(Mammalia, Artiodactyla) from the late of Akkas ¸ dag ˘ ı, Turkey

Liping LIU Institute of Vertebrate Paleontology and Paleoanthropology, P.O. Box 643, Beijing, 100044 (China) [email protected]

Dimitris S. KOSTOPOULOS Department of Geology, Laboratory of Paleontology, University of Thessaloniki, Thessaloniki, GR-54124 (Greece) [email protected]

Mikael FORTELIUS Department of Geology, University of Helsinki, P.O. Box 64, Helsinki, FIN-00014 (Finland) [email protected]

Liu L., Kostopoulos D. S. & Fortelius M. 2005. — Suidae (Mammalia, Artiodactyla) from the late Miocene of Akkas ¸ dag ˘ ı, Turkey, in Sen S. (ed.), Geology, and environments at Akkas ¸ dag ˘ ı, late Miocene of Central Anatolia. Geodiversitas 27 (4) : 715-733.

ABSTRACT The suid remains from Akkas¸ dag˘ ı, late Miocene of Central Anatolia (Turkey), represent the widespread, long-ranging, and polymorphic Microstonyx major (Gervais, 1848). The rich material represents at least 10 individuals, KEY WORDS two of which are juveniles, and comprises both postcranial and craniodental Mammalia, material, including one nearly complete skull. The Akkas¸ dag˘ ı population is Suidae, Microstonyx, characterised by medium size, strong elongation of the skull, and moderate Miocene, reduction of premolar size. These characteristics are shared with other popu- Akkas¸ dag˘ ı, lations of late middle Turolian age (MN 12). The elongation of the skull Central Anatolia, Turkey, appears elsewhere to be associated with the arid end of the species’ ecological palaeoecology. range.

RÉSUMÉ Suidae (Mammalia, Artiodactyla) du Miocène supérieur d’Akka¸s dag˘ ı, Turquie. Le matériel de suidés du Miocène supérieur d’Akkas¸ dag˘ ı (Anatolie Centrale, Turquie) est attribué à l’espèce polymorphe Microstonyx major (Gervais, 1848), d’une extension géographique très vaste. Un crâne presque complet

GEODIVERSITAS • 2005 • 27 (4) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 715 Liu L. et al.

MOTS CLÉS fait partie de cette collection ; les éléments crânio-dentaires et postcraniens Mammalia, représentent au moins 10 individus : deux jeunes et huit adultes. La popu- Suidae, lation d’Akkas¸ dag˘ ı est caractérisée par sa taille moyenne, l’allongement du Microstonyx, Miocène, museau et la réduction de la série prémolaire, morphologie habituelle des Akkas¸ dag˘ ı, populations du Turolien moyen terminal (MN 12). L’allongement du crâne, Anatolie Centrale, Turquie, caractéristique du point extrême de la variation écologique de l’espèce, est paléoécologie. probablement associé aux conditions arides.

INTRODUCTION One is the Hippopotamodon-Microstonyx group, the other the Korynochoerus-Propotamochoerus The late Miocene fossil land locality group (Pickford 1988, 1993; Van der Made & Akkas¸ dag˘ ı is situated in Central Anatolia Hussain 1989; Bonis & Bouvrain 1996; Fortelius (Turkey), in the Çankırı-Çorum Basin, NNW of et al . 1996). The rich and well preserved collec- Kaman, in a thick volcanic tuff layer in the mesa- tion from the well dated Akkas¸ dag˘ ı locality type Akkas¸ dag˘ ı Hill (Kazancı et al . 1999). The allows us to explore previous hypothesis (Liu et sedimentological and taphonomical evidence al . 2004) about the morphological variability of suggests that the tuff and the fossil material pre- Microstonyx further. served within represent a single depositional event, radiometrically dated to 7.1 ±0.15 Ma C OMPARATIVE MATERIAL AND DATA (Karadenizli et al . 2005), an age close to the MN The comparative material of Microstonyx (casts 12-13 boundary of recent calibrations of the and originals) used in the study is stored in the European land mammal chronology (Steininger following institutions: IVPP (Institute of Verte- et al . 1996; Sen 1997; Daams et al . 1998; brate Paleontology and Paleoanthropology, Chi- Steininger 1999; Agustí et al . 2001). nese Academy of Science, Beijing), BMNH (The Since no taxa characteristic of MN 13 are known Natural History Museum, London), MNHN from the locality, and most of the studied fami- (Muséum national d’Histoire naturelle, Paris), lies (e.g., hipparions, bovids and giraffids) rather LGPUT (Laboratory of Geology and Paleontol- agree with a middle-late Turolian age, the ogy of the University of Thessaloniki), LMNHA Akkas¸ dag˘ ı fauna can be referred to late MN 12 (Local Museum of Natural History, Assenovgrad, with certain confidence (Kazancı et al. 1999; Bulgaria). Other comparative material was stud- Koufos & Vlachou 2005; Kostopoulos 2005; ied mainly from figures in the published litera- Kostopoulos & Saraç 2005). ture. Some of our localities data are gained from Previous material collected by Heintz and co- the ongoing database NOW (NOW database workers during the 1970s (labelled GOK and 2003). stored in the Muséum national d’Histoire naturelle, Paris) does not include suid remains. The material is stored at the Natural History SYSTEMATICS Museum in Ankara. The Akkas¸ dag˘ ı suid materi- al studied here derives from 14 fossiliferous pock- Family S UIDAE Gray, 1821 ets, excavated during the 1999-2001 field seasons Genus Microstonyx Pilgrim, 1926 (labelled AKA,B,K, AK2-AK7 and AK10-AK14) and is thought to represent a single taxon: Microstonyx major (Gervais, 1848) Microstonyx major . (Figs 1-5; Appendix: Tables 1-6) The suines of the continental late Miocene of Europe and western Asia comprise two distinct Sus major Gervais, 1848: pl. XII, fig. 2. clades, variably split between two to four genera. Microstonyx major – Kazancı et al . 1999: 507.

