The Maragheh bovids (Mammalia, Artiodactyla): systematic revision and biostratigraphiczoogeographic interpretation Author Kostopoulos, Dimitrios S. Author Bernor, Raymond L. text Geodiversitas 2011 2011-12-31 33 4 649 708 http://dx.doi.org/10.5252/g2011n4a6 journal article 10.5252/g2011n4a6 1638-9395 5377377 Skoufotragus laticeps ( Andree, 1926 ) ( Fig. 19 ) Protoryx carolinae var. laticeps Andree, 1926: 153 , pl. 12, figs 5, 9. Protoryx longiceps Pilgrim & Hopwood, 1928: 34 . Protoryx carolinae – Mecquenem 1925: 33 . — Bosscha- Erdbrink 1988: 130-137 . Pachytragus laticeps – Gentry 1971: 244 . — Solounias 1981: 199 . — Bernor et al . 1996 : table 10.2. TYPE LOCALITY. — Samos, Greece (late Miocene). MATERIAL EXAMINED. — MNHN .F: Part of cranium, MAR3206 ( Mecquenem 1925 : pl. V, fig. 3 as Protoryx carolinae ); frontlet, MAR1321 ( Mecquenem 1925 : pl. V, fig. 4 as Pr. carolinae ); part of right horn-core, MAR2092 +2122; left P2-P4, MAR1079 ( Mecquenem 1925 : pl. IV, fig. 5 as Pr. carolinae ); right P2-P4, MAR2991 ( LP = 37.8 mm ); left M2-M3, MAR3002; right p3-m3, MAR1063 ( Mecquenem 1925 : pl. IV, fig. 4 as Pr. carolinae ; Lm = 63.9 mm ); left p4-m3, MAR2972 (Lm = 61.1 mm ); right m1-m3part, 2985+3005. — NHML : part of cranium M3878 ( Pilgrim & Hopwood 1928 : pl. III, figs 2, 2a labelled as M3841). — HUW : p2-m1dex, MMTT13/1397(Lp = 39.6 mm ); m2-m3sin, MMTT13/2504; MMTT7/1987; m3sin. DESCRIPTION AND REMARKS Large antelopes are mainly represented in the Maragheh fauna by the so-called protoryxoid bovids. Two partial crania and a mandible from Maragheh identified by Mecquenem (1925 : pl. IV, fig. 4; pl. V, figs 3, 4) as Protoryx carolinae Major, 1891 were later attributed to a new species Protoryx longiceps Pilgrim & Hopwood, 1928 together with the Maragheh cranium NHML M3878 ( Fig. 19 ). After revising the systematics of this bovid group, Gentry (1971: 245) ascribed the Maragheh sample to Pachytragus laticeps Schlosser, 1904 . Descriptions and illustrations of both the London cranium and the Paris cranial sample are already given by Mecquenem (1925) , Pilgrim & Hopwood (1928) and Gentry (1971) (see also Fig. 19 , Table 14). In summary, the braincase is narrow and long ( Fig. 19B, C ) with the cranial roof facing partly laterally, the occipital condyles are rather small, the basioccipital is long with very strong posterior and strong, welllocalized anterior tuberosities, the dorsal orbital rims project weakly laterally, the fronto-parietal and midfrontal sutures are open, the supraorbital foramina are small, oval and not sunken into depressions, there are no postcornual fossae, the fronto-nasal suture ends to a single point, the frontals are moderately elevated and hollowed, the anterior rim of the orbit is placed behind M3, the ethmoidal fissure is long and narrow, the lacrimal fossa is long, rather shallow and without sharp limits, and the infraorbital foramen opens above P3-P4 limit. The horn-cores are inserted above the orbits, gently and uniformly curved backwards, with increasing upward divergence, flat medial surface and elliptical cross-section that may become semicircular to spindle-shaped towards the tips ( Fig. 19A, B ). The MNHN.F Maragheh sample includes some additional toothrow specimens attributable to this species; a few toothrows in HUW should be also referred here. The upper dentition of this species is characterized by bilobed P2 with an anteriorly placed paracone rib and a weak parastyle. The P3 has a strong parastyle, a thin but strong metastyle and a strong, sharp paracone rib. The P4 shows a weakly developed hypocone, a strong parastyle and a blunt paracone rib. The protocone of the M1 protrudes strongly lingually. The upper molars show a central islet, thin and weak mesostyle