Late Miocene bovids from Şerefköy- 2 (SW Turkey) and their position within the sub-Paratethyan biogeographic province Author Kostopoulos, Dimitris S. Author Karakütük, Seval text Acta Palaeontologica Polonica 2015 2013-06-17 60 1 49 66 http://dx.doi.org/10.4202/app.2012.0129 journal article 294287 10.4202/app.2012.0129 c99fe802-403c-4cbf-bf37-6ef57f433f48 1732-2421 10626287 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 PV-1512, 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 PV-1434, 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 PV-1570, 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 , 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 foramen 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 m 1s, and 3 out of 13 m 2s. 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. 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	TDbase	APDbase	TD10	APD10	Lfpo	Wbrc	Wptbc	Watbc
MYŞE PV-547 dex	29.29	58.84	20.29	37.22	65.13	61.69	35.85	19.94
MYŞE PV-547 sin	32.2	55.52	22.6	38.3
MYŞE PV-1571 dex	34	59.5	22.56	43.36
MYŞE PV-1575 dex	33.36	61.38	22.58	39.84
MYŞE PV-1570 sin	25.92	64.9	22.65	45.54
MYŞE PV-1576 dex	22.42	26.64	11.18	14.69
MYŞE PV-1576 sin	23.16	26.77	9.77	13.95

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. 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 (B 1 ) and lateral (B 2 ) views. 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. 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 Kostopoulos2005), as well as from Samos, Greece and Maragheh, Iran ( Kostopoulos 2009a , Kostopoulos and Bernor 2011 ). Skoufotragus laticeps Skoufotragus sp. Skoufotragus Samos Maragheh Kemiklitepe-A zemalisorum Kinik Samos Skoufotragus schlosseri Samos Akkasdagi Protoryx carolinae Şerefköy-2 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, transverse diameter at the base. 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 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.