Rhinocerotidae (Mammalia, Perissodactyla) from the late Miocene of Akka ș da g ˘ ı, Turkey

Pierre-Olivier ANTOINE Laboratoire des Mécanismes de Transfert en Géologie, UMR 5563, Université Paul Sabatier, 14 avenue Édouard Belin, F-31400 Toulouse (France) [email protected]

Gerçek SARAÇ General Directorate of Mineral research and exploration, Natural History Museum, TR-06520 Balgat, Ankara (Turkey)

Antoine P.-O. & Saraç G. 2005. — Rhinocerotidae (Mammalia, Perissodactyla) from the late Miocene of Akka ș da g ˘ ı, Turkey, in Sen S. (ed.), Geology, mammals and environments at Akka ș da g ˘ ı, late Miocene of Central Anatolia. Geodiversitas 27 (4) : 601-632.

ABSTRACT More than 120 rhinocerotid remains unearthed in the middle Turolian locality of Akka ș dag˘ ı(Central Anatolia) are described. The fauna is diversified, with a pair of large two-horned species, Ceratotherium neumayri (Osborn, 1900) and Stephanorhinus pikermiensis (Toula, 1906), and two smaller species of short limbed aceratheriines ( Chilotherium sp. and Acerorhinus sp.). Ceratotherium neumayri is by far the most common species, with a complete skull, 114 specimens and at least 11 individuals. The cranial, dental and postcranial remains of C . neumayri are among the largest ones described so far for this species. The coexistence of C . neumayri , KEY WORDS S . pikermiensis ,and chilotheres is common in the Turolian of Eastern Mammalia, Rhinocerotidae, Mediterranean: comparable rhinocerotid associations are known at Kavakdere Ceratotherium neumayri , (MN 12, Turkey) and Samos (MN 12, Greece). The large size of the middle Turolian, C . neumayri specimens is consistent with the middle Turolian age for Akka ș dag˘ ı, Central Anatolia, Akka ș dag˘ ı(MN 12), as stated on the whole mammalian fauna and Turkey. radiometric data.

RÉSUMÉ Rhinocerotidae (Mammalia, Perissodactyla) du Miocène supérieur d’Akka ș da˘g ı, Turquie. Plus de 120 restes de rhinocérotidés découverts dans le gisement Turolien moyen d’Akka ș dag˘ ı (Anatolie centrale) sont décrits. La faune est diversifiée, avec deux espèces de grands rhinocéros bicornes, Ceratotherium neumayri (Osborn, 1900) et Stephanorhinus pikermiensis (Toula, 1906), et deux espèces

GEODIVERSITAS • 2005 • 27 (4) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 601 Antoine P.-O. & Saraç G.

de petits acérathères aux membres trapus ( Chilotherium sp. et Acerorhinus sp.). Ceratotherium neumayri est de loin l’espèce la mieux représentée, avec un crâne complet, 114 spécimens et au moins 11 individus. Les restes crâniens, dentaires et postcrâniens comptent parmi les plus grands jamais attribués à MOTS CLÉS cette espèce. La coexistence de C . neumayri , S . pikermiensis et de chilothères Mammalia, Rhinocerotidae, est fréquente dans le Turolien de la Méditerranée orientale : des associations Ceratotherium neumayri , comparables de rhinocérotidés sont connues à Kavakdere (MN 12, Turquie) Turolien moyen, et Samos (MN 12, Grèce). La grande taille des restes de C . neumayri est Akka ș dag˘ ı, Anatolie Centrale, conforme à l’âge du gisement (MN 12), établi sur l’ensemble de la faune Turquie. mammalienne et sur la base de données radiométriques.

INTRODUCTION MNHN Muséum national d’Histoire natu- relle, Paris; MTA General Directorate of Mineral The middle Turolian locality of Akka ș dag˘ ı, be- research and exploration, Ankara; tween Kaman and Keskin (Central Anatolia; NHM The Natural History Museum, Kazancı et al. 1999), has yielded a diversified London. mammalian fauna thanks to the large excavations Anatomy under the leadership of S. Sen (Kazancı et al. TD transverse diameter; APD antero-posterior diameter; 1999). Among the mammals, the Perissodactyla Hheight; play a leading part, with four species of Equidae Llength; (Koufos & Vlachou 2005) and as many species of Wwidth (when different from TD); Rhinocerotidae. More than 100 specimens have ant.anterior; post.posterior; been attributed to this family. Two large two- w.n. without number. horned and two small hornless rhinocerotids are present: (Osborn, 1900) In the text, the generic names Ceratotherium , Stepha- Ceratotherium neumayri norhinus, Chilotherium , Acerorhinus, and Alicornops (114 specimens), Stephanorhinus pikermiensis will be abbreviated in C ., S ., Ch ., A ., and Al . respec- (Toula, 1906) (nine specimens), Chilotherium sp. tively. (one specimen) and Acerorhinus sp. (seven speci- mens), respectively. These taxa occur more or less frequently in the Eastern Mediterranean area MATERIAL AND METHODS during Turolian times (Heissig 1975a, b, 1996, 1999; Geraads 1988; Geraads & Koufos 1990; All the specimens from Akka ș dag˘ ı described Bonis et al . 1992a, b; Saraç 1994; Antoine et al . hereunder are stored in the Natural History 2003; Fortelius et al . 2003). Museum of the MTA, in Ankara. The specimens GOK-1 to GOK-14 and GOK-16 to GOK-19 A BBREVIATIONS are deposited in the Département Histoire de la Akka ș da˘g ıspecimens, inventory Terre of the MNHN. AK 2000-2001 excavations; AKA and AKB 1997 excavation; Capital letters are used for upper teeth (D, P, M), AKK surface collects and unknown pockets; and lower-case for lower teeth (d, p, m). For the GOK Heintz-Ginsburg excavation (former- astragalus/calcaneus facets, the terminology is ly Gökeș me). that of Heissig (1972: pl. 13). The suprageneric Institutions GDAU Geological Department of the Ankara systematics follows that proposed by Antoine University; (2002) and Antoine et al . (2003).

602 GEODIVERSITAS • 2005 • 27 (4) Late Miocene rhinocerotids from Akka ș dag˘ ı

SYSTEMATICS GOK-4; McIV, GOK-3) and right metacarpus (McII, GOK-6; McIII, GOK-1; McIV, GOK-5) from the Order PERISSODACTYLA Owen, 1848 same individual; left magnum, AK5-632; right mag- num, AK7-38; left unciform, AK5-633; left unciform Superfamily R HINOCEROTOIDEA Owen, 1845 lacking posterior tuberosity, AK6-58; left unciform Family R HINOCEROTIDAE Owen, 1845 lacking posterior tuberosity, AKK-157; right unci- Subfamily R HINOCEROTINAE Owen, 1845 form, AK4-74; right unciform, AK7-147; posterior tuberosity of a right unciform, AKK-156; posterior Tribe R HINOCEROTINI Owen, 1845 tuberosity of a right unciform, AKK-282; left McII, AK6-55; proximal part of a left McII, AK7-37; proxi- Genus Ceratotherium Gray, 1867 mal part of a left McII (juvenile), AK5-182; left McIII, AK5-68; proximal part of a left McIII, AK3-202; T YPE SPECIES. — Ceratotherium simum (Burchell, proximal part of a left McIII, AK14-23; distal part of a 1817) by original designation (Gray 1867: 1027). left McIII, AKB-83; left McIV, AK3-230; right McIV, AK5-436; proximal fragment of a right McIV, AK13-2; left femur, AK4-253; left femur (diaphysis), Ceratotherium neumayri (Osborn, 1900) AK5-367; left femur (diaphysis), AK6-153; distal part of a left femur, AK7-39; femoral head, AK5-323; dis- Atelodus neumayri Osborn, 1900: 263, text-fig. 16. tal end of a right femur, AK3-63; left patella, AK11-82; fused left tibia and fibula, AK7-40; distal Diceros pachygnathus Guérin, 1980: 202-400, text- end of a right tibia, AK5-366; distal end of a right figs30, 31, 33, 39-43, 46, 51-56, tabl. 45-49, 51-56, tibia, AK7-63; distal end of a right tibia, AKK-154; 59-63, 65, 67-72, 75-78. distal epiphysis of a left fibula, AK13-3; left astragalus Ceratotherium neumayri – Geraads 1988: 13-41, text- (young), AK2-438; left astragalus, AK4-75; left astra- figs 1-5, pl. 2; 1994: 82-85, text-figs 1, 2, pl. 2, fig. 5. — galus, AK5-523; left astragalus, AK5-423; lateral part Geraads & Koufos 1990: 151-154, pl. 1. —Kaya of a left astragalus (young), AK5-319; left astragalus 1994: 13-22, pls 1, 2. — Heissig 1996: 341-342, 347, (young), AK6-56; left astragalus, AK7-148; left astra- text-fig. 27.1. galus, AK11-1; right astragalus, AK5-69; right astra- galus, AK5-634; medial fragment of a right astragalus, For synonymy anterior to 1980, see Geraads (1988: AK13-4; left calcaneus, AK3-66; right calcaneus, 36). AK7-36; left navicular, AK4-76, and broken left ecto- M ATERIAL. — Complete skull with P2-M3, AK4-212; cuneiform, AK4-77, from the same individual; right right M1, much worn, AK2-294; fragment of left navicular, AK5-439; right cuboid, AK5-636; left ento- M1/2, AK4-243; left p4, AK2-295; left maxilla with cuneiform, AK13-5; right mesocuneiform, AK5-637; D1-3 and D4 erupting, AK5-502; right maxilla with left ectocuneiform, GOK-13; right ectocuneiform, D3 and alveolus of D4, AK6-61; fragment of a left AK5-635; left metatarsus (MtII, GOK-9; MtIII, juvenile mandible, with d3 and the alveolus of d2, GOK-7; MtIV, GOK-8); proximal end of a right AK2-296; right D1, AK5-424; fragment of left D2, MtII, AK5-440; left MtIII, AK2-163; right MtIII AK2-435; anterior fragment of left d3, AK5-425 and lacking distal epiphysis (young), AKA-44; distal end of right d3, AK6-134 (same individual); left ulna lacking a right MtIII, AK4-238; proximal end of a right MtIV, distal end, AK6-132; right humerus, right radius and AK5-437; distal fragment of a metapodial, AK4-214. right ulna from the same individual, AK4-w.n.; proxi- Additional material:cervical vertebra, AK3-61; cervi- mal part of a right humerus, AK6-301; distal part of a cal vertebra, AK3-67; cervical vertebra, AK4-182; cer- right humerus, AK-w.n.; fragment of diaphysis of a vical vertebra, without caudal epiphysis, AK5-435; left humerus (juvenile), AK4-183; distal epiphyses of a thoracic vertebra, AK7-186; thoracic vertebra, left (GOK-17) and a right radius (GOK-16) from the AK6-300; thoracic vertebra, AK3-133; thoracic verte- same young individual; left radius in two parts, GOK- bra, AK5-324; thoracic vertebra, AK14-22. These 18 and GOK-19; left radius, AK5-627; proximal end specimens are tentatively referred to C . neumayri , of a left radius, AK4-213; left radius lacking distal end, based on their large size and the domination of that AK4-241; proximal end of a left radius, AK5-67; left very species with respect to other associated rhinos in ulna, AK6-133; left ulna lacking distal end (juvenile), Akka ș dag˘ ı. However, their assignment to S . pikermi- AK6-131; left ulna lacking distal end, AK7-156; right ensis cannot be ruled out. ulna, AK3-118; right ulna, AK4-184; right ulna (pre- dated olecranon), AK6-302; proximal part of a right ulna, AK2-95; proximal part of a right ulna, AK6-303; D ESCRIPTION left scaphoid, AK2-437; right scaphoid, AK5-630; right pyramidal (fragment), AK5-631; left magnum Skull (Fig . 1; Appendix: Table 1) without posterior tuberosity (GOK-11), left unciform The adult skull AK4-212, complete and not de- (GOK-10), left metacarpus (McII, GOK-2; McIII, formed, is large and dolichocephalic (width/length

GEODIVERSITAS • 2005 • 27 (4) 603 Antoine P.-O. & Saraç G.

