Late Miocene bovids from Şerefköy-2 (SW Turkey) and their position within the sub-Paratethyan biogeographic province DIMITRIS S. KOSTOPOULOS and SEVAL KARAKÜTÜK Kostopoulos, D.S. and Karakütük, S. 2015. Late Miocene bovids from Şerefköy-2 (SW Turkey) and their position within the sub-Paratethyan biogeographic province. Acta Palaeontologica Polonica 60 (1): 49–66. We describe new fossil bovid craniodental remains from the Upper Miocene fossil site of Şerefköy-2, Yatağan Basin, SW Turkey. The new material belongs to six species: Gazella cf. G. capricornis, Palaeoryx pallasi, Sporadotragus parvidens, Skoufotragus cf. Sk. schlosseri, Urmiatherium rugosifrons, and ?Sinotragus sp., which together indicate a latest middle–early Late Turolian (Late Miocene) age. Medium-to-large bovid taxa prevail over small ones, and protoryxoid bovids clearly dominate the assemblage. An analysis of the taxonomic structure, size and diet spectra of several Turolian bovid assemblages from Greece and Turkey reveals Şerefköy-2 to be a member of a mammalian palaeocommunity particular to southwestern Anatolia, which in turn forms part of the sub-Paratethyan biogeographic province. Ke y w o rd s : Mammalia, Bovidae, Antilopinae, biogeography, Miocene, Turolian, Turkey, Anatolia. Dimitris S. Kostopoulos [dkostop@geo.auth.gr], Aristotle University of Thessaloniki, School of Geology, 54124 Thessaloniki, Greece. Seval Karakütük [sevaloruc.ege.univ@hotmail.com], Natural History Museum, Ege University, 35100 Izmir, Turkey. Received 7 November 2012, accepted 10 June 2013, available online 17 June 2013. Copyright © 2015 D.S. Kostopoulos and S. Karakütük. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Introduction Anatolia, located in modern-day Turkey, plays a crucial role in the reconstruction of late Neogene Eurasian palaeozoogeography and palaeoenvironments (e.g., Costeur and Legendre 2008; Eronen et al. 2009; Kostopoulos 2009b), because it represents the natural link between the western and eastern extremes of the so-called Late Miocene sub-Paratethyan biogeographic province (Bernor et al. 1979). There area is rich in Late Miocene fossil sites, but in most cases their contents have only been reported in the form of preliminary faunal lists (e.g., Sickenberg et al. 1975; Alan 1997; Saraç 2003). Pioneer works by Ozansoy (1965), Tekkaya (1973a, b), Senyürek (1952, 1953), Boscha-Erdbrink (1978), and Köhler (1987) only gave a fragmentary view of the rich and abundant Late Miocene bovid assemblage of Anatolia, which were partially, but never entirely, filled by later studies on newly discovered or re-examined material (e.g., Bouvrain 1994; Gentry 2003; Kostopoulos 2005; Geraads and Güleç 1999; Bibi and Güleç 2008). Thee field seasons at the new locality of Şerefköy-2, discovered in the summer of 2007 in Yatağan Basin, SW Turkey (Fig. 1; coordinates: N 37°21’47.80’’, E 28°14’11.10’’; see Kaya et al. 2012 for further information on the geological context), yielded more than 1200 identifiable fossil speciActa Palaeontol. Pol. 60 (1): 49–66, 2015 mens belonging to more than 25 species of mammals. Together, these represent one of the richest Turolian faunas from Anatolia, and offer a chance to (i) test faunal correlations within the sub-Parathethyan biogeographic province and (ii) further develop the local biochronological framework. Here, we provide a detailed description of the bovids forming part of the Şerefköy-2 bovid assemblage, preliminary reported by Kaya et al. (2012). The remaining taxa recovered from this site will be described elsewhere. Institutional abbreviations.—AMNH, American Museum of Natural History, New York, USA; MYŞE PV, collection of the Natural History Museum of Ege University, Izmir, Turkey; KNUA, Kapodistrian National University of Athens, Greece; LGPUT, Laboratory of Geology and Palaeontology, Aristotle University of Thessaloniki, Thessaloniki, Greece; MNHN, Museum national d’Histoire naturelle, Paris, France; NHMA, Aegean Museum of the Natural History-Zimalis Foundation, Samos Island, Greece; NHMW; Naturhistorisches Museum Wien, Austria. Other abbreviations.—APD, anteroposterior diameter; CFA, correspondence analysis; L, length; n, number of specimens; TD, transverse diameter; W, width. Upper and lower case letters denote upper and lower teeth, respectively. http://dx.doi.org/10.4202/app.2012.0129 ACTA PALAEONTOLOGICA POLONICA 60 (1), 2015 A B Black Sea Milet Fm. 50 Ankara TURKEY Mediterranean Sea marl breccia Eskihisar Fm. silty marl conglomerates ˆ fossil site Şerefköy-2 coal deposits Yatagan Fm. Muğla C This allowed us to (i) overcome differing taxonomic opinions; (ii) incorporate genera with low abundances; and (iii) emphasise general patterns in taxonomic structure. Finally, we compared the results the CFA analyses with the size distribution and diet spectra of the involved bovid faunas. The latter reflect crucial ecological parameters (body size and feeding prefeences) that are relevant at the level of both the organism and the community as a whole (e.g., Western 1979; McNaughton and Georgiadis 1986; Eisenberg 1990). Multivariate statistics were performed using PAST version 2.16 (Hammer et al. 2001). Skull and horn core nomenclature follows Gentry (1971, 1992). Dental nomenclature follows Bärmann and Rössner (2011). All measurements are in millimetres (mm). Further details on all the analyses and the data themselves are provided as Supplementary Online Material (SOM 1, SOM 2, available at http://app.pan.pl/ SOM/app60-Kostopoulos_Karakutuk_SOM.pdf). Systematic palaeontology Class Mammalia Linnaeus, 1758 Order Artiodactyla Owen, 1848 Family Bovidae Gray, 1821 Subfamily Antilopinae Gray, 1821 (including all nonBovinae taxa sensu Kingdon 1982; e.g., Groves and Grubb 2011) Genus Gazella Blainville, 1816 Type species: Gazella dorcas Linnaeus, 1758, Recent. Gazella cf. G. capricornis (Wagner, 1848) Fig. 2. D Fig. 1. The fossil mammal site of Şerefköy-2 (Turkey). A. Geographical location. B. Local lithostratigraphy (modified from Kaya et al. 2012). C. Typical fossil accumulation at Şerefköy-2. D. View of the fossiliferous site. Material and methods Horn core and cranial proportions of different bovids were explored using discriminant and Principal Components analyses. Following Bibi and Güleç (2008), Kostopoulos (2009b), Koufos et al. (2006, 2009a), and Kostopoulos and Bernor (2011), we performed a correspondence analysis (CFA) to investigate the structure of various Turolian (8.7–5.4 Ma; Late Miocene) bovid assemblages from Greece, Turkey and Iran. Following Bibi and Güleç (2008), we first analysed bovid assemblages according to their relative abundances at the genus level. Next, we repeated the analysis by grouping all genera into five units reflective of the basic taxonomic structure of each assemblage. Material.—MYŞE PV-2501, frontlet; MYŞE PV-1577, PV1834, right horn cores; MYŞE PV-560, PV-1578, left horn cores; MYŞE PV-1831, PV-1832, partial right horn cores; MYŞE PV-1359, PV-1632, PV-1828, partial left horn cores; MYŞE PV-1360, partial horn core; MYŞE PV-1830, 1633, 1393, distal portions of horn cores; MYŞE PV-2572, palate; MYŞE PV-2575 right upper tooth row P2–M3; MYŞE PV2000, PV-2558, PV-2562, PV-2564, right mandibular body with p2–m3; MYŞE PV-2557, PV-2563, left mandibular body with p2–m3; MYŞE PV-2565, left mandibular body with p3–m3; MYŞE PV-1526, right m3. All from Şerefköy-2, Turkey, Late Turolian (Late Miocene). Description.—Gazella cf. G. capricornis is the only gazelle present at Şerefköy-2, and represented by at least 5 individuals. The supraorbital foramen is located close to the pedicle, within an elliptical and rather narrow depression (Fig. 2C1). The postcornual fossa is oval, small, and moderately deep. The pedicle is moderately high anteriorly (Fig. 2C). The horn cores are short (maximum length: 100 mm along the anterior surface), inserted moderately far apart from each other, and weakly divergent distally (Fig. 2C1). In lateral view, the horn core is slightly curved and moderately inclined posteriorly (Fig. 2A, B, C2), while in cross section it changes from be- KOSTOPOULOS AND KARAKÜTÜK—LATE MIOCENE BOVIDS FROM TURKEY A B 51 C1 C2 5 cm (A–C) D1 E 5 cm D2 (D–F) F Fig. 2. The bovid artiodactyl Gazella cf. G. capricornis (Wagner, 1848) from Şerefköy-2 (Turkey), Late Turolian (Late Miocene). A. Left horn core (MYŞE PV-1578) in lateral view. B. Left horn core (MYŞE PV-560) in lateral view. C. Frontlet (MYŞE PV-2501) in frontal (C1) and lateral (C2) views. D. Left mandibular body (MYŞE PV-2557) in occlusal (D1) and lingual (D2) views. E. Right mandibular body (MYŞE PV-2564) in occlusal view. F. Right P2-M3 of the palate (MYŞE PV-2572) in occlusal view. ing symmetrically elliptical near the base (owing to weak mediolateral compression) to a more rounded shape towards the tip (TD × 100/APD = 70.4–73.2 at the base, n = 4, and 84.6–104.6 at 7 cm above the base, n = 3; Table 1). Strong, deep, and continuous furrows run along both the anterior and posterior surfaces of the horn core. The premolars are moderately long compared to the molars (Fig. 2D–F), with the upper and lower premolar/molar ratios equalling about 71% (n = 1) and 54–60% (n = 6), respectively (SOM 3: Tables 1, 2). There is no entostyle on the upper molars. M3 has a strong metastyle and a weak mesostyle (Fig. 2F). The anterior conid of p3 and p4 is weak, but separated from the anterior stylid (MYŞE PV-2000). The meTable 1. Horn core measurements (in mm) of Gazella cf. capricornis from Şerefköy-2 (Turkey). TD, transverse diameter at the base and at 7 cm above the base; APD, anteroposterior diameter at the base and at 7 cm above the base. Specimen MYŞE PV-2501 MYŞE PV-560 MYŞE PV-1577 MYŞE PV-1578 TDbase 16.4 17.8 15.3 16.4 APDbase 23.1 24.3 21.7 22.2 TD7 12.8 14.7 10.0 APD7 14.8 14.1 11.8 solingual conid is elongated and oriented posteriorly on p3, but more triangular (during early wear) and located relatively further posteriorly on p4 (Fig. 2D, E). The anterior valley of p3 and p4 is widely open (Fig. 2D, E). The posterior valley of p4 is open in the single unworn specimen (MYŞE PV-2000) and still visible even at advanced stages of wear (Fig. 2D, E). A small ectostylid is present on m1. The mesostylids on m2 and m3 are strong. Remarks.