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Dorcatherium naui and pecoran ruminants from the late Middle Miocene Gratkorn locality (Austria)

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Abstract

One of the rare records of a rich ruminant fauna of late Middle Miocene age (Sarmatian sensu stricto; 12.2–12.0 Ma) was discovered at the Gratkorn locality (Styria, Austria). It comprises, besides Micromeryx flourensianus, ?Hispanomeryx sp., Euprox furcatus, Palaeomerycidae gen. et sp. indet., and Tethytragus sp., one of the oldest records of Dorcatherium naui. Gratkorn specimens of the latter species are in metric and morphologic accordance (e.g. selenodont teeth, bicuspid p2, non-fusion of malleolus lateralis and tibia) with type material from Eppelsheim (Germany) and conspecific material from Atzelsdorf (Austria), and do not show an intermediate morphology between Late Miocene Dorcatherium naui and Middle Miocene Dorcatherium crassum, thus enforcing the clear separation of the two species. It furthermore confirms the assignation of Dorcatherium naui to a selenodont lineage (together with Dorcatherium guntianum) distinct from a bunoselenodont lineage (including Dorcatherium crassum). The record of ?Hispanomeryx sp. is the first of this genus in Central Europe. While Tethytragus sp. could also be a new bovid representative for the Sarmatian of Central Europe, Micromeryx flourensianus and Euprox furcatus are well-known taxa in the Middle Miocene of Central Europe, but comprise their first records from Styria. Morphological data from this work in combination with isotopic measurements (δ18OCO3, δ13C; Aiglstorfer et al. 2014a, this issue) indicate a niche partitioning for the ruminants from Gratkorn with subcanopy browsing (Euprox furcatus), top canopy browsing (Tethytragus sp.) and even a certain amount of frugivory (Dorcatherium naui and Micromeryx flourensianus).

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Acknowledgements

The authors are indebted to P. Havlik for fruitful discussions and correction of the manuscript. M.A. wants to thank S. Kötter, A. Tröscher and A. Beckmann for their help in the search for comparison material. J van der Made, J. Morales, I. Sánchez (Museo Nacional de Ciencias Naturales, Madrid), E. Heizmann (SMNS), B. Azanza (Universidad de Zaragoza), N. Spassov (NMNHS), N. Heckeberg (SNSB-BSPG and LMU München, Department of Zoology Cambridge), F. Fack, G. Metais, B. Mennecart, M. Pickford (MNHN) and T. Wiedl (University of Graz) are thanked for help and fruitful discussions. D. Vasilyan, J. Prieto (both University Tübingen) and F. Witzmann (Museum für Naturkunde Berlin) are thanked for taking pictures of specimens for comparison. M. Gross (UMJGP), U. Göhlich (NHMW), D. Nagel (IPUW), L. Costeur (NMB), C. Argot (MNHN), G. Scharfe (IGM), D. Alba (ICP), R. Ziegler (SMNS), M. Rummel (NMA), H. Lutz (NHMM), T. Engel (NHMM), H. Stapf (SSN), J. Rösinger and E. Weber (both University Tübingen, Zoological Collection), N. Spassov and L. Hristova (NMNHS), E. Bernard (BMNH), and P. Havlik (GPIT) are thanked for access to comparison material. W. Gerber (University Tübingen) is thanked for taking pictures, P. Havlik, M. Gross, H. Luginsland (GPIT), R. Ellenbracht (GPIT), H. Stöhr (GPIT) N. Winkler (UMJGP), J. Fuß (GPIT) are furthermore thanked for preparation or great support in preparation of the material. This research received support from the SYNTHESYS Project (FR-TAF-1892) which is financed by the European Community Research Infrastructure Action under the FP7 “Capacities” Program. B. Azanza and L. Costeur greatly helped to improve the manuscript with careful reviews and helpful advice. And last but not least the authors want to thank students and volunteers from Graz, Munich and Tübingen for their help in excavations from 2005 to 2013.

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Authors and Affiliations

  1. Fachbereich Geowissenschaften, Eberhard Karls Universität Tübingen, Sigwartstraße 10, 72076, Tübingen, Germany

    Manuela Aiglstorfer & Madelaine Böhme

  2. Senckenberg Center for Human Evolution and Palaeoenvironment (HEP), Sigwartstraße 10, 72076, Tübingen, Germany

    Manuela Aiglstorfer & Madelaine Böhme

  3. SNSB—Bayerische Staatssammlung für Paläontologie und Geologie, Richard-Wagner-Str. 10, 80333, München, Germany

    Gertrud E. Rössner

  4. Department für Geo-und Umweltwissenschaften, Paläontologie & Geobiologie, Ludwig-Maximilians-Universität München, Richard-Wagner-Str. 10, 80333, München, Germany

    Gertrud E. Rössner

  5. GeoBio-Center LMU, Richard-Wagner-Str. 10, 80333, München, Germany

    Gertrud E. Rössner

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  1. Manuela Aiglstorfer
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  2. Gertrud E. Rössner
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Correspondence to Manuela Aiglstorfer.