F IG. 1. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı (Anatolia, Turkey), middle Turolian, skull (AK4-212); A , left lateral view; B , dorsal view; C , palatine view. Scale bar: 20 cm.

604 GEODIVERSITAS • 2005 • 27 (4) Late Miocene rhinocerotids from Akka ș dag˘ ı

ratio = 0.46). The premaxillae are rather long In distal view, the anterior start of the processus (length = 75 mm) and edentulous. The nasal zygomaticus maxillari is progressive, following the bones do not bear any lateral apophysis. The curvature of the teeth row. The palate is narrow foramen infraorbitalis is open above the P3/P4 (Appendix: Table 1). The palatine fossa reaches limit. The nasal notch reaches the middle of P3 the posterior part of the M2. The vomer is thick while the anterior border of the orbit is above and rounded. The articular tubercle for the man- the middle of M2. There is no nasal septum dible, transversally concave, forms a high and ossification. The only preserved suture is the salient semi-cylindre in lateral view. The foramen jugal/squamosal one, which is straight and postglenoideum is not visible. The processus postgle- smooth. The processus lacrymalis is absent, but a noidalis is long, strong and narrow transversally. strong lateral projection of the orbit widens the The articular surface of the latter defines a right skull (zygomatic width/frontal width ratio = dihedron in cross section. The foramen nervi 1.47). The processus postorbitalis is absent on the hypoglossi is open in the middle of the condylar frontal. The base of the processus zygomaticus fossa. A sagittal crest runs all along the basilar maxillari is high: it begins several centimetres process of the basioccipital. The posterior part of above the neck of M2/3. The zygomatic arch is the processus zygomaticus of the squamosal is low and poorly developed. It forms a thin stripe, concave, due to a transverse groove. The processus without any processus postorbitalis . The dorsal posttympanicus is curved forward and very short, profile of the skull is mainly flat, only rising in while the processus paraoccipitalis is long and well its posterior third. The foramen sphenorbitale and developed. Their bases are fused. The foramen the foramen rotundum are fused. The temporal magnum is circular. A median transverse ridge crest is short, so that the area between the latter runs all over the occipital condyle, but there is no and the nuchal crest is flat. The external auditory axial truncation on the condyle (at least at adult pseudomeatus is partially closed. The occipital stage). side is inclined backward: the occipital condyle Two juvenile maxillae (with D4 erupting) are is anterior to the occipital crest. The nuchal also preserved (AK5-502; AK6-61). They show tubercle is poorly developed. Yet, there is a deep afew morphological features: the foramen infra- axial fossa reaching the occipital crest. The orbitalis is located above the posterior third of D2 toothrow is restricted to the anterior half of the and the nasal notch reaches the middle of D2 on skull. The thin and straight hamulus pterygoideus both specimens. The processus zygomaticus is very close to the M3. The posterior margin of maxillari begins at the level of D4. The alveolus the pterygoid is nearly horizontal. The rostral forM1 is preserved on AK5-502. The palate is end of the nasal bones is very broad and round- narrow. ed. The nasal dome and the rough vascular The only mandible referable to C . neumayri prints testify the presence of a well developed is a fragmentary mandible with an unworn median nasal horn. The nasal bones are totally d3 (i.e. new born individual; Hillman-Smith fused, but there is a shallow median groove from et al . 1986). The inferior border is convex. the tip of the nasals until the top of the horn The foramen mentale is located under the dome. They are long (about the third of the total d2/d3 limit. The sulcus mylohyoideus is very skull length). A wide and low dome on the deep. frontals further indicates the presence of a smaller frontal horn. The fronto-parietal crests Dentition are smooth and widely separate (minimum dis- Except for the complete series of the skull AK4- tance = 50 mm). The occipital crest is strongly 212 (Fig. 2A) and the milk series of the juvenile concave, nearly forked. The temporal fossa maxillae (AK5-502; AK6-61; Fig. 2B-D), dental ishugely developed, forming a platform at each remains are very rare in Akka ș dag˘ ı, with respect side of the braincase. to postcranial specimens.

GEODIVERSITAS • 2005 • 27 (4) 605 Antoine P.-O. & Saraç G.

F IG. 2. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı (Anatolia, Turkey), middle Turolian; A , upper left series (P2-M3) from the skull AK4-212, occlusal view; B - D , left juvenile maxilla with D1-D3 and erupting D4 (AK5-502); B , occlusal view; C , labial view; D ,lingual view; E - G , right astragalus (AK5-69); E , anterior view; F , posterior view; G , distal view. Scale bars: 5 cm.

606 GEODIVERSITAS • 2005 • 27 (4) Late Miocene rhinocerotids from Akka ș dag˘ ı

The upper dental formula is 3P-3M (there is between antecrochet and hypocone). There is no neither alveolus nor any trace of D1/P1 on P2). lingual groove on the protocone of M2. The The premolar series is long when compared to mesostyle is thick on M1 and M2. M3 has a tri- the molar series (L P3-4/L M1-3 ratio = 0.62). There angular outline, with fused ectoloph and meta- are no enamel foldings on the crowns. A thin loph. The protoloph is transverse, even if curved layer of cement is present on the ectolophs. backwards. There is no posterior groove on the Elsewhere, the cement is scarcely preserved. The ectometaloph. enamel is thin, wrinkled at the neck and corrugat- Lower series. The only permanent lower tooth ed on the top of the crowns (permanent and unearthed in Akka ș dag˘ ı and referred to C . neu- milk teeth). The crowns are high but neither mayri is a left p4. The ectolophid groove is deve- hypsodont nor subhypsodont ( sensu Antoine loped until the neck. The paraconid and the 2002). No isolated permanent tooth has been protoconid are angulous. Thus, the trigonid is unearthed, thus the morphology of the roots is angulous and it forms a right angle. There is no unknown. The dental structures are very simple. constriction on the metaconid and the ento- There is neither antecrochet nor anterior conid. The posterior valley is V-shaped in lingual constriction on the protoloph on upper molars view. There is neither lingual nor labial cingu- and premolars. The protoloph is curved back- lum. wards on the whole upper series. The crochet and Upper milk teeth. Two juvenile maxillae are the crista are always present. Both are simple, preserved. The first one bears D1-3 and erupting acute and sharp. The former is sagittal, while the D4 (AK5-502) whilst AK6-61 bears D3. Isolated crista is transverse. There is no medifossette, des- D1 (AK5-424) and D2 (AK2-435) have also pite the constant strong development of the cro- been unearthed. The teeth have large dimensions chet and the crista. They nearly join, especially (Appendix: Table 2). The crowns are high, but on P4. The paracone fold is present but weak. not so. The parastyle is sagittal. The metacone fold is The upper milk molars bear a mesostyle and a absent from the whole series, except on P4, where long sagittal parastyle. The mesostyle is sharper it is restricted to the basal half of the teeth. and stronger in D2. The paracone fold is thick Upper premolars. The premolars are molari- and salient while the metacone fold is lacking. form, with separate lingual cusps. There is no The protoloph is curved backwards, without labial cingulum, but a reduced lingual cingulum anterior constriction or antecrochet. The crista is is always present. The metaloph is not constrict- long and transverse, reaching the lingual half of ed. The postfossette is deep and narrow. On P2, the tooth (AK5-502). The crochet is strong and the protocone is less developed than the hypoco- straight, sagittal (D1-4), getting longer from D1 ne. The protoloph is thin but continuous and to D4. The metaloph is lacking any antero- connected with the ectoloph. The hypocone is lingual groove. There is no labial cingulum. The posterior to the metacone on P2-4. There is no lingual cingulum is variably developed: present pseudometaloph on P3. and continuous on both D2, it is weak on the D3 Upper molars. The molars are lacking labial and AK5-502 (one spur at the entrance of the median lingual cingula, except on M3, where a wide and valley). There is no trace of lingual cingulum on low tubercle is laying at the entrance of the the D3 AK6-61. The postfossette is narrow and median valley. The metastyle is long. The meta- deeper than the median valley. The posterior loph is short and the posterior part of the ecto- cingulum is low while the anterior one is thick loph is concave on M1-2. There is no cristella. and high. The posterior cingulum is low and reduced. D1 is triangular, with a lingual wall. Its postfos- There is no antero-lingual groove on the hypo- sette is open backwards. D2 has no lingual wall cone of M1-2: the metaloph is continuous. The nor mesoloph, but secondary folds and small central valley is open lingually (no junction structures (medifossette-like). D3-4 may present

GEODIVERSITAS • 2005 • 27 (4) 607 Antoine P.-O. & Saraç G.

F IG. 3. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı (Anatolia, Turkey), middle Turolian; A , right humerus, distal end (AK4- w.n.), posterior view; B , left radius, anterior view (AK5-627); C , D , right ulna (AK3-118); C , anterior view; D , medial view; E , right mag- num, lacking the posterior tuberosity (AK7-38), medial view; F , left fused tibia and fibula (AK7-40), anterior view; G , H , left calcaneus (AK7-36); G , anteromedial view; H , posterolateral view; I , J , left navicular (AK4-76); I , proximal view; J , distal view; K , left ento- cuneiform (AK13-5), anterior view. Scale bars: 5 cm. a small vertical ridge posteriorly to the crochet, weak (AK2-296). There are no vertical rough- on the posterior side of the metaloph (AK5-502). nesses, nor anterior groove nor median fold on Lower milk teeth. Three germs of d3 are pre- the ectolophid. The external groove is developed served, among which two belong to the same but smooth. The paralophid is double. It deter- individual (AK5-425 and AK6-134). The enamel mines a small and wide closed pit in front of the is corrugated. The metaconid and entoconid are tooth. Only the reduced cingula exist on the constricted. The protoconid fold is lacking or anterior and posterior faces. The posterior valley

608 GEODIVERSITAS • 2005 • 27 (4) Late Miocene rhinocerotids from Akka ș dag˘ ı

is widely open, as is the trigonid valley. There is lateral epicondyle from the trochlea. The lateral no lingual groove on the entoconid. The crown is epicondyle is the lowest tip of the humerus. moderately high. Radius (Fig. 3B). The bone is strong, with thick ends. The proximal cochlea is higher in its poste- Postcranial skeleton rior border. The proximal end is wide and deep The bones from Akka ș dag˘ ı are large and robust, (APD), much deeper medially. The anterior bor- with thick long bones and broad and low manus der is straight in proximal view. The posterior and pes. The dimensions and proportions corres- facets, for the ulna, are separate. The medial one pond essentially to those of the large/largest indi- is low (10 mm) while the lateral one is very high viduals of Ceratotherium simum as listed by and hugely developed (triangular). The insertion Guérin (1980, 2000); they even sometimes reach for the m . biceps brachii is wide and marked by the average of Rhinoceros unicornis (Guérin roughnesses, but there is no deep depression. The 1980). On the other hand, they are 5 to 30% diaphysis is straight medially and strongly conca- smaller than corresponding remains referred to ve laterally. The contact between the radius and the Miocene African rhinocerotine Dicerosaus- the ulna extends all along the diaphysis, except tralis Guérin, 2000. for a short spatium interosseum , located at the Atlas. AK5-w.n. is badly preserved, lacking the proximal third of the bone. These observations processi transversi . The bone is thick and the arti- indicate a twisted diaphysis for the ulna. In lateral cular width fits the skull AK4-212, which leads view, the diaphysis is curved, with a convex ante- us to assign it tentatively to the same individual. rior border. The distal widening is strong The foramen vertebrale has a wide piriform out- (Appendix: Table 4). No radius-ulna fusion has line. The foramen transversarium is large, only been observed in Akka ș dag˘ ı. The gutter for the connected with the foramen vertebrale. The m . extensor carpi is deep and wide, deepened by condylar facets are kidney-shaped, while the axis- the latero-distal tubercle ( tuberculum dorsale) facets are slightly concave transversally. laying close to it. The lateral ulna-facet is unique, Cervical and thoracic vertebrae. They bear no crescentiform and low. It is vertical and sagittally significant character. orientated. The distal articulation is only for the Humerus (Appendix: Table 3; Fig. 3A). This scaphoid and semilunate. The distal end is get- bone is robust. The length is approximately ting lower medially than laterally. The scaphoid- 440mm, owing to the proximal end AK6-30 and facet is visible in anterior view on a considerable the distal end AK4-w.n. The proximal end is height. This facet has a sigmoid sagittal cross thick and deep (APD), with a high and strong section. Its posterior expansion forms a high tuberculum majus. The caput humeri is wide, rounded triangle. The semilunate-facet is wide, weakly rounded. The tuberculum minus is low. concave antero-posteriorly and flat transversally. Both are separate by a shallow depression. The There is no pyramidal-facet. deltoid tuberosity is wide and high, thickly deve- Ulna (Fig. 3C, D). There is an unexpected num- loped, with a strong and extended insertion for ber of preserved ulnae (10 more or less complete the m . deltoideus. The diaphysis is narrowing specimens). They are as long as the femora, with below the deltoid tuberosity. The lateral epicon- a variable length (Appendix: Tables 5; 12). The dyle is again wide, but rather low, forming a right olecranon is thick, and developed with variable angle at its proximal end in anterior view. The shapes. It is always long, forming a closed angle trochlea is huge, especially the medial lip. The with the diaphysis. The posterior tip of the latter is much more developed (APD) than the olecranon (insertion of the m . triceps brachii) is lateral one. The median narrowing is strong. The salient with respect to the distal border of the fossa olecrani is wide and low. There is no syno- process. The humeral cochlea is wide, as high vial fossette on the antero-proximal part of the medially as laterally. The median constriction is trochlea. A shallow distal gutter separates the particularly marked. Distally to this articular