—The gazelle from Şerefköy-2 is considerably larger than Gazella ancyrensis Tekkaya, 1973 from Middle Sinap, as well as G. cf. ancyrensis from Maragheh (Iran), Samos (Greece), and Kemiklitepe-D (Turkey), but smaller than Gazella mytilinii Pilgrim, 1926 from Samos (Bouvrain 1994; Kostopoulos 2009a; Kostopoulos and Bernor 2011). Gazella pilgrimi Bohlin, 1935 from Samos and the Axios Valley (Greece) and Akkaşdağı (Turkey) differs from the Şerefköy-2 specimens in having a shorter pedicle, as well as a longer, more mediolaterally compressed, and more strongly inclined horn core (Bouvrain 1996; Kostopoulos 2005, 2009a). The overall shape of the horn core of the Şerefköy-2 gazelle recalls both Gazella deperdita (Gervais, 1847) from Western Europe and the SE European Gazella capricornis (Wagner, 1848). However, the horn core of G. deperdita 52 ACTA PALAEONTOLOGICA POLONICA 60 (1), 2015 Palaeoryx pallasi (Wagner, 1857) Discriminant function: D = -0.75TDbase + 0.21APDbase - 1.14TD7 + 1.31APD7 Fig. 4. 18 G.caprinornis, Pikermi: #55 G. deperdita, Luberon: #31 Hotelling t2: 108.53 P (same): 6.1E-14 16 Frequency 14 12 10 8 6 4 2 0 -8 -6.4 -4.8 -3.2 -1.6 0 1.6 3.2 4.8 6.4 Discriminant Fig. 3. Discriminant analysis of the horn core proportions of Gazella deperdita from Luberon (= Cucuron), Late Miocene, SW Europe and Gazella capricornis from Pikermi, Late Miocene, Greece, based on four variables: TD at the base (TDbase), APD at the base (APDbase), TD at 7 cm above the base (TD7) and APD at 7 cm above the base (APD7). Arrows indicate values for the Şerefköy-2 gazelle according to the discriminant function. Abbreviations: APD, anteroposterior diameter; TD, transverse diameter. is larger (Fig. 3), much more curved in lateral view, more inclined posteriorly, and more mediolaterally compressed along its distal portion. Gazella capricornis was originally described from Pikermi (Greece), but similar forms are widely distributed between the Balkans and Iran (Kostopoulos 2005, 2009a; Kostopoulos and Bernor 2011). Both Gazella cf. G. capricornis from Samos (particularly specimens from Mytilinii-1A) and the Şerefköy-2 gazelle are characterised by an anterior conid distinct from the anterior stylid on p3 and p4, and the closing of the posterior valley of p4 at late wear stages. Together with G. cf. G. capricornis from Akkaşdağı and Maragheh, they further differ from typical specimens of G. capricornis from Pikermi in having slightly shorter premolars compared to the molars (Table 2). Genus Palaeoryx Gaudry, 1861 Type species: Antilope pallasi (Wagner, 1857), Pikermi, Greece; Late Miocene. Material.—MYŞE PV-2573, right upper tooth row with P2– M3; MYŞE PV-1293, right upper tooth row with P3–M3; MYŞE PV-1294/B, partial left upper tooth row with M2–M3; MYŞE PV-1295, partial left upper tooth row with P4–M1; MYŞE PV-1599, left mandibular body with p2–m3; MYŞE PV2574, right mandibular body with p2–m3; MYŞE PV2552, partial of right mandibular body with m2–m3. All from Şerefköy-2, Turkey, Late Turolian (Late Miocene). Description.—Specimens from Şerefköy-2 referable to this taxon are limited to some upper and lower tooth rows, representing at least two individuals. The premolars are long compared to the molars (Fig. 4, SOM 3: Tables 1, 2), with upper and lower premolar/molar ratios of 70.8% (n = 1) and 64.3–66.7% (n = 2), respectively. P4 is narrower than both P3 and P2, and P2 is shorter than P3. The anterior style is weakly developed on P2, but sharp, posteriorly curved and joined to the anterolabial cone on P3 (Fig. 4C). The “metaconule” is barely recognisable on P2 and P3, which have a distinctly convex lingual wall, and absent on P4, in which the anterolingual and posterolingual crista of the lingual cone are symmetrically developed (Fig. 4C). The anterior style of P4 is more pronounced than the anterolabial cone, with the latter having a more central position on the labial wall of the tooth than on P3 (Fig. 4C). All upper molars bear a fossetta (central islet) and low, double entostyles, originating from both the protocone and the metaconule (Fig. 4C). The anterior lobe of M1 and M2 is narrower than the posterior one. The development of the mesostyle on the upper molars ranges from weak in MYŞE PV-2573 to strong in PV-1293 and PV-1294. The p2 has a simple and strong anterior stylid and no anterior conid. The mesolingual conid is also simple, short and placed both anteriorly and almost perpendicularly to the longitudinal axis of the tooth (Fig. 4A, B). The posterior stylid on p3 is much less developed than the posterolingual conid, with both structures becoming fused during late stages of wear (Fig. 4A, B). The anterior conid and the anterior stylid are indistinct, thus forming a robust anterior cuspid. The mesolingual conid is elongated and slants distally, joining the talonid at advanced stages of wear (Fig. 4A). The p4 resembles p3, but is larger, and retains and open posterior valley until the onset of late stages of wear (Fig. 4A, B). The posterolabial conid is indistinct on p2 and p3, and weakly developed on p4. The lower molars have a strong mesostylid, Table 2. Comparison of Gazella cf. G. capricornis from Şerefköy-2 with G. cf. G. capricornis from different localities and with G. capricornis from Pikermi. Data are from Kostopoulos (2005, 2009a) and Kostopoulos and Bernor (2011). APD, anteroposterior diameter; L M/m, length of upper/lower molar row; L P/p, length of upper/lower premolar row; TD, transverse diameter at the base and at 7 cm above the base. APD × 100/TDbase APD × 100/TD7 LP/LM × 100 Lp/Lm × 100 Gazella cf. G. capricornis Şerefköy-2 (Turkey) Akkaşdağı (Turkey) Samos (Greece) 135.4–141.8 103.3–137.6 102.6–139.7 95.9–118.0 100.0–128.9 100.0–125.0 71.1–71.2 67.7–74.6 67.0–73.0 54.0–60.2 55.2–56.4 58.0 Maragheh (Iran) 112.2–147.4 103.8–128.7 76.5 58.8–61.8 Gazella capricornis Pikermi (Greece) 102.5–139.2 85.0–136.4 75.4–78.4 59.7–70.8 KOSTOPOULOS AND KARAKÜTÜK—LATE MIOCENE BOVIDS FROM TURKEY 53 A B 5 cm C Fig. 4. The bovid artiodactyl Palaeoryx pallasi (Wagner, 1848) from Şerefköy-2 (Turkey), Late Turolian (Late Miocene). A. Left mandible (MYŞE PV1599) in occlusal view. B. Right mandible (MYŞE PV-2574) in lingual view. C. Right upper tooth row (MYŞE PV-2573) in occlusal view. moderate ectostylids (basal pillars), and no anterior cingulid (goat fold). The third lobe of m3 is rather large and bears a flat entoconulid, as well as a bulbous hypoconulid (Fig. 4A). Remarks.—Palaeoryx Gaudry, 1861 is mainly known from Turolian sites of the sub-Paratethyan zoogeographic province (e.g., Kostopoulos and Bernor 2011 and literature therein), although it may have appeared during the Vallesian (e.g., Vislobokova 2005). In Turkey, records of the genus are restricted to the localities of Kayadibi, Eski Bayırköy, and Mahmutgazi (Köhler 1987), as well as Kemiklitepe-A, B and Akkaşdağı (Bouvrain 1994; Kostopoulos 2005). Kostopoulos (2005, 2009a) recognised two species in the Aegean region, P. pallasi and P. majori Schlosser, 1904, which differ from each other mainly in terms of their cranial and horn core morphology. In addition, a larger species or variety might exist in Maragheh (Iran) and Ukraine (Kostopoulos and Bernor 2011). The specimens from Şerefköy-2 differ from P. majori from Samos and Akkaşdağı in having a relatively (compared to the premolars) and absolutely longer upper molar row (60.7–62.0 mm vs. 64.8–65.4 mm, based on the material from Akkaşdağı), as well as in lacking a central fold (hypoconal spur) on P3 and P4, and an anterior cingulid on the lower molars (Fig. 5; Kostopoulos 2009a). In addition, P. majori from Samos differs from the Şerefköy-2 material in the presence of basal pillars on the upper molars and in having a p4 bearing a centrally placed mesolingual conid and a fused posterolingual conid and posterior stylid. By contrast, the Şerefköy-2 specimens resemble P. pallasi from Samos and Pikermi in all of these characters, as well as in the presence of a posteriorly curved anterior style on P3, a more centrally placed anterolabial cone on P4 (relative to P3), a strong mesostyle on the upper molars, a weak posterolabial conid on p3 and p4, and a tendency for the mesolingual conid of p4 to be oriented anteroposteriorly. P. pallasi from Pikermi differs, however, in having distinct posterolingual and mesolingual conids on p3, as well as distinct anterior and mesolingual 54 ACTA PALAEONTOLOGICA POLONICA 60 (1), 2015 Length PM 140 130 120 110 100 70 72 80 78 76 74 Index P/M Length pm 140 130 120 62 64 66 68 70 Index p/m Palaeoryx pallasi Şerefköy-2 Samos Pikermi Palaeoryx sp. Kemiklitepe Maragheh Dytiko-2 Perivolaki Palaeoryx majori Samos Fig. 5. Scatter plot comparing the upper and lower premolar (P, p) / molar (M, m) ratio against the complete tooth row length of Palaeoryx from several localities. conids on p4, although the degree of variability of these features within P. pallasi is not well known. There is considerable size variation of both the upper and lower tooth rows within Palaeoryx. The lower tooth row from Şerefköy-2 (Lpm = 138.9 mm) appears close to that from Maragheh (LPM = 139.3 mm). By contrast, the upper tooth row (LPM = 111.5 mm) is considerably smaller than both that from Maragheh (LPM = 140.8 mm) and a particularly large specimen from Pikermi (MNHN PIK2447; e.g., Kostopoulos and Bernor 2011), but close to P. pallasi from Samos and other specimens from Pikermi. The available data are insufficient for statistically sound conclusions; nevertheless, in the absence of marked morphological differences, we suggest that the observed size variation may be intraspecific. Stratigraphic and geographic range.