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The Diet of Metaschizotherium Bavaricum (Chalicotheriidae, Mammalia) from the MN 5 of Sandelzhausen (Germany) Implied by the Mesowear Method

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The Diet of Metaschizotherium Bavaricum (Chalicotheriidae, Mammalia) from the MN 5 of Sandelzhausen (Germany) Implied by the Mesowear Method View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by RERO DOC Digital Library Pala¨ontol Z (2009) 83:175–181 DOI 10.1007/s12542-009-0007-7 RESEARCH PAPER The diet of Metaschizotherium bavaricum (Chalicotheriidae, Mammalia) from the MN 5 of Sandelzhausen (Germany) implied by the mesowear method Ellen Schulz Æ Julia M. Fahlke Received: 27 February 2007 / Accepted: 7 August 2007 / Published online: 10 February 2009 Ó Springer-Verlag 2009 Abstract The genus Metaschizotherium is a Miocene MN 6 of the Franconian Alb. This is interpreted as reflecting member of the Schizotheriinae (Chalicotheriidae, Perisso- a similar dietary niche of the two species. dactyla) and appears in the MN 5 of Southern Germany with the species M. bavaricum. The Chalicotheriidae have Keywords Metaschizotherium bavaricum Á mostly been reconstructed as browsers. In this study, the Chalicotheriidae Á Sandelzhausen Á Mesowear Á mesowear method is applied to 11 upper premolars and Feeding adaptation Á Middle Miocene molars of M. bavaricum from the Upper Freshwater Molasse locality of Sandelzhausen (MN 5). With this Kurzfassung Die Gattung Metaschizotherium ist ein mio- method the amount of abrasive and attritive dental wear is za¨ner Vertreter der Schizotheriinae (Chalicotheriidae, investigated and thus it provides a time-averaged signature of Perissodactyla) und tritt in der MN 5 von Su¨ddeutschland food abrasiveness of ungulates. Principal components anal- mit der Art M. bavaricum auf. Generell werden die ysis is performed on mesowear variables of M. bavaricum. Chalicotheriidae meist als Browser rekonstruiert. In der This species is found to classify closest to extant vorliegenden Studie wird die Mesowearmethode auf 11 mixed feeding ruminants. This indicates that the diet of obere Pra¨molaren und Molaren von M. bavaricum aus M. bavaricum essentially included nonabrasive browse but Sandelzhausen (MN 5) angewandt. Mit dieser Methode wird also a certain amount of abrasive plant material. The com- der Anteil von abrasiver und attritiver Zahnabnutzung position of the diet of extant reference species indicates that untersucht und dadurch ein zeitlich gemitteltes Abra- this abrasiveness was most likely imposed by bark and whole sivita¨tssignal der der Nahrung von Ungulaten bestimmt. Mit branches. In the palaeoecological context of Sandelzhausen den Mesowearvariablen von M. bavaricum wird eine M. bavaricum occupied the dietary niche of a mixed feeder Hauptkomponentenanalyse durchgefu¨hrt. Die Art wird bei or an abrasion-dominated browser. A high degree of simi- den rezenten Mischko¨stlern nahe den rezenten Ruminantiern larity is recognised between the mesowear patterns of eingestuft. Dies zeigt eine Nahrung an, die im Wesentlichen M. bavaricum from Sandelzhausen and M. fraasi from the aus einer nichtabrasiven Blattnahrung und zusa¨tzlich einem bedeutenden Anteil von abrasivem Pflanzenmaterial bestand. Aufgrund der Zusammensetzung der Nahrung der rezenten Vergleichsarten wird geschlussfolgert, dass die Abrasivita¨tho¨chstwahrscheinlich auf Rinde und ganze A¨ ste E. Schulz (&) zuru¨ckzufu¨hren ist. Im pala¨oo¨kologischen Kontext der Biozentrum Grindel und Zoologisches Museum, Fundstelle Sandelzhausen besetzt M. bavaricum somit die Universita¨t Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany erna¨hrungso¨kologische Nische eines Mischkostfressers e-mail: [email protected] bzw. abrasionsdominierten Konzentratselektierers. Eine hohe A¨ hnlichkeit wird zwischen dem Mesowearsignal von J. M. Fahlke M. bavaricum aus Sandelzhausen und M. fraasi aus der MN Steinmann-Institut fu¨r Geologie, Mineralogie und Pala¨ontologie, Universita¨t Bonn, Nussallee 8, 53115 Bonn, Germany 6 der Fra¨nkischen Alb festgestellt. Dies wird als a¨hnliche e-mail: [email protected] Nahrungsanpassung der zwei Arten interpretiert. 123 176 E. Schulz, J. M. Fahlke Schlu¨sselwo¨rter Metaschizotherium bavaricum Á (MN 5), was first described as Chalicotherium (Fahlbusch Chalicotheriidae Á Sandelzhausen Á Mesowear Á and Gall 1970). Later, its schizotheriine characters were Nahrungsanpassung Á Mittelmioza¨n recognised by Coombs (1974) and Fahlbusch (2003). In her revised diagnosis, Coombs (2009) reintroduces the species name Metaschizotherium bavaricum Koenigswald, 1932 Introduction for the Sandelzhausen chalicothere and gives a detailed morphological description of the cranial and postcranial The Chalicotheriidae belong to the Ancylopoda within the material. For further details concerning the Chalicotherii- Perissodactyla (Hooker and Dashzeveg 2004). Ever since dae and the exact systematic position