716 GEODIVERSITAS • 2005 • 27 (4) Late Miocene Suidae from Akkas¸ dag˘ ı

M ATERIAL EXAMINED. — Skull: AK3-131, skull with preserved skull AK3-131 (Fig. 1). The latter skull all cheek teeth; AK5-501, broken skull with partial is almost complete but dorso-ventrally com- face and all cheek teeth. Maxilla: AK7-153, maxilla with all cheek teeth; AK11- pressed, especially in the occipital region (Fig. 1A). 66, right maxilla with P2-M3; AK5-443, juvenile right Judging from the completely erupted M3 it maxilla with DP3-DP4 and M1; AK5-623, left maxil- belongs to an adult individual. In dorsal view la with left P3-M3; AK7-100, right maxilla with DP4 (Fig. 1B), the caudal part of the skull is broken at and M1. Mandible: AK2-112, left mandible with p2-m3; the posterior part of the parietal, and much of the AK11-72, mandible with both rami and complete braincase is not preserved. The left zygomatic arch dentition; AK11-67, juvenile mandible with both is missing, while the right side is well preserved. rami, which bears i1 (right), i2 (left) and both side The nasals are long, of nearly constant width dp3, dp4, m1; AK3-126, left mandible with p3-m3; AK4-187, mandible with both rami, with both side almost to the tip. In ventral view (Fig. 1C), the p2-m3, incisor and canine are missing; AK4-251, left specimen is almost complete from the apices of the mandible with p3-m3; AK5-270, juvenile left premaxillae to the occipital condyles, but the ptery- mandible with dp2-dp4, m1-m2; AK5-442, left mandible with p2-m3; AKB-51, right mandible with goid process is heavily deformed by compression p3-m2; AKK-120, female(?) mandible with both rami and difficult to investigate. The right zygomatic and all incisors (di2), canine, and p2-p4 (half left p4); arch is robust and extends strongly towards lateral. AKK-121, left mandible with m1-m3. There are no facial crests, and the anterior rim of Isolated teeth: AK12-5, right I1; AK2-488, left i3; AK4-186, right M3; AK5-624, left I1; AK6-85, a right the zygomatic arch originates at the anterior end broken m3; AK7-154, left m2; AK7-183, right m3; of M3. The orbit is small and far behind M3. The AKA-1, right M3; AKK-192, germ fragment of left rim of the orbit is incomplete but there is a dis- M2; AKK-286, left m1; AKK-287, left m2; AKK-288, tinctive, deep lachrymal notch (infraorbital fossa). right m2; AKK-83, right I2. Postcranial: AK3, AK5-236, AK5-258, AK5-346, The occipital condyles and sphenoid surfaces are AK6-91, AK5-628, AK5b-838, atlases; AK6-96, verte- of typical suid form. Although broken, the jugular bra centrum; AK4-109, left humerus; AK7-184, left processes seem robust and the tympanic bullae are distal humerus; AK11-52, right radius; AK3-302, left oriented downwards, both suggesting a modern radius and ulna; AK5-188, left distal radius; AK5-570, AK6-258, left proximal radii; AK5-625, right proxi- suid form. The choanae open posteriorly, far mal metacarpal III; AKB-54, left metacarpal III; AKK- behind M3. The cheek dentition (P1-M3) is well 82, right metacarpal III; AK5-48, right metacarpal IV; preserved, but all the canines and incisors are miss- AK2-489, left tibia; AK4-88, left astragalus; AK5-149, ing. The small and shallow canine alveolus suggests right metatarsal IV; AK5-199, metapodial. that the canine was small. The alveolar crest is A GE. — Late Miocene, radiometric age 7.1 ±0.15 Ma elongated and relatively slender. (Karadenizli et al . 2005). L OCALITY. — Akkas¸ dag˘ ı, Çankırı-Çorum Basin, Mandible Turkey. The mandible is the most common element in the Akkas¸ dag ˘ ı suid collection. The best pre- D ESCRIPTION served specimen is labelled as AK3-126 According to the mandibular data, seven adult, (Fig. 2A), and is certainly associated with one young adult and two young individuals the skull AK3-131. The mandible is almost compose the local population, which includes one complete except for minor damage, but the mature male, and at least two mature females. canine is unfortunately missing. Two mandibles preserve canines, AK11-72 (Fig. 2B) and AKK- Skull 120 (Fig. 2C). The description of the mandibu- There are two skulls in the Akkas¸ dag˘ ı suid collec- lar morphology of the Akkas¸ dag˘ ı suid is mainly tion, AK3-131 and AK5-501. The latter is only a based on specimens AK3-126, AK11-72, and middle part of a skull with the face partially pre- AKK-120. served. Apart from showing clearly that the occiput The horizontal ramus is shallow and slim, while is very high, it adds little to what is seen in the well the ascending ramus is high, about three times the

GEODIVERSITAS • 2005 • 27 (4) 717 Liu L. et al.

F IG. 1. — Microstonyx major (Gervais, 1848) from Akkas¸ dag ˘ ı, skull AK3-131; A , lateral view; B , dorsal view; C , ventral view. Scale bar: 10 cm.

718 GEODIVERSITAS • 2005 • 27 (4) Late Miocene Suidae from Akkas¸ dag˘ ı

F IG. 2. — Microstonyx major (Gervais, 1848) from Akkas¸ dag ˘ ı; A , lateral view of mandible AK3-126; B , occlusal view of mandible AK11-72; C , occlusal view of mandible AKK-120. Scale bars: 10 cm.