—Upper Miocene; Balkans to Iran. Genus Sporadotragus Kretzoi, 1968 Type species: Microtragus schafferi (Andree, 1926), Samos, Greece; Late Miocene. Sporadotragus parvidens (Gaudry, 1861) Fig. 6. Material.—MYŞE PV-2502, partial skull; MYŞE PV-1573, frontlet preserving the basal part of the horn cores; MYŞE PV-1300, partial left horn core; MYŞE PV-1522, right upper tooth row with P2–M3; MYŞE PV-1412. right M3; MYŞE PV-1423, left M3; MYŞE PV-1533, left M1; MYŞE PV-1511, right mandibular body with p3–m3; MYŞE PV-2561, right mandibular body with p2–m2; MYŞE PV-1429, right mandibular body with p4–m2; MYŞE PV-2556, right mandibular body with m1–m3; MYŞE PV-1630, left mandibular body with p3–m3; MYŞE PV-1407, PV-1574, left mandibular body with p2–m2; MYŞE PV-2569, left mandibular body with p4– m3; MYŞE PV-1406, PV-2559, left mandibular ramus with p2–m3; MYŞE PV-1311, left mandibular ramus with p4–m2. All from Şerefköy-2, Turkey, Late Turolian (Late Miocene). Description.—This species is represented by at least 7 individuals. The frontals form a 105–115° angle along the sagittal plane and appear moderately pneumatised above the orbits. The supraorbital foramen (doubled on the left side in both MYŞE PV-1573 and PV-2502) is small and round, placed well below the pedicle, and opens directly into the orbit (Fig. 6A). The interfrontal suture is complex in outline and slightly pinched between the horn cores (Fig 6A). The frontoparietal suture is also complex, Y-shaped, and runs very close to the base of the horn core. The postcornual fossa is shallow and round. As preserved, the horn core is long (maximum length: 210 mm along the anterior surface) and gently curved posteriorly. It furthermore bears thin and discontinuous grooves on its surface and shows no evidence of torsion (Fig. 6A). In MYŞE PV-1300, a weak anterior keel runs along its basal portion. The cross section of the horn core is elliptical at the base, with its maximum transverse diameter located posteriorly (TD × 100/APD at the base = 75.3–76.4, n = 3; Table 3). The angle between the greatest anteroposterior diameter of the horn core base and the sagittal plane ranges from 42 to 50°. Towards the tip, the cross section of the horn core remains elliptical, but becomes symmetrical (TD × 100/APD at 7 cm above the base = 72.9–78.3, n = 3; Table 3). The upper premolars are moderately long compared to the molars, with a premolar/molar ratio 69.7% (n = 1) (Fig. 6C, SOM 3: Table 1). P2 and P3 have a strong anterolabial cone and bear a fossa unequally divided by a central fold (Fig. 6B). The anterior style of P3 and P4 is also strong, but the anterolabial cone of P4 is weak. The upper molars have a strongly developed paracone, parastyle and mesostyle, a weak metacone, and no basal pillars (entostyle) (Fig. 6B). A metaconule fold is present on M2. The lower premolar row is moderately short compared to the molars, with a premolar/molar ratio of 58.3–62.4%, n = 2 Table 3. Horn core measurements (in mm) at the base and at 7 cm above the base of Sporadotragus parvidens from Şerefköy-2 (Turkey). APD, anteroposterior diameter; TD, transverse diameter. Specimen MYŞE PV-2502 MYŞE PV-1573 MYŞE PV-1300 TDbase 35.5 34.6 34.5 APDbase 44.3 46.0 45.1 TD7 28.1 27.8 29.0 APD7 35.9 35.6 39.8 KOSTOPOULOS AND KARAKÜTÜK—LATE MIOCENE BOVIDS FROM TURKEY 5 cm A B 55 conid is well developed (Fig. 6C). Both the metastylid and especially the entostylid of the lower molars are well marked during early wear. There is no anterior cingulid. A low ectostylid is present on m1 (Fig. 6D). Remarks.—Both the size and the overall horn core and dental morphology of the material from Şerefköy-2 match those of Sporadotragus, the taxonomic status of which was recently revised by Geraads et al. (2006) and Kostopoulos (2009a). The specimens from Şerefköy-2 have larger horn cores than Sporadotragus vasili Geraads, Spassov, and Kovachev, 2006 from the SW Bulgarian localities of Kalimantsi and Strumyani (Geraads et al. 2006, 2011; Fig. 7), which furthermore differs from the present material in having a fused interfrontal suture, as well as a less curved horn core with a flat medial and a wide anterior surface, strong longitudinal grooves, and an anteromedial keel (Geraads et al. 2006). By contrast, the specimens from Şerefköy-2 have several features in common with Sporadotragus parvidens, mainly known from Pikermi, Samos (Greece) and Kemiklitepe-D (Turkey) (Bouvrain 1994; Kostopoulos 2009a). These include a small supraorbital foramen not located inside a pit, a complex and slightly raised interfrontal suture, and a Y-shaped frontoparietal suture. In addition, both have a horn core that is long, sub-cylindrical and gently (but markedly) curved posteriorly, bears irregular grooves, and shows no evidence of torsion or keels. The flexion of the frontals along the sagittal plane appears slightly stronger in the material from Şerefköy-2 (105–115°) than in Sp. parvidens from Samos (100–105°); however, this, as well as other minor differences (such as somewhat longer 51 C 47 5 cm D APDbase (B–D) 43 39 Fig. 6. The bovid artiodactyl Sporadotragus parvidens (Gaudry, 1861) from Şerefköy-2 (Turkey), Late Turolian (Late Miocene). A. Frontlet (MYŞE PV2502) in anterior view. B. Right P2–M3 (MYŞE PV-1522) in occlusal view. C. Right mandibular ramus with p3–m3 (MYŞE PV-1311) in lingual view. D. Left mandibular ramus with p2–m3 (MYŞE PV-1406) in labial view. (Fig. 6C, D, SOM 3: Table 2). The p3 and p4 have a strong anterior stylid and a barely developed anterior conid (Fig. 6C). The mesolingual conid is simple and elongated on p3, but rounded towards the base and slightly curved anteriorly on p4. On both p3 and p4, the posterolingual conid fuses with the posterior stylid during early wear and the posterolabial 35 30 35 TDbase Sporadotragus parvidens Sporadotragus vasili Strumyani Şerefköy-2 Samos Kalimantsi Pikermi 25 40 Sporadotragus sp. Kemiklitepe-D Fig. 7. Scatter plot comparing the basal proportions of the horn cores of Sporadotragus parvidens from Şerefköy-2 (Turkey), Samos and Pikermi (Late Miocene, Greece) with Sporadotragus vasili from Kalimantsi and Strumyani (Late Miocene, Bulgaria), and Sporadotragus from Kemiklitepe D (Late Miocene, Turkey). APDbase, anteroposterior diameter at the base; TDbase, transverse diameter at the base. 56 ACTA PALAEONTOLOGICA POLONICA 60 (1), 2015 horn cores in some specimens from Samos or the degree of p4 molarisation) may well reflect intraspecific variation. Stratigraphic and geographic range.—Upper Miocene; Aegean region. Genus Skoufotragus Kostopoulos, 2009b Type species: Pachytragus schlosseri (Andree, 1926), Samos, Greece; Late Miocene. Skoufotragus cf. Sk. schlosseri (Andree, 1926) Figs. 8–10. Material.—MYŞE PV-547, partial cranium with horn cores; MYŞE PV-1571, 1576, female frontlet; MYŞE PV-1570, left female horn core; MYŞE PV-1575, right horn core; MYŞE PV-1579, partial left female horn core; MYŞE PV-2606, partial female horn core; MYŞE PV-1309, palate; MYŞE PV1512, PV-1516, PV-2570, right upper tooth row with P2–M3; MYŞE PV-1513, PV-1514, PV-1515, right upper tooth row with P3–M3; MYŞE PV-1622, right upper tooth row with M1–M3; MYŞE PV-1491, right upper tooth row with M2– M3; MYŞE PV-1523, right M3; MYŞE PV-1520, PV-1521, left upper tooth row with M2–M3; MYŞE PV-1519, left upper tooth row with P2–M2; MYŞE PV-1525, left upper tooth row with P4–M1; MYŞE PV-2571, left upper tooth row with P3–M3; MYŞE PV-1410, left upper tooth row with M1–M3; MYŞE PV-1517, left M2–M3; MYŞE PV-1312, left upper tooth row with P2–P4; MYŞE PV-1315, left M3; MYŞE PV1434, PV-1436, PV-1313, M1 or M2; MYŞE PV-1532, left P2; MYŞE PV-1510, PV-1542, PV-2560, right mandibular body with p2–m3; MYŞE PV-1541, right mandibular body with p3–m3; MYŞE PV-2551, right mandibular body with p4–m2; MYŞE PV-2568, right mandibular body with m1–m3; MYŞE PV-1546, left mandibular body with m2–m3; MYŞE PV-1543, PV-1544, PV-2554, PV-2566, left mandibular ramus with p2–m3; MYŞE PV-1540, left mandibular ramus with p4–m3; MYŞE PV-2001, left mandibular ramus with p3–m1; MYŞE PV-1156, left mandibular ramus with m2–m3. All from Şerefköy-2, Turkey, Late Turolian (Late Miocene). Description.