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This article is a contribution to the special issue “The Sarmatian vertebrate locality Gratkorn, Styrian Basin.”

Electronic supplementary material

Below is the link to the electronic supplementary material.

Online Resource 1

References for stratigraphic range of Dorcatherium species in Central Europe and reference localities (PDF 61.8 kb)

Online Resource 2

Dental and postcranial measurements for Dorcatherium naui from Gratkorn. (PDF 44.4 kb)

Online Resource 3

Dental measurements for Moschidae from Gratkorn, in comparison to literature data. (PDF 64 kb)

Online Resource 4

Dental, postcranial and antler measurements for Euprox furcatus from Gratkorn. (PDF 67.3 kb)

Online Resource 5

Dental and postcranial measurements for Tethytragus sp. from Gratkorn. (PDF 32.9 kb)

Appendix: Historical context for the description of the species Dorcatherium naui and considerations on species validity of different Dorcatherium species

Appendix: Historical context for the description of the species Dorcatherium naui and considerations on species validity of different Dorcatherium species

The genus Dorcatherium was erected by Kaup in 1833 in a letter to Prof. Bronn (published in Neues Jahrbuch für Mineralogie, Geognosie und Petrefaktenkunde, 1833, p. 419), on a ruminant mandibula with p3–m3 (and alveolae for p1–2) from Eppelsheim (Rheinland-Pfalz, Germany), based on the presence of four premolars, and the rostral extension of the premolar to the level of the symphysis. Due to the resemblance of tooth morphology to that of a deer, he chose the name Dorcatherium (ή δορκάς greek for gazella, deer). In the same letter, he erected the type species, which he named naui after his friend, Geheimrat von Nau. The catalogue of gypsum casts of the Palaeontological Collection in Darmstadt (Kaup and Scholl 1834) refers to the mandibula, described in 1833, and a fragment of a maxilla with P4–M3. Casts of both were sent to Berlin, Bonn, Frankfurt, London, Lyon, Paris, Strasburg, Stuttgart, and Zürich (Kaup and Scholl 1834).

The other medium-sized Dorcatherium species besides D. naui, Dorcatherium crassum, is more common in the Miocene of Europe. It was erected by Lartet (1851) as ?Dicrocerus crassus and he had already noticed the similarity of the upper canines with those in chevrotains (at that time seen as close relatives of Moschus and Moschus as a cervid genus; Milne Edwards erected the family Tragulidae in 1864). The first description is often cited as D. crassum (Lartet, 1839), but no indication can be found in any of the works of Lartet published in 1839 for the species name “Dorcatherium crassum” nor in the first mentioning of remains of the species by Blainville (1837): “Des dents canines supérieures d’un petit ruminant sans bois ou à bois pédonculé des sous-genres moschus ou cervulus;” (Blainville 1837, p. 425) (for detailed discussion of the scientific history concerning the species, see Morales et al. 2012). With D. crassum, Milne Edwards (1864) included a fossil species in his newly erected family Tragulidae, together with the modern genera Hyemoschus, and Tragulus (including T. meminna which is considered today to represent a third tragulid genus, Moschiola), but affiliated it to the genus Hyemoschus. Although he observed the similarity between his Hyemoschus crassus and Dorcatherium naui, he did not include the latter in the Tragulidae due to the presence of a p1, which is completely reduced in the modern representatives of the family. Finally, Schlosser (1916) found sufficient morphological accordance of both species to affiliate Hyemoschus crassus to the genus Dorcatherium.

