Palaeobio Palaeoenv (2014) 94:71–82 DOI 10.1007/s12549-013-0138-4 ORIGINAL PAPER Perissodactyla from the late Middle Miocene Gratkorn locality (Austria) Manuela Aiglstorfer & Kurt Heissig & Madelaine Böhme Received: 30 August 2013 /Revised: 5 November 2013 /Accepted: 20 November 2013 /Published online: 24 January 2014 # Senckenberg Gesellschaft für Naturforschung and Springer-Verlag Berlin Heidelberg 2014 Abstract Although quite rare in comparison to other large 12.2–12.0 Ma; Gross et al. 2011) comprises abundant small mammal groups, the Perissodactyla from Gratkorn show a and large vertebrate fossils and is one of very few qualitatively diverse assemblage. Besides the three rhinocerotid species, and quantitatively rich vertebrate localities of this time period Aceratherium sp., Brachypotherium brachypus (Lartet, of the Paratethys realm. While artiodactyls are abundant with- 1837), and Lartetotherium sansaniense (Lartet, in Laurillard in the large mammals from Gratkorn, perissodactyl remains 1848), the families Chalicotheriidae and Equidae are repre- are rare and comprise only some isolated teeth and some sented by Chalicotherium goldfussi Kaup, 1833 and fragmented bones. Taxonomic determination is therefore Anchitherium sp., respectively. The perissodactyl assemblage limited for the material and cannot supply much infor- fits well in a late Middle Miocene (Sarmatian) riparian wood- mation on general taxonomic and phylogenetic ques- land with diverse habitats from active rivers to drier more tions. For palaeoenvironmental considerations on the open environments, as were present at the Gratkorn locality. Gratkorn locality, the perissodactyls are essential ele- ments. In contrast to many other, though richer, locali- Keywords Chalicotherium . Aceratherium . ties, they can furthermore provide confidently dated Brachypotherium . Lartetotherium . Anchitherium . material for stratigraphic range estimations. late Middle Miocene . Sarmatian . Central Europe Materials and methods Introduction Rhinocerotidae vel Chalicotheriidae: UMJGP 204701 (distal The Gratkorn locality (clay pit St. Stefan) is located 10 km fragment of humerus sin.?), UMJGP 204719 (petrosum) NNW of Graz (Styria, Austria). The fossil-bearing palaeosol Rhinocerotidae indet: UMJGP 203705 (distal fragment of of late Middle Miocene age (late Sarmatian sensu stricto; tibia dex.?), GPIT/MA/2400 (costa sin.?; proximal fragment) Chalicotherium goldfussi: UMJGP 204676 (M3 dex.) Aceratherium sp.: UMJGP 203711 (fragment of D2 sin.) This article is a contribution to the special issue “The Sarmatian vertebrate Brachypotherium brachypus: UMJGP 203434 (lateral half of locality Gratkorn, Styrian Basin.” astragalus sin.), UMJGP 204720 (Mt II sin.) M. Aiglstorfer (*) : M. Böhme Lartetotherium sansaniense: UMJGP 203459 (m1 sin.; frag- Fachbereich Geowissenschaften, Eberhard Karls Universität ment of m2 sin.) Tübingen, Sigwartstraße 10, 72076 Tübingen, Germany Anchitherium sp.: UMJGP 204694 (distal articulation of hu- e-mail: [email protected] merus sin.), UMJGP 203422 (distal articulation of radius dex.) M. Aiglstorfer : M. Böhme Terminology for dental material of Chalicotheriidae fol- Senckenberg Center for Human Evolution and Palaeoenvironment lows Fahlke et al. (2013; except for the term ectoloph, (HEP), Sigwartstraße 10, 72076 Tübingen, Germany which is understood sensu Zapfe 1979; postfossette is understood sensu Butler 1965; and labial is used instead of K. Heissig SNSB – Bayerische Staatssammlung für Paläontologie und buccal). For Rhinocerotidae, it is modified after Heissig Geologie, Richard-Wagner-Str. 10, 80333 München, Germany (1969, 1972), and Heissig and Fejfar (2007). For postcranial 72 Palaeobio Palaeoenv (2014) 94:71–82 elements, current anatomical terms are used. Measure- comparison to the Gratkorn specimen, the concavity in the distal ments are taken with a digital calliper (where possible trochlea is even stronger in the two rhinocerotid humeri (per- with a precision of 0.1 mm) in the way indicated on sonal observation). As no humerus of Chalicotherium goldfussi Figs. 2 and 3. The way of measurement follows modified was available for comparison and as the fragment resembles Heissig (1969), Zapfe (1979), Hünermann (1989)and both large perissodactyl families recorded from Gratkorn but Antoine (2002). slightly differs from both and cannot be assigned with certainty, it is assigned only to Rhinocerotidae vel Chalicotheriidae. Institutional abbreviations The size of an isolated petrosal fragment (UMJGP 204719) also corresponds with a rather large mammal. It is smaller than a BMNH British Museum of Natural History, London, proboscidean and larger than Anchitherium and all occurring Great Britain Artiodactyla from Gratkorn. Whether it represents a GPIT