GEODIVERSITAS • 2005 • 27 (4) 609 Antoine P.-O. & Saraç G.

surface, there is a broad oval area which is not facet, for ulna, is small. So is the pisiform-facet. articulated with the humerus. It forms a shallow There is a strong tubercle on the lateral side. The depression on the whole sample. The radius- medial facets for the semilunate are not preser- facets are separate: the medial facet is vertical and ved. On the distal side, the unciform-facet is flat transversally elongated, determining a low stripe, transversally and concave antero-posteriorly. while the lateral facet is higher than wide. The Magnum (Fig. 3E). The posterior tuberosity is medial facet is transversally convex, with a trans- lacking in the three specimens (GOK-11, AK5- verse lateral end and a sagittal medial end. The 632, AK7-38). The anterior side is broad and low fossa between both facets is deep. The roughness (Appendix: Table 7), with a salient central corresponding to the contact with the radius runs tubercle ( m . interossei dorsales ). Its lateral border all along the diaphysis, except for a short spatium is oblique and straight. In proximal view, the sca- interosseum , at the proximal third of the diaphy- phoid-facet is wide and transversally concave. sis. The diaphysis is curved and twisted. Its cross The semilunate-facet has a question mark outline section is triangular, with sharp ridges. The distal in lateral view. The proximal process is high, with end widens strongly. The distal radius-facet is a small diameter, and rounded. This facet reaches low and crescentiform. The distal articulation, the anterior side of the bone. It is difficult to dis- for the carpus, shows three facets. The smallest tinguish it from the unciform-facet, which is and medial one is for the semilunate (almond- drop-shaped. On the medial side, there is a shal- shaped); the rest of the surface corresponds to the low anterior indentation between the scaphoid- pyramidal-facet (quarter circle) and the pisiform- and the McII-facets. The latter forms a flat sagit- facet. The latter is large but restricted to the tal stripe, horizontal and nearly rectangular. postero-lateral side of the articulation. It is trian- Distally, the McIII-facet is wide and saddle- gular and nearly vertical. Both are separated by a shaped, tapering backwards. smooth ridge. There is no distal tubercle ( tuber- Unciform. The unciform is well represented culum dorsale) on the antero-lateral side of the (eight specimens). It is a large bone, with a broad ulna. All these features are also shared by the anterior side and a long posterior tuberosity (low juvenile ulnae (e.g., AK6-131). and wide). The anterior side is smooth, nearly Scaphoid. Two specimens are complete. They are flat, except on the medial corner where a sharp cubic (Appendix: Table 6). The anterior and pos- horizontal tubercle is salient (insertion of the terior heights are equal. The proximal facet is m . interosseus dorsalis ). The proximal facets are deep (ADP) and concave in medial view. There separate by an acute ridge. The semilunate-facet are only two facets for the semilunate. The proxi- determines a quarter-circle. It may be transversal- mal one is flat and wide. There is a large and ly flat (AK7-147, AK4-74) or concave (GOK-10, rounded tubercle in the vicinity of the latter. On AK6-58, AKK-157). The pyramidal-facet is lar- the distal side, the magnum-facet forms an equi- ger, flat transversally and regularly convex sagit- lateral triangle. Its surface is concave antero- tally. There is a wide and short postero-lateral posteriorly and slightly convex transversally. The expansion to this facet, which generally connects trapezoid-facet is wide (TD) and saddle-shaped. it to the McV-facet (except on AK7-147). The The trapezium-facet is triangular, narrow (APD) distal facets (magnum, McIII, McIV) are not dis- and very high (Appendix: Table 6). tinct, except the most lateral one, for the McV. No semilunate, trapezium, trapezoid nor pisi- In anterior view, the latero-distal border of the form have been discovered in Akka ș dag˘ ı. bone is straight (McV-facet) while the rest of the Pyramidal. The specimen AK5-631 is badly pre- distal border is rounded. This McV-facet forms served and broken. It is low and wide. The pro- an angle about 60°with the horizontal line. This portions and structures correspond to those of orientation points out a tridactyl manus. AK5- Ceratotherium neumayri from other Turolian 633 and GOK-10 are about 25% larger than localities of Turkey (Saraç 1994). The proximal other specimens (Appendix: Table 8).

610 GEODIVERSITAS • 2005 • 27 (4) Late Miocene rhinocerotids from Akka ș dag˘ ı

McII (Fig. 4A, B). The McII is wide on all its (Appendix: Table 10). It is deeper medially. The length (Appendix: Table 9). The proximal facet, anterior border is slightly convex while the for the trapezoid, is pentagonal and saddle- medial and lateral ones are straight. The interme- shaped, transversally concave and sagittally diate relief is sharp and high, visible in anterior convex. There is no trapezium-facet. The proxi- view. It separates the trochlea into two equal mo-lateral facet (magnum) forms a sagittal stripe halves. which widens backwards, at 45° from the McIV (Fig. 4A, B). The McIV is shorter than the horizontal line. There is only one McIII-facet, McII and McIII (Appendix: Table 11). The bone the anterior one, which is triangular, vertical and is wide, without median narrowing. The proximal sagittally elongated. This facet is slightly concave. side, exclusively devoted to the unciform, forms a A large tubercle widens the proximal end on the rectangle triangle. On the medial side, the McIII- medial side. The diaphysis is wide and deep, facets are separate by a few millimetres. The ante- slightly flattened sagittally. It is nearly straight, rior one is drop-shaped, while the posterior one is only a little bit curved inwards. The insertion for circular (45° angle between them). The diaphysis the m . interossei is short. There is no distal widen- is curved laterally, at the half of the bone. The ing. The distal trochlea is roughly square in m . interosseus area reaches the half of the diaphy- distal view. Its anterior border is straight, as the sis. It is very salient, determining a large pad. The lateral border (at right angle). The intermediate diaphysis is a little flattened. The distal trochlea is relief is low but sharp and restricted to the poste- nearly square in distal view. The intermediate rior side. The medial lip is wider than the lateral relief is low but sharp and restricted to the poste- one. The latter is transversally flat, while the rior side of the trochlea. This relief is displaced former is deeply concave. The medial part is medially, so that the lateral lip is wider than the deeper (APD) than the lateral lip. medial one. The latter is transversally flat whilst McIII (Fig. 4A, B). The McIII is a wide bone, the former is concave. slightly widened distally. The proximal end is wide, Femur. AK4-253 is the only femur almost com- due to the strong lateral development of the unci- plete. The only part lacking is the trochanter form-facet. The magnum-facet is visible in ante- major . Other specimens are distal ends, except for rior view. It is regularly concave transversally and AK5-323, which is a caput femoris from a juvenile separate from the unciform-facet by a sharp ridge individual. The femur is rather slender. The head (80°angle). The latter is wide (TD), slightly is wide and hemispheric. The surface of epiphysis convex sagittally and triangular. The McII-facet is marked by a sharp transversal ridge (nor flat, has large dimensions. It is semi-circular and near- nor crescentiform). The third trochanter is deve- ly vertical. The surface for the m . interossei is res- loped but not so much. It is high (Appendix: tricted to the proximal quarter of the diaphysis Table 12) and only forming a wide stripe. The on the medial side, and it extends down to the trochanter minor determines a thin and narrow distal third on the lateral side. The McIV-facets ridge, almond-shaped in anterior view. The are large, well developed and fused on AK5-68 proximal border of the wide patellar condyle is (close in AK3-202). The anterior one is drop- curved, in anterior view. The distal end is thickly shaped and the posterior one is circular and verti- developed, with well separate condyles for the cal. It is distally displaced with respect to the tibia. proximal end of the bone. The diaphysis is Patella. The only specimen is as high as wide straight and flattened. On the anterior side, the (Appendix: Table 13). It is thick, with rounded insertion for the m . extensor carpalis is broad but structures. The APD is high. On the articular without relief. There is no distal tubercle on the side, the medial lip is large and broad. The lateral posterior side of the diaphysis. The distal trochlea one is narrower and transversally concave, as is has an antero-proximal border regularly rounded. the former. The proximal process (for the tendon In distal view, this surface is wider than deep m . quadriceps ) is strongly developed but not very

GEODIVERSITAS • 2005 • 27 (4) 611 Antoine P.-O. & Saraç G.

F IG. 4. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı (Anatolia, Turkey), middle Turolian; A , B , left metacarpus (GOK-2 to GOK-4); A , proximal view; B , anterior view; C , D , left metatarsus (GOK-7 to GOK-9); C , proximal view; D , anterior view. Scale bar:5cm. salient. The latero-proximal border of the arti- but equally high. It is oblique (45° with respect cular surface is straight. to the horizontal line). The tuberositas tibiae is Tibia. The tibia is robust, with broad ends very developed, with a broad and rough surface. (Appendix: Table 14). AK7-40 corresponds to In posterior view, the fibula is totally fused proxi- fused tibia and fibula. The contact probably mally with the tibia. Its proximal end is high but occurs at early growth stages, because all the spe- it does not bear any articular facet for the femur. cimens bear traces of the contact between both The diaphysis is narrowing in its median part, bones. The fusion may occur later, at adult stage with a triangular cross section. The spatium (Fig. 3F). The proximal condyles are well separat- interosseum is the only area where the tibia and ed by a deep intercondylar fossa. The medial the fibula are not fused. It is restricted to the condyle is sagittally flat and transversally concave second proximal quarter of the bone. The tibia – horizontal in its medial part and nearly vertical thickens strongly in its distal part (TD, APD). in its lateral tip. The lateral condyle for the femur The antero-distal groove is wide and rather deep, is convex antero-posteriorly and slightly concave running obliquely to the medial lip of the astraga- transversally. The latter is smaller (TD, APD), lus cochlea.The sulcus malleolaris is much deeper

