GEOLOGICA BALCANICA, 34. 3-4, Sofia, Decemb. 2004, p. 89-109
Tragoportax Pilgrim, 1937 and Miotragocerus (Pikermicerus) Kretzoi, 1941 (Mammalia, Bovidae) from the Turolian of Bulgaria
1 2 3 .Vikolai Spassov , Denis Geraads , Dimitar Kovachev
Sational Museum of Natural History, 1 Tsar Osvoboditel Blvd., 1000 Sofia, Bulgaria; E-mail: [email protected] :uPR 2147 CNRS, 44 rue de l'Amiral Mouchez, 75014 Paris, France; E-mail: [email protected] 'Paleontological Branch of the National Museum of Natural History- Sofia, Assenovgrad, Bulgaria submitted: 06.08.2004; accepted for publication: 26.11.2004)
H. Cnaco6, /(. )J(epaaiJc, /(. Ko6a'le6 - Tragoportax Pil Abstract. The present paper is focused on the occurrence n-im 1937 and Miotragocerus (Pikermicerus) Kretzoi, 1941 and taxonomy of the Boselaphini (Bovidae) in Bulgaria . .\fammalia, Bovidae) u3 mypo11un Eo11zapuu. Pa6oia The very rich material from the Upper Miocene locali· :JOcBell\eHa pacnpocrpaHeHKIO K Haxa.n.KaM trib. ties at the town of Hadjidimovo is described as well as Boselaphini (Bovidae) a EonrapHH. IlpKae.n.eHo omtcaHHe the finding at Kocherinovo. Up to now, the genus 'leHb 6oraToro MaTepHana, Haii.n.eHHoro a nOJ.U.HeMKoue Tragoportax s. Jato has been known from the occur HOBbiX nopo.n.ax paiioHa r. Xa.u.)I(K.U.HMOBO (I03 EonrapHH), rences near the village of Kalimantsi. New finds from the a TaK)I(e - Haxo.n.oK OKano c. Ko'lepHHOBO. Po.u. latter localities allow for a revision also on the remains Tragoportax s.lato y)l(e HJBecTeH no paiioHy KanHMaHUbJ. in this fossil-bearing area. It seems that near Kalimantsi HoBbie Haxa.n.KH .u.a10T BOJMO)I(HOCTb c.u.enaTb peBKJHIO the upper levels of the section contain T. amalthea that acex Q>occHJibHbiX Haxo.n.oK B 3TOM paiioHe. TaM, B 6onee differs from T. rugosifrons in its more massive horns and ::J 03.11.HKX OTJIO)I(eHHHX BH.U.HMO npHCYTCTByeT T. amalthea, small intercornual frontal surface. One of the skulls has i:OTOpb!H OTJIH'!aeTCH OT T. rugosifrons 6onee 6JIK3KOnOC features more similar to the second species mentioned TaBJieHHblMH poraMH. O.n.KH H3 HaJIK'IHbiX KCKonaeMbiX that could be due to provenance from a different locality 'lepenOB OTJIH'IaeTCJI OC06eHHOCTJIMK, CXO.U.HbiMH C in the area. The material from Hadjidimovo contains 3TOpbiM H3 ynOMHl{YTbiX .U.BYX BH.U.OB. Era HaJIK'!He MO)I( Tragoportax rugosifrons (Schlosser) and Miotragocerus HO 06'bliCHHTb pa3JIH'lHeM B npOCTpaHCTBeHHOH npHypo (Pikermicerus) gaudryi (Kretzoi). These taxa are here 'leHHOCTK pa3HbiX Haxo.n.oK. B paiioHe Xa.n.)I(K.li.KMOBO yc· described for the first time from Bulgaria. The very good TaHOBJieHbi: Tragoportax rugosifrons (Schlosser) H state of the fossil material from Hadjidimovo and its .\fiotragocerus (Pikermicerus) gaudryi (Kretzoi). 3TH TaK- richness make possible a revision of the diagnostic fea OHbi onKCbiBaiOTCJI BnepBbre Ha TeppHTOpHH EonrapHH. tures of the genera Tragoportax and Miotragocerus. The ~1aTepHaJI KJ Xa.n.)I(H.li.HMOBO OTJIH'IaeTCJI O'leHb xopoweii two genera are well discernible after the post-hornual COXpaHHOCTbiO. 3TOT QJaKT, B CO'IeTaHHK C 6oraTCTBOM morphology (the presence of a fronto-parietal post :>otaTepHana, .n.aeT B03MO)I(HOCTb KpHTH'IeCKH nepeCMOT cornual depression in Tragoportax) and the relief of the peTb .U.HarHOCTK'IeCKHe oco6eHHOCTH po.n.oa Tragoportax basioccipital surface (presence of a longitudinal groove a \fiotragocerus. 3TH .n.Ba po.u.a xopowo pa3JIH'IaiOTCJI no in Tragoportax). These features allow to confirm the ge l>JOpQJOJIOrHH '!epena B era '!aCTH, H3XO.U.HIUeHCH Ja para neric rank o.f M iotragocerus and to specify the taxa of the loU! (y po.n.a Tragoportax 3Ta o6nacTb BOrHYTa) H no 6aJK- species group within Tragoportax. A skull of M. (P.) KUHnHTaJibHOH noBepxHOCTH (y po.n.a Tragoportax '!epe3 gaudryi determined as a female individuum supplies new Hee npoxo.n.HTb npo.u.onbHall 6opo3.U.Ka). B 3BOJIIOUHK evidence for the presence of a strong sexual dimorphism Bovidae 3TH oco6eHHOCTH OTJIK'iiOTCJI cTa6HJibHOCTbiO B in the representatives of the genus Miotragocerus. Based BJI3H C KOHCepBaTHBHOCTbiO HeBpOKpaHHyMa. 0HK .U.aiOT on the morphofunctional features of Tragoportax 303MO)I(HOCTb nO.U.TBep.n.HTb pa.U.OBbiH paHr Miotragocerus rugosifrons and Miotragocerus gaudryi a conclusion is a YTO'IHHTb TaKCOHbi BK.D.OBOH rpynnbi B paMKax po.n.a made that these two species have occupied different eco Tragoportax. 0.U.HH HJ '!epenoB M.(P.) gaudryi 6hiJI onpe logical niches in the palaeozoocoenosis . .JeneH KaK 'lepen )l(eHcKoro HH.li.HBH.U.a. Enaro.u.apH eMy yc raHOBJieHbi HOBble .U.aHHbie 0 HaJIH'IHH Bbipa)l(eHHOrO no .JOBOrO .U.HMOpQJRJMa y npe.n.cTaBHTeneii p. Miotragocerus. Ha ocHoBaHHH MapQ>ocpyH~HaHaJibHbiX oco6eHHOCTeii
12 Geologica Balcanica, 3-4/2004 89 Tragoportax rugosifrons K Miotragocerus gaudryi, c.nenaH BbiBO.ll 0 TOM, "'TO OHH JaHHManH pa3Hble 3KOflOfH"'eCKHe HHlliH naneoJooueHoJa.
Spassov, N., Geraads, D., Kovachev D. Tragoportax Pilgrim, 1937 and Miotragocerus (Pikermicerus) Kretzoi, 1941 (Mammalia, Bovidae) from the Turolian of Bulgaria.- Geologica Bale., 34, 3-4; 89-109. Key words: Tragoportax; Miotragocerus; M. (Pikermicerus); Late Miocene; Bulgaria.
