Nasho¨rner Plesiacer- Kohn et al. 2005). In contrast to bone, tooth enamel is atherium fahlbuschi, Lartetotherium sansaniense und suited for such geochemical reconstructions as it reliably Prosantorhinus germanicus zeigen die Nutzung verschie- reflects the original chemical and isotopic composition dener Nahrungspflanzen und/oder Habitate. Der over millions of years (Schoeninger and DeNiro 1982; Lee- Palaeomerycide Germanomeryx fahlbuschi war ein Unter- Thorp and van der Merwe 1987; Wang and Cerling 1994; holz-Blattfresser in weniger geschlossenen Habitaten Kohn et al. 1999). Most carbon isotope studies aiming to wa¨hrend Metaschizotherium bavaricum (Chalicotheriidae) reconstruct the palaeodiet and niche partitioning of mam- und Prosantorhinus germanicus (Rhinocerotidae) Laub- mals have focused on environments capable of fresser in eher geschlossenen Wa¨ldern waren. Das Pferd distinguishing C3 browsers and C4 grazers (e.g., Quade Anchitherium aurelianense war vermutlich ein mehr et al. 1992; Cerling et al. 1997b). However, recent studies generalistischer Pflanzenfresser als aufgrund seiner Zahn- have also shown enamel carbon isotopes useful in modern morphologie bisher angenommen. Das Waldschwein and ancient C3-dominated ecosystems to infer niche par- Hyotherium soemmeringi hat die ho¨chsten d13C und titioning (Quade et al. 1995; Cerling et al. 1997a, 2003b, niedrigsten d18O-Werte aller analysierten Taxa, was 2004; MacFadden and Higgins 2004; Feranec and Mac- mo¨glicherweise auf eine frugivore Erna¨hrungsweise hin- Fadden 2006; Feranec 2007; Nelson 2007; Zanazzi and deutet. Die meisten Taxa waren wasserabha¨ngige Kohn 2008). Due to climatic and environmental conditions 18 Blattfresser, die d O-Werte des meteorischen Wassers von C3 plant ecosystems prevailed in Europe during Palaeogene -5,6 ± 0,7% VSMOW aufgezeichnet haben. Unter Nut- and Neogene and no C4 grasslands developed (Blondel zung einer modernen mittleren Jahrestemperatur (MAT)- et al. 1997; Cerling et al. 1997b). Therefore C4 plants were 18 d OH2O-Relation kann eine MAT von 19,3 ± 1,5°Cfu¨r not a significant component of the flora and thus not an Sandelzhausen rekonstruiert werden. Ein seriell fu¨r important food source for herbivores. d18O-Werte beprobter Gomphotherium subtapiroideum The reconstructions of palaeodiet and palaeoenvironment Stoßzahn zeigt kein klares Saisonalita¨tsmuster. Daher for the late Early/early Middle Miocene vertebrate fossil waren die meisten Taxa C3-Blattfresser in einer bewaldeten, locality of Sandelzhausen, Bavaria, southern Germany, have feuchten U¨ berflutungsebene im Molasse Becken mit einem mainly focused on geological, palaeontological, and sedi- warmtemperierten bis subtropischen Klima mo¨glicher- mentological evidence (e.g., Fahlbusch et al. 1972, 1974; weise ohne stark ausgepra¨gte Saisonalita¨t. Fahlbusch and Liebreich 1996; Schmid 2002; Moser et al. 2009b, this issue) and dental wear (Kaiser and Ro¨ssner 2007; Schlu¨sselwo¨rter Kohlenstoffisotopie Á Schulz et al. 2007; Schulz and Fahlke 2009, this issue). In Sauerstoffisotopie Á Strontiumisotopie Á this study we attempt to infer resource use and partitioning Mittlere Jahrestemperatur Á Zahnschmelz Á Nahrung Á of taxa at the Sandelzhausen site using stable carbon, oxy- Trinkwasser Á Molasse Becken gen, and strontium isotopes of the enamel of different sympatric large herbivorous mammals such as proboscide- ans, rhinocerotids, equids, cervids, and suids. Introduction Next to palaeontological evidence such as dental features, Geological setting dental wear, stomach contents, and coprolithes, the geo- chemical evidence revealed in stable carbon and oxygen The Miocene fossil locality of Sandelzhausen, a former isotope analysis of fossil tooth enamel has become an gravel pit, is situated 70 km north of Munich in the North important tool in reconstructing palaeodiet and palaeoe- Alpine Foreland Basin (Fig. 1). The fossiliferous sediments cology (see overviews in Kohn and Cerling 2002; Koch are part of the limnofluvial Upper Freshwater Molasse. On 2007). These geochemical systems allow for inferences to the basis of biostratigraphical correlation using micro- be made on diet, feeding behavior, and resource parti- mammals Sandelzhausen is placed in the early part of the tioning among taxa as well as drinking behavior and water European Land Mammal Zone MN5 (Heissig 1997). This use efficiency of mammals within modern and ancient corresponds to an age (between 16 and 16.5 Ma) close to ecosystems (e.g., Longinelli 1984; Kohn et al. 1996, 1998, the Early/Middle Miocene boundary (Heissig 1997; Abdul- 2005; Cerling et al. 1997a, b, 2003a; Sponheimer and Lee- Aziz et al. 2008; Moser et al. 2009b, this issue). Thorp 1999a; MacFadden et al. 1999; Levin et al. 2006). The depositional environment was a low-relief, fluvial Studies have shown that mammalian skeletal remains (i.e., plain with sedimentary conditions varying between river tooth enamel) record in their isotope composition the channels, flood plains, and riparian ponds and lakes. The 123 Stable isotope ecology of Miocene large mammals 209 discovered and collected along with some plant and abundant invertebrate fossils, including 69 species of gas- tropods, bivalves, and ostracods (Witt 1998; Fahlbusch 2003; Moser et al. 2009a, this issue). Most of the fauna consists of micro- and macrovertebrates including teleost fish, amphibians, turtles, squamats, crocodilians, birds, and 66 mammal species (Fahlbusch