612 GEODIVERSITAS • 2005 • 27 (4) Late Miocene rhinocerotids from Akka ș dag˘ ı

and narrower. It occurs at the posterior third of with respect to the axis of the distal articulation. the bone, in medial view. The posterior apophy- This articulation (navicular and cuboid) is well sis is rounded and low, very broad. The medial developed. The navicular-facet is lozengic, while lip of the astragalus cochlea is deeper (APD), the cuboid-facet is wide and short (oval), with a more concave (transversally and sagittally) and smooth posterior break. narrower (transversally) than the lateral one. The Calcaneus (Fig. 3G, H). The specimens are latter is almost flat transversally. robust, with a massive tuber calcanei. The fibula- Fibula (Fig. 3F). The only specimens are the facet is lacking, while the tibia-facet is hugely fused tibia/fibula (AK7-40) and a distal epiphysis developed and drop-shaped. The astragalus-facet from a young individual (AK13-3). The proximal 1 is question mark-shaped in lateral view. The end is merely crushed. The thin diaphysis, with facet 2 is oval, higher than wide and transversally sharp ridges, thickens distally. The TD is twice les- concave. It is separate from the facet 3, which is ser than the APD (Appendix: Table 15), so it smaller and almond-shaped. The sustentaculum keeps a slender shape in front view. Latero-distally, tali is thick, but not wide (TD) when compared the sulcus malleolaris is deep, with sharp ridges, to the general dimensions of the bone (Appendix: especially on the adult fibula. It is located in the Table 17). The proximal tip of the tuber calcanei posterior third of the bone. The astragalus-facet is deeper (APD) than the processus in lateral view, is flat, low, and nearly vertical. but it is still massive. The cuboid-facet, on the Astragalus (Fig. 2E-G). The astragalus is well distal side, is saddle-shaped and very wide. The represented (11 specimens). It is a large, wide and insertion for the m . fibularis longus forms a deep bone (TD/H ratio = 1.16; APD/H ratio = smooth tubercle, without any sharp ridge. 0.70; Appendix: Table 16). On the lateral side, Navicular (Fig. 3I, J). Two specimens are pre- the fibula-facet is flat and nearly vertical. It forms served (Appendix: Table 18). They have a lozen- a narrow stripe, tapering in its proximal quarter. gic but subrectangular proximal outline; so is the The trochlea is high, with acute borders. The outline of the proximal facet. The lateral facets medial lip is shorter and more convex transversal- are roughly separated. The proximal one runs all ly than the lateral one. On the medial side, the along the proximal border, while the distal one articular surface corresponding to the malleolus (hemicircular) is restricted to the posterior half of medialis is narrow and it tapers regularly proxi- the bone. The distal side bears three facets for the mally. The caudal border of the trochlea is nearly cuneiforms. The ectocuneiform-facet is L-shaped straight in proximal view. The collum tali is high and separated from the drop-shaped mesocunei- on adult specimens and even higher on juvenile form-facet by a shallow groove. The entocunei- specimens (AK2-438, AK6-56, AK5-319). The form-facet is smaller, semi-circular and oblique, medial tubercle is very high, nearly reaching the whilst the ectocuneiform- and mesocuneiform- mid-height of the bone, both in juveniles and facets are horizontal. adults. It is slightly salient from the trochlea, with Cuboid. The only specimen (AK5-636; a vertical medial border in anterior view. The Appendix: Table 19) lacks the distal part of the posterior side shows three facets for the calca- anterior side. The proximal articulation is oval neus, as usual. The facet 1 is strongly concave, and bears two facets. The medial one (astragalus) with a distal horizontal ridge determining a wide is oblique and flat transversally while the lateral latero-distal expansion. The latter forms a right one (calcaneus) is saddle-shaped: concave sagit- angle with respect to the rest of the facet, both in tally and convex transversally. They are separated juveniles and adults. The facet 2 is high, narrow, by an acute sagittal ridge. On the medial side, the and oval-shaped. Its surface is strongly convex posterior facets are not equally developed: the transversally. The facet 3 is always separate from proximal one is larger and much higher. The the facet 2. The former is wide and low, almond- preserved part of the MtIV-facet, on the distal shaped. In distal view, the trochlea is very oblique side, is flat and not elongated posteriorly. The

GEODIVERSITAS • 2005 • 27 (4) 613 Antoine P.-O. & Saraç G.

posterior tuberosity is very high, vertical and not The insertion for the m . interossei dorsalis reaches broad (TD) nor deep (APD). the distal half of the diaphysis. The diaphysis is Entocuneiform (Fig. 3K). The only specimen nearly straight, with a circular cross section. (AK13-5) is large (Appendix: Table 20), with a The distal widening is slight but present. The laterally projected distal tuberosity. The proximal distal trochlea is deeper than wide (Appendix: facet, for the navicular, is almond-shaped and Table 23), with parallel medial and lateral bor- biconcave. The ectocuneiform-facet is low and ders. In distal view, the anterior border is roun- crescentiform, while the MtII-facet is nearly cir- ded, while the posterior one is sigmoid: the cular and flat. The distal border is horizontal, intermediate relief is low and smooth, displaced except for the developed latero-distal tuberosity, into the lateral half of the trochlea. The medial which is particularly salient. The posterior side is lip is concave, with a lateral deepening, whilst the smooth, lacking any relief. lateral one is flat transversally, tapering laterally. Mesocuneiform. It is semi-circular in proximal MtIII (Fig. 4C, D). Two complete adult speci- view. The proximal facet is flat transversally and mens have been unearthed (GOK-7, AK2-163; concave sagittally. The anteromedial side is Appendix: Table 24). Another one, lacking the smooth. The entocuneiform-facet is semi- distal epiphysis, belongs to a juvenile individual circular. The ectocuneiform-facet is long (APD) (AKA-44). They have comparable structures and and low (Appendix: Table 21). The distal facet is proportions: they are rather slender, with straight flat sagittally and convex transversally, with a diaphyses. The distal widening is very slight, semi-circular outline. except for AKA-44, where it is stronger. In proxi- Ectocuneiform. The bone is low and wide mal view, the proximal end is trapezium-shaped, (Appendix: Table 22). The navicular-facet with a straight medial border, a nearly straight occupies the whole proximal side. A broad lateral border, and a convex anterior border tubercle for the m . interossei dorsales runs all along forming an open dihedron. There is no cuboid- the anterior side. In anterior view, this side has a facet. In anterior view, the proximal border is concave proximal border and a convex distal one. concave, higher laterally. The MtII-facets are On the medial side, three facets are preserved: the high, semi-circular, the anterior one being lower. antero-proximal one (for the mesocuneiform) is On the lateral side, the MtIV-facets are close to drop-shaped, with a higher posterior tip; the each other, only separate by a few millimetres. antero-distal facet (for the MtII) forms a distally The anterior one is vertical, while the posterior truncated circle; the postero-distal facet is badly one is visible in proximal view. Both are circular preserved. On the distal side, the MtIII-facet is and equally developed. The insertion for the convex transversally and flat sagittally. m . interossei is shorter for the MtII (proximal MtII (Fig. 4C, D). The proximal end is triangu- third) than for the MtIV (half of the diaphysis). lar, in proximal view. The antero-medial side is There is no posterior tubercle on the posterior lacking any proximal tubercle. The proximal side of the diaphysis. The distal trochlea is deep facet, for the mesocuneiform, is kidney-shaped. (APD) and rather narrow (TD). It is much deep- On the posterior side, the entocuneiform-facet is er medially. The intermediate relief is low and high and it forms a proximally truncated oval smooth, but it is visible in anterior view. This (AK5-440), in contact with the proximal facet. relief determines two equally developed halves. On the lateral side, there are two facets. Both are MtIV (Fig. 4C, D). There is a complete speci- high, separate from the proximal facet in AK5- men (GOK-8, from the left metatarsus) and a 440. The anterior facet is oval and vertical. The proximal end (AK5-437). It is more robust than posterior facet is rectangular and split into two that of the MtII (Appendix: Table 25). The equal halves. It is higher than the former and the proximal outline is triangular, much wider than sub-facets are rather distinct, owing to a horizontal deep. The proximal facet, for the cuboid, is trape- ridge. The distal sub-facet corresponds to the MtIII. zoid, slightly concave transversally and sigmoid

614 GEODIVERSITAS • 2005 • 27 (4) Late Miocene rhinocerotids from Akka ș dag˘ ı

sagittally. The postero-lateral tuberosity is hugely assumed as diagnostic (autapomorphies) for developed and continuous, pad-shaped. It is C . neumayri . partly destroyed on GOK-8. On the anterior According to age estimation of recent white rhi- side, the insertion for the m . fibularis is thick and nos ( C . simum ) by Hillman-Smith et al . (1986), salient. Between the cuboid-facet and the medial the juvenile maxillae belong to individuals aged facets, runs a narrow oblique sagittal stripe, for less than 1.5 year, meanwhile the adult skull cor- the ectocuneiform. A few millimetres separate the responds to a 10-15 years old individual. MtIII-facets, which are circular and equally deve- The dimensions of the remains found at loped. The diaphysis is strongly curved and Akka ș dag˘ ı and identified as C . neumayri are very distally widened. It has a semi-circular cross large (Appendix: Tables 1-25). The cranial and section and a thick antero-distal tubercle. The dental remains are equal in size and proportion to distal articulation is trapezoid in distal view, and the average of C . simum , while the postcranials fit it has a smooth and low intermediate relief; the with the large individuals of the latter taxon medial lip is flat, while the lateral lip is concave (Guérin 1980). Some bones are even larger transversely. The insertion for the m . interossei (scaphoid, McII, patella, astragalus, cuneiforms, forms a salient pad, which thickness almost MtII), reaching the average size observed in reaches 1 cm. Rhinoceros unicornis (Guérin 1980). In addition, No phalanx has been attributed to C . neumayri . several specimens of C . neumayri from Akka ș dag˘ ı exceed in size the conspecific remains from other D ISCUSSION late Miocene Eastern Mediterranean localities, Ceratotherium neumayri is very widespread in such as Samos, Maragha, Pikermi, Pentalophos-1, the late Miocene of Greece, Turkey, and Iran or Kemiklitepe (Guérin 1980, 2000; Geraads (Geraads 1988; Geraads & Koufos 1990; Saraç 1988, 1994; Geraads & Koufos 1990; direct 1994; Heissig 1996). The most abundant rhino- observation by P.-O. A.), being up to 10% cerotid taxon in Akka ș dag˘ ı bears cranial, dental, larger than the largest specimens ever described and postcranial features that are characteristic for for this species (radius, McII, astragalus, MtII; C . neumayri : dolichocephalic two-horned skull, Guérin 2000). According to Heissig (1975b) and with a very broad and rounded nasal tip, a nearly Kaya (1994), the size of C . neumayri increases flat dorsal profile, a forked occipital crest, a strong- from the late Vallesian to the late Turolian in ly inclined occipital side, long and edentulous Turkey and Greece. The same tendency might premaxillae, forwards bent processus postglenoi- exist between Kemiklitepe D (MN 11) and dalis , laterally projected orbits, short temporal Kemiklitepe A+B (MN 12), but the material is crests, teeth rows restricted to the anterior half of scant (Bonis et al . 1994; Geraads 1994). Even the skull; no anterior constriction on the proto- though latest Miocene (MN 12-13) localities cone nor antecrochet on the upper teeth, back- bearing C . neumayri are too scarce to make a wards bent protolophs, molariform upper consistent comparison, there seems to be no premolars, D1 lacking a metaloph, upper milk contradiction between the large size of C . neu- teeth with a crista; frequent absence of a trape- mayri and the late middle Turolian age of zium-facet on the McII, early fusion of tibia and Akka ș dag˘ ı (MN 12), as argued owing to the asso- fibula, high and narrow medial tubercle on the ciated fauna and radiometric ages (Karadenizli astragalus, low and transversally convex calca- etal . 2005). neus-facet 2 on the astragalus, low ectocunei- form, large articular facets between second and third metapodials, low and acute intermediate Genus Stephanorhinus Kretzoi, 1942 relieves on central metapodials; as a preliminary result of a cladistic analysis in process (Antoine T YPE SPECIES. — Stephanorhinus etruscus (Falconer, unpubl. data), these features are provisionally 1859) by original designation.

GEODIVERSITAS • 2005 • 27 (4) 615 Antoine P.-O. & Saraç G.

F IG.5. — A - E , Stephanorhinus pikermiensis (Toula, 1906), Akka ș da g ˘ ı (Anatolia, Turkey), middle Turolian; A , distal end of a right radius (GOK-14), posterior view, displaying the diagnostic horizontal ridge; B , C , right magnum, lacking the posterior tuberosity (GOK-12); B , medial view; C , anterior view; D , E , left MtIII (AKA-45); D , anterior view; E , lateral view. Notice the fusion of the MtIV- facets, visible in E; F , G , Chilotherium sp., Akka ș da g ˘ ı (Anatolia, Turkey), middle Turolian, left calcaneus (AK6-60); F , antero-medial view; G , posterior view. The Y-shaped vascular print is noticeable in G. Scale bar: 5 cm.