Introduction More than 19,000 bone remains from Hadji dimovo (D. K. collection) are presently stored The Bovids included in recent times in the ge in the Assenovgrad Museum - a paleonto nus Tragoportax Pilgrim, 1937, were among logical branch of the National Museum of the most widespread in Oriental and South Natural History (NMNH) - Sofia. This huge eastern Europe, as well as in Western Asia (the collection, which comes mainly from the lo "Balkano-Iranian" province), in the Late Mi cality Hadjidimovo-1 (Hadjidimovo-Girizite) ocene. Numerous European, Asiatic, and even and is still mostly unpublished, makes it the African sites, have yielded remains of various richest Upper Miocene site of Bulgaria and taxa linked to Tragoportax, that were described among the richest of the Eastern Mediterra during the last 150 years. These remains, un nean. Intensive investigations of the fauna and fortunately often incomplete · or poorly pre especially of the ungulates of the locality have served, are now stored in many scientific col started recently (Spassov, Ginsburg, 1999; lections. The opinions expressed by authors are Geraads et al., 200 I, 2003; Kostopoulos et al., often incomplete or contradictory, and numer 2001; Hristova et al., 2002). The faunal list of ous taxonomic and nomenclatorial problems the locality Hadjidimovo-1 includes about 30 remain unsolved. In spite of the wealth of sci mammal species. The faunal complex of entific papers on the subject, recent revisions Hadjidimovo-I shows similarities in taxonomic do not provide a satisfactory frame of the tax composition and faunal associations with sev onomy of this group of fossil Boselaphines. eral H ipparion localities of the Balkano-Ira We describe here the rich and well-pre nian region, and of the Northern Paratethys, served sample of boselaphines from the indicating a Middle/ Late Maeotian (Middle Turolian site of Hadjidimovo-1 (South-western Turolian) age. Hadjidimovo is somewhat ear Bulgaria) as well as from other Bulgarian lo lier than Pikermi and it could be placed at the calities (Kocherinovo and Kalimantsi) using MN11/12 boundary (Spassov, 2002). our new conceptions on the taxonomic frame of the "Tragoportax-Miotragocerus complex" (Spassov, Geraads, in press). Systematic description Order Artiodactyla Owen, 1848 The Boselaphini from Family Bovidae Gray, 1821 the locality Hadjidimovo-1 Subfamily Bovinae Gray, 1821 Tribe Boselaphini Simpson, 1945 Hadjidimovo-1. Location, Genus Tragoportax Pilgrim, 1937 fauna and biochronology Synonyms. Tragocerus Gaudry, 1861 (non Tragocerus de Jean, 1821 ); Pontoportax Kre The Hadjidimovo fossiliferous area is situated tzoi, 194I; ?Gazelloportax Kretzoi, 1941; ?Mi in the Mesta river valley near the Hadjidimovo rabilocerus Hadjiev, 1961; Tragoceridus Kre town (Gotse Delchev district) and the Bulgar tzoi, 1968; M esembriportax Gentry, 1974; ian-Greek border; it is a Upper Miocene fossil ? M esotragocerus Korotkevich, 1982 site with 4 localities of vertebrate fauna. The Type-species. Tragoportax salmontanus Pil sediments (light clayey sands) belong to the grim, 1937 Nevrokop Formation (Vatsev, 1980). The site Included species. Tragoportax amalthea was first mentioned by Nikolov (1973, 1985) (Roth & Wagner, 1854) (Capra amalthea Roth but its richness was only fully acknowledged & Wagner, I 854). Pikermi, most probably also after the excavations of one of us, the site in Samos and Halmyropotamos (Greece); = ?T. ventor (D. K.), between 1985 and 1998. frolovi (M. Pavlow) - Chobruchi (Moldova);
90 Fig. I. Post-cornual frontoparietal area in Tragoportax and Miotragocerus. Left : Tragoportax rugosijrons No HD 5125; right : M. (Pikermicerus) gaudryi No HD 5126. Hadjidimovo, coli. of the Assenovgrad Museum (Paleontological Branch of the N.M.N.H. - Sofia) (drawings - V. Simeonovsky) from the end (?) of the Early Turolian to the Stratigraphic and geographic distribution. beginning of the Late Turolian. From the Vallesian{furolian boundary (or La Tragoportax rugosifrons (Schlosser, 1904) [= te Vallesian ?) to the end of the Turolian, per T. parvidens (Schlosser, 1904); = T. recticornis haps earliest Pliocene in Africa. From South (Andree, 1926); = T. punjabicus (Pilgrim, Eastern Europe and the Northern Paratethys 191 0); = ? T. a(yengari (Pilgrim, 1939); = ? T. en region through Asia Minor and the Middle sicornis (K.retzoi, 1941)). Samos (lower levels?), East to Africa and the northern part of the In Prochoma, Ravin des Zouaves, Vathylakkos dian Subcontinent (and possibly central Asia). Ravin C (Greece), Hadjidimovo (Bulgaria), Ve Revised diagnosis. Size generally large, ap les-Karaslari (Republic of Macedonia), Tudo proximately that of European Cervus elaphus. rovo (Moldova) ? , Novoukrainka (Ukraine), The postcornual fronto-parietal surface is a Siwaliks (Pakistan) and possibly in: Vathy flat or slightly concave well-defined depressed lakkos 2, Nikiti 2 (Greece), Maragha (Iran); area, usually bordered laterally by well mar from the Vallesian{furolian boundary to the ked temporal ridges and caudally by a step Middle/Late Turolian boundary. leading to a slightly raised plateau (Fig. 1). ?Tragoportax curvicornis Andree, 1926. Sa The basi-occipital has a longitudinal groove mos (= T. browni Pilgrim, 1937, Siwaliks, Paki between the anterior and the posterior tuber tan, Turolian). osities, in the bottom of which often runs a Tragoportax salmontanus Pilgrim, 1937. weak sagittal crest (Fig. 2). Adult male horn Siwaliks (Pakistan). (Some investigations indi cores are long and slender, usually curved cate for the type an age of- 8.1 -7.9 Ma but backwards, with a triangular to subtriangular his absolute date is doubtful: Barry et al., 2002). cross-section, well marked postero-lateral keel Tragoportax maius Meladze, 1967. Bazaleti. and flattened lateral sides, but are less com Perhaps a synonym of Tragoportax eldaricus pressed than in Miotragocerus. Anterior rug (Gabashvili, 1956), type-species of the genus osities growing downwards from the anterior .\1irabilocerus Hadjiev, 1961. Eldar, Georgia: keel at the basis of horn-cores are absent or late Vallesian to Early/Middle Turolian ? weak, and usually do not extend onto the fron ?Tragoportax cyrenaicus Thomas, 1979, Sa tal. Demarcations (steps) on the anterior keel habi (Libya: probably Late Miocene). are often found, but are few when present. Tragoportax acrae (Gentry, 1974). Lange Horn-cores have a heteronymous torsion (anti aanweg (South Africa), Mio-Pliocene. clockwise on the right horn), so that the ante Tragoportax macedoniensis Bouvrain, 1988. rior keels first diverge in anterior view, but they Ditiko (Greece; MN13). re-approach towards the tips. The interco-
91 Fig. 2. Basi-occipital area in Tragoportax and Miotragocerus (Pikermicerus). Left: Tragoportax rugosifrons No HD 5125; right : M. (Pikermicerus) gaudryi No HD 5126 and 2010. Hadjidimovo (drawings- V Simeonovsky) rnual plateau is rather short antero-posteri trast to T. amalthea). The anterior keel is al orly, broad and almost rectangular between most straight to slightly twisted in front view. the horn -cores. The occipital is not much Description. The Hadjidimovo Material (Pl. broader ventrally than dorsally, giving it a I, 2). The Tragoportax sample from Hadji trapezoid (rather than triangular) outline. dimovo is more abundant than that of Mio Teeth rather hypsodont; labial walls of up tragocerus and is the third more abundant per teeth, and lingual ones of lower teeth with form of the site, together with Gazella, after less accentuated ribs and styles than in Mio Palaeoreas and Hipparion. It is represented tragocerus. Premolars relatively shorter than in mainly by mandibles (159 mandibles and man Miotragocerus. P2 short relatively to P3, espe dibular fragments), more than 40 skulls and cially its anterior part, and parastyle curved skull fragments, upper tooth rows and a lot of backwards. P3 with lingually inflated hypo isolated teeth, several isolated horn cores, sev cone. Metaconid of p3-p4 larger than in M io eral tens of metapodials and metapodial frag tragocerus, splayed lingually and T-shaped on ments, (see inventory numbers of the skull, p4, with an open anterior valley. mandibular and metapodial material in tables 1-4: Collection of the Assenovgrad Paleonto Tragoportax rugosifrons (Schlosser, 1904) logical Branch of National Museum of Natu Tragoportax sp. I (large form): Spassov, 2002 ral History - Sofia). Holotype. skull (Schlosser: 1904, taf. XII [IX], Skull. It is larger than that of Miotragocerus fig. 6) (Tables 1, 2). There is probably a contact be Type-locality. Sames tween the premaxilla and the nasal. There is Diagnosis. Tragoportax of large size. The a wide and deep lachrymal fossa. The infra fronto-parietal postcornual depression is as a orbital foramen opens above P2. The anterior rule distinctly surrounded by a continuous border of orbit remains behind the level of torus, formed laterally by the temporal ridges M3. The frontals and pedicles are hollowed. and caudally by a transverse ridge connecting The horn-cores are long, and the best pre the temporal ridges (Fig. 1). The intercornual served skulls suggest that they were inclined plateau is wide. The male horn cores are usu backwards. Their antero-posterior diameter is ally nearly straight with a moderately convex greater than in most other species of the ge anterior outline in lateral view, and a slightly nus. They are usually strongly divergent concave posterior one. The anterior keel is as a (about 40-60°), but the divergence decreases rule regularly curved, with at most a slight towards the tips. They are but slightly curved tendency to form a demarcation. The tip of the backwards, and the posterior border is almost horn core is not distinct from the general straight. In lateral view, the anterior border shape and direction of the horn core (in con- forms a regular curve, the tip not being dis-
92 Table 1 Skull dimensions and morphologic features of Tragoportax rugosifrons and Miotragocerus (Pikermicerus) gaudryi from Hadjidimovo (HD), and the fossil Boselaphini from Kalimantsi ((K) and Kocherinovo (call. of the Assenovgrad Paleont. Museum and N.M.N.H. -Sofia), Tragoportax amalthea and M. (Pikermicerus) gaudryi from Pikermi (PIK) (MNHN -Paris, University of Athens) and Tragoportax from Samos (NHM_ Wien; Pal. Inst.Univ. Munster).
