Suidae (Mammalia, Artiodactyla) from the late Miocene of Akkaşdağı, Turkey Liping LIU Institute of Vertebrate Paleontology and Paleoanthropology, P.O. Box 643, Beijing, 100044 (China) liuliping@ivpp.ac.cn Dimitris S. KOSTOPOULOS Department of Geology, Laboratory of Paleontology, University of Thessaloniki, Thessaloniki, GR-54124 (Greece) dkostop@geo.auth.gr Mikael FORTELIUS Department of Geology, University of Helsinki, P.O. Box 64, Helsinki, FIN-00014 (Finland) mikael.fortelius@helsinki.fi Liu L., Kostopoulos D. S. & Fortelius M. 2005. — Suidae (Mammalia, Artiodactyla) from the late Miocene of Akkaşdağı, Turkey, in Sen S. (ed.), Geology, mammals and environments at Akkaşdağı, late Miocene of Central Anatolia. Geodiversitas 27 (4) : 715-733. KEY WORDS Mammalia, Suidae, Microstonyx, Miocene, Akkaşdağı, Central Anatolia, Turkey, palaeoecology. ABSTRACT The suid remains from Akkaşdağı, late Miocene of Central Anatolia (Turkey), represent the widespread, long-ranging, and polymorphic species Microstonyx major (Gervais, 1848). The rich material represents at least 10 individuals, two of which are juveniles, and comprises both postcranial and craniodental material, including one nearly complete skull. The Akkaşdağı population is characterised by medium size, strong elongation of the skull, and moderate reduction of premolar size. These characteristics are shared with other populations of late middle Turolian age (MN 12). The elongation of the skull appears elsewhere to be associated with the arid end of the species’ ecological range. RÉSUMÉ Suidae (Mammalia, Artiodactyla) du Miocène supérieur d’Akkaşdağı, Turquie. Le matériel de suidés du Miocène supérieur d’Akkaşdağı (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 Mammalia, Suidae, Microstonyx, Miocène, Akkaşdağı, Anatolie Centrale, Turquie, paléoécologie. fait partie de cette collection ; les éléments crânio-dentaires et postcraniens représentent au moins 10 individus : deux jeunes et huit adultes. La population d’Akkaşdağı est caractérisée par sa taille moyenne, l’allongement du museau et la réduction de la série prémolaire, morphologie habituelle des populations du Turolien moyen terminal (MN 12). L’allongement du crâne, caractéristique du point extrême de la variation écologique de l’espèce, est probablement associé aux conditions arides. INTRODUCTION The late Miocene fossil land mammal locality Akkaşdağı is situated in Central Anatolia (Turkey), in the Çankırı-Çorum Basin, NNW of Kaman, in a thick volcanic tuff layer in the mesatype Akkaşdağı Hill (Kazancı et al. 1999). The sedimentological and taphonomical evidence suggests that the tuff and the fossil material preserved within represent a single depositional event, radiometrically dated to 7.1 ± 0.15 Ma (Karadenizli et al. 2005), an age close to the MN 12-13 boundary of recent calibrations of the European land mammal chronology (Steininger et al. 1996; Sen 1997; Daams et al. 1998; Steininger 1999; Agustí et al. 2001). Since no taxa characteristic of MN 13 are known from the locality, and most of the studied families (e.g., hipparions, bovids and giraffids) rather agree with a middle-late Turolian age, the Akkaşdağı fauna can be referred to late MN 12 with certain confidence (Kazancı et al. 1999; Koufos & Vlachou 2005; Kostopoulos 2005; Kostopoulos & Saraç 2005). Previous material collected by Heintz and coworkers during the 1970s (labelled GOK and stored in the Muséum national d’Histoire naturelle, Paris) does not include suid remains. The material is stored at the Natural History Museum in Ankara. The Akkaşdağı suid material studied here derives from 14 fossiliferous pockets, excavated during the 1999-2001 field seasons (labelled AKA,B,K, AK2-AK7 and AK10-AK14) and is thought to represent a single taxon: Microstonyx major. The suines of the continental late Miocene of Europe and western Asia comprise two distinct clades, variably split between two to four genera. 