Stephanorhinus pikermiensis (Toula, 1906) Stephanorhinus pikermiensis – Heissig 1996: 341-343, 347, text-fig. 27.2. Rhinoceros schleiermacheri pikermiensis Toula, 1906: 34, pl. 2, fig. 2. M ATERIAL. — Left worn M1, AK5-w.n.; distal end of a right radius, GOK-14; right broken and eroded Rhinoceros schleiermacheri samius Toula, 1906: pl. 2, magnum, GOK-12; medial part of a left astragalus, fig. 3. AK6-57; left mesocuneiform, AK5-197; left MtII and MtIII from the same individual, AKA-45; (?) distal Rhinoceros ( Ceratorhinus) schleiermacheri var. orientalis end of a left MtII (distal APD = 37), AK5-2; left Schlosser, 1921: pl. 1, fig. 8. MtIII, AK3-65.

616 GEODIVERSITAS • 2005 • 27 (4) Late Miocene rhinocerotids from Akka ș dag˘ ı

D ESCRIPTION AND COMPARISON C . neumayri ). In proximal view, the scaphoid- The remains belong to at least two individuals facet is narrow. The semilunate-facet has a semi- (e.g., two left MtIII are preserved). circular outline in lateral view: there is no anterior inflection as in C . neumayri . The proxi- Dentition mal process is very large, with a huge diameter, The M1 AK5-w.n. is much worn, so most of the and rounded. This facet reaches the anterior side morphological features have disappeared. The of the bone. It is separate from the unciform- enamel is thin and wrinkled at the neck. The facet by a shallow groove. On the medial side, roots are joined, but not as thickly as it could be there is a shallow anterior indentation between expected from an old individual. There is neither the scaphoid- and the McII-facets. The former labial nor lingual cingulum. The crochet is forms a flat and oblique sagittal stripe, nearly rec- simple. The lingual cusps are separate. There is tangular. Distally, the McIII-facet is wide and neither anterior constriction isolating the proto- saddle-shaped, without posterior tapering. cone, nor antecrochet. No crista is preserved at this stage of wear. The hypocone is devoid of any Astragalus groove. This tooth (L = 49; ant. W = 74; post. The only specimen is a broken astragalus, lacking W= 65; H = 13) is very wide with respect to the the lateral part (AK6-57). The proportions are M1 of the skull AK4-212 attributed to C . neu- similar to those from C . neumayri (see above; mayri (see above; Appendix: Table 2). Indeed, Appendix: Tables 17; 28). The size is comparable these dimensions and structures are similar to the (H, APD, TD) but some features differ anyway. M1 from the skull NHM M 10144 ( S . piker- The medial lip of the trochlea is identical to that miensis , Pikermi; Geraads 1988: pl. 2, fig. C). of C . neumayri , except in the facet for the malleo- lus medialis : this articular stripe for the tibia is Radius much wider in AK6-57. It forms a broad stripe, Only a distal end is preserved (GOK-14; with a brutal thinning in its proximal third. The Appendix: Table 26; Fig. 5A). It is large, with an lip is also smoother (sharp ridge in C . neumayri ). oblique distal border in anterior view (getting The collum tali is lower than in C . neumayri , lower medially than laterally). The tuberculum especially medially, where the trochlea nearly dorsale is very reduced and smooth. Thus, the reaches the proximal tip of the navicular-facet. anterior side is flat transversely. The single lateral The medial tubercle is very different in the two ulna-facet is crescent-like, vertical and oblique taxa: it is more laterally projected, forming a sagittally. The posterior side is keeled and deep- sharp and salient tubercle in AK6-57 (thin, with ened by a wide fossa beginning 3 cm above the a vertical medial border in C . neumayri ); the most distal articulation.The scaphoid-facet is visible in striking difference concerns the height of this anterior view on a considerable height. This facet tubercle. It does not reach the distal third of the has a sigmoid sagittal cross section. Its posterior bone in AK6-57, while it reaches the half of it in expansion forms a high rounded triangle. The C . neumayri . On the posterior side, some diffe- semilunate-facet is wide, concave antero-poste- rences appear. The calcaneus-facet 2 is large and riorly and slightly convex transversally. There is circular, nearly flat, whilst it is high, oval and no pyramidal-facet. transversally strongly convex in C . neumayri . Furthermore, this facet joins the facet 3, contrary Magnum to C . neumayri , where both are separate – the The posterior tuberosity is lacking in the only astragalus-facets 2 and 3 are also separate on the specimen (GOK-12; Appendix: Table 27; calcanei attributed to C . neumayri (see above). Fig. 5B, C), which is badly damaged. The anterior On the distal side, the navicular-facet has the side is roughly square, without a salient central same outline, but its posterior tip is marked by tubercle (contrary to GOK-11, referred to a high and brutal break (AK6-57), visible in

GEODIVERSITAS • 2005 • 27 (4) 617 Antoine P.-O. & Saraç G.

anterior view. This facet is concave transversally view, the anterior border of the proximal articula- whereas it is nearly flat in C . neumayri . The tion is depressed while the lateral border is strong- cuboid-facet is broad, short, and it tapers pos- ly concave; the proximal end is as deep (APD) as teriorly in AK6-57. The distal articulation is wide (TD); the MtII-facets are high and nearly narrower (TD) in AK6-57 than in C . neumayri . joined; the MtIV-facets are fused; the distal Further comparison with 30 astragali attributed trochlea is strongly thickening medially; the to C . neumayri , three specimens identified as intermediate relief is high and acute; there is no D . orientalis (Schlosser, 1921) by Saraç (1994) distal widening of the diaphysis. from several Turolian localities of Turkey (MTA collections), and the material from Pikermi D ISCUSSION (Gaudry Collection, MNHN), confirms the Toula (1906) established Rhinoceros schleierma- consistency of these differences. Furthermore, the cheri pikermiensis on the basis of some specimens astragalus of “ D . orientalis ” from the Turolian of from Pikermi. Later, Rhinoceros ( Ceratorhinus) Shanxi (Ringström 1924: 15, text-fig. 10) is iden- schleiermacheri var. orientalis was described on tical to AK6-57, at least in anterior view (dimen- similar remains discovered in Veles, Pikermi and sions, proportions, structures). Samos (Schlosser 1921). We agree with Geraads (1988), assuming that these fossils are conspeci- Mesocuneiform fic, especially those from Pikermi, the type locali- AK5-197 has the same size as AK5-637, attribut- ty of Rhinoceros schleiermacheri pikermiensis . ed to C . neumayri (Appendix: Tables 21; 29). Thus, and following the current International The main differences consist of the triangular Code of Zoological Nomenclature (ICZN 1999), proximal outline (semi-circular in AK5-637), the the principle of priority states that the valid name presence of a tubercle on the antero-medial side is R . pikermiensis Toula, 1906. (smooth in AK5-637), the low and drop-like On the other hand, we do follow the opinion of entocuneiform-facet (semi-circular in AK5-637), Heissig (1996) and Fortelius et al . (2003) concern- and the high and short ectocuneiform-facet (low ing its assignment to the genus Stephanorhinus and long in AK5-637). Kretzoi, 1942, under the name Stephanorhinus pikermiensis (Toula, 1906), rather than to the MtII genus Dicerorhinus Gloger, 1841. The type spe- Although bearing similar dimensions with the cies of Dicerorhinus and Stephanorhinus Kretzoi, MtII referred to C . neumayri (Appendix: Tables 1942 are D . sumatrensis (Fischer, 1814) and 23; 30), the specimens (AKA-45, AK5-2) some- S . etruscus (Falconer, 1859), respectively. As a how differ from the latter: the entocuneiform- preliminary result of an unpublished cladistic facet is lower and circular; the postero-lateral analysis including most one-horned and two- facet joins the proximal facet; both anterior and horned fossil and recent rhinocerotine species posterior lateral facets are split into two equal ( sensu Antoine et al . 2003), at least six synapo- parts by a median horizontal groove; the poste- morphies differentiate the clade [ S . etruscus, rior facet is higher and the sub-facets are more S . pikermiensis ] from Dicerorhinus sumatrensis : a distinct. Both distal halves (anterior and poste- small processus paraoccipitalis , a thick protocone rior) correspond to the MtIII, thus pointing out on P2, a constricted metaloph on P2-4, a spur- the presence of high MtII-facets on the MtIII. like paralophid and a reduced paraconid on p2, and the presence of vertical roughnesses on the MtIII (Fig. 5D, E) ectolophid of d2-3 support robustly the AK3-65 and AKA-45 display similar proportions Stephanorhinus clade and prevent any generic with the MtIII referred to C . neumayri assignment of R . pikermiensis Toula, 1906 to (Appendix: Tables 24; 31). Some morphological Dicerorhinus. However, the phylogenetic rela- differences can anyway be observed: in proximal tionships of Stephanorhinus pikermiensis and

618 GEODIVERSITAS • 2005 • 27 (4) Late Miocene rhinocerotids from Akka ș dag˘ ı

Ceratotherium neumayri would probably be 1994). Fortelius et al . (2003: 291) describe a resolved by a thorough study of the whole rhino single tooth and a juvenile mandible from Loc. material from Pikermi (e.g., the Gaudry collec- 49 (Igbek, North Central Anatolia; c .9.1 Ma), tions, MNHN). referring them to “ Chilotherium cf. C . habereri ” and assuming that this isolated tooth “probably represents the Anatolian form that Heissig Tribe A CERATHERIINI Dollo, 1885 (1975a, b, 1996) referred to C . habereri ”. On the other hand, Fortelius et al . (2003: 292) consider Genus Chilotherium Ringström, 1924 that younger remains “from the Upper Kavakdere Locs. 34 and 26” [ c .8.4-8.1 Ma, i.e. slightly ear- T YPE SPECIES. — Chilotherium anderssoni Ringström, lier than Akka ș dag˘ ı] belong to an “indeterminate 1924. Chilotherium ”, rather than to Ch . habereri as pre- viously assumed by Saraç (1994). Chilotherium sp. Specific features within Chilotherium are essen- tially based on cranial, mandibular, and dental M ATERIAL. — Left calcaneus, AK6-60. comparison (e.g., Fortelius et al . 2003). As the available material from Akka ș dag˘ ı is restricted to D ESCRIPTION a single calcaneus (AK6-60) and pending a revi- Calcaneus sion of the whole genus, we provisionally identify AK6-60 shares most of the characters with the it as Chilotherium sp., following the proposition two calcanei attributed to Acerorhinus sp. (AK6- of Fortelius et al . (2003). 130, AK7-98; see below), but it is thicker, shorter and smaller (Appendix: Tables 32; 35). AK6-60 bears a large and concave fibula-facet; the cuboid- Genus Acerorhinus Kretzoi, 1942 facet is circular and saddle-shaped, with a poste- ro-lateral salient tip. The presence of deep T YPE SPECIES. — Acerorhinus zernowi (Borissiak, vascular prints on the postero-distal side of the 1914). processus calcanei further distinguishes this speci- men from the other ones unearthed at Akka ș dag˘ ı Acerorhinus sp. (Fig. 5F, G). M ATERIAL. — Distal part of a left humerus, AKB-47; distal part of a left humerus, AK5-1; distal part of a D ISCUSSION right humerus, AK6-88; right radius, AK5-629; left All these features (size, proportions, thickness, calcaneus, AK7-98; right calcaneus, AK6-130; right fibula-facet and vascular prints) are common MtII, AK5-678. with the calcaneus 06-AKK-019 from Kavakdere (MN 12; Turkey), referred to Ch . habereri D ESCRIPTION Ringström, 1924 by Saraç (1994: pl. 14, fig. 1). Humerus Both specimens belong probably to the same Three distal parts have been unearthed. The taxon. Ch . habereri is a small and robust bones seem to be rather slender and long Chilotherium species, with strongly shortened (Appendix: Table 33). The diaphysis is triangular limbs and hypsodont teeth, originally described in cross section, below the lacking deltoid tubero- in the early Turolian of China (Ringström 1924; sity. The lateral epicondyle is high and wide, with Heissig 1975a). Some Turkish remains, ranging sharp ridges. In anterior view, its lateral border from the late Vallesian up to the middle forms a sharp dihedron (130°), with a proximal Turolian, were attributed to this species in the tubercle oriented upward. The trochlea is last decades (Kayadibi and Garkın faunal sets: narrow, with a broad (TD) lateral lip. The Heissig 1975a, 1996; Kınık faunal set: Saraç medial one is regularly conical, except for its last