GEODIVERSITAS • 2005 • 27 (4) 719 Liu L. et al.

F IG. 3. — Microstonyx major (Gervais, 1848) from Akkas ¸ dag ˘ ı, dorsal view of atlas AK5-258. Scale bar: 5 cm. height of the horizontal ramus. The ascending ramus rises gently upwards and backwards, with a mandibular angle of about 120°. The ascent begins well behind m3, so that this tooth is completely visible in lateral view. The glenoid condyle of the mandible is broken at the surface, but what remains clearly shows its robustness. The pointed coronoid process is a little higher than the glenoid condyle, and the mandibular notch is quite shallow. The symphysis is elongated, ending before p2. There F IG. 4. — Microstonyx major (Gervais, 1848) from Akkas ¸ dag ˘ ı, seem to be two kinds of canine, but the distinction astragalus AK4-88. Scale bar: 2 cm. is not very sharp. The canine in mandible AKK- 120 is short and narrow, almost symmetrical, with a very narrow posterior facet separated from the the main lingual cusp of p4 being placed as far lateral facets by two crests. Since the tooth is forward as the labial one. narrow, the boundary of the lateral facets forms a sharp anterior crest. Enamel covers the whole tooth Postcranials and ends above the alveolus. It is virtually identi- Late Miocene suid limb bones are rarely cal with the canine that we described from described and figured, making it difficult to sup- Hezheng as a female individual (Liu et al . 2004). ply a comparative study of the abundant postcra- The canine in mandible AK11-72 is slightly more nial material of the Akkas¸ dag˘ ı suid. In order to robust, with an oval transverse section and no obvi- facilitate future comparative work we here pres- ous crests separating the posterior facet from the ent figures and measurements of the material lateral ones. (Appendix: Tables 1; 2; Figs 3-5).

Dentition The dental morphology of Microstonyx is greatly COMPARISON AND DISCUSSION variable, and has been shown extensively in previ- ous publications (e.g., Van der Made et al . 1992; The general morphological characters of the Kostopoulos et al . 2001; Liu et al . 2004, and lit- Akkas¸ dag˘ ı suid, such as its large size, the elevated erature listed herein), and the Akkas¸ dag˘ ı suid fits occiput, the wide and flat frontoparietal region, well within the known range. Notable character- the elongated snout, the inflated and laterally istics of the Akkas¸ dag˘ ı population include a some- extended zygomatic arches, the wide and deep what complicated M3/m3 occlusal pattern, and lachrymal notch, and well developed alveolar

720 GEODIVERSITAS • 2005 • 27 (4) Late Miocene Suidae from Akkas¸ dag˘ ı

F IG. 5. — Microstonyx major (Gervais, 1848) from Akkas¸ dag ˘ ı, metapodials; A , AK5-199; B , AKK-82; C , AK5-48. Scale bar: 5 cm. crest show unambiguously that it belongs to the ally the case. The Akkas¸ dag˘ ı suine does have a Eurasian Hippopotamodon-Microstonyx large reduced canine, but since canine size in female suine group (Trofimov 1954; Pickford 1988; individuals of Hippopotamodon is unknown, this Van der Made & Moyà-Solà 1989; Kostopoulos is not in itself decisive. et al . 2001; Liu et al . 2004). The suines of the late In size and general proportions the Akkas¸ dag˘ ı suid Miocene Eurasian Hippopotamodon-Microstonyx skull is similar to the skull from Pikermi figured group are characterised by considerable morpho- by Gaudry (1862-1867), as well as to the skull spec- logical variability, with a confusing pattern of imen BMNH M9048 that probably represents a sexual, temporal, and regional differences. Hip- female individual. The Akkas¸ dag˘ ı skull is also close popotamodon (Lydekker, 1877) and Microstonyx to the Russian (Trofimov 1954) and Bulgarian are very similar in cranial and dental morpholo- skull samples (especially to the presumed female gy. The characteristics of Hippopotamodon skull specimen K5260 from Kalimanci; emphasized by Pickford (1988), such as the Kostopoulos et al . 2001) and is obviously larger gigantic size, the large and flaring canine, the and more elongated than any specimen from the short snout, the less developed alveolar crest, and Chinese latest Miocene locality Hezheng (Liu et al . the short diastema between C-P1-P2, do not 2004) (Appendix: Tables 3; 4). qualitatively distinguish the two genera, especial- Microstonyx, as conventionally conceived, has two ly not when the material is incomplete, as is usu- species, M. antiquus (Kaup, 1833) and M. major .

GEODIVERSITAS • 2005 • 27 (4) 721 Liu L. et al.