HO RN-CORES BRAIN-CASE
0! 0! :;:."' 0! 0! :§. -o ·"'G" (j c (j :§. 0 0 :§. 0 0 (j ~ 0 0 (j .... ·r; · ~ "' 0 0 . ~ .D J3 '"-' (j .D 0 u ...... "' B 0 0 .... ~ 0. "" N ~ "'0 0 0 0 0 " ~ 0. B ~ ~ c.. 'i' ... . ~ ~ "g. "'3 0! 0. " e B ~ ~ B 0 :i1 :~"' !l e .D 'il E 0! 0 .~ "5 0 '""~ c ... -5 ... c 0. u 0 " ... ~ " "'0 ..,e ...., ~ .., ~ -€ .D"' N c ... .s 0 § -5 .D "' 0 "'e- "' "'0 "' E E0 :::;; c.. !l ~ ..!l c a 0 0 J3 0 ' § -5 a ~ .c .c 5"' E 0! "'0 B B " 'il " 0. .~ ti .D ~0 ...... ~ ... ~ .§, " 0! § 0 "' 0 0 c c ~ u 'il ·c; 0. ~ ~ -€ ~ 0. 0 'il "' u 0 'il ~ 0 :e. " ·a " E 'C 'C · ~ ... (j i:2 .,... .0 ..,... :a ..c ... 0 0 c.. "' .."' ' (3 b 0 ... ~ > u > .. 0 > > ~ ~ <1:1 <1:1 .D .D 0 0 ..., 0 0 0 0" "' "' :::;; .... ~ 0 ~ "' "':::;; Kalimantsi (K 553) 50,8 I Kalimantsi (K757) - 50,3 I \0 w \0 ~ Continuation of table I T. ama/thea MNHN - PIK - 2285 80 70 33,4 240+ ------57 - - !57 ------MNHN - PIK - 2286 cf 90 76 33+ - - 38 - 78.4 98,5 57 50 - 28,5 42 174 ------MNHN- PIK - 2287 79,2 70,1 45,5 - 123 37 - 82,7 110 - 50 - - - - 164 ------MNHN- PIK - 2288, 88,8 63 30 - - - - - 95+ 59+ 53 - - - 179 ------MNHN - PIK - 2376 ------105 64 46 38 36 ------MNHN - PIK - 2379 ------100 58 45 34 26 40 ------60 - PIK -Athens 83 63,5 49 285 127 - 47 - 97,5 ------47 - PIK. mean cf 80 67 39 274 118 38 45 82 102 60 50 35 29 4 1 170 164 ------cf 1'. cf. amalthea Ra1imantsi (K759), 75-76 64,8 39,3 ------Ra1imantst, (K5560 I) 82 74 45 - - - - - 108 ------Ra1imantsi (K3725) ------62 43,4 - Ra1imantsi (K3768) ------59,5 41.8 - Ra1imantsi (K3727) ------66,9 - - Ralimantsi (K3731) ------61,5 - - Kahmantsi (K3742) ------. . - . - . ------62 Kalimantsi (K3728) 0 - . ------44 .8 - T . amalthea ? NHMW-SAM-V,12. 73,5 68,5 33 325 105 - 37 I 82 102 - 41+ - - - I 165 147 170 ------T . ama/th.ea PIM-SAM-66 - - - 1 - l 110 l - l 42 1 82 - L 58 _l - _1 32.3 23,7 - 148 _l __:_ ------Continuation of table I T. ~osi~rcins IID-5140, ~ ? o - 34,4 32,4 - - 48 - 77 85+ - 47 ------HD-3034, ~ ? - 35 ,5 33,8 - - - - 79 96 - - - 36,3 37 155 139 ------IID-2011, if 76 68 43 235+ - - - 81 - - - - - 35 - 156 ------IID-2327, ~? - 37 30 - 107 42,5 42 81 99 66,6 41,6 41 29 37 159 135 154 ------IID-2322,n 37? 34, 4 29,5 - 108,6 42 - 82 - - 40 36 28 37 - 122 ------IID-2006, if 60 51 38 - 116 50 - 83 103 68 46 42 25 37 ------HD-2027,if 61 56 35 210+ 127 61 - 83 105 72 50 44 25 38 165 153 183 ------IID-2326, ~ ? - 31 30 - 108,4 50 - 83 - - - - - 26 - 119 ------IID-5130,~ - 37 32 - - - - 84 - - - - - 37 145 137 - - - - 61 42,7 103,4 IID-2023, if 79 - 36 - - 61 - 86 115 - 48 42 26 42 - 161 ------IID-2001, if 71 64 44 - 123 - - 88 - - - - - 44 171 156 - - - - 62,5 42 103 HD-5129, if 67 56 38 200+ 117 - 36 88 - - 47 46 27 - 173 165 - 139 - - - - - IID-5132,rf 73 62 42 - 128 - 54 89 110 71 45 43 30 42 173 166 187 135 243 260 65,5 46 109 IID-5125, if iU 60 35 - 123 - 56 91 109 68 45 41 24 35 169 155 177 132 236 256 66,3 - 111 HD-5127, if 58 55 34 285 - - - - 110 72 41 39 23 4"3 - 145 183 134 232 - - 44,3 102,5 IID-2306, if ------107 69 42 43 28 ------liD, mean, if 68 59 38 243 122 59 49 87 108 70 46 43 26 40 1/U 157 183 135 237 258 - - - IID2339 ~ 35 29 IID2340 ~ 34,5 30 IID2377 ~ 39 29,5 IID2375 ~ 36 30 Tragcportax sp. K.ocherioovo-1, ? I - I 31 I 24 I - I - I - I - I - I - I - I - I - I - I - I - I - I - I - I - I - I - I - I - \0 Vl Table 2 pressed in adult males, often rugose, and sur Mandibular teeth dimensions of Tragoportax rounded by a strong ridge. The occipital sur rugosifrons and M. (Pikermicerus) gaudryi face is rectangular, its dorsal part being from Hadjidimovo broad. From the occipital foramen to the Measurements p2-m3 m3 m1-m3 p2-p4 sphenoid, a continuous groove runs along the No basioccipital, often with a weak sagittal keel TraJ(oportax ntJ!Osi{rons in its middle. The choanae open well behind HD 2506 - 27,7 67,9 - M3. The basicranial angle is quite open. HD 25 11 - - - - HD 25 13 - - 69,2 - Female and sub-adult skulls. There is no HD 3834 - - 72,9 - large Bovid hornless skull in Hadjidimovo. HD 2521 - - 77,8 - However, there are at least two skulls that we HD 3824 - - 69,7 - interpret as females of T. rugosifrons, HD HD 2443 110,2 - 66,4 43,4 5130 and 5138. Their teeth are well pre HD 2484 - - 70,7 - HD 25 18 - - 68 - served, and they are both fully adults. Their HD 2489 - - 68,2 - size and morphology is identical to those of HD 2519 - 27,7 67,9 - the male skulls, and there is no doubt that HD 2524 - - 70,3 - they belong to the same species. However, HD 3896 - - 69.1 - HD 2527 110,1 30 72,4 41 ,8 their horn-cores are much smaller, with a HD 2480 - - 71,7 - rounded oval basal cross-section, but with a HD 2484 - - 70,5 - postero-lateral keel in one of them (HD HD 2489 - - 68,2 - 5130), without anterior keels or flattening of HD 2440 121 3 1 73,6 47 the surfaces. Several separate horn cores (HD HD 2518 - - 68 6 - HD 2524 - - 70,5 - 2337; 2339; 2340, 2375) with suboval cross HD 2497 - - - - section, slightly flattened anterior surface HD 2445 - 27,5 65,4 - and similar dimensions (some of them with HD 2501 - - 67 1 - rudiments of postero-lateral keel) must also HD 2468 - - - - belong to female individuals (Table 1). HD 2565 - - 66,9 - HD 2493 - - 68,7 - We interpret as sub-adult male skulls a few HD 2520 - - 68,7 - frontlets and brain cases (HD 3034, 2327, HD 114,5 28,2 67,4 48,2 5137, 5140), with horn-cores smaller than in HD 108,5 27,7 65,3 43,3 adult males, but with a triangular basal cross M . (Pikermicen ts) J 96 Table 3 Metacarpal dimensions of Tragoportax rugosifrons from Hadjidimovo Transversal Antero-posterior Transversal Antero-posterior Diaphysal Length No diameter prox. diameter prox. diameter dist. diameter dist. transv. diameter 38001 40,5 29,1 41,5 30,8 25,1 254 38059 - - 39,8 30 24,4 - 38061 - - 42,3 30,3 23,9 - 38010 - - 42,7 30,7 23,4 - 38039 - - 44,7 30,9 25 - 38374 41,6 28,7 - - 24 - 38035 - - 38,4 30 23 1 - 38022 - - 41 29,6 24 - 38030 - - 41,2 30,6 - - 38034 - - 4 1,5 29,5 - - 38026 -- 42 28,5 - - 38071 - - 42 28,3 - - 38023 - - 41,8 30 - - 38047 - - 41,2 29 - - 38041 - - 40,3 29,8 - - Table 4 J1etatarsal dimensions ofTragoportax rugosifrons and M. (Pikermicerus) gaudryifrom Hadjidimovo Transversal Antero-posterior Transversal Antero-posterior Diaphysal Length I)Jo diameter prox. diameter prox. diameter dist. diameter dist. transv. diameter Tragoportax rugosifrons 38002 36 38,7 42,7 30,4 24,5 258 32004 35,7 36 40,8 29,8 24,3 265 38003 39 39,6 43,6 30 24,9 257 38043 - - - 28,5 23,5 - 38042 - - 41,2 29,5 - - - - - 42,9 28,9 24 - 38060 - - - 27,8 24,3 - 38036 - - 41 ,2 30 - - 38076 - - 43,5 30,1 - - 38057 -- 43,2 30,2 - - 38006 - - 38,6 28,5 - - 38055 - - 39,6 27,8 - - 138029 - - 41,2 30 - - M. (Pikermicerus' gaudryi 38370 32 29,7 - - 19 220? 38007 - - 31 5 23 17.7? - ,. - - 31 ,7 25,8 - - ]8015 - - 33,8 24,6 - - 13 8053 - - 33,5 25,3 - - Comparison diameter; their triangular cross-section and normal torsion; the hypsodont teeth; the rela There are many differences between the two tive length and morphology of the premolars Boselaphines from Hadjidimovo. Tragoportax (fig. 1-6, tab. 1-4). rugosifrons differs from Miotragocerus mainly The Tragoportax from Hadjidimovo has · y: the larger size; the broad rectangular similarities with T. amalthea (Roth & Wagner) - tercornual plateau; the presence of a fronto in its main features and general aspect, but ;>arietal postcornual depression; the shape of also differs clearly by: the greater intercornual :he basioccipital (with longitudinal groove of distance, the less robust horn cores (fig. 7), :en with a thin sagittal crest); the long horn lesser development of the rugosities on the an -ores with a relatively small antero-posterior terior horn-core keel, lesser concavity of the ! ' Geologica Balcanica, 3-4/2004 97 75 0 0 0 70 0 :§ 0 0 "i 0 c 65 • e M. (P.) gaudryi ~ 0 T. rugosifrons -=c ...C) 60 :.0 • • 55 I i 50 ! 