Today, five Dorcatherium species are generally accepted from the Miocene of Europe, differing in dimensions, dental and postcranial morphology and stratigraphic range (Fig. 1): the small-sized D. guntianum von Meyer, 1846 (late Early to Middle Miocene; MN 4–7/8; Seehuber 2008; Sach and Heizmann 2001; Rössner and Heissig 2013), the medium-sized D. naui (late Middle to Late Miocene; MN 7/8–11; Czyzewska and Stefaniak 1994; Rössner 2007, 2010; Alba et al. 2011; this publication) and D. crassum (Lartet,1851) (late Early to Middle Miocene; MN 4–7/8; Eronen and Rössner 2007; Alba et al. 2011; Rössner and Heissig 2013), the larger-sized D. vindebonense von Meyer, 1846 (late Early to Middle Miocene; MN4–6; Thenius 1952; Sach and Heizmann 2001; Rössner 2007, 2010; Rössner and Heissig 2013), and the large-sized D. peneckei (Hofmann 1893) (early Middle Miocene; MN5–6; Rössner 2007, 2010; Rössner and Heissig 2013).

D. puyhauberti, Arambourg and Piveteau, 1929 (Late Miocene; MN9–13; Gentry et al. 1999; Rössner and Heissig 2013) and D. jourdani (Déperet, 1887) (Late Miocene; MN 9–11; Gentry et al. 1999; Rössner and Heissig 2013) have been documented only rarely, with only a few specimens, which possess no unambiguous features distinguishing them from other European species and could be synonymous to D. guntianum and D. naui, respectively (for further information, see discussion in the section on Dorcatherium naui of this publication).

D. rogeri, erected by Hofmann in 1909 due to a misunderstanding in von Meyer (1846), must be considered synonymous with D. vindebonense (Thenius 1952). The quite small D. bulgaricum Bakalov and Nikolov, 1962 from the West-Mariza-Basin (?Pliocene, Bulgaria; Rössner 2007) was erected on two mandibulae with m1–3. Although both specimens show clearly developed internal postmeta- and postprotocristids, the assignation to the genus Dorcatherium is ambiguous. As far as it could be observed on the figures in Bakalov and Nikolov (1962; originals supposed to be lost), the specimens possess a strongly split posthypocristid, especially in the m3. This has so far been described only in the m3 of Dorcatherium cf. pigotti from Arrisdrift (basal Middle Miocene; Morales et al. 2003). In any case, the splitting of the posthypocristid is much stronger in D. bulgaricum than the minor splitting observed in D. crassum (Alba et al. 2013) and in D. naui from Gratkorn. Furthermore, D. bulgaricum possesses a rudimentary paraconid. “A small accessory cusplet” is described by Pickford (2002, p. 97) only in the Early Miocene D. iririensis from Africa. However, with a small hypoconulid in m2 and a rounded lingual wall in P4, the latter also differs significantly from other Dorcatherium species. A clearly developed paraconid can be observed, for example, in the lophiomerycid Zhailimeryx (Guo et al. 2000). Morales et al. (2012) also observed more similarities in D. bulgaricum to the Oligocene genera Lophiomeryx Pomel, 1853 and Cryptomeryx Schlosser 1886 (synonymised with Iberomeryx; Métais et al. 2001; Mennecart et al. 2011) than to other Dorcatherium species. The stratigraphic age of D. bulgaricum is furthermore ambiguous and could also be Paleogene (M. Böhme, personal opinion). We thus did not consider the species Dorcatherium bulgaricum in our discussions.

The Miocene tragulid genus Dorcabune Pilgrim, 1910 is so far only known, but with several species, from Asia (Rössner 2007). As Dorcatherium and Dorcabune overlap in morphological key features, a revision of the two genera would probably result in at least two morphotypes of Miocene tragulids with a differentiation into more bunodont (including D. crassum, vindebonense and peneckei) and more selenodont forms (including D. naui and guntianum; Rössner 2007 referring also to Mottl 1961; Fahlbusch 1985; Qui and Gu 1991). Other Miocene tragulid genera described from Asia are Siamotragulus Thomas et al., 1990 and Yunnanotherium Han, 1986.

Five Dorcatherium species have been recorded from the Miocene of Africa : D. songhorensis Whitworth, 1958, D. pigotti Whitworth, 1958, D. iririensis Pickford 2002 and D. chappuisi Arambourg, 1933, as well as a second tragulid genus, Afrotragulus, with the species A. parvus (Witworth, 1958) and A. moruorotensis (Witworth, 1958) (Sánchez et al. 2010b).

To get a better idea about the relationships of and faunal exchanges between Asian, African and European Miocene tragulids, a revision of the different taxa and lineages as also proposed in the section on Dorcatherium naui in this publication and by Sánchez et al. 2010b is surely needed.

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Aiglstorfer, M., Rössner, G.E. & Böhme, M. Dorcatherium naui and pecoran ruminants from the late Middle Miocene Gratkorn locality (Austria). Palaeobio Palaeoenv 94, 83–123 (2014). https://doi.org/10.1007/s12549-013-0141-9

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