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DSA10 Giourtsakis Ioannis X. Giaourtsakis Ludwig-Maximilians-Universität, München Late Neogene Rhinocerotidae of Greece: distribution, diversity and stratigraphical range Giaourtsakis, I.X., 2003 - Late Neogene Rhinocerotidae of Greece: distribution, diversity and stra- tigraphical range - in: Reumer, J.W.F. & Wessels, W. (eds.) - DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA. A VOLUME IN HONOUR OF HANS DE BRUIJN - DEINSEA 10: 235- 253 [ISSN 0923-9308] Published 1 December 2003 The present contribution provides a comprehensive overview of the Neogene rhinoceroses of Greece and evaluates their affinities with the European and Asian species. The distribution, diver- sity and dispersal of the family in the Eastern Mediterranean are discussed in detail. Knowledge of rhinos during the Middle Miocene in Greece is still very limited, because of the scarce mate- rial. During the Late Miocene the family was diverse and abundant in Greece. The dominant form, Ceratotherium neumayri, was probably an African immigrant that invaded Southeastern Europe, Anatolia and the Middle East during the Late Miocene. This rather subhypsodont tandem-horned rhino has already achieved most of the apomorphic characters of the living Dicerotina. The con- temporary species Dihoplus pikermiensis is also common, but the origin and evolutionary rela- tionships among its Miocene and Plio-Pleistocene relatives are still controversial. The Greek Late Miocene representatives of the subfamily Aceratheriinae are obviously closer related to the Asian genera Chilotherium and Acerorhinus, than to the typical aceratheres Aceratherium, Alicornops and Hoploaceratherium found in Central and Western Europe at the same time. Chilotherium and Acerorhinus migrated into Greece from Central Asia trough Anatolia at the beginning of the Vallesian. Chilotherium dominated Samos, where it was represented by at least three species: Ch. samium, Ch. kowalevskii and Ch. schlosseri. In continental Greece, the richest acerathere material comes from the recently discovered locality of Pentalophos-1, where the species Aceratherium kiliasi has been described. Later, this material was assigned partly to Chilotherium and partly to Acerorhinus. The Late Miocene rhino assemblages of Greece are in conformity with the ones from Southeastern Europe and the Eastern Mediterranean, especially with the ones from Anatolia. By the end of the Turolian, all these species became extinct in Greece, unable to survive the paleoen- vironmental changes and the faunal turnover at the Miocene-Pliocene boundary. Pliocene rhino- ceroses are poorly documented in Greece. Stephanorhinus cf. etruscus has been reported with more or less certainty from several localities, but the available material is not sufficient for useful comparisons. Correspondence: Ioannis X. Giaourtsakis, Institut für Paläontologie und Historische Geologie, Ludwig-Maximilians-Universität München, Richard-Wagner-Str. 10, D-80333 Munich, Germany, [email protected] Keywords: Rhinocerotidae, Dihoplus, Greece, Eastern Mediterranean, Miocene, Pliocene INTRODUCTION Wagner (1848) described and illustrated the The study of fossil Rhinocerotidae in Greece first rhino specimens from Pikermi introdu- has already begun at the middle of the last cing a new species, Rhinoceros pachygnathus. century. The first remains of a fossil rhinoce- In subsequent studies, Roth & Wagner (1854) ros from Greece were mentioned by Goldfuss and Wagner (1857) recognised the presence (1841) in a brief note announcing the disco- of a second species at Pikermi, Rhinoceros very of fossil mammals at a new locality schleiermacheri. Gaudry’s (1862-67) monu- north of Athens, Pikermi. A few years later, mental monograph on the Pikermi fauna 235 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003 documented adequately both species. In addi- created the species Aceratherium kiliasi. tion, Gaudry described an adult mandible These two papers mark also the last signifi- with well developed incisors as Aceratherium cant contributions on the Neogene fossil sp. Weber (1904, 1905) was the first one who Rhinocerotidae of Greece. studied thoroughly rhino material from More details on the research chronicle as Samos, although the island was known long well as on the geological setting and the ago for its wealth of Neogene mammals. In faunal history of each locality (Fig. 1) are his first contribution, Weber studied the given in the author’s Diploma Thesis, which remains of Rhinoceros pachygnathus and has been accomplished at the University of Rhinoceros schleiermacheri, while in his Athens (Giaourtsakis 1999). A Ph.D. study second work he published a remarkable dealing with an extensive systematic revision sample of acerathere material and described of the fossil