Microstonyx antiquus was larger and distributed what reduced premolars. Under our ecotype earlier in time than M. major . Fortelius et al . hypothesis, the great elongation of the skull of (1996) tentatively assigned M. antiquus to Hip- the Akkas¸ dag˘ ı form would indicate the arid end popotamodon and we follow this taxonomy here. of the habitat spectrum inhabited by the species. Under this usage, Hippopotamodon differs from In the Balkans, similar ecotypes with long muz- Microstonyx primarily in the premolar propor- zles and medium sized molars occur mainly in tions, especially the relative stoutness of the the middle Turolian, MN 12 (Kostopoulos et al . fourth premolar in the former (Liu et al . 2004). 2001). As the dental logarithmic ratio diagrams show (Fig. 6), it is easy to assign the Akkas¸ dag˘ ı suid to M ICROSTONYX IN T URKEY M. major as opposed to Hippopotamodon. Microstonyx is poorly documented in the late Microstonyx major was a common element of the Miocene of Turkey, even if references are late Miocene mammal faunas of Europe and numerous. S¸ enyürek (1952) described as Sus West Asia. It is also known from East Asia, but erymanthius (= Microstonyx major ) a suid was a rare element there, with evidence of sub- mandible with p3-m3 from the undated site stantial size decrease (Liu 2003). Some workers Akkırma II near Gökdere. The specimen (Van der Made & Moyà-Solà 1989; Van der belongs to a large form with stout p4, correctly Made et al . 1992; Pickford 1993; Kostopoulos recognized by Bonis & Bouvrain (1996) as 1994; Bonis & Bouvrain 1996; Van der Made belonging to Microstonyx ( Limnostonyx) antiquus. 1997) have interpreted the spatiotemporal vari- Dicoryphochoerus meteai Ozansoy, 1965 from ability of M. major as possible evidence of multi- the Vallesian locality of Yassıören (middle ple evolving lineages. In contrast, we have Sinap) is mainly based on a left mandibular recently argued (Liu et al . 2004) that the mor- ramus, which has been later considered by Pick- phological variability of M. major s.l. is best inter- ford & Ertürk (1979) as belonging to Hip- preted as polymorphism within a single evolving popotamodon and assigned by Bonis & Bouvrain species. The substantial inter-population size (1996) to Microstonyx ( Limnostonyx) antiquus. variability in M. major , is owing to intra-specific Both the Yassıören and Akkırma II suids, polymorphism, representing local ecotypes (Liu referred here to Hippopotamodon antiquus, differ et al . 2004). We have specifically proposed that from the Akkas¸ dag ˘ ı one in the larger dimen- elongation of the skull in M. major tends to be sions, stronger canine, persistence of p1 and associated with arid conditions and a dietary shift more robust p4. towards herbivory (Liu et al . 2004). Following Microstonyx major (including M. erymanthius) this view here, we recognise only one formal has been reported from several Turkish localities species-level taxon, M. major . ranging from MN 10 to MN 12 (Gülpınar, The tooth dimensions of Microstonyx from Çorak Yerler, Çevril, Mug˘ la Garkın, Kayadibi, Akkas¸ dag˘ ı (Appendix: Tables 5; 6) are well with- upper Kavakdere, Çoban Pınar, Mahmutgazi in the range of the middle-late Turolian Euro- and Kınık; Ozansoy 1965; Sickenberg 1975; pean M. major , and larger than those of other Pickford & Ertürk 1979; Fortelius et al . 1996; Asian samples (Maragha and China; Bonis & NOW database 2003) but we usually deal with Bouvrain 1996; Liu et al . 2004) or from some faunal lists or fragmentary and isolated speci- early Turolian populations of SE Europe (Vathy- mens which preclude a direct comparison with lakkos, Kerassia, Perivolaki; Kostopoulos et al . the Akkas¸ dag ˘ ı form. As far we know the 2001) (Fig. 7). Logarithmic ratio diagrams illus- Akkas¸ dag˘ ı M. major provides the clearest evi- trating the relative proportions of the Eurasian dence of the species in Turkey. The absence of Microstonyx major toothrows are shown in the species in the isochronous and very similar Figure8. The Akkas¸ dag˘ ı M. major is close to the fauna of Kemiklitepe A, B looks rather anom- Pikermian and Bulgarian samples but with some- alous.

722 GEODIVERSITAS • 2005 • 27 (4) Late Miocene Suidae from Akkas¸ dag˘ ı

F IG. 6. — Logarithmic ratio diagram of dental proportions of Microstonyx major (Gervais, 1848), M. antiquus (Kaup, 1833) and Hippopotamodon (Lydekker, 1877); standard: Pikermi. Data sources: Siwaliks (Pickford 1988); Eppelsheim (Hünermann 1968); Pikermi (Pearson 1928; Hellmund 1995; Bonis & Bouvrain 1996); Bulgaria (Kostopoulos et al . 2001); China (Pearson 1928; Liu et al . 1978, 2004; Tang et al . 1985); Maragha (Bonis & Bouvrain 1996).

GEODIVERSITAS • 2005 • 27 (4) 723 Liu L. et al.

P4M2

p4 m2

F IG. 7. — Bivariate scatter plots of Microstonyx major (Gervais, 1848) from different localities. Data sources: Lantian (Liu et al . 1978); Binxian (Tang et al . 1985); Pikermi (Pearson 1928; Hellmund 1995; Bonis & Bouvrain 1996); Spain (Van der Made et al . 1992); Bulgaria (Kostopoulos et al . 2001); Vathylakkos (Bonis & Bouvrain 1996); Lubéron (Bonis & Bouvrain 1996); Maragha (Bonis & Bouvrain 1996); Dorn-Dürkheim (Van der Made 1997); Dytiko (Bonis & Bouvrain 1996); Loc-114 (Pearson 1928); Hezheng (Liu et al . 2004).

724 GEODIVERSITAS • 2005 • 27 (4) Late Miocene Suidae from Akkas¸ dag˘ ı

F IG. 8. — Logarithmic ratio diagram of dental proportions of Microstonyx major (Gervais, 1848); standard: Pikermi. Data sources: Siwaliks (Pickford 1988); Eppelsheim (Hünermann 1968); Pikermi (Pearson 1928; Hellmund 1995; Bonis & Bouvrain 1996); Bulgaria (Kostopoulos et al . 2001); China (Pearson 1928; Liu et al . 1978, 2004; Tang et al . 1985); Maragha (Bonis & Bouvrain 1996).

GEODIVERSITAS • 2005 • 27 (4) 725 Liu L. et al.

F IG. 9. — Bivariate scatter plots of the alveolar crest development in Microstonyx major (Gervais, 1848) (anteroposterior diameter [DAP] against height) from several localities (from Kostopoulos et al . 2001, modified; two specimens from Pikermi from the BMNH collection added).