and strong parastyle and metastyle, the latter one being directed distally on M3. In the lower toothrow, the p3 and p4 are simple with strong hypoconid, a paraconid well separated from the parastylid (less so on p4), an entoconid quickly merged with the entostylid and a distally placed metaconid that directs backwards on p3, where it is associated by an anterior lingual flange instead of being curved to the front on p4, closing the anterior valley in the lower half of the crown. The lower molars have a strong parastylid, no goat folds, moderate lingual ribs, a basal pillar is usually present on the m1 and less often on the m2, and the third lob of the m3 is semicircular. FIG. 18. — Cf. Palaeoreas sp. from Maragheh: A -C , right mandible (MMTT7/2338) in occlusal ( A ), labial ( B ) and lingual ( C ) views; D , E , left P2-M3 (MMTT7/2159) in labial ( D ) and occlusal ( E ) views. Scale bar: 5 cm. Bosscha-Erdbrink (1988) described an important sample of crania and toothrows recovered in 1973 from the localities Guzguneh, K1 and K2 of Maragheh; disagreeing with Gentry’s (1971) view, he referred them to Protoryx carolinae . Bernor (1978) attributed a partial cranium with horn-cores and several toothrows of the LRE to Pachytragus crassicornis Schlosser, 1904 , adopting Gentry’s (1971) systematic views. Solounias (1981) following only the species-level taxonomy proposed by Gentry (1971) recognized two species of Protoryx at Maragheh, while some years later (in Bernor et al. 1996 ) he accepted Gentry’s (1971) distinction of genera. As a total, 12 more or less complete crania of the so-called “ Protoryx- Pachytragus group” have been so far described from Maragheh, indicating that the relevant taxon (or taxa) was of rather great importance in the structure of the local ruminant community. Given that the systematics of the “ Protoryx- Pachytragus group” has been recently reviewed Kostopoulos D. S. & Bernor R. L. TABLE 15. — Comparison of Palaeoryx Gaudry, 1861 dental characters from Maragheh and Pikermi. TYPE SPECIES. — Palaeoryx pallasi ( Wagner, 1857 ) .

Pikermi	Maragheh
P2 parastyle	strong	stronger
P4 paracone	weaker than para-	equal to parastyle
style and more	and similar with
centrally placed	P3
than in P3
P4 metastyle	strong	stronger
Upper molars	moderate paracone strong paracone rib
rib	and more
quadrangular posterior
lobe in lateral
view
M2	without hypoconal	with weak
hypospur	conal spur
p3	metaconid quickly	metaconid staying
fused with ento-	more
independconid	ent
p4	paraconid fused	open second valley
with metaconid	and less devel-
and well-devel-	oped hypoconid
oped hypoconid
m3	rounded-squarish	angular
protoconidprotoconid and	hypoconid
hypoconid

( Kostopoulos 2009a ), there is no need to repeat all details. According to this viewpoint, Protoryx is generically distinct from Pachytragus (in accordance with Gentry [1971] ), which in its turn is an invalid name replaced by a new one, Skoufotragus . Skoufotragus , originally known from Samos, Greece is represented by three species that succeed one another through time ( Kostopoulos 2009a ). Although the presence of a second Skoufotragus species at Maragheh (i.e. Bernor 1978 ; Bernor et al. 1996 ) cannot be excluded, the currently examined sample from Paris and London falls within the range of Sk. laticeps from Samos both morphologically and metrically; it is therefore referred to this species in agreement with Gentry (1971) . Some of the specimens of the K2 site illustrated by Bosscha-Erdbrink (1988: pl. IV, V) look similar to Sk. schlosseri ( Andree, 1926 ) (i.e. part of Pachytragus crassicornis ; Kostopoulos 2009a ) in their horn-core morphology but again their cranial aspect and basioccipital structure indicate closer affinities with Sk. laticeps .