rhinoceroses of Greece, inclu- two new species, Aceratherium schlosseri ding old and new unpublished material, is and Aceratherium samium. Andree (1920) currently in progress at the University of later created two additional species based on Munich. The essence of this contribution is to material from Samos, Aceratherium wegneri provide a comprehensive updated overview and Aceratherium angustifrons. Arambourg & of the present knowledge on the Neogene rhi- Piveteau (1929) documented the presence of noceroses of Greece and to evaluate their Rhinoceros pachygnathus and Aceratherium affinities with the European and Asian record. sp. at the Neogene deposits of the Axios Errors often repeated in the past are emended, Valley, near Thessaloniki. Naturally, modern while some new aspects on old controversies workers have gradually modified the generic are introduced. and some of the specific names used by these During the Neogene we can recognise three first authors. successive rhino assemblages in Greece and During the years after the Second World the Eastern Mediterranean, which are inter- War and up to the beginning of the 1980’s rupted by major faunal turnovers at the very little systematic work was undertaken on Middle/Late Miocene and at the Miocene/ the fossil rhinoceroses of Greece. However, Pliocene boundary. during this period quite a few occasional papers and many references in faunal lists of MIDDLE MIOCENE preliminary reports revealed in silence the Middle Miocene localities with large mam- presence of the family in numerous Neogene mals are poorly documented in Greece (De localities of Greece. The rhinoceroses of Bonis & Koufos 1999). The richest and best Pikermi and Samos continued to be mentio- known site from this time period is the locali- ned frequently in major reviews and global ty of Thymiana on the island of Chios. From works on the family, but in most cases neither the older excavations on Chios, the presence new material was introduced nor was the old of Rhinocerotidae indet. has been reported one properly revised. A bright exception to only in a preliminary report (Tobien 1968). A the aforementioned was the important contri- recent Greek-French field campaign has yiel- bution of Geraads (1988), who reviewed a lot ded many new specimens, but no rhinoceros of the known Rhinocerotinae material of was present (De Bonis & Koufos 1999). In an Pikermi and Samos. Geraads established overview of the family Rhinocerotidae, excellent criteria for distinguishing cranial Heissig (1989) stated the presence of "early and dental remains of Ceratotherium neumay- Dicerotini specimens from the Middle ri and Dihoplus pikermiensis. Two years Miocene of Anatolia and Chios". The author later, Geraads & Koufos (1990) studied in considered these specimens as being earlier detail the rich material discovered at the loca- even than the first named species of this line- lity Pentalophos-1, near Thessaloniki, and age, the African Paradiceros mukirii HOOIJER, 236 GIAOURTSAKIS: Neogene Rhinocerotidae of Greece 1968. During a conversation, Prof. K. Heissig constricted protocones and the wider, oblique explained that the referred specimens from ectometalophe in M3, often observed in Chios belong to the collections of Prof. H. Chilotherium. The affinities of this material Tobien, but their present status is uncertain recall the morphology of the "West European" (K. Heissig, pers. comm. 2001). Thus these aceratheres, which were present in the references will be considered here as Eastern Mediterranean region at that time Rhinocerotidae indet. (Heissig, 1976). This work was published Two lower premolars from a lignite mine before the modern revision that separated the near the village of Chrysavgi (Mygdonia various "Aceratherium" species in several basin, N. Greece) were described as "Diceros new genera and therefore Symeonidis (1974) pachygnathus" by Psarianos (1958). Dimo- used provisionally the name "Aceratherium poulos (1972) studied also these teeth sp." for the remains of Atalanti. together with three upper molars and some Currently three genera (Aceratherium ribs and placed them to "Dicerorhinus orien- KAUP, 1832; Alicornops GINSBURG & GUÉRIN, talis". Both authors implied a Late Miocene 1979; Hoploaceratherium GINSBURG & age for the fossils. In our opinion, these few HEISSIG, 1989) with about six species are teeth remains are insufficient to establish an considered valid from the later Middle and accurate taxonomic ascription at the present. Late Miocene of Western Europe (MN6 – De Bonis & Koufos (1999), in their review of MN13). Cerdeño (1996) proposed a synony- the Miocene large mammal succession in my between Hoploaceratherium and Greece, considered also this material as Acerorhinus KRETZOI, 1942, but we believe
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