S EXUAL DIMORPHISM It is not possible to establish beyond all doubt The Akkas¸ dag˘ ı suid collection does not at present that the minor differences observed in the lower add decisively to our understanding of sexual di- canines from Akkas¸ dag ˘ ı reflect sexual dimor- morphism in M. major (Kostopoulos et al . 2001; phism, but the pattern observed supports the sug- Liu et al . 2004). The weaker alveolar crest and the gestion of weak sexual bimodality (Liu et al . lesser width of the skull AK3-131 compared to 2004). those of other samples suggest that the Akkas¸ dag˘ ı skull belongs to a female individual (Kostopoulos et al . 2001; Liu et al . 2004). Figure9 summarizes CONCLUSIONS the available data on the distribution of the alveo- lar crest dimensions according to sex. Evidently, The study of the suid material from Akkas¸ dag˘ ı the Akkas¸ dag˘ ı suid is located among the female in- allows its attribution to Microstonyx major , as this dividuals of M. major from Pikermi (Greece) and species is considered in the frame of our last Kalimanci (Bulgaria) while the Hezheng form works (Kostopoulos et al . 2001; Liu 2003; Liu et represents a shift toward smaller overall size. al . 2004). The Akkas¸ dag˘ ı form represents a typi- The fact that the presumed female skull retains cal middle Turolian (MN 12) eastern Mediter- P1 supports our previous suggestion that females ranean population with close similarities to the retained P1 more frequently than males (Liu et al . Greek and Bulgarian samples. Morphological 2004). In contrast to the presumed male skull cranial characters indicate relatively arid condi- from Hezheng (Liu et al . 2004), the presumed tions, still within the range of a generalist species. female skulls from Hezheng and Akkas¸ dag˘ ı have Although the presence of the long-ranging almost no development of facial crests. Similarly, M.major cannot as such provide certain the deep preorbital fossa found in the Hezheng biochronological conclusions, the population presumed male individual is not as strongly characteristics do appear to support a middle-late developed in the presumed females. Turolian age for the Akkas¸ dag˘ ı assemblage.

726 GEODIVERSITAS • 2005 • 27 (4) Late Miocene Suidae from Akkas¸ dag˘ ı

Acknowledgements Evolution of Western Eurasian Neogene Mammal Geological studies and excavations at Akkas¸ dag˘ ı Faunas . Columbia University Press, New York: 348-377. have been authorized by the General Directorate G AUDRY A. 1862-1867. — Animaux fossiles et géologie of Mineral Research and Exploration (MTA) and de l’Attique . F. Savy, Paris, 2 vols, 472 p. Ankara University, Faculty of Sciences. The work G ERVAIS P. 1848-1852. — Zoologie et paléontologie was done thanks to grants from the CNRS francaises . A. Bertrand, Paris, 2 vols, 3 vols. H ELLMUND M. 1995. — The vertebrate locality (ECLIPSE Program and DRI), the Muséum na- Maramena (Macedonia, Greece) at the Turolian- tional d’Histoire naturelle, Paris, MTA and Ruscinian boundary (Neogene) 13. Suidae (Artio- TUBITAK. They have financially supported field- dactyla, Mammalia). Münchener Geowissenschaftliche work and reciprocal visits to the concerned institu- Abhandlungen, series A (28): 143-156. H ÜNERMANN K. A. 1968. — Die Suidae (Mammalia, tions. The authors are grateful to G. Bouvrain, Artiodactyla) aus den Dinotheriensanden L.de Bonis, L. Ginsburg, E. Heintz and all partici- (Unterpliozän+pont) Rheinhessens (Südwest- pants of 2000-2001 excavations. The authors deutschland). Schweizerische Paläontologische thank Sevket Sen for having invited them to dig Abhandlung 86: 1-96. K ARADENIZLI L., S EYITOG˘ LU G., S EN S., A RNAUD N., and study the Akkas¸ dag˘ ı material. We are grateful K AZANCı N., S ARAÇ G. & A LÇIÇEK C. 2005. — to Dr. Gerçek Saraç for generous assistance during Mammal bearing late Miocene tuffs of the our visit and to Denis Serrette (MNHN) for the Akkas¸ dag˘ ı region; distribution, age, petrographical and geochemical characteristics, in S EN S. (ed.), photographs used in the figures. We express our Geology, mammals and environments at Akkas¸ dag˘ ı, sincere gratitude to Dr. Jan Van der Made (Museo late Miocene of Central Anatolia. Geodiversitas 27 Nacional de Ciencias Naturales, Madrid) and (4): 553-566 (this volume). Dr.Martin Pickford (MNHN) for their reviews K AZANCı N., S EN S., S EYITOG˘ LU G., B ONIS L. DE, B OUVRAIN G., A RAZ H., V AROL B. & K ARADENIZLI and comments. This work was also supported L. 1999. — Geology of a new late Miocene mam- bythe Academy of Finland (200915), the mal locality in Central Anatolia, Turkey. Comptes Ministry of Science and Technology of China Rendus de l’Académie des Sciences , Paris, Sciences de (2002CB714002) and Natural Science la Terre et des Planètes 329: 503-510. K OSTOPOULOS D. S. 1994. — Microstonyx major Foundation of China (40102004). (Suidae, Artiodactyla) from the late Miocene locali- ty of “Nikiti-1”, Macedonia, Greece; some remarks about the species. Bulletin of the Geological Society of REFERENCES Greece 30 (1): 341-355. K OSTOPOULOS D.S. 2005. — The Bovidae (Mammalia, Artiodactyla) from the late Miocene of Akkas¸ dag˘ ı, A GUSTÍ J., C ABRERA L., G ARCÉS M., K RIJGSMAN W., Turkey, in S EN S. (ed.), Geology, mammals and envi- O MS O. & P ARÉS J. M. 2001. — A calibrated mam- ronments at Akkas¸ dag˘ ı, late Miocene of Central mal scale for the Neogene of Western Europe. State Anatolia. Geodiversitas 27 (4): 747-791 (this volume). of the art. Earth-Science Review 52: 247-260. K OSTOPOULOS D. S. & S ARAÇ G. 2005. — Giraffidae B ONIS L. DE & B OUVRAIN G. 1996. — Suidae du (Mammalia, Artiodactyla) from the late Miocene of Miocène supérieur de Grèce. Bulletin du Muséum Akkas¸ dag˘ ı, Turkey, in S EN S. (ed.), Geology, mam- national d’Histoire naturelle, Paris, 4 e sér., section C, mals and environments at Akkas¸ dag˘ ı, late Miocene of 18 (1): 107-132. Central Anatolia. Geodiversitas 27 (4): 735-745 (this D AAMS R., A LCALA L., A LVAREZ S IERRA M. D. L. A., volume). A ZANZA B., D AM J. V., M EULEN A. J. V. D., K OSTOPOULOS D. S., S PASSOV N. & K OVACHEV D. M ORALES J., N IETO M., P ELÁEZ-C AMPOMANES P. & 2001. — Contribution to the study of Microstonyx: S ORIA D.1998. — A stratigraphical framework for evidence from Bulgaria and the SE European popu- Miocene (MN 4-MN 13) continental sediments of lations. Geodiversitas 23 (3): 411-437. central Spain. Comptes Rendus de l’Académie des K OUFOS G. D. & V LACHOU T. D. 2005. — Equidae Sciences , Paris, Sciences de la Terre et des Planètes (Mammalia, Perissodactyla) from the late Miocene 327: 625-631. of Akkas¸ dag˘ ı, Turkey, in S EN S. (ed.), Geology, F ORTELIUS M., V AN DER M ADE J. & B ERNOR R. 1996. mammals and environments at Akkas¸ dag ˘ ı, late — Middle and late Miocene Suoidea of central Miocene of Central Anatolia. Geodiversitas 27 (4): Europe and the eastern Mediterranean: evolution, 633-705 (this volume). biogeography and paleoecology, in B ERNOR R., L IU D. S., L I C. K. & Z HAI R. J. 1978. — [Pliocene F AHLBUSCH V. & M ITTMANN H. (eds), The vertebrates of Lantian, Shensi]. Professional Papers