716 One is the Hippopotamodon-Microstonyx group, the other the Korynochoerus-Propotamochoerus group (Pickford 1988, 1993; Van der Made & Hussain 1989; Bonis & Bouvrain 1996; Fortelius et al. 1996). The rich and well preserved collection from the well dated Akkaşdağı locality allows us to explore previous hypothesis (Liu et al. 2004) about the morphological variability of Microstonyx further. COMPARATIVE MATERIAL AND DATA The comparative material of Microstonyx (casts and originals) used in the study is stored in the following institutions: IVPP (Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Science, Beijing), BMNH (The Natural History Museum, London), MNHN (Muséum national d’Histoire naturelle, Paris), LGPUT (Laboratory of Geology and Paleontology of the University of Thessaloniki), LMNHA (Local Museum of Natural History, Assenovgrad, Bulgaria). Other comparative material was studied mainly from figures in the published literature. Some of our localities data are gained from the ongoing database NOW (NOW database 2003). SYSTEMATICS Family SUIDAE Gray, 1821 Genus Microstonyx Pilgrim, 1926 Microstonyx major (Gervais, 1848) (Figs 1-5; Appendix: Tables 1-6) Sus major Gervais, 1848: pl. XII, fig. 2. Microstonyx major – Kazancı et al. 1999: 507. GEODIVERSITAS • 2005 • 27 (4) Late Miocene Suidae from Akkaşdağı MATERIAL EXAMINED. — Skull: AK3-131, skull with all cheek teeth; AK5-501, broken skull with partial face and all cheek teeth. Maxilla: AK7-153, maxilla with all cheek teeth; AK1166, right maxilla with P2-M3; AK5-443, juvenile right maxilla with DP3-DP4 and M1; AK5-623, left maxilla with left P3-M3; AK7-100, right maxilla with DP4 and M1. Mandible: AK2-112, left mandible with p2-m3; AK11-72, mandible with both rami and complete dentition; AK11-67, juvenile mandible with both rami, which bears i1 (right), i2 (left) and both side dp3, dp4, m1; AK3-126, left mandible with p3-m3; AK4-187, mandible with both rami, with both side p2-m3, incisor and canine are missing; AK4-251, left mandible with p3-m3; AK5-270, juvenile left mandible with dp2-dp4, m1-m2; AK5-442, left mandible with p2-m3; AKB-51, right mandible with p3-m2; AKK-120, female(?) mandible with both rami and all incisors (di2), canine, and p2-p4 (half left p4); AKK-121, left mandible with m1-m3. Isolated teeth: AK12-5, right I1; AK2-488, left i3; AK4-186, right M3; AK5-624, left I1; AK6-85, a right broken m3; AK7-154, left m2; AK7-183, right m3; AKA-1, right M3; AKK-192, germ fragment of left M2; AKK-286, left m1; AKK-287, left m2; AKK-288, right m2; AKK-83, right I2. Postcranial: AK3, AK5-236, AK5-258, AK5-346, AK6-91, AK5-628, AK5b-838, atlases; AK6-96, vertebra centrum; AK4-109, left humerus; AK7-184, left distal humerus; AK11-52, right radius; AK3-302, left radius and ulna; AK5-188, left distal radius; AK5-570, AK6-258, left proximal radii; AK5-625, right proximal metacarpal III; AKB-54, left metacarpal III; AKK82, right metacarpal III; AK5-48, right metacarpal IV; AK2-489, left tibia; AK4-88, left astragalus; AK5-149, right metatarsal IV; AK5-199, metapodial. AGE. — Late Miocene, radiometric age 7.1 ± 0.15 Ma (Karadenizli et al. 2005). L OCALITY . — Akkaşdağı, Çankırı-Çorum Basin, Turkey. DESCRIPTION According to the mandibular data, seven adult, one young adult and two young individuals compose the local population, which includes one mature male, and at least two mature females. Skull There are two skulls in the Akkaşdağı suid collection, AK3-131 and AK5-501. The latter is only a middle part of a skull with the face partially preserved. Apart from showing clearly that the