—This is by far the most abundant bovid found at Şerefköy-2 and represented by at least 10 individuals. The opisthocranium is high, narrow, dolichocephalic (sensu Bosscha-Erdbrink 1978) and has a straight dorsal profile in lateral view (Fig. 8C, Table 4). The temporal lines are moderately developed and run parallel to each other posteriorly. In posterior view, the occiput is triangular and bears a sharp occipital crest (Fig. 8B, C) that ends dorsally in a strong protuberance surrounded by deep scars. The nuchal crest is well developed. The mastoid faces posterolaterally and the paroccipital process is large and flattened. In lateral view, the occipital condyles project posteroventrally, thus forming a very acute angle with the occipital level. The basioccipital is long, relatively narrow and bears a shallow, narrow longitudinal groove (Fig. 8D). The sharp and prominent (crestlike) posterior tuberosities of the basioccipital are oriented perpendicular to the sagittal plane, whereas the weak anterior tuberosities are oriented anteroposteriorly. The oval foramen faces laterally. The preserved outline of the auditory bulla indicates that it was large and bulbous. The frontal contains large sinuses, one of which occupies the pedicle and even reaches the base of the horn core. There is no postcornual fossa. The horn core, inserted above the orbit, is sabre-like without keels or torsion (Figs. 8A, C, 9A). It is moderately long, moderately curved posteriorly in lateral view and strongly compressed mediolaterally along its entire length (TD × 100/APD at the base: 40–57, n = 5; Table 4). In cross section, the horn core forms an elongated ellipse, which becomes narrower towards the tip. A deep furrow occasionally runs along the upper half of the posterior surface, and, in combination with the strong mediolateral compression, may give the impression of a distal posterior keel. Two frontlets and three horn core specimens (MYŞE PV1570, PV-1571, PV-1576, PV-1579, and PV-2606; Fig. 9) likely represent female individuals. The supraorbital foramen is large and round, and placed far anterior to the pedicle. The interfrontal suture is open and simple in outline. The horn core is thin and long (~140 mm) and inserted above the back of the orbit. It is far removed from its counterpart, weakly curved posteriorly and barely twisted homonymously (Fig. 9B). In cross section, the horn core is elliptical at the base but becomes more compressed mediolaterally towards the tip (Table 4). In anterior view, the divergence of the horn cores is weak up to their mid-height, but stronger above. The premolars are moderately long compared to the molars, with upper and lower premolar/molar ratios of 59.6– 65.6% (n = 5) and 60.3–66.6% (n = 4), respectively (Fig. 10, Table 4. Cranial and horn core measurements (in mm) of Skoufotragus cf. Sk. schlosseri from Şerefköy-2 (Turkey). APD, anteroposterior diameter at the base and at 10 cm above the base; Lfpo, sagittal length from the frontoparietal suture to the occipital protuberance; TD, transverse diameter at the base and at 10 cm above the base; Watbc, width of the skull at the anterior tuberosities of the basioccipital; Wbrc, maximum width of the braincase; Wptbc, width of the skull at the posterior tuberosities of the basioccipital. sin, left; dex, right. Specimen MYŞE PV-547 dex MYŞE PV-547 sin MYŞE PV-1571 dex MYŞE PV-1575 dex MYŞE PV-1570 sin MYŞE PV-1576 dex MYŞE PV-1576 sin TDbase 29.29 32.2 34 33.36 25.92 22.42 23.16 APDbase 58.84 55.52 59.5 61.38 64.9 26.64 26.77 TD10 20.29 22.6 22.56 22.58 22.65 11.18 9.77 APD10 37.22 38.3 43.36 39.84 45.54 14.69 13.95 Lfpo 65.13 Wbrc 61.69 Wptbc 35.85 Watbc 19.94 KOSTOPOULOS AND KARAKÜTÜK—LATE MIOCENE BOVIDS FROM TURKEY A 57 C B 5 cm 5 cm (A, C) D 5 cm Fig. 8. The bovid artiodactyl Skoufotragus cf. Sk. schlosseri (Andree, 1926) from Şerefköy-2 (Turkey), Late Turolian (Late Miocene). Partial skull (MYŞE PV-547) in anterior (A), posterior (B), lateral (C), and ventral (D) views. SOM 3: Tables 1, 2). The upper molars have strong styles, bear a fossetta (central islet) and lack entostyles (Fig. 10A, B, D). P2 is bilobed lingually, whereas P3 has a trapezoidal occlusal outline. Both have a strong anterolabial cone (Fig. 10A). The protocone of M1 protrudes lingually. The metastyle of M3 is strong and, in one specimen (MYŞE PV-1622), flares distally (Fig. 10D). The mandibular body is shallow and long (Fig. 10C). On the labial face, a second mental fora- 58 ACTA PALAEONTOLOGICA POLONICA 60 (1), 2015 A B1 B2 5 cm 5 cm Fig. 9. The bovid artiodactyl Skoufotragus cf. Sk. schlosseri (Andree, 1926) from Şerefköy-2 (Turkey), Late Turolian (Late Miocene). A. Frontlet (MYŞE PV-1571) in anterior view. B. Female frontlet (MYŞE PV-1576) in anterior (B1) and lateral (B2) views. A B 5 cm (A–D) C D E 5 cm (E, F) F Fig. 10. The bovid artiodactyl Skoufotragus cf. Sk. schlosseri (Andree, 1926) from Şerefköy-2 (Turkey), Late Turolian (Late Miocene). A. Palate (MYŞE PV-1309) in occlusal view. B. Left upper tooth row P2–M3 (MYŞE PV-1312) in buccal view. C. Right upper tooth row M1–M3 (MYŞE PV-1622) in occlusal view. D. Right mandibular ramus with p2–m3 (MYŞE PV-1510) in labial view. E. Left mandibular ramus with p2–m3 (MYŞE PV-1544) in occlusal view. F. Right mandibular ramus with p2–m3 (MYŞE PV-1510) in occlusal view. KOSTOPOULOS AND KARAKÜTÜK—LATE MIOCENE BOVIDS FROM TURKEY Remarks.—Protoryxoid bovids appear in the Eastern Mediterranean area as early as the Late Astaracian (e.g., Gentry 2000). Although they remained rare during the Vallesian, they strongly radiated and dispersed during the Turolian, especially in Anatolia and adjacent territories. They include small to medium-sized antelopes of caprine/hippotragine cranial appearance, but their taxonomy and evolutionary relationships remain debated. Kostopoulos (2009a) partly revised the Turolian protoryxoids from SE Europe assigned to the so-called “Protoryx/Pachytragus complex” and recognised two distinct genera: Protoryx Major, 1891 and Skoufotragus Kostopoulos, 2009 (= partim Pachytragus Schlosser, 1904). The Şerefköy-2 specimens resemble Skoufotragus in their dental morphology and in having (i) a narrow, long braincase with a straight dorsal profile and parallel sides, (ii) a triangular occiput and (iii) a sabre-like, mediolaterally compressed and uprightly inserted horn core (Kostopoulos 2009a: 364). Skoufotragus is known from the Turkish Late Miocene assemblages of Kinik (Protoryx sp. of Köhler 1987), Kemiklitepe-A (Protoryx laticeps of Bouvrain 1994), and Akkaşdağı (Pachytragus crassicornis of Kostopoulos 75 70 APD base men appears below p3 (Fig. 10C). The p2 is simple without an anterior conid, but with a strong mesolingual conid and anterior stylid (Fig. 10E, F). The p3 has a well-developed anterior conid and stylid, which become fused together with wear. The mesolingual conid of p3 is oriented parallel to the posterolingual conid, which in turn fuses with the weaker and posterolingually directed posterior stylid during early wear (Fig. 10E, F). The p4 resembles p3, but has an anteroposteriorly developed mesolingual conid (Fig. 10E). The lower molars have a strong meta- and mesostylids, but both elements disappear with wear. There is no anterior cingulid. A basal pillar appears on 5 out of 13 m1s, and 3 out of 13 m2s. The third lobe of m3 is labially displaced and has an elongated, semicircular occlusal outline; it bears a strong posterolingual stylid on the upper half of the crown. 59 65 60 55 50 45 25 30 35 40 45 50 TD base Skoufotragus laticeps Maragheh Samos Skoufotragus zemalisorum Samos Skoufotragus sp. Kemiklitepe-A Kinik Skoufotragus schlosseri Akkasdagi Samos Şerefköy-2 Protoryx carolinae Pikermi Fig. 11. Scatter plot comparing the basal horn core proportions of Skoufotragus from several sites and Protoryx from Pikermi (Late Miocene, Greece). APDbase, anteroposterior diameter at the base; TDbase, trans- verse diameter at the base. 2005), as well as from Samos, Greece and Maragheh, Iran (Kostopoulos 2009a, Kostopoulos and Bernor 2011). The Şerefköy-2 specimens differ from Skoufotragus laticeps from Samos, Kemiklitepe-A and Maragheh in having a narrower braincase, as well as a shorter dorsal parietal sector, less developed anterior tuberosities of the basioccipital, shorter, more slender, and more mediolaterally compressed horn cores, and, on average, smaller tooth rows (but with a similar premolar/molar ratio) (Figs. 11, 12). They are more Component 2 (24%) 6 3 -20 -16 -12 -8 4 -4 8 12 -3 -6 Fig. 12. Principal component analysis of Skoufotragus from Samos (Late Miocene, Greece), Akkaşdağı and Şerefköy-2 (both Late Miocene, Turkey), based on four cranial and horn core variables (braincase width, dorsal length of parietal, transverse diameter of the horn cores at the base, and anteroposterior diameter of the horn cores at the base) (modified from Kostopoulos 2009a). -9 -12 -15 Component 1 (62%) Skoufotragus laticeps Skoufotragus schlosseri Skoufotragus zemalisorum Akkasdagi Serefkoy 16 60 similar to Sk. zemalisorum from Samos in terms of their cranial proportions, but differ in having a narrower braincase (61.7 mm vs. 70–78 mm; Kostopoulos 2009a) and a more anteroposteriorly expanded horn core (Figs. 11, 12). Except for a somewhat longer lower premolar row (relative to the molars), the Şerefköy-2 specimens closely resemble Sk. schlosseri from Samos and Akkaşdağı in their size and morphology (Figs. 11, 12), although the material from Samos (Q5) is characterised by slightly longer and more divergent horn cores bearing an anterior keel (the latter seems to be more common in short-horned individuals; cf. Kostopoulos 2005: 777). Female individuals of Skoufotragus are rare, but Gentry (1971: 252, pl. 3: 3) interpreted AMNH 20687 as a female individual of Sk. laticeps. The frontlet MYŞE PV-1576 from Şerefköy-2 is very similar to this specimen, suggesting that females of Sk. schlosseri were likely horned. Genus Urmiatherium Rodler, 1889 Type species: Urmiatherium polaki Rodler, 1889, Maragheh, Iran; Late Miocene. Urmiatherium rugosifrons (Sickenberg, 1932) Fig. 13. Material.—MYŞE PV-1182, partial opisthocranium; MYŞE PV-2503, frontlet; MYŞE PV-2504, frontlet of young individual; MYŞE PV-2599, axis+3rd cervical vertebra; MYŞE PV-2506, axis; MYŞE PV-2505, 3rd cervical vertebra (possibly representing the same individual as MYŞE PV-2505). All from Şerefköy-2, Turkey, Late Turolian (Late Miocene). Description.