Tragoportax Pilgrim, 1937 and Miotragocerus Stromer, 1928 (Mammalia, Bovidae) from the Turolian of Hadjidimovo, Bulgaria, and a revision of the late Miocene Mediterranean Boselaphini Nikolai SPASSOV National Museum of Natural History, 1 blvd Tsar Osvoboditel, 1000 Sofia (Bulgaria) [email protected] Denis GERAADS UPR 2147 CNRS, 44 rue de l’Amiral Mouchez, F-75014 Paris (France) [email protected] Spassov N. & Geraads D. 2004. — Tragoportax Pilgrim, 1937 and Miotragocerus Stromer, 1928 (Mammalia, Bovidae) from the Turolian of Hadjidimovo, Bulgaria, and a revision of the late Miocene Mediterranean Boselaphini. Geodiversitas 26 (2) : 339-370. ABSTRACT KEY WORDS A taxonomic revision of the late Miocene Boselaphini is proposed on the basis Mammalia, Bovidae, of the description of abundant Turolian material from the locality of Boselaphini, Hadjidimovo, Bulgaria. The genus Tragoportax Pilgrim, 1937 as usually Tragoportax, Miotragocerus, understood is divided into two genera – Tragoportax and Miotragocerus Pikermicerus, Stromer, 1928 – the latter itself divided into two subgenera – M.(Mio- late Miocene, tragocerus) Stromer, 1928 and M. (Pikermicerus) Kretzoi, 1941. The sexual Bulgaria, taphonomy, dimorphism and the paleoecology of the taxa are discussed as well as the ecology. taphonomy of Tragoportax from Hadjidimovo. RÉSUMÉ Tragoportax Pilgrim, 1937 et Miotragocerus Stromer, 1928 (Mammalia, Bovidae) du Turolien de Hadjidimovo, Bulgarie, et révision des Boselaphini du Miocène supérieur de Méditerranée. MOTS CLÉS À partir de la description de l’abondant matériel turolien de la localité de Mammalia, Bovidae, Hadjidimovo en Bulgarie, nous proposons une révision des Boselaphini du Boselaphini, Miocène supérieur. Le genre Tragoportax Pilgrim, 1937 tel qu’il est habituel- Tragoportax, Miotragocerus, lement compris est divisé en deux genres – Tragoportax et Miotragocerus Pikermicerus, Stromer, 1928 – ce dernier lui-même divisé en deux sous-genres M.(Mio- Miocène supérieur, tragocerus) Stromer, 1928 and M. (Pikermicerus) Kretzoi, 1941. Le dimor- Bulgarie, taphonomie, phisme sexuel et la paléoécologie des taxons, ainsi que la taphonomie de écologie. Tragoportax de Hadjidimovo sont aussi discutés. GEODIVERSITAS • 2004 • 26 (2) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 339 Spassov N. & Geraads D. INTRODUCTION which comes mainly from the locality Hadjidimovo-1 (Hadjidimovo-Girizite) and is The Bovids included in recent times in the genus still mostly unpublished, makes it the richest Tragoportax Pilgrim, 1937 were among the most upper Miocene site of Bulgaria and among the widespread in oriental and southeastern Europe, richest of the Eastern Mediterranean. Intensive as well as in Western Asia (the “Balkano-Iranian” investigations of the fauna and especially of the province: Spassov 2002; Geraads et al. 2003), in ungulates of the locality have started recently the upper Miocene. Numerous European, (Spassov & Ginsburg 1999; Geraads et al. 2001; Asiatic, and even African sites, have yielded Kostopoulos et al. 2001; Hristova et al. 2002; remains of various taxa linked to Tragoportax, Geraads et al. 2003). The faunal list of the loca- that were described during the last 150 years. lity Hadjidimovo-1 includes about 30 mammal These remains, unfortunately often incomplete species. The faunal complex of Hadjidimovo-1 or poorly preserved, are now stored in many shows similarities in taxonomic composition and scientific collections. The opinions expressed by faunal associations with several Hipparion locali- authors are often incomplete or contradictory, ties of the Balkano-Iranian region, and of the and numerous taxonomic and nomenclatorial Northern Paratethys, indicating a middle/late problems remain unsolved. In spite of the wealth Maeotian (middle Turolian) age. Hadjidimovo is of scientific papers on the subject, recent revi- somewhat earlier than Pikermi and it could be sions do not provide a satisfactory frame of the placed at the MN11/12 boundary, or perhaps at taxonomy of this group of fossil boselaphines. the beginning of MN12 (Spassov 2002). We describe here the rich and well preserved sample of boselaphines from the Turolian site of ABBREVIATIONS Hadjidimovo-1 (southwestern Bulgaria). This HD Hadjidimovo; MNHN Muséum national d’Histoire naturelle, also gives us the opportunity to review critically Paris; the taxonomic features mentioned in the litera- NHMW Naturhistorisches Museum Wien; ture, and to propose our own conceptions on NMNH National Museum of Natural History, the taxonomic frame of the “Tragoportax- Sofia; PIK Pikermi; Miotragocerus complex” as well as a new revision PIM Paläontologisches Institut Münster; of the late Miocene Mediterranean Boselaphini. SAM Samos. Upper teeth are in