occiput is very high, it adds little to what is seen in the well GEODIVERSITAS • 2005 • 27 (4) preserved skull AK3-131 (Fig. 1). The latter skull is almost complete but dorso-ventrally compressed, especially in the occipital region (Fig. 1A). Judging from the completely erupted M3 it belongs to an adult individual. In dorsal view (Fig. 1B), the caudal part of the skull is broken at the posterior part of the parietal, and much of the braincase is not preserved. The left zygomatic arch is missing, while the right side is well preserved. The nasals are long, of nearly constant width almost to the tip. In ventral view (Fig. 1C), the specimen is almost complete from the apices of the premaxillae to the occipital condyles, but the pterygoid process is heavily deformed by compression and difficult to investigate. The right zygomatic arch is robust and extends strongly towards lateral. There are no facial crests, and the anterior rim of the zygomatic arch originates at the anterior end of M3. The orbit is small and far behind M3. The rim of the orbit is incomplete but there is a distinctive, deep lachrymal notch (infraorbital fossa). The occipital condyles and sphenoid surfaces are of typical suid form. Although broken, the jugular processes seem robust and the tympanic bullae are oriented downwards, both suggesting a modern suid form. The choanae open posteriorly, far behind M3. The cheek dentition (P1-M3) is well preserved, but all the canines and incisors are missing. The small and shallow canine alveolus suggests that the canine was small. The alveolar crest is elongated and relatively slender. Mandible The mandible is the most common element in the Akkaşdağı suid collection. The best preserved specimen is labelled as AK3-126 (Fig. 2A), and is certainly associated with the skull AK3-131. The mandible is almost complete except for minor damage, but the canine is unfortunately missing. Two mandibles preserve canines, AK11-72 (Fig. 2B) and AKK120 (Fig. 2C). The description of the mandibular morphology of the Akkaşdağı suid is mainly based on specimens AK3-126, AK11-72, and AKK-120. The horizontal ramus is shallow and slim, while the ascending ramus is high, about three times the 717 Liu L. et al. FIG. 1. — Microstonyx major (Gervais, 1848) from Akkaşdağı, 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 Akkaşdağı FIG. 2. — Microstonyx major (Gervais, 1848) from Akkaşdağı; 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. FIG. 3. — Microstonyx major (Gervais, 1848) from Akkaşdağı, 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 seem to be two kinds of canine, but the distinction is not very sharp. The canine in mandible AKK120 is short and narrow, almost symmetrical, with a very narrow posterior facet separated from the lateral facets by two crests. Since the tooth is narrow, the boundary of the lateral facets forms a sharp anterior crest. Enamel covers the whole tooth and ends above the alveolus. It is virtually identical with the canine that we described from Hezheng as a female individual (Liu et al. 2004). The canine in mandible AK11-72 is slightly more robust, with an oval transverse section and no obvious crests separating the posterior facet from the lateral ones. Dentition The dental morphology of Microstonyx is greatly variable, and has been shown extensively in previous publications (e.g., Van der Made et al. 1992; Kostopoulos et al. 2001; Liu et al. 2004, and literature listed herein), and the Akkaşdağı suid fits well within the known range. Notable characteristics of the Akkaşdağı population include a somewhat complicated M3/m3 occlusal pattern, and 720 FIG. 4. — Microstonyx major (Gervais, 1848) from Akkaşdağı, astragalus AK4-88. Scale bar: 2 cm. the main lingual cusp of p4 being placed as