GEODIVERSITAS • 2005 • 27 (4) 619 Antoine P.-O. & Saraç G.

centimetre, where it further tapers. The median (Appendix: Table 35). The tuber calcanei is trian- strangulation is not very marked. The trochlea is gular and massive. The insertion for the m . fibu- eggcup-like. The lateral lip is first conical, laris longus is salient. The sustentaculum tali is widening laterally, and then it tapers until its broad (TD > APD) and thick. The astragalus- lateral tip. The frontier between these two parts is facet 1 is flattened in its proximal part. A distal smooth. There is no synovial fossa on the tro- circular expansion is always present, separate chlea. The fossa olecrani is wide and low. A distal from the main facet 1 by a shallow groove. There gutter isolates the lateral epicondyle. In anterior is no fibula-facet, but a large tibia-facet on both view, the distal end is lower on the lateral side. specimens (AK6-130; AK7-98). The astragalus- facets 2 and 3 are widely separate. The facet 2 is Radius oval and concave. The facet 3 forms a narrow AK5-629 is complete. It is high and slender, stripe. The distal facet (for the cuboid) is saddle- thickening downward (Appendix: Table 34). In shaped, but the postero-lateral tip is broken on proximal view, the proximal articulation has a both specimens. straight anterior border. This articulation is sepa- rate into two equally wide cochleae, the medial MtII one being deeper (APD). The frontier between AK5-w.n. is rather slender, with a slightly curved both cochleae is a smooth ridge, slightly higher diaphysis and distal widening. The proximal posteriorly. The insertion for the m . biceps bra- end is hemicircular in proximal view. The meso- chii is developed, but shallow, and medially dis- cuneiform-facet occupies most of the surface. It placed. The ulna-facets are vertical and widely is also hemicircular and nearly flat. The ento- separate. The medial one is low, crescent-like. cuneiform-facet is high and narrow, in contact The lateral one is higher, but still small. The dia- with the mesocuneiform-facet. On the lateral physis is twisted, with a convex medial border side, the anterior facet is broken; the posterior and a concave lateral border. The posterior side one is circular and split into two equivalent sub- of the diaphysis shows a long trace corresponding facets. The proximal subfacet corresponds to the to the contact with the ulna. It is essentially ectocuneiform and the distal one to the MtIII. concentrated in the distal half of the bone, down The insertion for the m . interosseus is very long. from the spatium interosseum . This surface forms It nearly reaches the bottom of the diaphysis. a rough triangle. The diaphysis widens distally, An oblique groove runs on the antero-lateral but it does not get deeper (APD). There is only side of the diaphysis. The distal trochlea is deep- one distal ulna-facet, almond-shaped. The gutter er than wide (APD > TD; Appendix: Table 36), for the m . extensor carpi is shallow. There is no wider posteriorly. The depth is equal medially tuberculum dorsale laying on the anterior side. and laterally. The intermediate relief is nearly The distal end is lower on the medial side. The absent but it determines a large medial lip and a scaphoid-facet is just a little visible in anterior narrow lateral lip. The former is slightly concave. view. The distal articulation bears three facets. The scaphoid-facet is not deep (APD), but strong- D ISCUSSION ly furrowed. The median one, for the semilunate, These postcranial specimens display characteristic is transversally flat and sagittally concave. The traits of several late Miocene aceratheriine genera pyramidal-facet forms a long and straight oblique from Eurasia, i.e. Alicornops Ginsburg & Guérin, stripe. 1979, Acerorhinus Kretzoi, 1942,and Chilo- therium (Antoine et al . 2003). Their representa- Calcaneus tives are convergent with Neogene teleoceratines The bone is very thick and robust, with a very in that they have acquired independently more or short tuber and processus calcanei. The dimensions less shortened limbs (“hippo-like”; e.g., Heissig are variable but the proportions are similar 1999; Antoine et al . 2003).

620 GEODIVERSITAS • 2005 • 27 (4) Late Miocene rhinocerotids from Akka ș dag˘ ı

The relatively slender proportions of these seven Qiu& Qiu 1995; Cerdeño 1996; Deng 2000; postcranials – which are likely to document a Fortelius et al . 2003). single taxon – prevent any assignment to Chilotherium , the most “hippo-like” aceratheriine genus. In return, these proportions and dimen- CONCLUSION sions (Appendix: Tables 33-36) recall those observed in Acerorhinus (Ringström 1924; Among the four rhinocerotid species recognised Cerdeño 1996; Fortelius et al . 2003),and at a in Akka ș dag˘ ı, C . neumayri is by far the most lesser degree Alicornops (Guérin 1980, 1988; abundant (114 specimens, at least 11 indivi- Cerdeño 1997; Cerdeño & Sanchez 2000; duals). Besides, these specimens are among the Antoine et al . 2003). For instance, the humeri largest ones attributed so far to C . neumayri . The and the radius from Akka ș dag˘ ı are strongly simi- comparison with other Turolian rhinocerotid lar to the specimens referred to Acerorhinus zerno- faunas from Eastern Mediterranean reveals wi (Cerdeño 1996) but much larger than the astrong similarity between Akka ș dag˘ ıand largest ones identified as Al . simorrense and Al . Kavakdere (Anatolia, MN 12; Saraç 1994; complanatum (Appendix: Tables 33; 34; e.g., Fortelius et al . 2003), with three out of four taxa APDs). The calcanei are wider and more massive in common ( C . neumayri , S . pikermiensis , than any calcaneus assigned so far to Alicornops or Chilotherium sp.). Furthermore, the large size of to Acerorhinus, which is especially visible on the C . neumayri is consistent with the Ar/Ar dating tuber calcanei (Appendix: Table 35). However, for the Akka ș dag˘ ı tuff (7.1 ±0.1 Ma; Karadenizli the closest affinities exist with A . zernowi et al . 2005). (Cerdeño 1996) and “ Alicornops ” alfambrense Cerdeño & Alcalá, 1989, the generic assignment of which has already been challenged by Antoine Acknowledgements et al . (2003), who tentatively referred it to The authors are grateful to Geneviève Bouvrain, Acerorhinus. In that respect, the characteristics of Emile Heintz, Léonard Ginsburg, Louis de the MtII AK5-w.n. (morphology, proportions, Bonis, and all the people who participated in the and dimensions) are quite puzzling, in that they 2000-2001 excavations. The MNHN, CNRS match perfectly those of Alicornops simorrense (ECLIPSE Program and DRI), MTA, and (Appendix: Table 36). The MtII is at the same TUBITAK have financially supported the field- time more robust than those of A . zernowi work, excavations, and reciprocal visits to the (middle Miocene; Cerdeño 1996) and much concerned institutions for the authors. The more slender than the McIV assigned to authors warmly thank Sevket Sen for having Acerorhinus sp. by Fortelius et al . (2003: 289, invited them to dig and study such tremendous fig. 12.6; c .8.1 Ma). The trend toward shorter specimens. This article has widely benefited from metapodials is generalised – though probably fair and constructive criticisms by Claude Guérin independently acquired – within Chilotherium and Mikael Fortelius. The photographs were and Acerorhinus (Fortelius et al . 2003). Thus, and taken by Denis Serrette (MNHN). in the case of a linear trend, such morphology for the MtII would rather point to an intermediate age for Akka ș dag˘ ı. REFERENCES However and pending new discoveries – notably associated cranial, mandibular and/or dental A NTOINE P.-O. 2002. — Phylogénie et évolution des material in Akka ș dag˘ ı –, we propose to assign Elasmotheriina (Mammalia, Rhinocerotidae). these postcranials to sp. It should be Mémoires du Muséum national d’Histoire naturelle Acerorhinus 188: 1-359. added that Akka ș dag˘ ı counts among the latest A NTOINE P.-O., D URANTHON F. & W ELCOMME J.-L. occurrences of Acerorhinus (MN 7-MN 12; 2003. — Alicornops (Mammalia, Rhinocerotidae)

GEODIVERSITAS • 2005 • 27 (4) 621 Antoine P.-O. & Saraç G.

dans le Miocène supérieur des Collines Bugti G ERAADS D. & K OUFOS G. 1990. — Upper Miocene (Balouchistan, Pakistan) : implications phylogéné- Rhinocerotidae (Mammalia) from Pentalophos-1, tiques. Geodiversitas 25 (3): 575-603. Macedonia, Greece. Palaeontographica, A , Stuttgart A YAN M. (1963). — Contribution à l’étude pétro- 210 (4-6): 151-168. graphique et géologique de la région située au nord- G UÉRIN C. 1980. — Les Rhinocéros (Mammalia, est de Kaman (Turquie). Mining Research and Perissodactyla) du Miocène terminal au Pléistocène Exploration Institute of Turkey, Special Publications , supérieur en Europe occidentale. Comparaison avec Ankara 115: 1-332. les espèces actuelles. Documents du Laboratoire de B ONIS L. DE, B OUVRAIN G., G ERAADS D. & K OUFOS Géologie, Université de Lyon, Sciences de la Terre 79, G. D. 1992a. — Multivariate study of late 3 volumes: 1-1184. Cenozoic mammalian faunal compositions and G UÉRIN C. 1988. — 6 - Périssodactyles Rhino- paleoecology. Paleontologia i Evolucio, Sabadell cerotidae. Palaeovertebrata , Mémoire Extraordinaire 24-25: 93-101. « Contributions à l’étude du gisement miocène B ONIS L. DE, B RUNET M., H EINTZ E. & S EN S. supérieur de Montredon (Hérault). Les grands mam- 1992b. — La province gréco-irano-afghane et la mifères »: 97-134. répartition des faunes mammaliennes au Miocène G UÉRIN C. 2000. — The Neogene rhinoceroses of supérieur. Paleontologia i Evolucio, Sabadell 24-25: Namibia. Palaeontologia africana 36: 119-138. 103-112. G RAY J. E. 1867. — Observations on the preserved B ONIS L. DE, B OUVRAIN G., G ERAADS D., K OUFOS G. specimens and skeletons of the Rhinocerotidae in D., S EN S. & T ASSY P. 1994. — Les gisements de the collections of the British Museum and Royal mammifères du Miocène supérieur de Kemiklitepe, College of Surgeons, including the description of 3 Turquie : 11. Biochronologie, paléoécologie et rela- new species. Proceedings of the Zoological Society of tions paléobiogéographiques. Bulletin du Muséum London: 1003-1022. national d’Histoire naturelle, Paris, 4 e sér., section C, H EISSIG K. 1972. — Paläontologische und geologische 16 (1): 225-240. Untersuchungen im Tertiär von Pakistan. 5. C ERDEÑO E. 1996. — Rhinocerotidae from the Rhinocerotidae (Mamm.) aus den unteren und mit- Middle Miocene of the Tung-gur Formation, Inner tleren Siwalik-Schichten. Bayerische Akademie der Mongolia (China). American Museum Novitates Wisseschaften, Mathematisch-Naturwissenschaftliche New York 3184, 43 p. Klasse Abhandlungen 152: 1-112. C ERDEÑO E. 1997. — Rhinocerotidae from the H EISSIG K. 1975a. — Rhinocerotidae aus dem Turolian site of Dorn-Dürkheim 1 (Germany). Jungtertiär Anatoliens. Geologisches Jahrbuch B.15: Courier Forschungsinstitut Senckenberg 197: 145-151. 187-203. H EISSIG K. 1975b. — Rhinocerotidae aus dem C ERDEÑO E. & A LCALÁ L. 1989. — Aceratherium Jungtertiär Anatoliens . Munich, Institut für alfambrense sp. n., nuevo rinocerótido del Paläontologie und historische Geologie der Vallesiense Superior de Teruel (España). Revista Universität München. Unpublished manuscript, Española de Paleontologia 4: 39-51. 600 p. C ERDEÑO E. & S ÁNCHEZ B. 2000. — Intraspecific H EISSIG K. 1996. — 27. The stratigraphical range of variation and evolutionary trends of Alicornops fossil rhinoceroses in the Late Neogene of Europe simorrense (Rhinocerotidae) in Spain. Zoologica and the Eastern Mediterranean, in B ERNOR R. L., Scripta 29 (4): 275-305. F AHLBUSCH V. & M ITTMANN H.-W. (eds), The D ENG T AO 2000. — A new species of Acerorhinus Evolution of Western Eurasian Neogene Mammal (Perissodactyla, Rhinocerotidae) from the late Faunas . Columbia University Press, New York: Miocene in Fugu, Shaanxi, China. Vertebrata 339-347. PalAsiatica 38 (3): 203-217. H EISSIG K. 1999. — 16. Family Rhinocerotidae, in F ORTELIUS M., H EISSIG K., S ARAÇ G. & S EN S. 2003. R ÖSSNER G. E. & H EISSIG K. (eds), The Miocene — Rhinocerotidae (Perissodactyla), in F ORTELIUS Land Mammals of Europe. Pfeil, Munich: 175-188. M., K APPELMAN J., S EN S. & B ERNOR R. L. (eds), H ILLMAN-S MITH A. K. K., O WEN-S MITH N., Geology and Paleontology of the Miocene Sinap A NDERSON J. L., H ALL-M ARTIN A. J. & S ELALADI Formation, Turkey . Columbia University Press, J.P. 1986. — Age estimation of the White rhino- New York: 282-307. ceros ( Ceratotherium simum ). Journal of Zoology , G ERAADS D. 1988. — Révision des Rhinocerotinae London 210: 355-379. (Mammalia) du Turolien de Pikermi. Comparaison ICZN 1999. — International Code of Zoological avec les espèces voisines. Annales de Paléontologie 74: Nomenclature . 4th ed. The International Trust for 13-41. Zoological Nomenclature, London, 306 p. G ERAADS D. 1994. — Les gisements de mammifères K ARADENIZLI L., S EYITOG ˘ LU G., S EN S., A RNAUD N., du Miocène supérieur de Kemiklitepe, Turquie : 4. K AZANCı N., S ARAÇ G. & A LÇIÇEK C. 2005. — Rhinocerotidae. Bulletin du Muséum national d’His- Mammal bearing late Miocene tuffs of the toire naturelle, Paris, 4 e sér., section C, 16 (1): 81-95. Akka ș dag˘ ı region; distribution, age, petrographical