—This bizarre species is represented by at least two individuals and can unambiguously be identified based on (i) its small size (Table 5); (ii) strong cranio-facial flexion; (iii) strongly elevated, thick and pneumatised frontals; (iv) an extremely shortened opisthocranium; (v) a reduced parietal forming an obtuse angle with the occipital plane; (vi) an enlarged and thickened occipital with large occipital condyles and a dorsally-facing occiput; (vii) a thickened basioccipital with strong and completely fused posterior tuberosities, thus forming an additional, oval facet for the atlas (Fig. 13C); and (viii) thick, short, and moderately homonymously twisted and grooved horn cores, originating very close from each other and—in juveniles—bearing wide and well-defined lateral depressions (Fig. 13A, B). Remarks.—Urmiatherium rugosifrons is the only representative of its genus from Asia Minor. Besides Şerefköy-2, the species is only known from the neighbouring site of Salihpaşalar (Kaya et al. 2012) and the Turolian of Samos (e.g., Solounias 1981; Kostopoulos 2009a, 2014; Jafarzadeh et al. 2011; Table 5). Other possible occurrences of the genus at Garkın, Kınık (Afyon) and Kayadibi (Konya) in mid-western Anatolia (Alan 1997; Saraç 2003) still need to be confirmed. Stratigraphic and geographic range.—Upper Miocene; Asia Minor. ACTA PALAEONTOLOGICA POLONICA 60 (1), 2015 Genus Sinotragus Bohin, 1935 Type species: Sinotragus wimani Bohlin, 1935, Locality 30, Shanxi Province, North China. cf. Sinotragus sp. Fig. 14. Material.—MYŞE PV-1409, partial left upper tooth row preserving only P2–P3; MYŞE PV-2553, left mandibular body with p2–m3; MYŞE PV-2567, right mandibular body with p2–m3. All from Şerefköy-2, Turkey, Late Turolian (Late Miocene). Description.—The only maxillary fragment preserves P2 and P3 (Fig. 14C, SOM 3: Table 1). P2 is weekly bilobed lingually and has a trapezoidal occlusal outline, with a weak anterior style, strong anterolabial cone, posterolabially protruding posterior style, and distolingually broadened posterolingual crista. P3 is subrectangular in occlusal view and bears strong labial styles and ribs, as well as a weak lingual groove dividing the distolingually expanded lingual cone (Fig. 14C). A weak notch on the distal wall of the tooth emphasises the posterior style. Traces of enamel on the distal occlusal surface suggest that the tooth originally had an additional distal fossa or a strong central fold. The lower premolar row is moderately short compared to the molars, with premolar/molar ratios of 56.7% and 57.9% (n = 2) (Fig. 14A, B, SOM 3: Table 2). The lower premolars are narrow and short. The p2 lacks an anterior conid, but bears an anteriorly located mesolingual conid and a week anterior stylid. The mesolingual conid of p3 is elongated and directed posteriorly. The posterolingual conid of p3 is strong. The anterior conid and stylid are equally developed and fuse together during advanced stages of wear. The anterior valley is open. Unlike p3, p4 is distinctly molarised (Fig. 14A, B). On the lingual wall, the anterior valley closes quickly during wear and the mesolingual conid fuses with the posterior conid and stylid. Labially, there is a well-developed posterolabial conid. The lower molars bear a strong ectostylid and have a gently undulating lingual wall covered by cement. The Table 5. Cranial and horn core measurements (in mm) of Urmiatherium rugosifrons from Şerefköy-2 (Turkey) compared to Urmiatherium rugosifrons from Samos (Greece) and Urmiatherium polaki from Maragheh (Iran). Data are from Kostopoulos (2009a) and Kostopoulos and Bernor (2011). APDbase, anteroposterior diameter at the base; TDbase, transverse diameter at the base; Wbi-co, bicondylar width of the skull; Wbrc, maximum width of the braincase; Wptbc, width of the skull at the posterior tuberosities of the basioccipital. Specimen Wbrc MYŞE PV-1182 Wbi-co 57.6 Wptbc TDbase APDbase 28.1 MYŞE PV-2504 72.6 35.2 48.9 MYŞE PV-2503 92.7 ~46.8 ~70.8 42 78–130 Urmiatherium rugosifrons, Samos Urmiatherium polaki, Maragheh 95–96 50–62 24–30 98 77–83 39 KOSTOPOULOS AND KARAKÜTÜK—LATE MIOCENE BOVIDS FROM TURKEY A 61 B 5 cm C D Fig. 13. The bovid artiodactyl Urmiatherium rugosifrons (Sickenberg, 1932) from Şerefköy-2 (Turkey), Late Turolian (Late Miocene). A. Juvenile frontlet (MYŞE PV-2504) in anterior view. B. Frontlet (MYŞE PV-2503) in anterior view. C. Opisthocranium (MYŞE PV-1182) in ventral view. D. Axis + 3rd cervical vertebra (MYŞE PV-2599) in dorsal view. third lobe of m3 is relatively high and large, oval in occlusal view, and bears a convex entoconulid. Remarks.—The finely rippled enamel, advanced p4 morphology, reduced premolars, build-up of cement on the molars, large third lobe of the m3, strong basal pillars, and weak development of the lingual stylids and ribs exclude this taxon from the Late Miocene Antilopini sensu stricto. Together with the inferred additional posterior fossa on P3, the distolingual widening of the lingual cone and the distolabial development of the posterior style on P2 and P3 are reminiscent of the considerably larger Urmiatherium intermedium Bohlin, 1935, U. polaki Rodler, 1889, and Plesiaddax depereti Bohlin, 1935. However, the Şerefköy-2 taxon differs from the large-sized Late Miocene “ovibovines”, but resembles the smaller U. rugosifrons from Samos (Kostopoulos 2009a: 371), in the development of the third lobe on m3, as well as the weak buccolingual compression of the lower molars and the degree of fusion of their lobes. U. rugosifrons and the Şerefköy-2 species furthermore share a comparably reduced premolar row. However, the former species is still about 30–35% larger and characterised by less derived lower premolars. Although the available data are not sufficient for definite conclusions, it is worth mentioning that a right upper tooth row from Muğla (MNHN TRQ 974) resembles the material from Şerefköy-2 in its size and premolar morphology. The molars of TQR 974 considerably differ from those of Urmiatherium and Plesiaddax, but resemble those of Sinotragus wimani Bohlin, 1935 from China (Bohlin 1935: pl. 16: 8, 9). Unfortunately, the lower dentition of Si. wimani is unknown, preventing a direct comparison with Şerefköy-2. TRQ 974 is accompanied by a frontlet from the same region (MNHN TRQ 973), which, based on its morphology and size, likely represents a female Si. occidentalis Geraads, Güleç, and Kaya, 2002. This species is well-documented in Muğla by 62 ACTA PALAEONTOLOGICA POLONICA 60 (1), 2015 Karadenizli et al. 2005) and characterises the Final Bovid Assemblage of Samos (~7.0–6.8 Ma; Kostopoulos 2009a; Koufos et al. 2009b). The co-occurrence of U. rugosifrons and Skoufotragus cf. Sk. schlosseri at Şerefköy-2 thus places the latter close to the Middle–Late Turolian boundary (~6.8 Ma; e.g., Steininger 1999; Agusti et al. 2001). A1 A2 Palaeobiogeographical and palaeoenvironmental analysis 5 cm B C Fig. 14. The bovid artiodactyl cf. Sinotragus from Şerefköy-2 (Turkey), Late Turolian (Late Miocene). A. Left mandibular ramus (MYŞE PV2553) in labial (A1) and lingual (A2) views. B. Right mandibular body (MYŞE PV-2567) in occlusal view. C. Left upper P2–P3 (MYŞE PV-1409) in occlusal view. cranial, but not dental, elements (Geraads et al. 2002). Given the particular dental characters of TRQ 974, the similar size of TQR 974 and TRQ 973, and their common geographic provenance, we speculate that both specimens belong to Si. occidentalis. In addition, we tentatively refer the Şerefköy-2 specimens to Sinotragus based on their similar dental morphology. Biochronology of Şerefköy-2 Kaya et al. (2012) suggested a Middle Turolian age for the Şerefköy-2 assemblage and stressed its structural similarity to the mammalian palaeocommunity from Samos. The results of the present study help to refine this biochronological assessment. Among the bovid recovered from Şerefköy-2, Gazella capricornis and Palaeoryx pallasi are the most widespread, ranging from the Balkans to Iran and the northern coast of the Black Sea. Their co-existence at Şerefköy-2 is strongly indicative of a Middle–Late Turolian age for this faunal assemblage, as is the presence of Sporadotragus parvidens. The latter is so far only known from Samos and Pikermi (Greece), although the genus occurs from the southern Balkans all the way to Afghanistan and possibly China and Mongolia (e.g., Geraads et al. 2006). Urmiatherium rugosifrons certainly occurs at Mytilinii-1A and most likely also at Q5 of Samos (Kostopoulos 2009a), indicating an age younger than 7.1 Ma (Koufos et al. 2009b). By contrast, Skoufotragus schlosseri is present at Akkaşdağı (<7.1 Ma; Protoryxoid bovids represent more than 60% (more than 80% together with gazelles) of the Şerefköy-2 bovid community in terms of both the minimum number of individuals (n = 27) and the number of identifiable specimens (n = 102). In sharp contrast, boselaphines and spiral-horned antelopes, both of which are common in contemporaneous faunas of the sub-Paratethyan province, are absent from Şerefköy-2. Even though taphonomic and sampling biases may have exaggerated these results, the absence of these taxa indicates that they were at least rare in the local mammal community. On the other hand, the presence of Urmiatherium rugosifrons, so far only known from neighbouring Samos, and of a western Asian representative of Sinotragus, known mainly from the surroundings of Şerefköy (i.e., the Muğla-Yatağan Basin; e.g., Geraads et al. 2002), increase the local character of the Şerefköy-2 mammal assemblage. In our genus-level correspondence analysis, the first three