80 85 90 95 100 105 110 115 maximum brainc:ase Mdth Fig. 3. Neurocrania) dimensions in Tragoportax rugosifrons and M. (Pikermicerus) gaudryi from Hadjidimovo 300 285 270 .r:: ill 255 c ..!:! 240 ..... 0 e M. (P .) gaudryi C) 225 0 0 '( 0 T. rugosifrons ...c 210 C) .r:: 195 180 • 165 150 • • 55 60 65 70 75 80 horn-c:ore maximum antero-posterior diameter Fig. 4. Horn-core proportions and length in Tragoportax rugosifrons and M . (Pikermicerus) gaudryi from Hadjidimovo 50 -·-·---·----.----...... -·-·------·--·- ...... -- _,. ·- .. 45 .. 0 Q; 0 e 0 i' • 40 0 :a e M. (1' .) lli'udryi iii 0 I .. 0 0 T. rugosifrons ..> 35 ..c 0 0 • • • . o •• = 30 u • 25 • 55 60 65 70 75 80 85 maximum anter&-pOSterior diameter Fig. 5. Basal horn-core diameters in Tragoportax rugosifrons and M. (Pikermicerus) gaudryi from Hadjidimovo 98 125 ] ...... _,.,_ ...... _, ___ ...... ______...... --..... ___ ,______,...... ·----.. ! 120 - 0 j I liS I 'I ,.., .... 110 I E "' • M . ( P. ) gaudryi ' I ~ lOS 0 T . rugosifrons • 100 • I I 95 • • 90 38 40 42 44 46 48 p2-p4 Fig. 6. Mandibular teeth proportions in Tragoportax rugosifrons and M . (Pikermicerus) gaudryi from Hadjidimovo Ql 80 ~ - ·- -··-····-- --········· 'Z ! ..E 75 • :s 70 ... 0 • .. 65 =:s"' • • •• OHD = 60 0 • PIK .. + e- 55 0 0 A DT K ..c. ... ! ... 50 +RZO 0 ! ·;:: I + SAM ~ 45 ...0 ~ 40 ... .. 35 u =.. 30 100 150 200 250 300 350 horn-core length Fig. 7. Horn-core proportions in various Tragoportax taxa from various localities (HD - Hadjidimovo, PIK. - Pikermi, DTK - Ditiko, RZO - Ravin des Zouaves, SAM - Samos) 76 ...... ____ ... _,______.....- ...... _ .., ____ , ___, ______l 74 T------_~o~------_,~ 0 72 o HD .... t>. MAR E 70 I o SLQ -e 68 • PIK 0 0 - NIK 66 • 64 0 0 ! • j 62 41 42 43 44 45 46 47 48 49 p2-p4 Fig. 8. Teeth proportions and size in Tragoportax from various localities. (SLQ - "Thessaloniki", Arambourg's collection; other abbreviations as in fig. 7) 99 caudal face of the horn core in lateral view, Compared with T. salmontanus from Siwaliks usually weaker torsion of the horn-core ante (type-species of the genus) the horns of the rior keel, as well as by the virtual lack of steps Hadjidimovo Tragoportax are longer, less in (demarcations) on the keel. The tip of the clined and not so twisted. There are less rugosi horn-core is usually not distinct from the gen ties on the fronto-parietal postcornual surface eral outline of the horn core, or at least less so (see Pilgrim 1937, 1939). The horn-cores of the than in the Pikermi T. amalthea, where this tip poorly known T. perimensis from the Midlle is demarcated by a marked concavity of the Siwaliks (Pilgrim, 1939) are much shorter. An anterior edge in lateral view. The cheek-teeth other poorly known species from Siwaliks, sample from Hadjidimovo show also some dif T. islami (a possible synonym ofT. salmontanus), ferences, having somewhat shorter premolar has also more rugosities on the fronto-parietal row in relation of the molars (fig. 8). postcornual surface, which extends farther From the specimens described as T. curvi back than in the Hadjidimovo form. cornis (Andree, 1926: Pl. XI -6,7 and Solou The sample of Tragoportax skulls from nias, 1981: fig. 30-H) the sample from Hadji Hadjidimovo show all the main features of the dimovo differs by the very slight backward Tragoportax rugosifrons morphology, by com curve of the horn core (reflected in the very parison with the type specimen and the other slight concavity of the posterior border of the Samos specimens figured and described by horn core in lateral view). Schlosser (1904). A small sample of Trago "Miotragocerus" cyrenaicus Thomas, from portax skulls from Veles-Karaslari from the Sahabi (Libya) differs by its curved and diver Skopje Museum of Natural History (R. of gent horn-cores, but the species is illustrated Macedonia) is identical with the skulls from by a single specimen. Hadjidimovo and represent the same species. The single specimen from Novaya Emetovka The skulls from Hadjidimovo are similar to the (type of M esotragocerus citus (Korotkevich) ) preserved skulls with horn cores of T. that could be referred to Tragoportax shows rugosifrons from Prochoma (Spassov, Geraads, more twisted and more inclined horn cores in press). than the Hadjidimovo form. From T. ("Mirabilocerus") maius Meladze, Genus Miotragocerus Stromer, 1928 of which we have seen good photos kindly pro Synonyms. Pikermicerus Kretzoi, 1941; ?Indo vided by A. Vekua and D. Lordkipanidze (see tragus Kretzoi, 1941; Dystychoceras Kretzoi, also Meladze, 1967: Pl. XXVII), the sample 1941 from Hadjidimovo differs by the larger and Type species. Miotragocerus monacensis more quadrangular inter-cornual surface, the Stromer, 1928 much stronger temporal lines and torus that Included sub-genera. Miotragocerus (Mio surrounds the fronto-parietal postcornual de tragocerus) Stromer, 1928; Miotragocerus (Pi pression, lesser inclination of the horn core kermicerus) Kretzoi, 1941 and the totally different profile of the horn Included species. Miotragocerus (Miotrago core, with the antero-posterior diameter de cerus) monacensis Stromer, 1928 - Ober creasing gradually, in contrast of the abrupt fohring (Austria); ?Hostalets, ? Ballestar diminishing in "Mirabilocerus maius" above (Spain); approx. MN8f9. the first half of the horn core, with a change of ?Miotragocerus (Miotragocerus) pannoniae curve in its second half, thus forming a sinu Kretzoi, 1941 - Sopron (Hungary); Altmanns soidal contour. "M." maius is perhaps a junior dorf, Mistelbach, Inzersdorf (Austria); most synonym of T. ("Mirabilocerus") eldaricus possibly also Eppelsheim and Howenegg (Ger (Gabashvili, 1956), the type of which is an iso many). The species has also been listed in sev lated horn-core. eral localities from the Vallesian of Spain (Mo The horn cores of the specimens from Hadji rales et al., 1999), from Kalfa in Moldavia dimovo differ by the same features from those (Pevzner and Vangengeim, 1993) and Grizev ofT. acrae (Gentry, 1974, 1980). The latter dif in Ukraine (Korotkevich, 1988); Late Middle fer also by their convex posterior profile (in Miocene-Valles ian. stead of concave in the Hadjidimovo sample). Miotragocerus (Pikermicerus) gaudryi (Kre The Tragoportax from Hadjidimovo is much tzoi, 1941) with three subspecies (see below). larger than T. macedoniensis (Bouvrain, 1988), Mostly at: Pikermi, Samos, Halmyropotamos, and has straighter and more robust horns in Nikiti 1 (Greece, Turolian); Hadjidimovo (Bul males, more rectangular inter-cornual surface garia, Early/Middle Turolian); Veles - Karas and more rectangular (wider) occipital surface. lari (R. of Macedonia), Le Coiron (France, 100 early Turolian); possibly at Belka (Ukraine, short and massive, with a concave caudal Turolian); Piera, Early Turolian and Venta del edge. Long premolar row. Metapodials of the Moro, Late Turolian (Spain); Mt Luberon size of the fallow deer, without lateral longitu (France, Middle/Late Turolian); ? Maragha dinal groove; abrupt widening from diaphysis (Iran, ?E. Turolian), Nikiti-2 (Greece, early to epiphysis. Turolian), Maramena (Greece, Turolian/ Stratigraphic and geographic distribution. Ruscinian boundary), Kalimantsi (see below). Whole Turolian; possibly late Vallesian. Eu The presence of the species at Nikiti-1 (Gree rope and perhaps Middle East (see above the ce, end of Vallesian ?) is uncertain. distribution of the Genus) Possibly also a fourth species: see "Tra goportax (Pikermicerus) afT. gaudryi": Maya Miotragocerus (Pikermicerus) gaudryi (Kre Sola, 1983 (see Discussion). Spain, France, tzoi, 1941) Late Vallesian. Pikermicerus gaudryi Kretzoi, 1941 Stratigraphic and geographic distribution. Miotragocerus monacensis: Solounias, 1981- From MN8/9 to MN13 of Europe and possibly pars the Middle East (from W.E. Europe to C. Asia). Tragoportax gaudryi: Moya -Sola, 1983 Revised diagnosis. (The shape of horn-cores Holotype. skull illustrated in Gaudry (1865, Pl.49, fig. I) and those of associated structures are those of Type-locality. Pikermi adult males). Size small (about that of fallow Diagnosis. That of the subgenus deer). The post-cornual area of the skull is not depressed or raised as a low plateau (Fig. 1). M. (Pikermicerus) gaudryi gaudryi (Kretzoi, Basioccipital without median longitudinal 1941) groove, but with a faint sagittal keel (Fig. 2). Tragoportax sp. II: Spassov, 2002 Strong temporal crests in early forms (Mio tragocerus), weaker in more recent ones (Pi Holotype. skull illustrated in Gaudry (1865, kermicerus). Horn-cores short, media-laterally Pl. 49, Fig. 1) compressed, with flattened lateral and medial Type-locality. Pikermi surfaces. Sharp anterior keel, but postero-lat Diagnosis. A subspecies of M. (P.) gaudryi eral keel absent or poorly marked, and poste with horn cores straighter than in M. (P.) rior face not well delimited. The section is gaudryi andancensis from the Early Turolian of therefore sub-elliptic. Anterior rugosities at France, larger than M. (P.) gaudryi crusafonti base of horn-cores usually strong, extending from the Early Turolian of Spain and France, onto the frontal along the keel, which often has teeth smaller than those of M. (P.) gaudryi several demarcations (steps) along its course. leberonensis from the Late Turolian of the In front view, due to a slight torsion of the same area. horn-cores bases, the keels are often slightly Description. The Hadjidimovo material (pl. convergent upwards in the basal portion, then 1-3). (Coll. of the Assenovgrad paleontologi diverge towards the tips. The inter-cornual cal branch of the N.M.N.H. - Sofia ): com area is much longer than broad, especially plete and partial skulls (HD 2007, 2010, 2015- narrow anteriorly. The occipital surface is 2016, 2019, 2039, 2325, 3704, 5519); several high, much broader basally than at its top. identified mandibles and metapodials (see in Teeth brachyodont, with strongly folded walls. ventory numbers in tables 2-4). Compared to Tragoportax, well documented Skull. It is larger than those of M. mona forms have a long premolar row, with espe censis, but smaller than those of Tragoportax cially long P2 compared to P3, due to length from Hadjidimovo (Fig. 3). The ante-orbital ening of its anterior part. Hypocone of P3 fossa is small and shallow. The anterior border poorly expanded lingually. Metaconid of p3- of orbit does not reach the level of M3 (one p4 weak, anterior valley with incipient lingual specimen observed). The frontals, and prob wall (Spassov, Geraads, in press). ably also the pedicles, are hollowed. The horn cores are short, slightly inclined backwards, Subgenus Miotragocerus (Pikermicerus) and very compressed transversally. They look Kretzoi, 1941 stout and massive in lateral view: their antero Type species. Pikermicerus gaudryi Kretzoi, posterior diameter is greater than in other spe 1941: 342. cies of the genus (Fig. 9) and this dimension is Diagnosis. Cranial features as for genus greater relatively to the length of the horn -core (fig. 2). Weak temporal lines. Compared to M. than in Tragoportax from the same locality. (\!iotragocerus), the horn-cores are relatively Horn-cores are only slightly divergent. In lat- I 01 ~ 41 e HD i ~ ] -~- - -- ·------.. ------0 --~ OI'IK • - NIK ] 60 +---=·=------··----*- g'-'------' I+M . monacensis t 55 +------+ ______..'------; x M . monac. Spain ~ 50 i 16 SAM · ~ 45 g o I ~M . g. crusafonti 41 ~ ~ +------~ 0 X • t;jMl. Leberon b 35 • - ;JM. cf. gaudryi ~ 30 +----.-----.-----.----.---~ =.. 15 20 25 30 35 40 horn-core trans~rsal diameter Fig. 9. Horn-cores' proportions and dimensions in various taxa of Miotragocerus 95 -··- .... __ ------...... 0 w+------0 ----- ... 0 0 .; 0 'i 85 0 e M. (P .) gaudryi ~ 0 0 .. 0 70 110 120 130 I~ 150 160 170 180 braincase length Fig. 10. Neurocranial proportions of Tragoportax and M. (Pikermicerus) from Hadjidimovo. Length of brain-case from front of horn-cores to top of occipital eral view the anterior contour is strongly rocranium is long compared to Tragoportax curved, but the posterior one is only slightly (Fig. I 0). The fronto-parietal post-cornual curved backwards. The demarcations are well area is not depressed, and shows rugosities in represented along the anterior keel. The keel one specimen. The occipital surface is sub torsion is weak (in some specimens almost ab triangular, its dorsal part being relatively nar sent) and homonymous: in front view the keels row with bulging top. The basioccipital has no are often slightly convergent upwards at base, median longitudinal groove, but bears a faint then diverge towards the tips. The basal cross sagittal keel (Fig. 2). The basicranial angle section in adult specimens of both sexes is (one specimen observed) is quite well marked subtriangular, less triangular than in Tra (but less than in Palaeoreas). goportax from the same locality, with a Teeth. Premolars and molars are relatively rounded posterior face, without longitudinal small and brachyodont with pronounced re groove, nor postero-lateral keel (only very lief of the posttrite walls. The premolar row is weakly distinct in a few specimens ). The ante relatively long. P2 is long relatively to P3, due rior keel between the demarcations is well to lengthening of its anterior part. Its parastyle marked. The rugosities on the anterior keel are is straight, not curved distally. P3 has a narrow more common than in Tragoportax from the parastyle, not protruding labially. The meta same locality and in most cases they extend conid of p3-p4 is weak, the anterior valley has onto the frontal. The inter-cornual plateau is an incipient lingual wall. Length Ml-M3 (53 usually long, narrow and triangular. The neu- mm) falls within the size range of Pikermi (48.6 102 36 .. 0 ~ 34 = DO • • :6 32 •• ~ 30 • ..... 