GEODIVERSITAS • 2005 • 27 (4) 727 Liu L. et al.

on Stratigraphy and Paleontology 7: 149-200 (in tionale Neogen-Stratigraphie. Geologisches Jahrbuch Chinese). B, 15: 1-167. L IU L. P. 2003. — Chinese Fossil Suidea: Systematics, S TEININGER F. F. 1999. — Chronostratigraphy, Evolution and Paleoecology . University Printing geochronology and biochronology of the Miocene House, Helsinki, 41 p. “European Land Mammal Mega-Zones” L IU L. P., K OSTOPOULOS D. S. & F ORTELIUS M. (ELMMZ) and the Miocene “Mammal-Zones 2004. — Late Miocene Microstonyx remains (MN-Zones)”, in R ÖSSNER G. E. & H EISSIG K. (Suidae, Artiodactyla) from Hezheng areas. Geobios (eds), The Miocene Land Mammals of Europe. Verlag 37 (1): 49-64. Dr. Friedrich Pfeil, München: 9-24. NOW DATABASE 2003. — Neogene of the Old S TEININGER F. F., B ERGGREN W. A., K ENT D. V., World. Database of fossil mammals: www.helsinki. B ERNOR R. L., S EN S. & A GUSTÍ J.1996. — Circum- fi/science/now Mediterranean Neogene (Miocene-Pliocene) O ZANSOY F.1965. — Étude des gisements continen- marine-continental chronologic correlations of taux et de mammifères du Cénozoïque de Turquie. European mammal units, in B ERNOR R. L., Mémoires de la Societé géologique de France N. S. F AHLBUSCH V. & M ITTMANN H. W. (eds), The 102: 5-92. Evolution of Western Eurasian Neogene Mammal P EARSON H. 1928. — Chinese fossil Suidae. Faunas . Columbia University Press, New York: 7-46. Palaeontologia Sinica C (5): 1-75. T ANG Y. J., L IU Z. Q., C HEN D. & C HEN L. Q. 1985. P ICKFORD M. 1988. — Revision of the Miocene —The fossil Suidae from late Miocene of Binxian, Suidae of the Indian Subcontinent. Münchener Shaanxi. Vertebrata PalAsiatica 23 (1): 60-68. Geowissenschaftliche Abhandlungen, series A, 12 (1): T ROFIMOV B. A. 1954. — The fossil suids of the genus 1-95. Microstonyx, in Tertiary Mammals, part 2: on the P ICKFORD M. 1993. — Old world suoid systematics, Mammalia of the southern SSSR and Mongolia. phylogeny, biogeography and biostratigraphy. Doklady Akademii Nauk SSSR 47: 61-99. Paleontologia i Evolucio 26-27: 237-269. V AN DER M ADE J. 1997. — The fossil pig from the P ICKFORD M. & E RTÜRK C. 1979. — Suidae and late Miocene of Dorn-Dürkheim 1 in Germany. Tayassuidae from Turkey. Bulletin of the Geological Courier Forschungs-Institut Senckenberg 197: 205- Society of Turkey 22: 141-154. 230. S EN S.1997. — Magnetostratigraphic calibration of V AN DER M ADE J. & H USSAIN S. T. 1989. — the European Neogene mammal chronology. “ Microstonyx” major (Suidae, Artiodactyla) from Palaeogeography Palaeoclimatology Palaeoecology 133: Nagri. Estudios Geologicos 45: 409-416. 181-20. V AN DER M ADE J. & M OYÀ-S OLÀ S. 1989. — S¸ ENYÜREK M. S.1952. — A study of the Pontian European (Artiodactyla) from the late fauna of Gökdere (Elmadagi), southeast of Ankara. Miocene onwards. Bollettino della Società Türk Tarih Kurumu Belleten 16: 449-492. Paleontologica Italiana 28 (2-3): 329-339. S ICKENBERG O. 1975.—Die Gliederung des höheren V AN DER M ADE J., M ONTOYA P. & A LCALÁ L. 1992. Jungtertiärs und Altquartärs in der Türkei nach — Microstonyx (Suidae, Mammalia) from the upper Vertebraten und ihre Bedeutung für die interna- Miocene of Spain. Geobios 25 (3): 395-413.