components (out of 15) account for 23.2%, 20.8%, and 14.1% of the inertia, respectively (Fig. 15A). Both in the first (axes 1 and 2; Fig. 15A) and the second (axes 1 and 3; not shown) plane, the continental Greek Turolian bovid associations are sharply distinguished in taxonomic composition and relative abundances from most Anatolian bovid assemblages (including Samos), whereas the Turkish assemblages of Mahmutgazi, Sivas and Akkaşdağı, and the Iranian fauna of Maragheh, occupy an intermediate position, rather close to the Greek bovid faunas (Fig. 15A). The primary polarizing taxa include Paraoioceros, Plesiaddax, Protragelaphus, Nisidorcas, Majoreas, and Urmiatherium. There is a clear, time-controlled taxonomic shift in the Greek bovid assemblages from Nikiti-2 (Early Turolian) to Dytiko (Late Turolian), with an increase of the magnitude of Miotragocerus, Gazella, and Protragelaphus against Nisidorcas, and Tragoportax (Fig. 15A). In the east, the situation seems much more complicated than in continental Greece. The Middle Turolian bovid assemblages of Asia Minor, Mytilinii-1A, B, C, Mytilinii-3, Kemiklitepe-A, B, Kinik, and Şerefköy-2 form a distinct cluster, characterised by Skoufotragus, Sporadotragus, Palaeoryx, Urmiatherium, and Paraoioceros. The older (Early Turolian) but geographically close assemblages of Kemiklitepe-D and Mahmutgazi are separated from this group mainly because of the high percentages of Criotherium, Majoreas, Oioceros, and Plesiaddax. The central-eastern Anatolian bovid faunas KOSTOPOULOS AND KARAKÜTÜK—LATE MIOCENE BOVIDS FROM TURKEY A 63 16 2.0 15 KTD 1.5 2.0 1.5 Cc Axis 2 1.0 1 NIK 0.5 0.0 14 PXM 2 RZO PER MAR 13 34 Sivas -0.5 5 Mahm PIK DYTI 6 7 Akkas 12 8 MTLA 9 Seref KTAB MYT 10 11 Kinik -1.0 -1.5 -2.0 -2.0 -1.5 -1.0 0.0 -0.5 0.5 RZO B 1.0 2.5 Axis 1 Cc NIK DYTI Akkas Sivas PXM MAR PER PIK Kinik MYT Mahm Seref KTD MTLABC KTAB Fig. 15. Correspondence analysis of Turolian (Late Miocene) bovid associations from the sub-Paratethyan zoogeographic province, based on relative taxonomic abundances at the genus level (A) and relative size abundances at each site (B). See text for explanation, SOM 2 for details of the methodology, and SOM 1: Table 1 for the dataset. Analysed variables: 1, Nisidorcas; 2, Tragoportax; 3, Prostrepsiceros; 4, Palaeoreas; 5, Protragelaphus; 6, Miotragocerus; 7, Gazella; 8, Urmiatherium; 9, Palaeoryx; 10, Paraoioceros; 11, Skoufotragus/Protoryx; 12, Sporadotragus; 13, Plesiaddax; 14, Oioceros; 15, Majoreas; 16, Criotherium. Squares represent Early Turolian assemblages; circles represent Middle–Late Turolian assemblages. In the spectra of relative size abundances (pie charts) grey represents small (<50 kg), white medium, and black large-sized bovids (>150 kg). Abbreviations: Akkas, Akkaşdağı; Cc, Çorak-yerler; DYTI, Dytiko-1, 2, 3; KTAB, Kemiklitepe A, B; KTD, Kemiklitepe D; Mahm, Mahmutgazi; MAR, Maragheh (mainly levels MMTT7-13); MTLABC, Mytilinii-1A, B, C; MYT, Mytilinii-3; NIK, Nikiti-2; PER, Perivolaki; PIK, Pikermi; PXM, Prochoma; RZO, Ravin de Zouaves-5; Seref, Şerefköy-2. exhibit a similar pattern, with the Early Turolian Çorak-yerler assemblage being separated from those of Sivas and Akkaşdağı (late Early and late Middle Turolian, respectively). The bovid assemblages of Çorak-yerler and Kemiklitepe-D are much more alike in their taxonomic structure than are the roughly contemporaneous faunas of continental Greece (Nikiti-2 and Ravin de Zouanes-5) (Fig. 15A). By contrast, Sivas and, to a lesser degree, Akkaşdağı, more closely re- semble assemblages from continental Greece to the west and Maragheh to the east than contemporary faunas from Asia Minor. In the correspondence analysis using broader taxonomic bins, the first two components (out of four) account for 51.8%, and 25.2% of the inertia, respectively (Fig. 16A). The results are similar to those of the genus-level analysis and show no major changes between the Early and mid-Late Tur- 64 ACTA PALAEONTOLOGICA POLONICA 60 (1), 2015 A B 2.4 RZO PER PIK KTD KTAB MYT DYTI OVIB 2 KTD Axis 2 1.6 MTLABC 1.2 0.8 Seref Sivas 0.4 MAR 0 -0.4 SPIRA NIK RZO Akkas Mahm Cc PER DYTI PXM BOSE PIK Sivas GAZE Seref MTLABCPROTO Akkas KTAB MYT Kinik -0.8 -0.9 -0.6 -0.3 0 0.3 Axis 1 0.6 0.9 1.2 1.5 1.8 Fig. 16. Correspondence analysis of Turolian (Late Miocene) bovid associations from the sub-Paratethyan zoogeographic province based on taxonomic bins (A) and diet spectra of several local bovid associations based on dental wear patterns (B). See text for explanation, SOM 2 for details of the methodology, and SOM 1: Table 2 for the dataset. Abbreviations: Akkas, Akkaşdağı; Cc, Çorak-yerler; DYTI, Dytiko-1, 2, 3; KTAB, Kemiklitepe A, B; KTD, Kemiklitepe D; Mahm, Mahmutgazi; MAR, Maragheh (mainly levels MMTT7-13); MTLABC, Mytilinii-1A, B, C; MYT, Mytilinii-3; NIK, Nikiti-2; PER, Perivolaki; PIK, Pikermi; PXM, Prochoma; RZO, Ravin de Zouaves-5; Seref, Şerefköy-2. Dashed-line represents a convex hull of Early Turolian bovid assemblages; BOSE, GAZE, PROTO, OVIB, SPIRA, represent analysed variables (SOM 1: Table 2). Pie charts represent the spectra of feeding preferences: black, browsers; grey, mixed feeders; white, grazers. olian assemblages from continental Greece, all of which are dominated by spiral-horned antelopes and boselaphines (Fig. 16A). The same taxa fare less well in contemporary Anatolia, where they and “ovibovines” are increasingly replaced by gazelles and protoryxoids. This trend is much sharper in the western part of the region; in central-eastern Anatolia, assemlages instead remain more similar to those of continental Greece and Iran (Fig. 16A). Although based on a much more limited dataset lacking Turolian faunas from Greece, Bibi, and Güleç (2008: fig. 11) found a similar taxonomic trend in the Anatolian bovid assemblages, including the separation of Çorak-yerler and Kemiklitepe-D from the remaining assemblages. However, they also identified a cluster comprising Kemiklitepe-A, B and Sivas, despite their marked taxonomic differences (Tragoportax constitutes 41.5% at Sivas but is absent from KTAB; “Pachytragus/Protoryx” constitutes 3.8% at Sivas vs. 39.6% at KTAB; Bibi and Güleç 2008), size distribution and diet spectra (see below). This grouping thus seems rather improbable and probably stems from the inclusion of the Greek Vallesian assemblage of Nikiti-1. This may have resulted in the compression of the remaining faunas within multispace, as in de Bonis et al. (1994: fig. 4). Faunal body size distributions generally support the results of the CFA (Fig. 15B). Thus, the Turolian faunas of continental Greece contain very few large taxa (<4%) and, with the exception of the mostly small (<50 kg) species constituting the earliest Turolian assemblage of Nikiti-2, show a balance between small and medium-sized forms (Fig. 15B). A similar pattern occurs in central-eastern Anatolia (Fig. 15B), although the dataset is much more restricted here. By contrast, large and small-sized bovids are balanced and predominate at Kemiklitepe-D, whereas from Mahmutgazi to Şerefköy-2 large taxa decline and medium-sized bovids increase considerably, with percentages reaching 80% in some cases (Fig. 15B). The diet spectra of the assemblages compared here do not follow the regional pattern revealed by the CFA and size distribution analyses. Instead, they point to a general rise of mixed feeders at the expense of grazers in younger associations, as shown by comparisons of, for instance, KTD with KTAB and Şerefköy-2 in western Anatolia, Sivas with Akkaşdağı in eastern Anatolia, and Ravin de Zouaves-5 with Perivolaki, Pikermi, and Dytiko in Greece (Fig. 16B). In addition, several roughly contemporaneous faunas show similar diet spectra despite marked differences in their geographical location, taxonomic composition and size distribution (e.g., Nikiti-2 vs. Çorak-yerler, Dytiko vs. Akkaşdağı, and Pikermi vs. Kemiklitepe-A, B and Mytilinii-1A, B, C; Figs 15A, B, 16B). KOSTOPOULOS AND KARAKÜTÜK—LATE MIOCENE BOVIDS FROM TURKEY 65 Discussion and conclusions Acknowledgements As in the study by Bibi and Güleç (2008: 516), there is no straightforward correlation between the component axes arising from the taxonomy-based CFA and environmental trends, at least at a generalised Greco-Anatolian level. Instead, the age and geographical location of the sites, two parameters not directly involved in the analysis, seem to control much of the distribution of the data. Thus, in line with preliminary suggestions by Kostopoulos and Bernor (2011), our analysis revealed a “Greek” cluster and a “SW Anatolian” cluster, whereas the eastern Anatolian Turolian bovid assemblages possibly form a third group more closely related to Maragheh in Iran. Like in continental Greece, the general increase of mixed feeders relative to grazers throughout the Turolian in eastern Anatolia is not accompanied by a major reorganization of the structure of the bovid community, with the exception of an increase in the number of protoryxoids. Instead, size ratios generally remain stable, with small taxa always contributing more than 50% (based on the number of identifiable specimens). During approximately the same time period, small taxa slightly decrease in number (by about 10%) in SW Anatolia, whereas medium-sized taxa considerably increase (by about 30%). Protoryxoids at least quadruple at the expense of boselaphines and “ovibovines”, while the relative abundances of gazelles and spiral-horned antelopes remain stable at a moderate and low level, respectively. These results generally agree with ungulate regional endemicity along the Greco-Iranian longitudinal