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(Mammalia, Perissodactyla) from the Late Miocene of Akkas¸Dag˘I, Turkey

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(Mammalia, Perissodactyla) from the Late Miocene of Akkas¸Dag˘I, Turkey Chalicotheriidae (Mammalia, Perissodactyla) from the late Miocene of Akkas ¸ dag ˘ ı, Turkey Gerçek SARAÇ General Directorate of Mineral Research and Exploration, Natural History Museum, TR-06520 Balgat, Ankara (Turkey) Sevket SEN Muséum national d’Histoire naturelle, Département Histoire de la Terre, UMR 5143 du CNRS, Paléobiodiversité et Paléoenvironnements, case postale 38, 8 rue Buffon, F-75231 Paris cedex 05 (France) [email protected] Saraç G. & Sen S. 2005. — Chalicotheriidae (Mammalia, Perissodactyla) from the late Miocene of Akkas ¸ dag ˘ ı, Turkey, in Sen S. (ed.), Geology, mammals and environments at Akkas ¸ dag ˘ ı, late Miocene of Central Anatolia. Geodiversitas 27 (4) : 591-600. ABSTRACT Among over 5000 large mammalian remains collected at Akkas¸ dag˘ ı, only three specimens belong to a Chalicotheriidae, Ancylotherium pentelicum (Gaudry & Lartet, 1856). The juvenile skull resembles the one from Hadjidimovo-1 in Bulgaria. A duplex bone and a calcaneum were also found. KEY WORDS Chalicotheriidae, All these specimens, although similar in morphology, are about 10% smaller Ancylotherium , than the adult individuals from Pikermi (Greece) which is the type locality of late Miocene, A . pentelicum . The occurrence of this species at Akkas¸ dag˘ ı confirms once Akkas¸ dag˘ ı, Central Anatolia, more the presence of an open woodland bioprovince from Greece to Turkey. Afghanistan during the Turolian. RÉSUMÉ Chalicotheriidae (Mammalia, Perissodactyla) du Miocène supérieur d’Akkas¸ dag˘ ı, Turquie. Sur plus de 5000 restes de grands mammifères récoltés à Akkas¸ dag˘ ı, seuls trois spécimens appartiennent à un Chalicotheriidae, Ancylotherium pentelicum (Gaudry & Lartet, 1856). Le crâne juvénile ressemble par tous ses caractères à celui de Hadjidimovo-1 en Bulgarie. Un os duplex et un calcanéum ont aussi été trouvés. Les dimensions de ces deux spécimens sont environ 10% plus petites que celles des individus adultes de Pikermi (Grèce) qui est la localité MOTS CLÉS Chalicotheriidae, type de A . pentelicum ,mais leur morphologie ne montre pas de différence Ancylotherium , notable. La présence de cette espèce à Akkas¸ dag˘ ı et son aire de dispersion des Miocène supérieur, Balkans à l’Afghanistan confirme une fois encore la réalité de la bioprovince Akkas¸ dag˘ ı, Anatolie Centrale, gréco-irano-afghane, en particulier durant le Turolien, avec des milieux relati- Turquie. vement ouverts et plutôt secs par rapport à d’autres régions en Europe. GEODIVERSITAS • 2005 • 27 (4) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 591 Saraç G. & Sen S. INTRODUCTION In the past, the species pentelicum was often refer- red to other genera including Macrotherium Remains of Chalicotheriidae Gill, 1872 are gene- Lartet in Blainville, 1837, Nestoritherium Kaup, rally scarce in Eurasian late Miocene mammal loca- 1859 and Colodus Wagner, 1861 before Schaub lities, even in those which are richly documented. (1943) revised its systematic status after an exten- The only exception is probably the classical sive study of all postcranial material available at Pikermilocality inGreece which yielded the richest that time (see history in Roussiakis & Theodorou record with over 100 specimens of dental and post- 2001). cranial remains (Roussiakis & Theodorou 2001). In Turkey the occurrence of A . pentelicum was At Akkas¸ dag˘ ı (in the southern Çankırı basin be- noted in several localities, but it is rare: Gülpınar tween the towns of Keskin and Kaman), over 5000 (Kaya 1986), Kemiklitepe B (Kaya 1988; Sen specimens of mammals have been unearthed 1994), Salihpas¸ alar and Pınaryaka (Saraç et al . during thefieldcampaigns of1997, 2000 and 2001 2002), Konya-Kızılören (Saraç 1994). All these but only three specimens of Chalicotheriidae. This localities are correlated to MN11-12. confirms once more that this group is quite mar- The terminology used in this paper follows ginal in late Miocene mammalian faunas. Butler (1965) and Coombs (1978). The material Among the order Perissodactyla, the members of is preserved in the Natural History Museum of the family Chalicotheriidae have a peculiar status the MTA in Ankara. The measurements are having laterally compressed and deeply cleaved given in millimetres. AK2 and AK5 are the num- strong claws on hands and feet. In addition, bers of bone pockets with Ancylotherium . members of the subfamily Schizotheriinae Holland & Peterson, 1913 show an unusual A BBREVIATIONS fusion of the proximal and middle phalanges to Llength; Wwidth; form a single bone named the duplex. Their den- DT transverse diameter; tition is also peculiar in having a W-pattern of maxmaximum; lophs in the cheek teeth, allowing a chisel-like MTA General Directorate of Mineral Research action on food, indicating a browser diet. and Exploration, Ankara. Ancylotherium pentelicum (Gaudry & Lartet, 1856) is the typical representative of the subfami- ly Schizotheriinae, and is only known in localities SYSTEMATICS correlated to the zones MN11-12 in eastern Europe and SW Asia. This species is known from an area extending from Macedonia and Bulgaria Order PERISSODACTYLA Owen, 1848 up to Afghanistan (see Geraads et al . 