Submitted on 13 June 2003; accepted on 2 June 2004.

728 GEODIVERSITAS • 2005 • 27 (4) Late Miocene Suidae from Akkas¸ dag˘ ı

APPENDIX

T ABLE 1. — Measurements of atlases and one indetermined vertebra of Microstonyx major (Gervais, 1848) from Akkas¸ dag ˘ ı (in mm). M1 , centrum length; M2 , centrum width; M3 , centrum height; M4 , maximal length; M5 , maximal width; M6 , maximal height; M7 ,proximal articulation width; M8 , proximal articulation height; M9 , distal articulation width; M10 , distal articulation height.

Specimen ElementM1M2M3M4M5M6M7M8M9M10

AK3Atlas66.077.042.076.036.0 AK5-236 Atlas77.076.039.033.0 AK5-258 Atlas148.063.083.038.5 76.4 32.0 AK5-346Atlas60.077.034.0 AK5-628Atlas81.8 153.070.081.238.075.034.0 AK5b-838Atlas67.085.038.077.035.0 AK6-91 Atlas67.0 AK6-96Vertebra55.045.037.0

T ABLE 2. — Measurements of limb bones of Microstonyx major (Gervais, 1848) from Akkas¸ dag ˘ ı (in mm). M1 , maximal length; M2 ,functional length; M3 , proximal width; M4 , proximal antero-posterior depth; M5 , shaft width; M6 , antero-posterior depth of shaft; M7 , distal width; M8 , distal antero-posterior depth; M9 , distal articulation width; M10 , antero-posterior depth of distal articulation.

Specimen ElementSide M1M2M3M4M5M6M7M8M9M10

AK4-109Distal humerus left57.5 62.4 47.5 43.3 AK7-184 Distal humerus left66.8 69.051.247.8 AK11-52Proximal radius right51.9 33.634.627.2 AK3-302 Distal radius left33.060.040.052.229.4 AK5-188 Distal radius left28.024.054.037.044.4 29.5 AK5-570 Proximal radius left44.1 30.9 AK6-258 Proximal radius left49.035.0 AK3-302 Distal ulnaleft23.317.1 AKB-54 Metacarpal III left109.0105.030.4 29.1 21.9 15.726.9 29.625.329.6 AKK-82Metacarpal III right107.0103.032.630.622.616.1 29.8 27.1 26.727.0 AK5-625Proximalright30.024.4 metacarpal III AK5-48 Metacarpal IV right105.099.030.024.4 21.313.325.8 27.226.1 27.2 AK2-489 Distal tibialeft43.739.635.4 34.1 AK4-88 Astragalus left62.235.1 34.9 30.4 34.1 34.726.3 AK5-149 Metatarsal IV right113.0109.025.034.320.015.725.1 27.9 24.4 28.2

GEODIVERSITAS • 2005 • 27 (4) 729 Liu L. et al.

T ABLE 3. — Skull measurements of Microstonyx major (Gervais, 1848) from Akkas¸ dag ˘ ı compared with specimens from Pikermi (Greece) and Hezheng (China) (in mm). Data from Gaudry (1862-1867), Liu et al. (2004) and personal data.

Skull Akkas ¸ dag ˘ ı Pikermi Hezheng

AK3-131 Gaudry’sBMNH HMV 0976 HMV 0977 female specimen M9048 male female

Basal length of the skull 468470 420 < 380– Condyles to posterior border of 374310350– palatine fissure Condyles to posterior border114 125.9 123 – of palate Posterior border of palatine fissure260 227 – to posterior border of palate Posterior border of palatine fissure234202 195 to the end of M3 Posterior border of palate to M3263325– Length of the diastema between 40.8 44 51.744 canine and P2 Length of P2-M3153141 143123.6121.6 P2-P4/M1-M30.570.58 0.530.52 Length of alveolar crest 648759 67 50 M3 to the anterior end of orbit6450– Maximal length of the orbit–42.5 – Maximal width of the skull 262 310272 < 225– Width of palate at alveolar crest 109140133 122 89.5 Width of palate at anterior end of M34238.332 Breadth of frontal at the supraorbital150128.6 process Nasal width above preorbital foramen 8375– Nasal width above P266.5 56– Nasal width at the end of incisor notch6959– Width of the palatine fissure2122 –

T ABLE 4. — Mandible measurements of Microstonyx major (Gervais, 1848) from Akkas¸ dag ˘ ı compared with specimens from Hezheng (China) and Kalimanci (Bulgaria) (in mm). Data from Kostopoulos et al. (2001), Liu et al. (2004).