622 GEODIVERSITAS • 2005 • 27 (4) Late Miocene rhinocerotids from Akka ș dag˘ ı

and geochemical characteristics, in S EN S. (ed.), O SBORN H. F. 1900. — Phylogeny of Rhinoceroses of Geology, mammals and environments at Akka ș dag˘ ı, Europe. Memoirs of the American Museum of late Miocene of Central Anatolia. Geodiversitas Natural History, New York 13: 229-267. 27 (4): 553-566 (this volume). R INGSTRÖM T. J. 1924. — Nashörner der Hipparion- K AYA T. 1994. — Ceratotherium neumayri (Rhino- Fauna Nord-Chinas. Geological Survey of China , cerotidae, Mammalia) in the Upper Miocene of Series C 11 (4): 1-156. Western Anatolia. Turkish Journal of Earth Sciences S ARAÇ G. 1994. — [ The Biostratigraphy and 3: 13-22. ˘ Palaeontology of the Rhinocerotidae (Mammalia- K AZANCı N., S EN S., S EYITOG LU G., B ONIS L. DE, Perissodactyla) of the Continental Neogene Sediments B OUVRAIN G., A RAZ H., V AROL B. & K ARADENIZLI in the Ankara Region]. Ph.D. Thesis,Ankara L. 1999. — Geology of a new Late Miocene mam- University, Turkey, 214 p. (in Turkish). mal locality in central Anatolia, Turkey. Comptes Rendus de l’Académie des Sciences , Sciences de la S CHLOSSER M. 1921. — Die Hipparionenfauna von Terre et des Planètes, Paris 329: 503-510. Veles in Mazedonien. Abhandlungen der K OUFOS G. D. & V LACHOU T. D. 2005. — Equidae Bayerischen Akademie der Wissenschaften, Munich (Mammalia, Perissodactyla) from the late Miocene 29 (4): 1-55. of Akka ș dag˘ ı, Turkey, in S EN S. (ed.), Geology, T OULA F. 1906. — Das Gebiss und Reste der mammals and environments at Akka ș dag˘ ı, late Nasenbeine von Rhinoceros ( Ceratorhinus Osborn) Miocene of Central Anatolia. Geodiversitas 27 (4): hundsheimensis . Abhandlungen der Kaiser-Königlichen 633-705(this volume). Geologische Reichs-Ausz., Vienna 20: 1-38.

Submitted on 7 August 2003; accepted on 25 May 2004.

GEODIVERSITAS • 2005 • 27 (4) 623 Antoine P.-O. & Saraç G.

APPENDIX Abbreviations: AK, 2000-2001 excavations; AKA and AKB, 1997 excavation; AKK, surface collects; ant. , anterior; APD, antero-pos- terior diameter; astrag., astragalus; art. , articulation; Calc. , calcaneus; Cub. , cuboid; D , distance (between two elements); dia. , dia- physis; diag., diagonal length; delt. , deltoide; dist. , distal; Ectocun., ectocuneiform; Entocun., entocuneiform; epiph., epiphysis; ext. , extremity; fac. , facet; GOK, Heintz-Ginsburg excavation (formerly Gökes, me); H , height; j , juvenile; l ,left; L , length; lat. , lateral; mag., magnum; max. , maximum; med., medial; mesocf., mesocuneiform; mid., middle; min., minimum; Navic. , navicular; post. , posterior; prox. , proximal; Pyram., pyramidal; r , right; Rad., radius; sust. , sustentaculum; S.-L. , semilunate; TD, transverse diame- ter; Trap., trapezium; Trapzd., trapezoid; trochl., trochlea; tuber. , tuberosity; Uncif., unciform; W ,width (when different from TD); w.n., without number. Values between brackets are doubtful.

T ABLE 1. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, T ABLE 2. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the adult skull middle Turolian of Central Anatolia, dental measurements AK4-212 (in mm). (in mm).

Specimen Lant. Wpost. WH Length occipital crest/tip of nasals755 Length occipital crest/tip of premaxilla770 l P242393835 Length occipital crest/caudal end of M3380 l P348 54 51 40 Length occipital condyle/front of orbit390 l P4515952.5 49 l M15862.5 57.5 42 Length occipital condyle/tip of premaxilla690 l M26466 54 65 Length of nasal notch140 l M354 60 61.5 (l max.) >60 Maximum zygomatic width345 r P2AK4-21240393934 Maximum frontal width235 r P349 53.5 5042 Occipital crest width234 r P45160 53.5 50 r M15863 59 43 Occipital condyle width130 r M26567 55 > 55 Foramen magnum width46 r M3536162.5 (l max.) > 60 Palate width (at P2 level) 56 r M1AK2-294 48 63 57.5 8 Palate width (at P4-M1 level) 80 l p4 AK2-295 43263138 Palate width (at M3 level) 67 r D1AK5-42424.5 18 24.5 21 Occipital face height220 l D12420 2525 l D2AK5-502 43.5 37 38.5 33 Length of P2-M3 (left/right)294/292 l D35045 44 42 Length of P2-P4 (left/right)136/137 r D3AK6-615047.5 45 34 Length of P3-P4 (left/right)105/104 l d3AK2-29642–– – Length of M1-M3 (left/right)170/166 r d3AK6-13444.5 18.5 21.5 31

T ABLE 3. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the humeri (in mm).

Number prox. ext.tuber. delt.diaphysisdist. ext.APD trochl.

TD APD TD TD min. APD TD APD med. mid.

AK4-w.n. ––(> 150)7977 (160)139120 (58) AK6-301197158 160 73 72 –––– AK-w.n. ––(> 138) 77 –168–10553 AK4-183–––5756––––

624 GEODIVERSITAS • 2005 • 27 (4) Late Miocene rhinocerotids from Akka ș dag˘ ı

T ABLE 4. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the radii (in mm).

NumberLprox. ext.prox. art.diaphysisdist. ext.dist. art.

TD APD TD APD APD APD TD APD TD APD TD APD med. mid. lat.

GOK-18/19 > 405133 88 12573 50(48) 7550127 84 11253 GOK-16 (j) –– –– –––––108758949 GOK-17 (j) –– –– –––––107 (73)9149 AK4-w.n. 4121258612181(54) (48) 70 40126 8311653 AK4-213–>123 –(> 119) ––––––––– AK4-241 –119 (74) 116–––6845– ––– AK5-67 –123 (71) 118 644848–37 –––– AK5-w.n. 380120 79116765045 6539114 70 99 52

T ABLE 5. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the ulnae (in mm). Abbreviation: L*, L without olecranon.

Number L L* olecranon humerus cochlea diaphysis dist. ext. Pyram. fac.

TD HTD HHHTD APD TD APD TD APD max.med. mid. lat.

AK4-w.n. ––––1137544825359 –––– AK6-132 ––801171127744 825259 –––– AK6-133 470 40891112107 63 4378515962 60 45 58 AK6-131––––(> 87)62 43–3943–––– AK7-156–––(120)(> 112)70 4281 5258 –––– AK3-118 497430 95 126 121754083526267654457 AK4-184 –423 ––10575–81435654 694157 AK6-302 –385 ––107 7537 77 465757624253 AK2-95 ––––(110)77 4082–––––– AK6-303 –––120 10473 41 81 ––––––

T ABLE 6. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the scaphoids (in mm).

NumberTD APD HRad. fac.Trap. fac. Trapzd. fac. mag. fac.D S.-L. fac.

ant.mid. post.APD HAPD TD APD TD APD

AK2-436 –8865496556281438.5 33.5 3133 (13) AK5-630 6586674762532914393430 3510

Number TD H ant.HS.-L.McIII fac. fac.

APD TD APD

GOK-11 58 45 72605352 T ABLE 7. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, AK7-3851.5 426862 (38) (57) middle Turolian of Central Anatolia, dimensions of the magnums (in mm). AK5-632 54 4267–49–

GEODIVERSITAS • 2005 • 27 (4) 625 Antoine P.-O. & Saraç G.

T ABLE 8. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the unciforms (in mm).

NumberTD HAPD max.APD post. tuber.S.-L. fac.Pyram. fac.McV fac.

TD HTDAPD TD APD TD APD

GOK-108372––––433950502331 AK5-633 (76)73 109100 45 29–4448.5 42–40 AK6-158 76 69– –––37 3847(40)– – AKK-1577462 ––––36 37 44 44 –– AK4-7475659576 37 2534384245 2537 AK7-1477567 9779382533 38384122 36 AKK-156––––36 26 –––––– AKK-280––––(37)(24) ––––––

T ABLE 9. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the McII (in mm).

NumberLprox. ext.Trapzd. fac.H lat. fac.diaphysisdist. art.

TD APD TD APD ant.mid. post.TD APD TD APD

GOK-2167 5257.5 33 51 27 20 20 46275048 GOK-61685257355128181848285047 AK6-55 17953573552–1719 51 285048 AK7-37 –5056355422 15 20 51 24– – AK5-182–44523248 14 12123821– –

T ABLE 10. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the McIII (in mm).

NumberLprox. art.D fac. Uncif. fac.mag. fac.diaphysisdist. ext.dist. art. McIV

TD APD TD APD TD APD TD APD TD TD APD

GOK-1 194 7559652602833 63 30 76 60 51 GOK-4 1937560 85258 283465317860 53 AKB-83––––––––––8259 55 AK5-681907455248 532937 58.5 2876.5 60 51 AK3-202 ––58 7––––62 27 ––– AK14-w.n. ––––––––59 28– ––

T ABLE 11. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the McIV (in mm).

NumberLprox. art.Uncif. fac.McV fac.diaphysisdist. art.

TD APD TD APD HAPD TD APD TD APD

GOK-3158 575649 471020463045 43 GOK-5 159 58 55 49 4711 –46304644 AK3-230 14249 48 40467244523 4643 AK5-436 144 48 504048 725432344 44 AK13-2–515443529244122 ––

626 GEODIVERSITAS • 2005 • 27 (4) Late Miocene rhinocerotids from Akka ș dag˘ ı

T ABLE 12. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the femora (in mm).