axis, as suggested by Costeur (2009: fig. 4), and are fully compatible with the conclusions of de Bonis et al. (1994). The partial isolation of SW Anatolia from continental Greece was thoroughly discussed by Kostopoulos (2009b, and literature therein) and likely resulted from both the mid-Tortonian/ Maeotian transgression into the Aegean domain and the latest Miocene invasion of the Paratethyan Sea. By contrast, the partial faunal isolation of SW Anatolia from adjacent regions to the east may be related to the broad Mio-Pliocene Central Anatolian lake system stretching between the Pontides in the north and the Taurides-Isparta Angle barrier in the south (e.g., Görür et al. 1995; Veen et al. 2009; Alçiçek 2010; Mehmet Cihat Alçiçek personal communication 2012). Our study demonstrates that similar ecological signals need not necessarily imply identical size distributions or taxonomic content, even for roughly contemporaneous, neighbouring faunas. This implies that the faunal composition of herbivores at various points of a supposedly homogeneous biogeographic province may depend more on historical (i.e., phylogeographic) relationships and limitations than the type of vegetation cover. Assuming that the diet spectrum of a local herbivore assemblage directly reflects vegetational and/ or climatic conditions, it therefore seems that local bovid communities adjust to the available habitat, rather than (or prior to) being reorganised by migratory movements and replacements. We thank Tanju Kaya and Serdar Mayda (both Ege University, Izmir, Turkey), Robert Scott (Rutgers University, New Brunswick, New Jersey, USA), and Gildas Merceron (Poitiers University, France) for their help and support in the field and lab, and Mehmet Cihat Alçiçek (Pamukkale University, Denizli, Turkey) for helpful discussions on SW Anatolia geodynamics. Thanks are also due to Kaye Reed (Arizona State University, Tempe, USA), Denis Geraads (CNRS MNHN, Paris, France) and Felix G. Marx (University of Otago, New Zealand) for their fruitful comments and suggestions, as well as to Katerina Vasileiadou (Natural History Museum of the Petrified Forest of Lesvos, Greece) for linguistic improvements. 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Kostopoulos and Seval Karakütük Late Miocene bovids from Şerefköy-2, SW Turkey, and their position within the sub-Paratethyan biogeographic province Published in Acta Palaeontologica Polonica 2015 60 (X): xxx-xxx. http://dx.doi.org/10.4202/app.2012.0019 SOM 1 Table 1. Relative abundances of bovid genera from Greek, Turkish, and Iranian Turolian assemblages. Table 2. Proportions of size, taxonomic, and feeding categories of bovid assemblages from Greek, Turkish and Iranian Turolian assemblages. SOM 2 Correspondence analysis, size and diet spectra. References SOM 3 Table 1. Upper tooth measurements (in mm) of Gazella cf. capricornis, Palaeoryx pallasi, Sporadotragus parvidens, Skoufotragus cf. Sk. schlosseri, and cf. Sinotragus from Şerefköy-2 (Turkey). Table 2. Lower tooth measurements (in mm) of Gazella cf. capricornis, Palaeoryx pallasi, Sporadotragus parvidens, Skoufotragus cf. Sk. schlosseri, and cf. Sinotragus from Şerefköy-2 (Turkey). 3.00 70.83 39.6 6.81 68.2 1.16 4.11 41.72 43.14 22.17 7.50 12.50 4.55 2.62 5.96 17.65 1.25 4.17 12.50 2.09 3.49 1.58 8.61 6.86 3.30 2.02 2.50 1.00 42.86 41.51 33.75 57.53 4.00 23.00 50.00 28.57 4.00 16.98 16.25 15.07 1.25 1.37 5.00 1.25 15.07 4.17 8.33 1.05 6.82 10.17 3.16 6.62 35.08 18.90 8.86 0.66 3.30 34.41 2.36 2.02 4.11 1.99 5.88 4.65 32.91 21.47 2.27 3.20 21.84 15.12 0.95 21.47 7.85 0.95 1.74 2.85 23.58 9.72 21.86 Gazella Protragelaphus 66.67 10.66 3.00 Paraoioceros 7.58 2.54 Majoreas Prostrepsiceros Oioceros Plesiaddax 22.84 14.29 3.77 3.75 Criotherium Urmiatherium 19.70 54.82 Nisidorcas 3.00 Tragoportax Miotragocerus Palaeoryx 1.52 Palaeoreas NIK RZO Cc KTD Sivas Mahm PXM MYT KTAB PER Kinik PIK MAR7-13 MTLABC Seref Akkas DYTI Sporadotragus Skoufotragus/ Protoryx SOM 1. Table 1. Relative abundances of bovid genera from Greek, Turkish, and Iranian Turolian assemblages. 4.55 9.14 10.00 14.29 30.19 27.50 10.96 8.33 39.58 3.66 18.18 38.95 10.44 34.44 23.53 45.28 19.43 SOM 1. Table 2. Proportions of size, taxonomic, and feeding categories of bovid assemblages from Greek, Turkish, and Iranian Turolian assemblages. NIK RZO Cc KTD Sivas Mahm PXM MYT KTAB PER Kinik PIK MAR MTLABC Seref Akkas DYTI PROTO 1,5 0 3 14,3 3,8 12,5 0 87,5 53 8,9 72,7 7,3 5,7 56,3 67,6 25,5 2 BOSE 19,7 54,8 2 0 41,5 36,2 57,5 0 0 36,1 6,8 29,1 14,2 7,3 2,9 5,7 36,5 OVIB 0 0 5 43 0 16,3 0 0 0 0 0 0 4,1 2 5,9 0 0 SPIRA 74,2 36 80 28,6 17 7,5 31,5 4,2 8,3 51,3 2,3 24,7 59,5 0 0 23,6 42,1 GAZE 4,6 9,1 10 14,3 37,7 27,5 11 8,3 39,6 3,7 18,2 39 16,5 34,5 23,5 45,3 19,4 Small 78,8 45,2 86 42,8 54,7 33,8 42,5 12,5 39,6 47,2 20,4 61,9 73,1 34,4 23,5 68,9 39,7 Medium 19,7 54,8 9 14,3 45,3 48,7 57,5 83,3 47,9 50,8 79,6 34,6 19 57 69,6 27,8 58,3 Large 1,5 0 5 42,9 0 17,5 0 4,2 12,5 2 3,5 7,9 8,6 6,9 3,3 2 Br Mix Gr 35,6 ? 42,9 17 0 ? 0 11,3 64,4 ? 57,1 71,7 87,5 20,8 30,6 4,2 39,6 45,3 8,3 39,6 24,1 21,8 76,5 1,7 30,5 24,5 6,6 6 48,3 43,1 50 47,8 21,2 32,4 43,4 46,2 NISP 66 197 100 56 53 80 73 24 48 191 44 344 316 151 102 212 247 Abbreviations.—Greek faunas: NIK, Nikiti-2; RZO, Ravin de Zouaves-5; PXM, Prochoma; MYT, Mytilinii-3; PER, Perivolaki; PIK, Pikermi; MTLABC, Mytilinii-1A,B,C; DYTI, Dytiko-1,2,3. Turkish faunas: Cc, Çorak-yerler; KTD, Kemiklitepe D; Sivas, Sivas; Mahm, Mahmutgazi; KTAB, Kemiklitepe A, B; Kinik, Kinik; Seref, Şerefköy-2; Akkas, Akkaşdağı. Iranian faunas: MAR, Maragheh (mainly levels MMTT7-13). NISP, number of identified specimens. Size categories: small, <50 kg; medium, 50–150 kg; large, > 150 kg. Diet categories: Br, browsers; Mix, intermediate feeders; Gr, grazers. See SOM 2 for further details. SOM 2. Correspondence analysis, size and diet spectra We performed two separate correspondence analyses. The first followed the basic concepts of Bibi and Güleç (2008: 515, 519), based on the relative abundances of bovid genera per site for Turolian (Late Miocene; 8.7–5.0 Ma; Steininger 1999) assemblages from Greece (n = 8), Turkey (n = 8) and Iran (n = 1). Relative abundances of bovid genera were estimated based on the number of identified specimens (NISP), which is closely related to the minimum number of individuals (MNI) [a regression analysis of MNI-NISP pairs of 60 bovid genera from 10 of the Greek and Turkish faunas resulted r2 = 0.93 and p (uncorr) = 2.8E - 33; analysis not shown, but available on request], and weakly related to the number of bovid genera per site [r2 = 0.49; p (uncorr) = 0.001]. Counts for Sivas and Çorak-yerler are from Bibi and Güleç (2008), for Kinik and Mahmutgazi from Köhler (1987), for Akkaşdağı, Perivolaki, Mytilinii-3, and Mytilinii-1A,B,C from Kostopoulos (2005, 2006, 2009b), for Nikiti-2 from Kostopoulos and Koufos (1999) and personal data, for Maragheh from Kostopoulos and Bernor (2011), and for Kemiklitepe D and A,B from Bouvrain (1994). Data for Pikermi include specimens from the NHML, MNHN, and KNUA collections. Counts for Ravin de Zouaves-5, Prochoma, and Dytiko are based on personally collected data. Single occurrences at the genus level were excluded from the analysis. In contrast to Bibi and Güleҫ (2008), we grouped bovid genera into five taxonomic units in our second correspondence analysis: PROTO, including protoryxoid bovids (Skoufotragus/Pachytragus, Protoryx, Sporadotragus, and Palaeoryx); BOSE, including Late Miocene boselaphines (Miotragocerus, Tragoportax, and Samokeros); GAZE, including Late Miocene representatives of Gazella and its allies; OVIB, including Late Miocene “ovibovines” (Criotherium, Urmiatherium, Sinotragus, and Plesiaddax); and SPIRA, including Late Miocene spiral-horned antelopes (Oioceros, Paraoioceros, Dytikodorcas, Hispanodorcas, Prostrepsiceros, Palaeoreas, Majoreas, Nisidorcas, Protragelaphus, and Pheraios). These taxonomic bins, based principally on phenetic similarity (but in some cases also phylogenetic relationships), simply reflect the basic taxonomic structure of the bovid assemblages and allowed us to (1) emphasize general patterns in the taxonomic structure; (2) incorporate genera with low abundances that otherwise require exclusion from the analysis; and (3) overcome differing taxonomic opinions (e.g., Bibi and Güleç 2008). We do not follow Bibi and Güleç (2008) in assuming that all species of a genus tend to have similar environmental responses, but agree with their assumption that the absolute abundance of a taxon or group of taxa through time is itself timeindependent. As body size appears to be a crucial ecological parameter both at the level of the organism and the community level (e.g., Western 1979; Eisenberg 1990), we assume that the size structure of a bovid assemblage is important in understanding its paleoecological context and paleogeographic relationships. We therefore also performed an analysis of the relative abundances of small (less than 50 kg), medium, and large bovids (more than 150 kg) per site. All size categories include taxa of different tribal affiliations, thus making them sufficiently independent from a taxonomic point of view. Finally, we classified all of bovid genera into three main groups of feeding preferences (based on their relative abundances), so as to be able to show the diet spectra of several local bovid associations. These groups consisted of grazers (including intermediate grazers), mixed feeders, and browsers (including intermediate browsers). However, because a single genus may include species with rather distinct feeding behaviors (e.g., Prostrepsiceros; see Merceron et al. 2004, 2005a; Solounias et al. 2010), bovid genera from Ravin de Zouaves 5 (RZO), Perivolaki (PER), Pikermi (PIK), Dytiko (DYTI), Mytiliniii-3 (MYT) and Mytilinii-1A,B,C (MTLABC) were classified strictly based on the results of the dental-wear pattern analyses (including both microwear and mesowear) conducted by Koufos et al. (2006, 2009), Merceron et al. (2005a, b), and Solounias et al. (2010). This means that the same genus may appear as a grazer in one bovid association, but as a browser in another, depending on the local species record (e.g., Merceron et al. 2005a; Solounias et al. 2010). When available, we also considered differences within the same taxonomic unit, as in Tragopotrax rugosifrons from RZO and PER (Merceron et al. 2005a: 480; Koufos et al. 2006: 207), and Gazella from MTLAB (including three species with different ecological profiles; Koufos et al. 2009b). Owing to the absence of similar studies on Turkish faunas, we classified the bovids from Sivas, Kemiklitepe D and A-B, Şerefköy-2, and Akkaşdağı by assuming their feeding preferences to resemble those of taxa from other sites close in time and geography. We did not analyse the ecological spectra of Maragheh, Kinik, Corak-yerler, Mahmutgazi, and Nikiti-2, either because of a lack of data or because of other, similar studies on these sites currently in progress. References Bibi, F. and Güleç, E.S. 2008. Bovidae (Mammalia: Artiodactyla) from the late Miocene of Sivas, Turkey. Journal of Vertebrate Paleontology 28: 501–519. Bouvrain, G. 1994. Les gisements de mammifères du Miocène supérieur de Kemiklitepe, Turquie: 9. Bovidae. Bulletin du Muséum National d’Histoire Naturelle 4, sect. C. 16 (1): 175–209. Eisenberg, J.F. 1990. The behavioral/ecological significance of body size in the Mammalia. In: J. Damuth and B. MacFadden (eds), Body Size in Mammalian Paleobiology Estimation and Biological Implications, 25–37. Cambridge University Press, Cambridge. Köhler, M. 1987. Boviden des Türkischen Miozäns (Kanozoikum und Braunkohlen der Türkei). Paleontologia i Evolucio 28: 133–246. Koufos, G.D., Merceron, G., Kostopoulos, D.S., Vlachou, T., and Sylvestrou, I. 2006. The late Miocene vertebrate locality of Perivolaki, Thessaly, Greece. 11. Palaeoecology and Palaeobiogeography. Palaeontographica Abteilung A 276: 201–221. Koufos, G.D., Kostopoulos, D.S., and Vlachou, T. 2009. The Late Miocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece: New collection. 16. Chronology. Beiträge zur Paläontologie 31: 397–408. Kostopoulos, D.S. 2005. The Bovidae (Artiodactyla, Mammalia) from the Late Miocene mammal locality of Akkaşdağı (Central Anatolia, Turkey). Geodiversitas 27: 747–791 Kostopoulos, D.S. 2006. The late Miocene vertebrate locality of Perivolaki, Thessaly, Greece. 9. Cervidae and Bovidae. Palaeontographica Abteilung A 276: 151–183. Kostopoulos, D.S. 2009b. The Pikermian Event: temporal and spatial resolution of the Turolian large mammal fauna in SE Europe. Palaeogeography, Palaeoclimatology, Palaeoecology 274: 82–95. Kostopoulos, D.S. and Bernor, R.L. 2011. The Maragheh bovids (Mammalia, Artiodactyla): systematic revision interpretation. Geodiversitas 33: 649–708 and biostratigraphic-zoogeographic Kostopoulos, D.S. and Koufos, G.D. 1999. The Bovidae (Mammalia, Artiodactyla) of the Nikiti-2 [NIK] faunal assemblage (Chalkidiki peninsula, N. Greece). Annales de Paléontologie 85: 193–218. Merceron, G., Blondel, C., Brunet, M., Sen, S., Solounias, N., Viriot, L., and Heintz, E. 2004. The late Miocene paleoenvironment of Afganistan inferred from dental microwear in artiodactyls. Palaeogeography, Palaeoclimatology, Palaeoecology 207: 143–163. Merceron, G., Bonis, L. de, Viriot, L., and Blondel, C. 2005a. Dental microwear of fossil bovids from northern Greece: paleoenvironmental conditions in the eastern Mediterranean during the Messinian. Palaeogeography, Palaeoclimatology, Palaeoecology 217: 173–185. Merceron, G., Bonis, L. de, Viriot, L., and Blondel, C. 2005b. Dental microwear of the late Miocene bovids of northern Greece: Vallesian/Turolian environmental changes and disappearance of Ouranopithecus macedoniansis? Bulletin de la Societé géologique de France 176: 475–484. Solounias, N., Rivals, F., and Semprebon, G.M. 2010. Dietary interpretation and paleoecology of herbivores from Pikermi and Samos (late Miocene of Greece). Paleobiology 36: 113–136. Steininger, F. 1999. Chronostratigraphy, geochronology and biochronology of the Miocene “European land mammal mega-zones” (ELMMZ) and the Miocene “mammal-zones” (MN-zones). In: G.E. Rössner, K. Heissig, and V. Fahlbusch, (eds.), The Miocene Land Mammals of Europe, 9–24. Verlag Friedrich Pfeil, Munich. Western, D. 1979. Size, life history and ecology in mammals. African Journal of Ecology 17: 185–204. SOM 3 SOM 3. Table 1. Upper tooth measurements (in mm) of Gazella cf. capricornis, Palaeoryx pallasi, Sporadotragus parvidens, Skoufotragus cf. Sk. schlosseri, and cf. Sinotragus from Şerefköy-2 (Turkey). L, length; W, width; sin, left; dex, right. Specimen LPM Gazella cf. capricornis MYŞE PV-2575 MYŞE PV-2572sin 48.9 MYŞE PV-2572dex 48.0 Palaeoryx pallasi MYŞE PV-2573 111.5 MYŞE PV-1295 MYŞE PV-1293 MYŞE PV-1294 Sporadotragus parvidens MYŞE PV-1533 MYŞE PV-1412 MYŞE PV-1423 MYŞE PV-1522 69.7 Skoufotragus cf. Sk. schlosseri MYŞE PV-1519 MYŞE PV-1513 MYŞE PV-1512 96.3 MYŞE PV-1516 86.3 MYŞE PV-1514 MYŞE PV-1520 MYŞE PV-1515 MYŞE PV-1521 MYŞE PV 1309 dex 90.2 MYŞE PV 1309 sin 88.9 MYŞE PV-1312 MYŞE PV-1525 MYŞE PV-2570 86.2 MYŞE PV-2571 MYŞE PV-1410 MYŞE PV-1491 MYŞE PV-1622 MYŞE PV-1532 MYŞE PV-1312 MYŞE PV-1517 MYŞE PV-1315 MYŞE PV-1523 MYŞE PV-1434 MYŞE PV-1436 MYŞE PV-1313 ?Sinotragus sp. MYŞE PV-1409 LP LM LP2 WP2 LP3 WP3 LP4 WP4 21.5 20.5 20.4 28.7 28.6 7.7 7.6 6.9 6.6 5.7 5.5 7.3 7.1 6.8 6.8 6.3 6.1 6.4 6.5 6.7 7.3 7.7 6.9 46.8 65.4 15.1 13.0 16.8 16.7 18.0 18.1 15.2 14.7 15.1 16.9 17.0 17.6 64.8 LM1 WM1 43.6 32.0 35.6 33.7 57.2 59.7 54.2 59.0 WM2 LM3 WM3 7.6 7.4 9.9 9.2 10.0 10.5 9.5 9.6 11.0 10.8 8.9 9.2 20.3 20.4 22.9 19.8 24.8 21.4 22.4 21.8 26.7 27.1 25.3 24.7 26.8 26.4 15.7 18.8 14.7 12.2 11.3 9.7 12.7 13.6 14.8 12.2 12.8 14.0 12.2 14.9 14.7 14.5 30.5 LM2 12.4 9.9 7.7 9.8 7.9 9.7 8.5 14.8 12.2 16.0 11.4 9.3 9.1 9.8 10.1 11.8 9.9 8.8 11.05 9.6 10.6 12.2 11.1 11.9 12.6 12.4 11.2 13.2 11.9 12.0 11.7 11.4 7.9 9.8 11.4 8.2 15.1 18.3 19.0 16.6 18.9 17.4 16.2 14.8 17.7 13.9 18.7 20.7 20.32 16.4 15.4 15.2 12.3 11.0 10.4 12.2 16.5 16.8 16.7 17.6 21.2 21.7 18.4 21.3 19.6 19.9 14.1 14.3 16.0 15.1 16.0 16.5 20.9 22.8 20.0 21.7 21.5 18.6 20.6 20.8 19.7 18.8 17.5 15.3 20.0 14.7 14.5 17.3 14.6 17.8 16.7 19.8 18.6 21.0 21.1 20.2 14.3 18.3 14.0 14.8 16.1 20.4 19.6 19.1 20.0 23.0 11.8 11.6 12.8 13.2 12.5 18.5 15.0 22.1 21.0 22.2 11.7 13.7 13.7 19.9 19.5 14.3 14.7 15.0 52.0 36.6 35.7 55.8 55.3 12.4 11.5 9.3 9.85 10.5 8.8 12.8 12.5 11.3 11.4 11.4 12.0 12.4 11.7 10.6 11.0 9.5 34.8 53.9 9.9 6.8 11.5 13.0 8.9 11.6 11.8 11.5 8.1 13.2 10.2 9.1 8.8 8.2 54.0 8.5 8.4 11.2 8.9 7.4 9.1 11.6 8.1 SOM 3. Table 2. Lower tooth measurements (in mm) of Gazella cf. capricornis, Palaeoryx pallasi, Sporadotragus parvidens, Skoufotragus cf. Sk. schlosseri, and cf. Sinotragus from Şerefköy-2 (Turkey). L, length; W, width. Specimen Lpm Gazella cf. capricornis MYŞE PV-2562 54.8 MYŞE PV-2557 54.6 MYŞE PV-2563 54.7 MYŞE PV-2558 55.0 MYŞE PV-2564 52.6 MYŞE PV-2565 MYŞE PV-2000 56.7 MYŞE PV-1528 Palaeoryx pallasi MYŞE PV-2574 138.9 MYŞE PV-1599 137.0 MYŞE PV-2552 Sporadotragus parvidens MYŞE PV-1574 MYŞE PV-2561 MYŞE PV-1511 MYŞE PV-1311 MYŞE PV-2556 MYŞE PV-1630 MYŞE PV-1407 MYŞE PV-1406 72.6 MYŞE PV-1429 MYŞE PV-2569 MYŞE PV-2559 Skoufotragus cf. Sk. schlosseri MYŞE PV-1543 90.0 MYŞE PV-1540 MYŞE PV-1541 MYŞE PV-1510 90.0 MYŞE PV-1542 101.2 MYŞE PV-1156 MYŞE PV-2001 MYŞE PV-1546 MYŞE PV-2566 MYŞE PV-2560 MYŞE PV-2568 MYŞE PV-2554 92.3 MYŞE PV-2551 ?Sinotragus sp. MYŞE PV-2553 65.3 MYŞE PV-2567 65.7 Lp Lm Lp2 Wp2 Lp3 Wp3 Lp4 Wp4 Lm1 Wm1 Lm2 Wm2 Lm3 Wm3 20.3 20.1 20.8 20.2 18.7 5.1 5.3 4.8 5.7 5.0 3.0 3.1 2.8 3.1 3.1 2.9 3.8 4.1 3.8 4.0 3.8 3.8 4.1 8.9 7.5 7.6 7.6 8.9 4.7 4.2 4.8 4.8 4.9 4.6 4.4 8.8 9.1 8.7 8.2 8.5 8.1 9.3 6.0 6.0 5.9 5.5 7.6 6.9 7.2 7.2 6.7 6.9 7.8 8.7 21.5 35.7 34.4 34.9 34.4 34.6 32.0 35.7 11.4 10.5 10.5 10.8 10.6 9.9 11.5 6.3 6.3 6.1 6.6 5.9 6.4 6.1 15.9 14.9 16.0 15.4 15.3 14.1 15.5 15.9 5.4 6.2 5.3 6.6 5.7 6.2 5.8 6.1 54.0 55.1 84.0 82.7 14.2 13.7 7.9 8.6 18.9 18.6 10.7 11.9 20.7 21.4 11.6 14.8 22.2 18.7 12.2 27.0 25.9 23.4 13.5 18.5 33.7 37.7 29.6 13.5 16.4 6.8 6.7 4.4 3.8 10.4 10.5 9.9 6.3 5.3 6.4 10.6 11.0 10.9 11.2 6.5 6.4 6.5 6.5 12.2 13.1 12.6 12.0 11.1 7.3 7.9 7.8 8.3 7.3 8.2 7.1 8.4 7.9 17.4 7.1 5.4 6.3 6.3 19.5 19.5 8.0 8.3 13.2 11.1 11.9 8.5 9.4 9.6 9.6 10.3 19.5 9.2 11.4 7.8 8.8 9.6 9.5 8.6 14.6 15.3 13.8 14.8 14.3 15.2 14.8 13.6 16.6 16.1 14.5 20.8 22.9 9.1 9.8 9.3 10.1 11.5 9.9 17.8 18.9 19.3 17.6 19.6 18.3 11.2 9.0 9.9 11.2 10.3 10.2 24.9 26.0 25.9 24.5 27.4 26.3 10.6 8.7 10.3 10.3 9.9 9.5 18.0 17.5 18.4 17.6 16.8 17.4 10.0 10.6 11.7 12.3 12.5 25.3 25.1 10.3 10.1 10.7 11.3 11.3 12.7 13.3 9.7 9.2 19.9 18.6 9.0 8.7 27.3 28.5 43.9 43.5 27.0 27.8 44.5 6.8 8.0 4.2 4.6 10.5 10.0 10.0 27.5 47.0 47.2 7.0 4.0 10.2 5.2 10.1 5.8 11.4 6.6 34.8 35.0 38.5 56.0 61.8 63.8 56.2 63.8 35.1 35.2 9.9 10.5 5.6 5.9 12.1 13.2 6.2 6.7 12.0 6.0 5.8 5.7 10.8 10.9 11.2 11.2 12.2 7.1 7.1 7.2 7.1 7.0 13.6 12.9 8.1 7.4 14.2 14.0 7.6 8.0 16.2 16.9 13.3 15.9 7.5 16.9 9.5 10.0 10.4 5.3 5.3 11.6 11.9 6.5 7.0 13.4 13.4 8.1 8.2 15.7 15.0 10.3 12.2 37.1 55.6 55.6 11.1 5.9 13.0 7.0 13.7 13.7 8.6 8.0 13.7 13.0 11.2 10.5 24.3 24.0 42.0 42.3 7.0 7.5 4.9 4.8 8.5 8.3 5.8 5.8 8.8 8.5 6.4 9.4 8.9