far forward as the labial one. Postcranials Late Miocene suid limb bones are rarely described and figured, making it difficult to supply a comparative study of the abundant postcranial material of the Akkaşdağı suid. In order to facilitate future comparative work we here present figures and measurements of the material (Appendix: Tables 1; 2; Figs 3-5). COMPARISON AND DISCUSSION The general morphological characters of the Akkaşdağı suid, such as its large size, the elevated occiput, the wide and flat frontoparietal region, the elongated snout, the inflated and laterally extended zygomatic arches, the wide and deep lachrymal notch, and well developed alveolar GEODIVERSITAS • 2005 • 27 (4) Late Miocene Suidae from Akkaşdağı FIG. 5. — Microstonyx major (Gervais, 1848) from Akkaşdağı, metapodials; A, AK5-199; B, AKK-82; C, AK5-48. Scale bar: 5 cm. crest show unambiguously that it belongs to the Eurasian Hippopotamodon-Microstonyx large suine group (Trofimov 1954; Pickford 1988; Van der Made & Moyà-Solà 1989; Kostopoulos et al. 2001; Liu et al. 2004). The suines of the late Miocene Eurasian Hippopotamodon-Microstonyx group are characterised by considerable morphological variability, with a confusing pattern of sexual, temporal, and regional differences. Hippopotamodon (Lydekker, 1877) and Microstonyx are very similar in cranial and dental morphology. The characteristics of Hippopotamodon emphasized by Pickford (1988), such as the gigantic size, the large and flaring canine, the short snout, the less developed alveolar crest, and the short diastema between C-P1-P2, do not qualitatively distinguish the two genera, especially not when the material is incomplete, as is usuGEODIVERSITAS • 2005 • 27 (4) ally the case. The Akkaşdağı suine does have a reduced canine, but since canine size in female individuals of Hippopotamodon is unknown, this is not in itself decisive. In size and general proportions the Akkaşdağı suid skull is similar to the skull from Pikermi figured by Gaudry (1862-1867), as well as to the skull specimen BMNH M9048 that probably represents a female individual. The Akkaşdağı skull is also close to the Russian (Trofimov 1954) and Bulgarian skull samples (especially to the presumed female skull specimen K5260 from Kalimanci; Kostopoulos et al. 2001) and is obviously larger and more elongated than any specimen from the Chinese latest Miocene locality Hezheng (Liu et al. 2004) (Appendix: Tables 3; 4). Microstonyx, as conventionally conceived, has two species, M. antiquus (Kaup, 1833) and M. major. 721 Liu L. et al. Microstonyx antiquus was larger and distributed earlier in time than M. major. Fortelius et al. (1996) tentatively assigned M. antiquus to Hippopotamodon and we follow this taxonomy here. Under this usage, Hippopotamodon differs from Microstonyx primarily in the premolar proportions, especially the relative stoutness of the fourth premolar in the former (Liu et al. 2004). As the dental logarithmic ratio diagrams show (Fig. 6), it is easy to assign the Akkaşdağı suid to M. major as opposed to Hippopotamodon. Microstonyx major was a common element of the late Miocene mammal faunas of Europe and West Asia. It is also known from East Asia, but was a rare element there, with evidence of substantial size decrease (Liu 2003). Some workers (Van der Made & Moyà-Solà 1989; Van der Made et al. 1992; Pickford 1993; Kostopoulos 1994; Bonis & Bouvrain 1996; Van der Made 1997) have interpreted the spatiotemporal variability of M. major as possible evidence of multiple evolving lineages. In contrast, we have recently argued (Liu et al. 2004) that the morphological variability of M. major s.l. is best interpreted as polymorphism within a single evolving species. The substantial inter-population size variability