Paläontologische Sammlung der Universität chalicothere or a large rhinocerotid cannot be determined. It is Tübingen, Tübingen, Germany therefore also assigned to Rhinocerotidae vel Chalicotheriidae. SNSB-BSPG Staatliche Naturwissenschaftliche Sammlungen Bayerns - Bayerische Family Chalicotheriidae Gill, 1872 Staatssammlung für Paläontologie und Subfamily Chalicotheriinae Gill, 1872 Geologie, München, Germany Genus Chalicotherium Kaup, 1833 NHMW Naturhistorisches Museum Wien, Wien, Type species: Chalicotherium goldfussi Kaup, 1833 Austria NMA Naturmuseum Augsburg, Augsburg, Chalicotherium goldfussi Kaup, 1833 Germany UMJGP Universalmuseum Joanneum, Graz, Austria Lectotype: M3 dex. (Kaup 1833,tab.VII,fig.3) Type locality: Eppelsheim, Rheinhessen, Germany Anatomical abbreviations L Anteroposterior length of tooth Description and comparison l max Maximum anteroposterior length of tooth l basally Basal length of tooth at the base of the tooth The M3 dex. (UMJGP 204676; l: 41.5 mm, w ant: 44.2 mm, w ∼ – crown (sensu Heissig 1969) post: 37 mm; Figs. 1, 2a e) is well preserved. It is low l ling Lingual anteroposterior length of tooth crowned and possesses a trapezoid shape decreasing in width w ant Anterior linguolabial width of tooth posteriorly. Paracone, protocone, metacone, and hypocone are w post Posterior linguolabial width of tooth well developed. The paracone is the dominant cusp with a strongly inclined labial wall. The protocone is large and as strong as the hypocone. It is located slightly more posterior than the paracone. Para- and mesostyle are distinct, the Systematic palaeontology metastyle is little developed and nearly fused with the metacone. The ectoloph is triconcave with a small depres- Order Perissodactyla Owen, 1848 sion anterior to the parastyle; its largest depression is at the Rhinocerotidae vel Chalicotheriidae labial wall of the paracone. A protoloph and paraconule are present and are connected to the paracone but only by a faint Description and comparison basal ridge to the protocone. The protoloph is short and the paraconule is lower than the protocone. The anterior valley is In size and morphology, specimen UMJGP 204701 fits well to shallow, while the central valley is strongly developed, lin- the distal articulation of a left humerus from a large perissodac- gually open, and deeply incised into the lingual wall of the tyl. It resembles Anisodon grande (see, e.g. Zapfe 1979, fig. 69, mesostyle. The posterior part of the ectoloph, comprising the and Guérin 2012, fig. 1), but differs from it in the stronger distal metacone and metastyle, is directed posterolingually. The concavity of the trochlea and the more rounded medial condyle. short metastyle is bent to the rear becoming almost longitudi- Furthermore it is slightly larger than Anisodon grande,which nal at its end. The posterior crest of the hypocone turns would fit well to Chalicotherium goldfussi (Zapfe 1989). How- labially. Basally, it is connected to the metastyle forming a ever the morphology also resembles the humeri of short posterior cingulum enclosing with the metacone a ba- Rhinocerotidae [see, e.g. Heissig 2012,figs.65–67, sally narrow and occlusally more open postfossette. A strong Brachypotherium from Petersbuch (SNSB-BSPG 1969; anterior and a weak lingual cingulum are present, while labi- Germany; Miocene), or Rhinocerotidae indet. from Mering ally no cingulum is developed. At the protocone, the lingual (SNSB-BSPG 1960 I 121; Germany; Middle Miocene)]. In cingulum rises and is less distinct, but is clearly still present. Palaeobio Palaeoenv (2014) 94:71–82 73 wide and lingually open central valley (Schaefer and Zapfe 50 1971;Zapfe1979), and the course of the labial wall of metacone–metastyle and hypocone (fig. 30 in Schaefer and Zapfe 1971; Anquetin et al. 2007). In A. grande, the metacone 45 and metastyle are differently shaped and aligned to the anteroposterior axis of the tooth (see, e.g. fig. 31 in Schaefer and Zapfe 1971; Anquetin et al. 2007) and the central valley is 40 narrower and lingually closed (Schaefer and Zapfe 1971; Zapfe 1979). In the M3 of Anisodon sp. from Dorn- w ant [mm] Dürkheim 1 (Germany; Late Miocene; MN 11), which pos- 35 sesses a wider central valley than observed in A. grande (Fahlke et al. 2013), the central valley is still narrower than in the specimen from Gratkorn. From the M3 of Kalimantsia 30 Geraads et al., 2001, the specimen differs by a more subsquare 30 35 40 45 50 shape and the pattern and morphology of meta-, hypocone, and metastyle (Geraads et al. 2001). l [mm] C. goldfussi M3 (UMJGP204676) Family Rhinocerotidae Gray, 1821 C. goldfussi M3 (from Wehrli 1939) A. grande M3 (from Zapfe 1979) Rhinocerotidae indet. A. grande M3 (from Guerin 2012) Anisodon sp. M3 (from Fahlke et