0 ~ 28 01 0 0 .:: 26 • .,... e 24 .: ::; 22 .c 20 30 40 50 60 70 80 90 100 manimum horn-core antero-posterior diameter Fig. 11. Comparison of the horn-cores' proportions and dimensions of M. (Pikermiccrus) from Hadjidimovo and Pikermi - 53.9 mm), it is slightly smaller than at Nikiti abrupt change of width. The trochlear keels (55.1 mm), but much larger than at Piera (45 are prominent and sharp. The lateral longitu mm). dinal groove specific to the metapodials of Female skull. HD 55611 (Pl. 3.1, Table 1). ?M. pannoniae is absent here. The skull is of a subadult animal (dP3-dP4 present, frontal suture visible). It is well pre served, (except horn cores) incl. the rostral Comparisons bones, but it is crushed laterally (incl. the frontals anteriorly to the horn core bases). The Comparison with the ante-Turolian forms. The mandible, which is still attached, is also well horn core of the type of M. (Miotragocerus) preserved except the angular processes. The monacensis from the AstaracianfVallesian basioccipital surface is damaged. The skull is (Stromer, 1928) is smaller than the horn cores smaller than a Tragoportax skull (total length of the sample from Hadjidimovo, and the tem about 305 mm) and the fronto-parietal poral lines are stronger (Fig. 9). The compari postcornual area is not depressed. The angle son with M. pannoniae is difficult because the between the splanchnocranium and the neu type-material is scarce, and moreover belongs rocranium is large (about 1500) - a condition to a subadult individual, and because referral typical for the Boselaphini. The P2 (length of material from other localities is uncertain. about 15 mm) and the premolar row are very The differences of the M. (Pikermicerus) long. The P2 parastyle is high and not curved gaudryi .sample from Hadjidimovo with the backward. The labial relief of the upper teeth graceful horn-cores of the type of M. (P.) is strong, the ml-m2 have strong ectostylids. monacensis are clear. From the horn-core from All these features as well as the tooth row Inzersdorf supposed to represent an adult length are the same as in M. (P.) gaudryi. The male of this species (Thenius, 1948) it differs anterior border of the orbits are above M2. The mainly by the less straight posterior edge of the praemaxillae are slender, not at all broadened horn-core and probably by the smaller number anteriorly. The upper profile of the splanchno of demarcations. From the Howenegg sample cranial is convex but this is possibly somewhat described as M. pannoniae by Romaggi (1987) exaggerated because of the transverse crush the Hadjidimovo skull material differs also by ing. The mandibular corpus is strongly convex the curved (no straight) surface of the posterior ventrally. The frontals under the horn cores horn core wall as well as by the shorter horn are pneumatized. The preserved horn core cores. The metatarsals of theM. (Pikermicerus) bases are small. The horn core shape in spite from Hadjidimovo lack the proximal lateral of some deformation suggest a nearly oval (and in some cases lateral and medial - at cross-section, wider and more rounded poste Howenegg) longitudinal depression descri riorly and more angular anteriorly. bed for some Vallesian bones referred to M. Metapodials. The metapodials, of the size of pannoniae (see below). The comparison with the fallow deer, are relatively small compared the material of M. (Pikermicerus) "cf. gaud with Tragoportax from the same locality (see ryi" from the Late Vallesian of La Croix tab. 4) and slender. The widening from dia Rousse in France (data in Moya-Sola, 1983: physis to epiphysis is clearly bovid-like with 166) shows that both the metapodials and I 03 PLATE I I. M.(Pikermicerus) gaudryi, lower tooth-row, no N•. 2. Tragoportax rugosifrons, lower tooth-row, HD-2443. 3. Tragoportax rugosifrons, upper tooth-row, HD-3664. 4. M.(Pikermicerus) gaudryi, upper tooth-row (P2 missing), HD- 3672. 5. Tragoportax rugosifrons, skull HD-2040, lateral view. 6. Same specimen, front view. 7. Tragoportax rugosifrons, skull, HD-5127, lateral view. 8. M.(Pikermicerus) gaudryi, frontlet HD-2010, front view. 9. M .(Pikermicerus) gaudryi, braincase with horn-cores, HD-5519, lateral view. Scale= 5 em for Figs. 1-4, 15 em for Figs. 5-6, 8-9, 20 em for Fig. 7. PLATE II 1. Tragoportax rugos1jrons, occipital, HD-5132. 2. M.(Pikermicerus) gaudryi, occipital, HD-2039. 3. Tragoportax rugosifrons, female skull, HD-5138. 4. Tragoportax sp., female skull from Kocherinovo (FM 2024 ). 5. M.(Pikermicerus) gaudryi, basi-occipital, HD-2007 (stereo). 6. Tragoportax rugosifrons, occipital, no N• (stereo). Scale = 10 em for Figs. 1, 2, 4-6; 15 em for Fig. 3. PLATE III 1. M. (Pikermicerus) gaudryi, female subadult skull from Hadjidimovo (HD 55611), lateral view. 2. Same specimen, dorsal view of neurocranium. 3. Tragoportax cf. amalthea, frontlet from Kalimantsi, K-759, antero-dorsal view. 4. T. ? rugosifrons, dorsal view of skull from Kalimantsi, K 55603. 5. Tragoportax cf. amalthea, incomplete skull from Kalimantsi, K-55601. Scale = 6 em for Fig. 2, 10 em for all others. horn cores of the Hadjidimovo material are Skull No. 55611 deserves special attention. clearly larger. The combination of features such as face/neu Comparison with the Turolian samples. M. rocranium angle, tooth dimensions, propor gaudryi - The differences between the three tions and morphology as well as the post accepted forms of M. gaudryi are also mostly cornual morphology fit the Boselaphini, espe metrical. So, M. (P.) gaudryi (= Tragoportax cially Miotragocerus (Pikermicerus) gaudryi. gaudryi sensu Moya-Sola, 1983 and Bouvrain, The facial bones of the genus and the species 1988) generally increase in size from Early to are very poorly known, but the shape of the Late Turolian (Moya-Sola, 1983; Bouvrain, mandible and of the splanchnocranial con 1988). The metrical data of the metapodials tour resemble the skulls determined by Ro (tab. 4) as well as those of the mandibular/ maggi (1987) as Miotragocerus pannoniae and maxillar teeth and horn-cores (Tab. 2, Fig. 9, Graecoryx andancensis (i.e. M. (P.) gaudryi 11) show that the Hadjidimovo sample has andancensis in our meaning). The small horn larger body and teeth than M. gaudryi cru core cross-section could correspond to a fe safonti from Piera (MN11, Spain; Moya-Sola, male specimen. The dimensions are somewhat 1983: 137-138). This locality has yielded a smaller than in the female of Graecoryx large number of Boselaphine crania. However, andancensis (see Romaggi, 1987) (i.e. of M. (P.) some details of Moya-Sola's description and gaudryi andancensis) but it could be explained figures suggest that the sample might not be by the subadult age of the Hadjidimovo speci homogeneous, and perhaps include some men. These dimensions (31 -32 x 21) are not Tragoportax. On the other hand, the lower very far from the horn cores dimensions (35.3- teeth from Hadjidimovo (Tab. 2) are smaller 38.7 x 22.2-24) of the skulls noted as females than the latest as well as largest known form, from the Nikiti-1 M. (Pikermicerus) (see Kosta M. g. leberonensis from Mt Luberon (MN 12- poulos & Koufos (1996). The presence of only 13, France) and Venta del Moro (MN13 of one female skull of M. (Pikermicerus) in the Spain) (data in Moya-Sola, 1983: 161). The Hadjidimovo taphocoenosis is not so aston horn cores from Hadjidimovo differ from those ished having in mind the difference in the from Le Coiron, France, described by Romaggi thickness of the female and male bones of the (1987) as Graecoryx andancensis by the lack of skull dome. We have also found only one fe a strong curve backward, especially in females. male skull in a very large sample of male skulls The Hadjidimovo material is very close in skull in the Palaeoreas collection from Hadjidimovo and horn-core morphology as well as in di as well (Geraads et al., 2003). mensions to the material from the Middle Turolian of the type-locality, Pikermi (Tab. 1, Fig. 15, 17). The Greek material from Sames Discussion (Solounias 1981 ), as well as two undescribed frontlets from Veles-Karaslari could be in The Hadjidimovo material and the