NumberLtrochl. 3rd trochl. diaphysisdist. ext.condyles major H

TDdia.HTDAPD TD APD max. TD D

AK4-253490–(> 150)1109763159 (160)13127 AK5-w.n. ––1439384 63 –––– AK6-153––138115 7859155 (155) 12519 AK7-39–101(> 125) 106 94 61– ––– AK3-63 –––––––183––

T ABLE 13. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the patella (in mm).

NumberTD APD Hart.TD med. art.lat.art.

max.min. TD HTDH

AK11-82117.5 72 53116113100 73 4081

T ABLE 14. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the tibias (in mm).

NumberLprox. ext.diaphysisdist. ext.astrag. cochlea

TD APD TD APD TD APD TD APD med. mid. lat.

AK7-40370 126 1426156102 8685 63 4659 AK5-366 –––7466 –88–6950– AK7-63 –––6959115 9092625068 AKK-154 –––––111 9387––(60)

T ABLE 15. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the fibulae (in mm).

NumberLprox. ext.diaphysisdist. ext.Tibia fac.astrag. fac.

TD APD TD APD TD APD APD HAPD H

AK7-40310284626303058 54 21– – AK13-3 (j) –––––26 52––44 19

T ABLE 16. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the astragali (in mm).

NumberTD TD APD HCalc. fac. 1Calc. fac. 2 dist. art. max.trochl. max.

med. mid. lat.TD HTDHTD max.APD

AK5-w.n. 108100 (> 63)92779753523245 9765 AK4-75106 96.5 6585– –51522738– –

GEODIVERSITAS • 2005 • 27 (4) 627 Antoine P.-O. & Saraç G.

T ABLE 16. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the astragali (in mm) (continuation).

NumberTD TD APD HCalc. fac. 1Calc. fac. 2 dist. art. max.trochl. max.

med. mid. lat.TD HTDHTD max.APD

AK5-423 119 104699379100 53603145–58 AK11-1 106 98 658876 9348 5232469260 AK7-148 ––––––––(32)48– – AK6-56(j) –86.5 54 73 687846422742(> 77)54 AK5-319 (j) –––––8248 52–––– AK5-6911394 64847494465328449255 AK5-63410193(63)807186475331498956 AK13-4 –––83––––––––

T ABLE 17. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the calcanei (in mm).

NumberHHart.tuber.beaksust.min. min. astrag. Cub. TD APD fac. 3 fac. post.post.

TD APD APD TD TD HTDH

AK7-36 137 7460 77 748442624011 44 32 AK3-66 136.5 67 63 74– –4967 ––––

T ABLE 18. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, T ABLE 19. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the naviculars middle Turolian of Central Anatolia, dimensions of the cuboid (in mm). (in mm).

NumberTD APD Hprox. NumberTD Hprox. med. art. art.face

ant.mid. post.APD ant.post.ant.post.TD APD DH ant.post. AK4-76 5682312537 68 AK5-439557836 27 36 63 AK5-636 45 54 –70 49 51 –38

T ABLE 20. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, T ABLE 21. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the ento- middle Turolian of Central Anatolia, dimensions of the meso- cuneiform (in mm). cuneiform (in mm).

NumberTD APD HNavic. fac.D fac. NumberTD APD HEctocun. fac.

TD APD ant. APD H

AK13-5 562366291911 AK5-637 2445.5 22 246

628 GEODIVERSITAS • 2005 • 27 (4) Late Miocene rhinocerotids from Akka ș dag˘ ı

T ABLE 22. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the ecto- cuneiforms (in mm).

NumberTD APD HNavic. fac.

TD APD

GOK-1358 532953(52) AK4-77 ––28– – AK5-63565(> 60)295652

T ABLE 23. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the MtII (in mm).

NumberLprox. art.mesocf. fac.lat. fac.diaphysis dist. art.

TD APD TD APD H ant.H post.DTDAPD TD APD

GOK-9 163 3553254322221332263745 AK5-40–3548214118–10––––

T ABLE 24. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the MtIII (in mm).

NumberLL without prox. art.TD dia.diaphysisTD max. dia.dist. art. dist. epiph.

TD APD TD APD TD APD

GOK-7183–66 61385528755951 AKA-44 (j) –146.5 63 60 37 49 28(69) –– AK2-163 175– 64594453.5 29695648 AK4-238– –––– –– –6153

T ABLE 25. — Ceratotherium neumayri (Osborn, 1900), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the MtIV (in mm).

NumberLprox. ext.prox. art.med. fac.diaphysisTD dist. max. dia.art.

TD APD TD APD APD HAPD HTDAPD TD APD ant.ant.post.post.

GOK-8 163 58 48 51 41 26 252120 37 29463446 AK5-437 –5050473721.5 21–20 –––––

T ABLE 26. — Stephanorhinus pikermiensis (Toula, 1906), T ABLE 27. — Stephanorhinus pikermiensis (Toula, 1906), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the radius (in mm). the magnum (in mm).

Numberdist. ext.dist. art. NumberTD H ant.HS.-L. fac.McIII fac.

TD APD TD APD APD TD APD

GOK-14 100 688645 GOK-12(43)33 63 60 (42)(53)

GEODIVERSITAS • 2005 • 27 (4) 629 Antoine P.-O. & Saraç G.

T ABLE 28. — Stephanorhinus pikermiensis (Toula, 1906), T ABLE 29. — Stephanorhinus pikermiensis (Toula, 1906), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the astragalus (in mm). the mesocuneiform (in mm).

NumberAPD H dist. art. NumberTD APD HEctocun. Entocun. max. fac.fac.

med. mid. lat.TD max.APD APD HTDH

AK6-57658871–8753 AK5-1972645 23 25820 6

T ABLE 30. — Stephanorhinus pikermiensis (Toula, 1906), Akka ș da g ˘ ıı, middle Turolian of Central Anatolia, dimensions of the MtII (in mm).

NumberLprox. art.mesocf. fac.lat. fac.diaphysisdist. art.

TD APD TD APD H ant.H post.DTDAPD TD APD

AKA-45 16437 51 22 422224734.5 24.5 41.5 50

T ABLE 31. — Stephanorhinus pikermiensis (Toula, 1906), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the MtIII (in mm).

NumberLprox. art.TD diag. diaphysisTD max. dia.dist. art.

TD APD TD APD TD APD

AKA-45 181 62 55 4051 2467 5649 AK3-65161595338492567 5244

T ABLE 32. — Chilotherium sp. (Schlosser, 1903), Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the calcaneus (in mm).

NumberHHart.tuber.sust.min. TD astrag. Cub. fac. TD post.fac. 3

TD APD TD HTDH

AK6-60 94 56476374382562342

T ABLE 33. — Acerorhinus sp., Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the humerus (in mm). Comparison with other late Miocene aceratheriines from Eurasia (data from Cerdeño [1996], Cerdeño & Sanchez [2000] and Fortelius et al. [2003]), for which values correspond to size range and numbers between brackets indicate the size of the sample.

Taxon Numberdiaphysisdist. ext.trochl. APD trochl.

TD min. APD TD APD TD med. mid. lat.

Acerorhinus sp. AKB-475252124–88 73 3953 Acerorhinus sp. AK5-1 5662126 95 91 7143.5 59 Acerorhinus sp. AK6-88 ––––(90)8045 – Acerorhinus –49.2-58.253.4 111-126.4 93.3 84-88.271.1-78.2–– zernowi (3)-67.1 (3)(3)-110 .8 (3)(3)(3) Alicornops –40-50 39-57 72-97.6 63.5-76.7 –––– simorrense (4) (4) (5) (4) cf. Chilotherium AS.90.1345358 1288992––– sp. (primitive)

630 GEODIVERSITAS • 2005 • 27 (4) Late Miocene rhinocerotids from Akka ș dag˘ ı

T ABLE 34. — Acerorhinus sp., Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the radius (in mm). Comparison with other late Miocene aceratheriines from Eurasia (data from Ringström [1924], Heissig [1972], Guérin [1980, 1988], Cerdeño [1996], Cerdeño & Sanchez [2000], and Fortelius et al. [2003]), for which values correspond to size range and numbers between brackets indicate the size of the sample.

Taxon NumberLprox. ext.diaphysisdist. ext.dist. art.

TD APD TD APD TD APD TD APD

Acerorhinus sp. AK5-629330 89 56473399 5783(42) Acerorhinus palaeosinense –285 ––39– –––– Acerorhinus zernowi –290.4 82.8 50.6 4230.8 88 55.2–– -320.6 (4) -87.3 (4) -55.9 (5) -49.5 (5) -38.7 (5) -90.2 (3)-62.2 (3) Alicornops simorrense –2426032.5 33.618 58.5 33.6–– -300 (4) -85 (16)-50.5 (16)-48 (7)-30 (7)-91.5 (12)-52.5 (12) Alicornops complanatum ––66 38– –64 (2)39-41(2)– – cf. Chilotherium AS.93.1210–77 40–––––– sp. (primitive)

T ABLE 35. — Acerorhinus sp., Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the calcanei (in mm). Comparison with other late Miocene aceratheriines from Eurasia (data from Heissig [1972], Guérin [1980, 1988], Cerdeño & Alcalá [1989], Cerdeño [1996, 1997], and Cerdeño & Sanchez [2000]), for which values correspond to size range and numbers between brackets indicate the size of the sample.

Taxon NumberHHart.tuber.beaksust.min. min. Cub. TD APD fac. H

TD APD APD TD post.post.

Acerorhinus sp. AK7-98 10162 47–587842(55) (42) Acerorhinus sp. AK6-130 109615674648049 60 (45) Acerorhinus –97.9 –39.354.1 50.9 56.4 31– – zernowi -111.7 (9) -57.8 (9) -65.4 (9) -58 (9) -68.2 (7)-38.4 (9) Alicornops –76.7–3141.5 4041.223–– simorrense -107 (27)-42 (31) -59.5 (32)-63 (29) -63 (22)-33.6 (23) Alicornops –78-98 (2)– 33-41 44-54 37-41 62 (j) 24-35 –– complanatum (2)(2)(2)(2) “ Alicornops ”–93-105.5 –36.8 57.3 54.5 67 30 –– alfambrense (5) -45 (7)-67.7 (8) -65.7 (4) -68.6 (2)-36.4 (8)

T ABLE 36. — Acerorhinus sp., Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the MtII (in mm). Comparison with otherlate Miocene aceratheriines from Eurasia (data from Heissig [1972], Guérin [1980], Cerdeño [1996], Cerdeño & Sanchez [2000], and Fortelius et al. [2003]), for which values correspond to size range and numbers between brackets indicate the size of the sample.

Taxon NumberLprox. art.mesocf. lat. fac. diaphysisdist. art. fac.H post.

TD APD TD APD TD APD TD APD

Acerorhinus sp. AK5-w.n. 111 2434.5 14.5 2916(25) (21) 2934 Acerorhinus AMNH 122.1 25.1-28.8 36.4 –––21.5 19.6 33.4- 35.8 zernowi 129903 -126.8 (3)(3)-41.3 (3)-28.4 (3)-20.8 (3)33.7 (2)-37.8 (3)

GEODIVERSITAS • 2005 • 27 (4) 631 Antoine P.-O. & Saraç G.

T ABLE 36. — Acerorhinus sp., Akka ș da g ˘ ı, middle Turolian of Central Anatolia, dimensions of the MtII (in mm). Comparison with otherlate Miocene aceratheriines from Eurasia (data from Heissig [1972], Guérin [1980], Cerdeño [1996], Cerdeño & Sanchez [2000], and Fortelius et al. [2003]), for which values correspond to size range and numbers between brackets indicate the size of the sample (continuation).

Taxon NumberLprox. art.mesocf. lat. fac. diaphysisdist. art. fac.H post.

TD APD TD APD TD APD TD APD

Alicornops –87.319-27.5 28– ––18 14-2124.222.2-36 simorrense -111.5 (12)(20)-37.5 (18) -26.5 (14) (15) -31 (11) (12) Alicornops ––2822 ––––––– complanatum Chilotherium indet.06-AKK-020 85.22633.2–––25.4 19 32.8 30.6 cf. Chilotherium –100.721.4 31.1 –––18.9 16.9 28.4 31.9 sp. (primitive)

632 GEODIVERSITAS • 2005 • 27 (4)