Mandible (sex)Akkas ¸ dag ˘ ıAkkas ¸ dag ˘ ıKalimanciHezheng AK11-72/ AK3-126 K-5277/K-5276 HMV 0576 AKK-120 (female) (male/female) (female) (male/female)

Mandible length from tip to m2212/––233/239192 Length of p2-m299/–102 111/111 98 p2-p4/m1-m20.56/–0.54 Length of symphysis100.5/85.5 100 102/8780 Diastema between c-p247/38.5 56.8 52/5040 Height of mandible at p252.6/43.653.8 56/47 Height of mandible at muscular–171– – process Height of mandible at articular–152–_ process

730 GEODIVERSITAS • 2005 • 27 (4) Late Miocene Suidae from Akkas¸ dag˘ ı

T ABLE 5. — Upper dentition measurements of Microstonyx major (Gervais, 1848) from Akkas¸ dag ˘ ı (in mm). L , maximum length of incisor (basal measured), canine and cheek teeth; W1 , maximum width of incisor, canine, premolar or first moist width of the molar; W2 , second moist width of molar or the height of incisor and canine; W3 , width of talon; upper data-line, left toothrow; lower data- line, right toothrow.

Specimen I1I2I3CP1P2P3P4M1M2M3

L16.016.614.319.727.1 39.5 W110.015.5 16.9 18.022.224.9 AK11-66 W218.4 22.8 22.2 W314.8 L AK12-5 W19.9 W216.1 L9.315.316.8 17.021.8 29.740.5 9.615.316.616.4 21.5 28.5 40.4 W14.5 9.5 14.9 18.219.5 24.5 26.1 4.69.314.4 17.9 19.5 23.5 26.1 AK3-131 W219.224.023.5 19.624.9 23.7 W313.4 13.8 L43.4 W128.1 AK4-186 W224.4 W312.0 L15.316.9 15.4 19.626.041.6 15.9 15.3 14.7 18.9 26.0 41.0 W19.5 15.5 18.720.1 24.227.2 8.715.4 18.4 20.2 24.7 27.3 AK5-501 W221.024.8 24.3 21.9 24.9 24.8 W315.0 15.8 L17.616.322.5 30.1 42.8 W116.8 19.4 19.624.8 27.4 AK5-623 W219.023.8 24.6 W317.1 L22.5 AK5-624 W110.6 W227.0 L21.6 AK7-100 W118.7 W218.3 L15.9 16.5 14.9 20.226.8 38.6 16.1 14.5 21.026.5 39.3 W110.1 14.9 18.1 18.723.024.3 14.5 18.018.722.8 24.2 AK7-153 W222.4 22.2 22.8 22.0 W311.3 11.3 L41.7 W126.1 AKA-1 W223.2 W312.8 L11.8 AKK-83 W18.2 W216.2

GEODIVERSITAS • 2005 • 27 (4) 731 Liu L. et al.

T ABLE 6. — Lower dentition measurements of Microstonyx major (Gervais, 1848) from Akkas¸ dag ˘ ı (in mm). L , maximum length of incisor (basal measured), canine and cheek teeth; W1 , maximum width of incisor, canine, premolar or first moist width of the molar; W2 , second moist width of molar or the height of incisor and canine; W3 , width of talonid; upper data-line, left toothrow; lower data-line, right toothrow.

Specimen i1 i2i3cp2p3p4 m1 m2m3

L6.5 22.9 22.8 W19.316.0 AK11-67 15.9 W215.0 15.8

L8.618.317.5 9.015.018.319.620.728.741.9 9.1 17.8 16.211.014.6 18.4 20.2 20.6 27.6 42.7 W114.014.1 6.36.06.69.314.5 14.9 18.4 21.5 14.5 12.1 6.5 6.5 6.68.5 14.4 14.718.221.2 AK11-72 W235.730.219.312.214.318.620.0 34.032.019.011.014.319.220.3 W315.9 16.0

L14.217.219.620.4 28.8 48.9 W16.4 9.014.4 14.1 19.221.5 AK2-112 W214.8 19.320.2 W316.1

L21.6 AK2-488 W17.3 W217.5

L18.319.8 21.8 29.046.1 W18.4 13.1 14.018.322.1 AK3-126 W214.8 18.5 20.6 W317.0

L14.017.5 19.8 18.5 25.5 43.4 17.8 19.1 18.4 25.242.3 W17.29.214.714.718.9 20.6 9.014.8 14.219.1 21.8 AK4-187 W215.220.8 20.9 14.620.020.3 W316.4 17.0

L19.4 20.4 21.628.5 46.0 AK4-251 W110.5 16.5 16.320.722.5 W215.9 20.4 21.0 W317.3

L21.5 AK5-270 W114.4 W214.3

L14.317.5 19.8 20.5 26.643.4 W17.5 9.9 14.4 14.618.9 22.2 AK5-442 W215.5 19.220.7 W316.1

L28.4 AK7-154 W120.4 W221.0

732 GEODIVERSITAS • 2005 • 27 (4) Late Miocene Suidae from Akkas¸ dag˘ ı

Specimen i1 i2i3cp2p3p4 m1 m2m3

L44.3 W122.5 AK7-183 W220.8 W316.9 L18.4 21.1 23.029.8 AKB-51 W110.012.4 15.9 20.5 W215.620.2 L10.1 16.5 17.215.219.221.5 9.5 17.216.5 15.8 19.5 W18.07.5 6.9 7.4 9.714.7 AKK-120 8.06.9 7.5 7.4 10.1 W212.318.218.2 13.4 18.4 18.4 L22.229.5 39.2 W115.1 18.9 20.9 AKK-121 W215.319.320.8 W316.3 L22.3 AKK-286 W114.2 W215.0 L29.0 AKK-287 W119.8 W220.1 L28.5 AKK-288 W118.5 W219.8

GEODIVERSITAS • 2005 • 27 (4) 733