taxonomy of cluded in the same form, M. (P.) gaudryi "Tragoportax" and the Late Miocene Bosela gaudryi. phini. Today, most authors agree that there are 104 PLATE I .V. Spassov, D. Geraads, D. Kovachev - Tragoportax Pilgrim, 1937 and Miotragocerus (Pikermicerus) Kretzoi ... Geologica Balcanica, 34, 3-4, 2004 PLATE II 5 6 PLATE III 2 two clearly distinct forms at Pikermi (So micerus) gaudryi and Tragoportax rugosifrons lounias, 1981; Moya-Sola, 1983; Bouvrain, from the Hadjidimovo taphocoenosis. We could 1994), one is smaller with shorter and more characterize both forms after the morphologic massive horn-cores, with an elliptical cross and taphonomic analysis -as follows: M. section, the other is larger with horn-cores (Pikermicerus) gaudryi is a medium sized bovid sub-triangular in section (Moya -Sola, 1983). (clearly smaller than Tragoportax rugosifrons). The smaller species - T. gaudryi Kretzoi - has The teeth are relatively brachyodont, with been recognized in other European sites strongly folded posttrite walls. The horns are (Moya-Sola, 1983; Bouvrain, 1988; Kostopou relatively short with clear sexual dimorphism. los, Koufos, 1996). Kretzoi (1941) insisted The species is clearly less abundant in the upon the very characteristic morphology of the taphocoenosis of the locality than Tragoportax smaller form, based upon a Pikermi skull de rugosifrons. scribed by Gaudry (1865), which led Kretzoi to Tragoportax rugosifrons is a large bovid (its create a new genus for it, Pikermicerus, with weight reaches 200 kg, as estimated from its me the type-species P. gaudryi. However, later au tapodials). Its cheek-teeth are hypsodont for a thors have been reluctant to accept this new Miocene bovid, with posttrite walls with rela genus, and most of them kept gaudryi as a spe tively weak relief. The horns are large, present in cies of Tragoportax (Moya-Sola, 1983; Bouv both sexes but with strong sexual dimorphism in rain, 1988, 1994). shape and size. The species is abundant - one Solounias (1981) had divided, correctly in our of the best represented species in the tapho opinion, the "Tragoportax" complex into two coenosis of the locality (its remains are more distinct genera: Miotragocerus Stromer (where than three times more abundant in the tapho he places the type of "Pikermicerus" gaudryi coenosis than M . (Pikermicerus) Spassov, 2002). Kretzoi) and Tragoportax Pilgrim. However, he According to the general conceptions exist used an unsuitable criterion for the separation ing till the recent discoveries, Tragoportax s. of the two genera, the "demarcation" of the str. has very strong sexual dimorphism (with horn core ("steps" after several other authors), a hornless females), stronger than the sexual di feature that is in fact present in both forms. As a morphism in M. (Pikermicerus) (see Bouvrain, result, the contents of his two genera are, in our 1994). Our data from Hadjidimovo show that opinion, incorrect. The rich and well-preserved at least in one of the Tragoportax species the new material from Hadjidimovo offered us the females possess horns. Romaggi (1987) has opportunity to confirm most of the typical fea demonstrated that in M. pannoniae and M . (P.) tures of the small form usually called "T." gaudryi andancensis ( = Graecoryx andancensis) gaudryi but also to identify a number of other the females have very weak horn cores and the morphological features setting this form apart sexual dimorphism is pronounced. Consider from Tragoportax s.str. Some of them are lo able sexual dimorphism demonstrate also the cated in the most conservative part of the skull, M. (Pikermicerus) sp. from Nikiti-1, Greece the neurocranium, and this supports the ge (see T. gaudryi in: Kostopoulos, Koufos, 1996). neric distinction proposed by Kretzoi. Thus, the Our data from Hadjidimovo show that the fe forms usually included in Tragoportax belong males of M. (Pikermicerus) gaudryi gaudryi in fact to two distinct genera (Spassov, Geraads, could also have very graceful horns. The in press). As noted above, the type material of sexual dimorphism in the horns of M. (Piker the type-species of M. (Miotragocerus) is at micerus) is probably weaker than in some present too incomplete to permit a good distinc Tragoportax species (hornless female individu tion from M. (Pikermicerus). Both subgenera als are known at least from Samos) but clearly have compressed horn-cores with a more or less pronounced, and more expressed than in mod oval basal cross-section. All these bovids stand ern highly gregarious animal of the open clearly apart from the group centred on spaces (see Janis, 1986). This particularity of Tragoportax amalthea, i.e. from Tragoportax, a M . (Pikermicerus) in combination with rela genus based on T. salmontanus. In our concep tively small (medium) body size, tooth mor tion, the genus Tragoportax includes the usually phology and relative rarity in the tapho large Turolian forms with marked fronto-pari coenosis could indicate relatively wooded etal postcornual d.epression and grooved basi habitat preferences· for this species. The rela occipital as well as with horns long and clearly tively strong horn sexual dimorphism in Trago triangular in section (Fig. 1, 2). portax characterizes this form as a modt?rately Morphofunctional characteristics, sexual di gregarious (Geist, 197 4) and partially territo morphism and ecology of Miotragocerus (Piker- rial animal with developed visual display sig- -i Geologica Balcanica, 3-4/2004 105 nals, probable one-male dominance in the per Miocene localities of vertebrate fauna. In herds, and possible herds of male individuals. fact under the name Kalimantsi refers to a 2 The combination of all above mentioned fea large area (- 6 km ) of Upper Miocene fossilif tures of this bovid indicates, following the erous deposits with more than I 0 known points Jarman principle (Janis, I986) that Trago (localities) of fossil mammalian fauna. This portax ecological niche was related to more fauna is best known by the classic monograph open spaces than the M. (Pikermicerus) habi of Bakalov and Nikolov (EaKaJIOB, HHKOJIOB, tats in probable mosaic landscapes. In this re 1962). The biochronology of the fauna from spect, it is comparable to modern Hippotragus. the localities was based until recently mostly on the latest papers of Nikolov (Nikolov, 1985) related to the analyses of the data known till The Boselaphini the 80s from the "classic" localities of the from Kocherinovo-1 Kalimantsi area as: Kalimantsi I (the old ex cavations at the river bank), Kalimantsi 2 ( The site of Kocherinovo-1 Peshternik), Kalimantsi 3 (Prehvarloka) and The site is discovered by I. Nikolov in the early Kalimantsi 4 (Beliovski pat) (U:aHKOB et al., in 70s in the green clays of Gradishte region be press). The excavations in the late 70s and the tween the villages of Kocherinovo and Mursa 80s organized by one of us (D.K.) led to the dis levo (SW Bulgaria). The very recent investiga covery of a rich fossil material as well as new tions consider the sites Kocherinovo I and 2 as Kalimantsi (KAL) localities as KAL-Pehtsata, localities of the newly established Gradishte KAL-Bukovets, KAL-Bardovski pat, KAL lithocomplex (Gradishte bench- mark group point "Atso" etc. of strata) (Tzankov et al., in press). The pro An opinion prevailed until recently (Kojum posed age of both localities is early Turolian djieva et al., I982) that the lower fossiliferous (Spassov et al., 2004; Cnacos et al., in press. level in the Kalimantsi area (KAL-I) is of Vallesian age. The very recent revision of the Tragoportax sp. biostratigraphy of the Kalimantsi localities Mate rial, description and comparison. A skull demonstrate that all Kalimantsi sites (local fragment (FM 2024 N.M.N.H.