2001; Family C HALICOTHERIIDAE Gill, 1872 Roussiakis & Theodorou 2001). It is worth men- Subfamily S CHIZOTHERIINAE tioning that A . aff. pentelicum was reported from Holland & Peterson, 1913 Concud (MN11), Nombrevilla (MN10) and Los Valles de Fuentidueña (MN10), all in Spain Genus Ancylotherium Gaudry, 1863 (Alberdi et al . 1981; Heissig 1999). The nature of the material from Concud and Nombrevilla is Ancylotheriumpentelicum not mentioned (Crusafont & Casanovas 1973). (Gaudry & Lartet, 1856) The material from Los Valles de Fuentidueña M ATERIAL. — A juvenile skull having the dorsal part consists of a left astragalus similar in morphology crushed and damaged and the ventral part and incom- to that of A . pentelicum but 1/4 smaller. To con- plete dentition including left P2, P3, M1, M2 and firm the occurrence of this species in Spain, more erupting M3, right M1, M2 and erupting M3 (AK2- 293); a left duplex (AK5-4) and a left calcaneum material and a more detailed study of the avail- (AK2-438). This material was unearthed from two dif- able specimens are needed. ferent bone pockets about 3 m apart. 592 GEODIVERSITAS • 2005 • 27 (4) Late Miocene Chalicotheriidae from Akkas¸ dag˘ ı D ESCRIPTION Dentition (Fig. 1B; Appendix: Table 1) Skull (Fig. 1A) On the Akkas¸ dag˘ ı specimen the premolars are The skull belongs to a young individual, onto- still erupting and are unworn, and thus they pre- genically similar to that of Hadjidimovo-1 in serve all details of the occlusal pattern. We Bulgaria (Geraads et al . 2001), but younger than emphasize that such fresh premolars of the one from Titov Veles in Macedonia Ancylotherium have not previously been des- (Garevski 1974). In the Akkas¸ dag˘ ı specimens, cribed. The premolars are not molarized, so their almost nothing is preserved from the top part of occlusal pattern is different from that of the the skull, while the ventral part is nearly com- molars. The P3 is much smaller than the P4. plete. The skull anatomy of this species is still Both have strong cingula on their anterior, lin- poorly known; apart from the material from gual and posterior faces. The lingual cingulum is Hadjidimovo-1 and Titov Veles, only some stronger in P4. Both premolars have a slightly upper jaw fragments were described from oblique protolophule which is shorter than the Pikermi (Gaudry 1862-1867; Thenius 1953). metalophule. The protocone is without any crest. However, an almost complete skull from It is relatively conical in P3, but rather pinched Gülpınar (Department of Çanakkale, northwest- mesio-distally and lined up with the protolophule ern Turkey) is preserved at the Ankara Natural in P4. Both premolars have the metalophule History Museum, and it belongs to an adult indi- thinner than the protolophule and oriented vidual. This skull has not been the subject of any rather transversely. The buccal surface of the pre- publication. We compared the Akkas¸ dag˘ ı skull molars is flat, thus distinguishing it from that of with this one in order to appreciate the propor- Chalicotherium Kaup, 1833 which has P3 and P4 tions of missing and damaged parts. with folded or depressed buccal surface (Zapfe On the skull AK2-293, the dorsal part is com- 1979). The width/length ratio is 125.3 for P3 pletely crushed preventing detailed observation and 127.3 for P4. of the nasal, frontal and parietal bones. Also, the The M1 is much larger than the P4 but notably posterior part of the skull is missing. Some parts smaller than the other molars. Its outline is of the temporal bones are preserved, as well as almost square with a W/L ratio of 98.9. the zygomatic arches, maxillae, palate, glenoid However, the anterior part of the M1 is narrower fossae, post-glenoid processes, and some cheek than the posterior part. The ectoloph is W-shaped. teeth (see Fig. 1A for details). The M3s are just Wear has affected only the anterior faces of the erupting. paracone and metacone crests. The protocone is In ventral view, the palate narrows slightly for- an isolated cusp situated close to the antero- ward (narrowest width of the palate between M2s lingual corner of the tooth; it is without any crest is 67 mm, and between M1s 52 mm). The choa- or furrow. The protolophule is short. The meta- nae are large and elliptical in outline. Their ante- cone is slightly higher than the paracone, and rior border is behind the M1s. The median keel their lingual tips protrude lingually beyond the of the basisphenoid forms a bony septum which median line. The paracone rib is weak and there narrows anteriorly and divides the choanae into is no metacone rib. The parastyle protrudes ante- two parts. The zygomatic arch is rounded, and it ro-buccally, the hypocone lingually. The median narrows anteriorly. It reaches the maxilla imme- valley is directed toward the mesostyle.
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