of M. bavaricum see they were first mentioned (cf. Cuvier 1823), their uncom- Coombs (2009). mon anatomy has caught the attention of biologists and The lack of modern analogue to the uniquely built palaeontologists. The Chalicotheriidae are known in the Old Chalicotheriidae has always complicated the reconstruction World and North America from the Palaeogene onwards to of their ecology. In their recent study Schulz et al. (2007) the early Pleistocene (Coombs 1989). Two subgroups, the tested the hypothesis of the browsing behaviour and shed Chalicotheriinae and the Schizotheriinae, are distinguish- some new light on the palaeoecology of this often aban- able by multiple morphological characters (e.g. Coombs doned fossil group. For this purpose the palaeodiets of 1989, 1998). In the Chalicotheriinae, the postcranial skele- different Central European chalicothere species, including ton is strongly modified. The ungual phalanges are M. bavaricum, were analysed, using the mesowear method compressed laterally. The dental characters of the Chalico- (after Fortelius and Solounias 2000), a powerful tool to theriinae are plesiomorphic, i.e. the molars are low-crowned reconstruct the lifetime feeding behaviour of ungulates. in all species of this group (Coombs 1989). In contrast, the Schulz et al. (2007) noted a substantial amount of abrasive Schizotheriinae show a trend from low-crowned to higher- plant material, like bark or whole branches, in the diet of crowned and to more elongated molars. The postcranium is the Chalicotheriidae and concluded a less abrasive diet for less derived than in the Chalicotheriinae, and a typical the Schizotheriinae compared with the Chalicotheriinae. perissodactyl proportion with approximately equally long forelimbs and hind limbs is retained (Coombs 1989). Despite these differences, all chalicotheres are generally Materials and methods reconstructed as leaf browsers, with at least the more gorilla- like Chalicotheriinae being able to erect themselves on their This study is based on two last upper premolars (P4) and nine hind limbs and protract their arms to reach into the higher upper molars (M1,M2,M3)ofM. bavaricum from the MN 5 crowns of trees. The huge claws that must have covered the of Sandelzhausen (Table 1). These teeth are also incorpo- ungual phalanges, though differing between both subfami- rated in the morphological study of this species (Coombs lies, are seen as a tool to obtain foliage by dragging or tearing 2009) and a microwear analysis (Coombs and Semprebon down branches (e.g. Abel 1922; Coombs 1983, 1989; 2005). The material is housed in the Bayerische Staats- Heissig 1999; Koenigswald 1932; Zapfe 1976, 1979). sammlung fu¨r Pala¨ontologie and Geologie in Munich (BSP). The chalicothere material from the Upper Freshwater The mesowear method was first introduced by Fortelius Molasse deposits of Sandelzhausen, lower middle Miocene and Solounias (2000) and improved by e.g. Kaiser et al. Table 1 Mesowear scorings of Specimen Tooth Side Occlusal Cusp shape Cusp shape Metaschizotherium bavaricum position relief anterior posterior from Sandelzhausen; upper molars and premolars examined BSPG 1959II11571 M2 Right High — Sharp by mesowear method BSPG 1959II11573 P4 Right High Round Sharp BSPG 1959II11612 M2 Left High Round Blunt BSPG 1959II11614 M2 or M3 Left High Sharp — BSPG 1959II11621 M2 Right High — Round BSPG 1959II11624 M2 Right High Round Round BSPG 1959II11630 M1 Right High Round Sharp BSPG 1959II2507 P4 Left High — — BSPG 1959II2507 M1 Left High Round Round BSPG 1959II2507 M2 Left High Round Round BSPG 1959II2507 M3 Left High — — 123 The diet of Metaschizotherium bavaricum (Chalicotheriidae, Mammalia) from Sandelzhausen 177 Fig. 1 Principal components analysis based on a subset of ‘‘typical’’ EI = Ammodorcas clarkei,GC= Giraffa cameloparlis,Gg= Gaz- and ‘‘no particular class’’ species of the ‘‘CONS’’ dietary classifica- ella granti,Gt= Gazella thomsoni,he= Hippotragus equinus, tion after Fortelius and Solounias 2000. The subset comprises groups hn = Hippotragus niger,ke= Kobus ellipsiprymnus,LW= Litoc- of browsing (upper case only), mixed feeding (upper and lower case) ranius walleri,Ma= Antidorcas marsupialis,Me= Aepyceros and grazing species (lower case only). meba, Metaschizotherium melampus,Oc= Ovis canadensis,OH= Odocoileus hemionus, bavaricum from Sandelzhausen (asterisk), AA = Alces alces, OJ = Okapia johnstoni,OL= Capreolus capreolus,Om= Ovibos ab = Alcelaphus buselaphus,al= Alcelaphus lichtensteini, moschatus,oo= Ourebia ourebi,OV= Odocoileus virginianus, AM = Antilocapra americana,Ap= Axis porcinus,Ax= Axis axis, rf = Redunca fulvorufula,rr= Redunca redunca,RS= Rhinoceros bb = Bison bison,BE= Tragelaphus eurycerus,Bt= Budorcas sondaicus,Ru= Rhinoceros unicornis,sc= Syncerus caffer, taxicolor,Ca= Capricornis sumatraensis,Cc= Cervus canadensis, St = Saiga tatarica,Ta= Tragelaphus angasi,Ti= Tragelaphus Cd = Cervus
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