- Sofia) with ized in more than IOO m thick deposits) are of part of the postcornual fronto-parietal sur Turolian age and that there is no evidence of face, part of the left maxillary and the bases of any Vallesian fauna in the Kalimantsi area. horn-cores as well as fragments of attached KAL-I, which seems to belong to the newly left and right mandible, probably from the established Gradishte genetic lithocomplex, is same adult specimen (Pl. 2.4). The preserved more likely to yield a fauna of early Turolian part of the postcornual surface indicate simi age, while other Kalimantsi faunas (referred to larities with the characteristic morphology of the Strumyani lithocomplex) are probably of Tragoportax (see the diagnosis of the genus). Middle Turolian age. (U:aHKOB et al., in press; The dimensions of the horn core cross-section Spassov et al., in press; Spassov et al., 2004). at the base are close to the female specimens of Tragoportax Pilgrim, 1937 T. rugosifrons from Hadjidimovo (Table 4a). As we have noted, the females of this species, pos Bakalov and Nikolov (I962) (EaKaJioB, HHKO sibly in contrast to some other Tragoportax JIOB, I962) have described from Kalimantsi species (see discussion in Spassov and Ge and Gorna Sushitsa (without mentioning raads, in press) were not hornless. However, which part of the material belongs to G. the dimensions of the preserved teeth ( L m3 = Sushitsa site) some maxillary and mandibular 23-24 mm) are clearly smaller than the dimen fragments with teeth and postcranial remains sions of the males ofT. rugosifrons. (partially stored in the Univ. of Sofia) as Tragoportax amalthea. Nikolov (I985) notes the presence of the species at Kalimantsi The Boselaphini Peshternik without description of the material. from Kalimantsi At least part of this material could indeed be long to Tragoportax (sensu our taxonomic The site of Kalimantsi concept - Spassov, Geraads, in press), from the tooth dimensions and premolar morphol The fossiliferous site of the Kalimantsi village ogy (see EaKaJioB, HHKOJI OB, I962, Pl. is the richest one in the basin of Middle Struma XXXIV-I and XXXV-I; and p. 80, specimens river and the best known of the Bulgarian Up- I-2, 4). On the other hand, it will be hardly I06 possible, because of the scarcity of the mate General conclusions rial, to determine the species with certainty. New Kalimantsi material in the Assenovgrad At present, at least three Bulgarian Upper Mi Paleontological Museum. Several skulls and ocene localities document the genus Trago skull fragments in poor condition of fossiliza portax: Hadjidimovo-1 (MN1lfl2 boundary) tion: K 759, 55601, 55602, 55604, 55605, - T. rugosifrons, Kocherinovo-1 (MN1l) - 55606, 55610, 55603 (Pl. 3; Table 1), maxillary Tragoportax sp. and Kalimantsi fossiliferous fragments with teeth: K 3725, 3727, 3728, area (the Kalimantsi sites from the MN12 3731, 3742, 3768 (Table 1): All the material is zone)- most possibly T. amalthea. In the latter most probably from the MN12 zone of the sites, it coexists with M. (Pikermicerus) gaudryi. Kalimantsi fossiliferous area; exact localities The presence of T. rugosifrons is probable in unknown. one of the MN12 zone Kalimantsi localities. Description and discussion. All skulls dis Our conclusions on the ecology and distri play Tragoportax features such as triangular bution of Tragoportax rugosifrons and M. horn core shape, depressed fronto-parietal (Pikermicerus) gaudryi confirm those of postcornual region, and (when preserved) Bouvrain (1994) that T. rugosifrons is mostly a basioccipital with longitudinal grove (fig. 1- grazer (or perhaps a mixed feeder; Solounias et 2) and almost rectangular intercornual fron al., 1999), and in any case a species of more tal surface. Compared to T. rugosifrons and open environment than M . (P.) gaudryi which T. amalthea almost all skulls have (if we inhabits more woody biotopes. However, the could judge after the poorly preserved mate occurrence of both species at Hadjidimovo rial): generally large horn core cross-sec (and possibly in Kalimantsi) demonstrate, in tion; relatively small intercornual frontal contrast to previous opinions, that Tragoportax surface; in most of the skulls well expressed and M. (Pikermicerus) can co-exist in East Eu rugosities of the anterior horn core crest, rope, where both genera were present. It is reaching in some cases the frontal bone in likely, therefore, that they co-existed also in front of the horns (Pl. 3.3, 3.5). These char Pikermi, as they did in Halmyropotamos, Veles acters are closer to the T. amalthea condi - Karaslari, Maraga and probably Nikiti - 2 tions and we could refer all these material to and Maramena (Spassov & Geraads, in press). Tragoportax cf. amalthea. One of the skulls After Solounias et al. ( 1999) the Turolian (No. K 55603) (Pl. 3.4) has relatively large Pikermian biome of the "Greco-Iranian" prov intercornual distance at the base of the ince were dominated by relatively homogenous horns and demonstrate more affinity with T. woodlands. According to another recently ex rugosifrons. This similarity might be related pressed opinion, the Pikermian biome and es to individual variations. On the other hand, pecially the landscape from Hadjidimovo was we need to note again that the exact locali rather represented by forest-savanna-like (in ties of the noted Kalimantsi Tragoportax terms of . physiognomy of the landscape) material is not known. Most probably the biotopes and not by hardly penetrable dense skulls were found in different local fossilif thickets and woods (Spassov, 2002). The above erous points. The last mentioned skull (that mentioned ethological interpretations of the we refer to T. ? rugos1jros) could originate morphological characteristics of M . (P.) gaudryi from a separate (lower?) local site. and T. rugosifrons support mostly the second Miotragocerus (Pikermicerus) opinion. If the occurrence of the two discussed gaudryi cf. M. (P.) bovids in one and the same stratigraphic level is gaudryi gaudryi Kretzoi, 1941 not proven in Pikermi (Bouvrain, 1994) it is not the case in Hadjidimovo, where they are both Material, description and comparison. (col!. of elements of the same taphocoenosis. They co the Assenovgrad Paleontological Museum). existed in the Hadjidimovo zoocoenosis occu Maxillaries with teeth: K 3730, 3745, 3748, pying different niches: more woody habitats for 3753, 3757, 3761, 3772; two frontlets (K 55608, M. (P.) gaudryi and more open habitats for T. K 55609) belong probably to juv.-subad. indi rugosifrons in a mosaic landscape. In both viduals of this species. forms females possess horns, but demonstrate The tooth features and measurements (see clear sexual dimorphism. Table 1) correspond to the noted description One of the latest representatives of Trago of M. gaudryi from Hadjidimovo (see above) portax , T. macedoniensis, could be a relic form morphology and dimensions. of the wooded habitats, or probably documents 107 a neotenic reversal. Living in woody local con ben.- Geologica Balkanika, 12, 3, 69-81. Korotkevich, 0. 1982. 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