in M. major, is owing to intra-specific polymorphism, representing local ecotypes (Liu et al. 2004). We have specifically proposed that elongation of the skull in M. major tends to be associated with arid conditions and a dietary shift towards herbivory (Liu et al. 2004). Following this view here, we recognise only one formal species-level taxon, M. major. The tooth dimensions of Microstonyx from Akkaşdağı (Appendix: Tables 5; 6) are well within the range of the middle-late Turolian European M. major, and larger than those of other Asian samples (Maragha and China; Bonis & Bouvrain 1996; Liu et al. 2004) or from some early Turolian populations of SE Europe (Vathylakkos, Kerassia, Perivolaki; Kostopoulos et al. 2001) (Fig. 7). Logarithmic ratio diagrams illustrating the relative proportions of the Eurasian Microstonyx major toothrows are shown in Figure 8. The Akkaşdağı M. major is close to the Pikermian and Bulgarian samples but with some722 what reduced premolars. Under our ecotype hypothesis, the great elongation of the skull of the Akkaşdağı form would indicate the arid end of the habitat spectrum inhabited by the species. In the Balkans, similar ecotypes with long muzzles and medium sized molars occur mainly in the middle Turolian, MN 12 (Kostopoulos et al. 2001). MICROSTONYX IN TURKEY Microstonyx is poorly documented in the late Miocene of Turkey, even if references are numerous. Şenyürek (1952) described as Sus erymanthius (= Microstonyx major) a suid mandible with p3-m3 from the undated site Akkırma II near Gökdere. The specimen belongs to a large form with stout p4, correctly recognized by Bonis & Bouvrain (1996) as belonging to Microstonyx (Limnostonyx) antiquus. Dicoryphochoerus meteai Ozansoy, 1965 from the Vallesian locality of Yassıören (middle Sinap) is mainly based on a left mandibular ramus, which has been later considered by Pickford & Ertürk (1979) as belonging to Hippopotamodon and assigned by Bonis & Bouvrain (1996) to Microstonyx (Limnostonyx) antiquus. Both the Yassıören and Akkırma II suids, referred here to Hippopotamodon antiquus, differ from the Akkaşdağı one in the larger dimensions, stronger canine, persistence of p1 and more robust p4. Microstonyx major (including M. erymanthius) has been reported from several Turkish localities ranging from MN 10 to MN 12 (Gülpınar, Çorak Yerler, Çevril, Muğla Garkın, Kayadibi, upper Kavakdere, Çoban Pınar, Mahmutgazi and Kınık; Ozansoy 1965; Sickenberg 1975; Pickford & Ertürk 1979; Fortelius et al. 1996; NOW database 2003) but we usually deal with faunal lists or fragmentary and isolated specimens which preclude a direct comparison with the Akkaş dağ ı form. As far we know the Akkaşdağı M. major provides the clearest evidence of the species in Turkey. The absence of the species in the isochronous and very similar fauna of Kemiklitepe A, B looks rather anomalous. GEODIVERSITAS • 2005 • 27 (4) Late Miocene Suidae from Akkaşdağı FIG. 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. P4 M2 p4 m2 FIG. 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 Akkaşdağı FIG. 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. FIG. 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). SEXUAL DIMORPHISM The Akkaşdağı suid collection does not at present add decisively to our understanding of sexual dimorphism in M. major (Kostopoulos et al. 2001; Liu et al. 2004). The weaker alveolar crest and the lesser width of the skull AK3-131 compared to those of other samples suggest that the Akkaşdağı skull belongs to a female individual (Kostopoulos et al. 2001; Liu et al. 2004). Figure 9 summarizes the available data on the distribution of the alveolar crest dimensions according to sex. Evidently, the Akkaşdağı suid is located among the female individuals