upper case, lower teeth in lower THE LOCALITY HADJIDIMOVO-1: case. LOCATION, FAUNA AND BIOCHRONOLOGY The Hadjidimovo fossiliferous area is situated in the Mesta river valley near the Hadjidimovo SYSTEMATICS town (Gotse Delchev district) and the Bulgarian- Greek border; it is a late Miocene fossil site with Order ARTIODACTYLA Owen, 1848 four localities of vertebrate fauna. The sediments Family BOVIDAE Gray, 1821 (light clayey sands) belong to the Nevrokop Subfamily BOVINAE Gray, 1821 Formation (Vatsev 1980). The site was first men- tioned by Nikolov (1973, 1985) but its richness Tribe BOSELAPHINI Knottnerus-Meyer, 1907 was only fully acknowledged after the excavations of Dimitar Kovachev between 1985 and 1998. Sokolov (1953) created a new subtribe Trago- More than 19,000 bone remains from cerina for the fossil antelopes Tragocerus Gaudry, Hadjidimovo (D. K. collection) are presently 1861, Miotragocerus Stromer, 1928, Para- stored in the Assenovgrad Museum – a paleonto- tragocerus Sokolov, 1949 and Sivaceros Pilgrim, logical division of the National Museum of 1937, but the monophyly of this group is Natural History – Sofia. This huge collection, questionable. Moyá-Solá (1983) defines the 340 GEODIVERSITAS • 2004 • 26 (2) Boselaphini (Mammalia, Bovidae) of Bulgaria Boselaphini by several features as follows: subtri- ?Tragoportax curvicornis Andree, 1926. Samos angular basal section on the horn-cores related to (= T. browni Pilgrim, 1937, Siwaliks, Pakistan, Turolian). the presence of distinct anterior and postero- Tragoportax salmontanus Pilgrim, 1937. Siwaliks external keels; presence of rugosities and crests on (Pakistan). (About 8.1 to 7.9 Ma, but this absolute the anterior part of the parietals. In fact, only the date is doubtful: Barry et al. 2002). presence of the anterior keel is a constant feature Tragoportax maius Meladze, 1967. Bazaleti, Georgia (late Turolian). Perhaps a synonym of the poorly for all taxa included in the Boselaphini. The known Tragoportax eldaricus (Gabashvili, 1956), type other ones are only strong tendencies that may no species of the genus Mirabilocerus Hadjiev, 1961, from be expressed in some species, or variably the late Vallesian/early Turolian of Eldar in expressed within the same species. Azerbaijan. ?Tragoportax cyrenaicus Thomas, 1979, Sahabi (Libya), probably late Miocene. Tragoportax acrae (Gentry, 1974). Langebaanweg Genus Tragoportax Pilgrim, 1937 (South Africa), Mio-Pliocene. Tragoportax macedoniensis Bouvrain, 1988. Dytiko Tragocerus Gaudry, 1861: 298 (type species: Capra (Greece; MN13). amalthea Roth & Wagner, 1854) (non Tragocerus de A number of other specific names, several of them Jean, 1821). from Asia, often based upon fragmentary material, are Tragoportax Pilgrim, 1937: 774. of doubtful validity; we will mention them in the comparison and discussion but we will not try to revise Pontoportax Kretzoi, 1941: 341 (type species: Trago- them here. cerus parvidens Schlosser, 1904). STRATIGRAPHIC AND GEOGRAPHIC DISTRIBUTION. — ?Gazelloportax Kretzoi, 1941: 341 (type species: G. gal- From the Vallesian/Turolian boundary (or late licus Kretzoi, 1941). Vallesian?) to the end of the Turolian, perhaps earli- est Pliocene in Africa. From southeastern Europe and ?Mirabilocerus Hadjiev, 1961: 3 (type species: the northern Paratethys region through Asia minor Tragocerus eldaricus Gabashvili, 1956). and the Middle East to Africa and the northern part Tragoceridus Kretzoi, 1968: 165 (type species: Capra of the Indian subcontinent (and possibly central amalthea Roth & Wagner, 1854). Asia). Mesembriportax Gentry, 1974: 146 (type species: NEW DIAGNOSIS. — Size generally large, approxi- M. acrae Gentry, 1974). mately that of European Cervus elaphus. The post- cornual fronto-parietal surface is a flat or slightly ?Mesotragocerus Korotkevich, 1982: 10 (type species: concave well defined depressed area, usually bor- M. citus Korotkevich, 1982). dered laterally by well marked temporal ridges and caudally by a step leading to a slightly raised plateau TYPE SPECIES OF TRAGOPORTAX. — Tragoportax salmontanus Pilgrim, 1937 (Pilgrim 1937: 774) by (Fig. 1). The basi-occipital has a longitudinal groove original designation. between the anterior and the posterior tuberosities, in the bottom of which often runs a weak sagittal INCLUDED SPECIES. — Tragoportax amalthea (Roth & crest (Fig. 2). Adult male horn-cores are long and Wagner, 1854). Pikermi, most probably also Samos slender, usually curved backwards, with a triangular and Halmyropotamos (Greece) (= ?T. frolovi to subtriangular cross-section, well marked postero- (M.Pavlow, 1913), Chobruchi [Moldova]); from the lateral keel and flattened lateral sides, but are less end(?) of the early Turolian to the beginning of the