of M. major from Pikermi (Greece) and Kalimanci (Bulgaria) while the Hezheng form represents a shift toward smaller overall size. The fact that the presumed female skull retains P1 supports our previous suggestion that females retained P1 more frequently than males (Liu et al. 2004). In contrast to the presumed male skull from Hezheng (Liu et al. 2004), the presumed female skulls from Hezheng and Akkaşdağı have almost no development of facial crests. Similarly, the deep preorbital fossa found in the Hezheng presumed male individual is not as strongly developed in the presumed females. 726 It is not possible to establish beyond all doubt that the minor differences observed in the lower canines from Akkaşdağı reflect sexual dimorphism, but the pattern observed supports the suggestion of weak sexual bimodality (Liu et al. 2004). CONCLUSIONS The study of the suid material from Akkaşdağı allows its attribution to Microstonyx major, as this species is considered in the frame of our last works (Kostopoulos et al. 2001; Liu 2003; Liu et al. 2004). The Akkaşdağı form represents a typical middle Turolian (MN 12) eastern Mediterranean population with close similarities to the Greek and Bulgarian samples. Morphological cranial characters indicate relatively arid conditions, still within the range of a generalist species. Although the presence of the long-ranging M. major cannot as such provide certain biochronological conclusions, the population characteristics do appear to support a middle-late Turolian age for the Akkaşdağı assemblage. GEODIVERSITAS • 2005 • 27 (4) Late Miocene Suidae from Akkaşdağı Acknowledgements Geological studies and excavations at Akkaşdağı have been authorized by the General Directorate of Mineral Research and Exploration (MTA) and Ankara University, Faculty of Sciences. The work was done thanks to grants from the CNRS (ECLIPSE Program and DRI), the Muséum national d’Histoire naturelle, Paris, MTA and TUBITAK. They have financially supported fieldwork and reciprocal visits to the concerned institutions. 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Geodiversitas 23 (3): 411-437. KOUFOS G. D. & VLACHOU T. D. 2005. — Equidae (Mammalia, Perissodactyla) from the late Miocene of Akkaşdağı, Turkey, in SEN S. (ed.), Geology, mammals and environments at Akkaşdağı, late Miocene of Central Anatolia. Geodiversitas 27 (4): 633-705 (this volume). LIU D. S., LI C. K. & ZHAI R. J. 1978. — [Pliocene vertebrates of Lantian, Shensi]. Professional Papers 727 Liu L. et al. on Stratigraphy and Paleontology 7: 149-200 (in Chinese). LIU L. P. 2003. — Chinese Fossil Suidea: Systematics, Evolution and Paleoecology. University Printing House, Helsinki, 41 p. L IU L. P., K OSTOPOULOS D. S. & F ORTELIUS M. 2004. — Late Miocene Microstonyx remains (Suidae, Artiodactyla) from Hezheng areas. Geobios 37 (1): 49-64. NOW DATABASE 2003. — Neogene of the Old World. Database of fossil mammals: www.helsinki. fi/science/now OZANSOY F. 1965. — Étude des gisements continentaux et de mammifères du Cénozoïque de Turquie. Mémoires de la Societé géologique de France N. 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Geologisches Jahrbuch B, 15: 1-167. S TEININGER F. F. 1999. — Chronostratigraphy, geochronology and biochronology of the Miocene “European Land Mammal Mega-Zones” (ELMMZ) and the Miocene “Mammal-Zones (MN-Zones)”, in R ÖSSNER G. E. & H EISSIG K. (eds), The Miocene Land Mammals of Europe. Verlag Dr. Friedrich Pfeil, München: 9-24. STEININGER F. F., BERGGREN W. A., KENT D. V., BERNOR R. L., SEN S. & AGUSTÍ J. 1996. — CircumMediterranean Neogene (Miocene-Pliocene) marine-continental chronologic correlations of European mammal units, in B ERNOR R. L., FAHLBUSCH V. & MITTMANN H. W. (eds), The Evolution of Western Eurasian Neogene Mammal Faunas. Columbia University Press, New York: 7-46. TANG Y. J., LIU Z. Q., CHEN D. & CHEN L. Q. 1985. — The fossil Suidae from late Miocene of Binxian, Shaanxi. Vertebrata PalAsiatica 23 (1): 60-68. TROFIMOV B. A. 1954. — The fossil suids of the genus Microstonyx, in Tertiary Mammals, part 2: on the Mammalia of the southern SSSR and Mongolia. Doklady Akademii Nauk SSSR 47: 61-99. VAN DER MADE J. 1997. — The fossil pig from the late Miocene of Dorn-Dürkheim 1 in Germany. Courier Forschungs-Institut Senckenberg 197: 205230. V A N D E R M A D E J. & H U S S A I N S. T. 1989. — “Microstonyx” major (Suidae, Artiodactyla) from Nagri. Estudios Geologicos 45: 409-416. V A N D E R M A D E J. & M O Y À -S O L À S. 1989. — European Suinae (Artiodactyla) from the late Miocene onwards. Bollettino della Società Paleontologica Italiana 28 (2-3): 329-339. VAN DER MADE J., MONTOYA P. & ALCALÁ L. 1992. — Microstonyx (Suidae, Mammalia) from the upper 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 Akkaşdağı APPENDIX TABLE 1. — Measurements of atlases and one indetermined vertebra of Microstonyx major (Gervais, 1848) from Akkaşdağı (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 Element M1 M2 M3 M4 M5 M6 M7 M8 M9 M10 76.0 36.0 AK3 Atlas 66.0 77.0 42.0 AK5-236 Atlas 77.0 76.0 39.0 AK5-258 Atlas AK5-346 Atlas AK5-628 Atlas 148.0 81.8 153.0 63.0 83.0 38.5 60.0 77.0 34.0 70.0 81.2 85.0 AK5b-838 Atlas 67.0 AK6-91 Atlas 67.0 AK6-96 Vertebra 55.0 45.0 33.0 76.4 32.0 38.0 75.0 34.0 38.0 77.0 35.0 37.0 TABLE 2. — Measurements of limb bones of Microstonyx major (Gervais, 1848) from Akkaşdağı (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 Element Side M1 M2 M3 M4 M5 M6 M7 M8 M9 M10 AK4-109 Distal humerus left 57.5 62.4 47.5 43.3 AK7-184 Distal humerus left 66.8 69.0 51.2 47.8 AK11-52 Proximal radius right 60.0 40.0 52.2 29.4 54.0 37.0 44.4 29.5 23.3 17.1 51.9 33.6 34.6 AK3-302 Distal radius left 33.0 AK5-188 Distal radius left 28.0 AK5-570 Proximal radius left 44.1 30.9 AK6-258 Proximal radius left 49.0 35.0 AK3-302 Distal ulna left AKB-54 Metacarpal III left AKK-82 Metacarpal III right AK5-625 Proximal right metacarpal III AK5-48 Metacarpal IV right AK2-489 AK4-88 AK5-149 Distal tibia left Astragalus left Metatarsal IV right GEODIVERSITAS • 2005 • 27 (4) 27.2 24.0 109.0 105.0 30.4 29.1 21.9 15.7 26.9 29.6 25.3 29.6 107.0 103.0 32.6 30.6 22.6 16.1 29.8 27.1 26.7 27.0 30.0 24.4 30.0 24.4 21.3 13.3 25.8 27.2 26.1 27.2 43.7 39.6 35.4 34.1 27.9 24.4 28.2 105.0 99.0 62.2 35.1 34.9 30.4 34.1 34.7 26.3 113.0 109.0 25.0 34.3 20.0 15.7 25.1 729 Liu L. et al. TABLE 3. — Skull measurements of Microstonyx major (Gervais, 1848) from Akkaşdağı compared with specimens from Pikermi (Greece) and Hezheng (China) (in mm). Data from Gaudry (1862-1867), Liu et al. (2004) and personal data. Pikermi Akkaşdağı Skull BMNH M9048 HMV 0976 male 420 < 380 350 – – 123 – 260 227 – 234 202 195 AK3-131 female Basal length of the skull Condyles to posterior border of palatine fissure Condyles to posterior border of palate Posterior border of palatine fissure to posterior border of palate Posterior border of palatine fissure to the end of M3 Posterior border of palate to M3 Length of the diastema between canine and P2 Length of P2-M3 P2-P4/M1-M3 Length of alveolar crest M3 to the anterior end of orbit Maximal length of the orbit Maximal width of the skull Width of palate at alveolar crest Width of palate at anterior end of M3 Breadth of frontal at the supraorbital process Nasal width above preorbital foramen Nasal width above P2 Nasal width at the end of incisor notch Width of the palatine fissure 468 374 Gaudry’s specimen Hezheng 470 310 114 125.9 26 40.8 153 0.57 64 64 – 262 109 42 33 44 141 87 310 140 25 51.7 143 0.58 59 272 133 HMV 0977 female – 44 123.6 0.53 67 50 42.5 < 225 122 38.3 150 121.6 0.52 50 – – – 89.5 32 128.6 75 56 59 22 – – – – 83 66.5 69 21 TABLE 4. — Mandible measurements of Microstonyx major (Gervais, 1848) from Akkaşdağı compared with specimens from Hezheng (China) and Kalimanci (Bulgaria) (in mm). Data from Kostopoulos et al. (2001), Liu et al. (2004). Mandible (sex) Mandible length from tip to m2 Length of p2-m2 p2-p4/m1-m2 Length of symphysis Diastema between c-p2 Height of mandible at p2 Height of mandible at muscular process Height of mandible at articular process 730 Akkaşdağı AK11-72/ AKK-120 (male/female) Akkaşdağı AK3-126 (female) Kalimanci K-5277/K-5276 (male/female) Hezheng HMV 0576 (female) 212/– 99/– 0.56/– 100.5/85.5 47/38.5 52.6/43.6 – – 102 0.54 100 56.8 53.8 171 233/239 111/111 192 98 102/87 52/50 56/47 – 80 40 – _ – 152 – GEODIVERSITAS • 2005 • 27 (4) Late Miocene Suidae from Akkaşdağı TABLE 5. — Upper dentition measurements of Microstonyx major (Gervais, 1848) from Akkaşdağı (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 dataline, right toothrow. Specimen I1 AK11-66 L W1 W2 W3 AK12-5 L W1 W2 I3 C P1 P2 P3 P4 M1 M2 M3 16.0 10.0 16.6 15.5 14.3 16.9 19.7 18.0 18.4 27.1 22.2 22.8 39.5 24.9 22.2 14.8 15.3 15.3 9.5 9.3 16.8 16.6 14.9 14.4 17.0 16.4 18.2 17.9 21.8 21.5 19.5 19.5 19.2 19.6 29.7 28.5 24.5 23.5 24.0 24.9 40.5 40.4 26.1 26.1 23.5 23.7 13.4 13.8 9.9 16.1 L 9.3 9.6 4.5 4.6 W1 AK3-131 I2 W2 W3 AK4-186 L W1 W2 W3 43.4 28.1 24.4 12.0 L 15.3 15.9 9.5 8.7 W1 AK5-501 16.9 15.3 15.5 15.4 15.4 14.7 18.7 18.4 19.6 18.9 20.1 20.2 21.0 21.9 26.0 26.0 24.2 24.7 24.8 24.9 41.6 41.0 27.2 27.3 24.3 24.8 15.0 15.8 17.6 16.8 16.3 19.4 22.5 19.6 19.0 30.1 24.8 23.8 42.8 27.4 24.6 17.1 26.8 26.5 23.0 22.8 22.4 22.8 38.6 39.3 24.3 24.2 22.2 22.0 11.3 11.3 W2 W3 AK5-623 L W1 W2 W3 AK5-624 L W1 W2 AK7-100 L W1 W2 AK7-153 22.5 10.6 27.0 21.6 18.7 18.3 L 15.9 W1 10.1 W2 W3 AKA-1 L W1 W2 W3 AKK-83 L W1 W2 GEODIVERSITAS • 2005 • 27 (4) 16.5 16.1 14.9 14.5 14.9 14.5 18.1 18.0 20.2 21.0 18.7 18.7 41.7 26.1 23.2 12.8 11.8 8.2 16.2 731 Liu L. et al. TABLE 6. — Lower dentition measurements of Microstonyx major (Gervais, 1848) from Akkaşdağı (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 AK11-67 i1 i2 L 6.5 W1 9.3 i3 c p2 p3 p4 m1 W1 AK11-72 W2 8.6 9.1 14.0 14.5 35.7 34.0 18.3 17.8 14.1 12.1 30.2 32.0 m3 22.9 22.8 16.0 15.9 15.0 15.8 W2 L m2 17.5 16.2 6.3 6.5 19.3 19.0 9.0 11.0 6.0 6.5 12.2 11.0 15.0 14.6 6.6 6.6 18.3 18.4 9.3 8.5 19.6 20.2 14.5 14.4 20.7 20.6 14.9 14.7 14.3 14.3 28.7 27.6 18.4 18.2 18.6 19.2 41.9 42.7 21.5 21.2 20.0 20.3 15.9 16.0 14.2 6.4 17.2 9.0 19.6 14.4 20.4 14.1 14.8 28.8 19.2 19.3 48.9 21.5 20.2 16.1 18.3 8.4 19.8 13.1 21.8 14.0 14.8 29.0 18.3 18.5 46.1 22.1 20.6 17.0 17.5 17.8 9.2 9.0 19.8 19.1 14.7 14.8 18.5 18.4 14.7 14.2 15.2 14.6 25.5 25.2 18.9 19.1 20.8 20.0 43.4 42.3 20.6 21.8 20.9 20.3 16.4 17.0 19.4 10.5 20.4 16.5 21.6 16.3 15.9 28.5 20.7 20.4 46.0 22.5 21.0 17.3 26.6 18.9 19.2 43.4 22.2 20.7 16.1 W3 AK2-112 L W1 W2 W3 AK2-488 L W1 W2 AK3-126 L W1 W2 W3 L W1 AK4-187 21.6 7.3 17.5 14.0 7.2 W2 W3 AK4-251 L W1 W2 W3 AK5-270 L W1 W2 AK5-442 L W1 W2 W3 AK7-154 L W1 W2 732 21.5 14.4 14.3 14.3 7.5 17.5 9.9 19.8 14.4 20.5 14.6 15.5 28.4 20.4 21.0 GEODIVERSITAS • 2005 • 27 (4) Late Miocene Suidae from Akkaşdağı Specimen i1 AK7-183 L W1 W2 W3 AKB-51 L W1 W2 L AKK-120 W1 W2 i2 i3 c p2 p3 p4 m1 m3 44.3 22.5 20.8 16.9 10.1 9.5 8.0 8.0 12.3 13.4 16.5 17.2 7.5 6.9 18.2 18.4 17.2 16.5 6.9 7.5 18.2 18.4 15.2 15.8 7.4 7.4 18.4 10.0 21.1 12.4 19.2 19.5 9.7 10.1 21.5 23.0 15.9 15.6 29.8 20.5 20.2 29.5 18.9 19.3 14.7 AKK-121 L W1 W2 W3 22.2 15.1 15.3 AKK-286 L W1 W2 22.3 14.2 15.0 AKK-287 L W1 W2 29.0 19.8 20.1 AKK-288 L W1 W2 28.5 18.5 19.8 GEODIVERSITAS • 2005 • 27 (4) m2 39.2 20.9 20.8 16.3 733