141

Greek bovids through time*

Dimitris S. Kostopoulos1 1 University of Thessaloniki, Department of Geology, 54124 Thessaloniki, Greece. e-mail: [email protected].

ABSTRACT: The Greek bovid record covers a time span of about 17Ma, from the early Middle Miocene to the Holocene and includes more than 70 species distributed in 5 tribes. A brief presentation of the Greek through time is given, focus on the taxonomic, phylogenetic and chronological aspects and questions that arise from their study. present a general increasing tendency from Miocene to Pleistocene supported by the gradual decrease of Boselaphini after middle Turolian. Caprinis . l . and their allies are always important but from the beginning of Pleistocene increase rapidly. predominate during Late Miocene but strongly decline at the end of Pliocene and finally disappear during Pleistocene. Key-words: Neogene, Quaternary, Bovidae, Greece.

ΠEPIΛHΨH: Το ελληνικό αρχείο των βοοειδών καλύπτει µια χρονική περίοδο περίπου 17Ma, από το κατώτερο Μέσο Μειόκαινο έως το Ολόκαινο και περιλαµβάνει περισσότερα από 70 διακριτά είδη διανεµηµένα σε 5 φυλές. Στο παρόν άρθρο δίνεται µια συνοπτική παρουσίαση των ελληνικών Bovidae διαµέσου του χρόνου, επικεντρώνοντας το ενδιαφέρον στις ταξονοµικές, φυλογενετικές και χρονολογικές όψεις αλλά και στα ερωτήµατα που προκύπτουν από τη µελέτη τους. Από το Άνω Μειόκαινο στο Πλειστόκαινο τα Bovini παρουσιάζουν µια γενική αυξητική τάση, η οποία υποστηρίζεται από την προοδευτική µείωση των Boselaphini µετά το µέσο Τουρόλιο. Η παρουσία των Caprini s . l . και των συγγενών τους είναι πάντα σηµαντική αλλά από την έναρξη του Πλειστοκαίνου αυξάνονται γρήγορα. Τα Antilopini υπερισχύουν κατά τη διάρκεια του Αν. Μειοκαίνου αλλά ελαττώνονται έντονα στο τέλος του Πλειοκαίνου και τελικά εξαφανίζονται στο Πλειστόκαινο. Λέξεις-κλειδιά: Nεογενές, Tεταρτογενές, Bovidae, Eλλάδα.

I N T R O D U C T I O N delicate process, especially in supra-specific level. Due to the high diversification into the family, the usual absence of Bovidae is certainly one of the most fascinating direct correlation between living and fossil bovid lineages families not only because of its complicated taxonomic and the scarcity of the recent “out of Africa” bovid record in structure, involving a plethora of living and fossil forms but contradiction to the past, the classification of several fossil also because of its multifarious evolutionary history expan- and living bovid taxa is far from being completely resolved ded into an extremely wide geographic domain and a great (see for example discussions about the late Miocene variety of natural environments. Hence, the study of fossil C r i o t h e r i u m in GE N T R Y (1971, 2000a), GE N T R Y et al. (1999) bovid associations could provide key information for the or about the living Aepyceros in GE N T R Y (1992), GA T E Z Y e t resolution of biochronological, paleoecological and paleo- a l. (1997). geographical problems. Evidently, the study of such a multidimensional problem, The aim of this work is to expose as briefly as possible the applying different theories and methodologies on fully richness and diversity of the Greek bovid association through distinct or partially comparable data sources (paleontological, time. Important information about taxonomical and zoological or molecular) could provide various taxonomic evolutionary problems concerning the Greek Bovidae, as well schemes. On the other hand, the implication of extinct taxa in as basic data on their chronological distribution is also the restoration of the Bovidae phylogeny is an essential and provided. As the bibliography on this topic is extremely large desirable approach but it can increase the vulnerability of the in order to be completely exposed in the frame of this article, produced reconstructions and evolutionary scenarios (RE I F, only the most fundamental and/or the more recent works 2 0 0 3) . have been spotlighted. Bovidae take part of the monophyletic Pecora infraor- der, which appears to be separated from rest artiodactyls at BOVIDAE SYSTEMATICS AND EVOLUTION Eocene-Oligocene times. The phylogenetic relation of the family into Pecora is rather unclear as molecular analyses Definitely, Bovidae systematics is an intricate and quite give contradicting results indicating either Bovidae as the

* Eλληνικά Bοοειδή διαµέσου του χρόνου. 142 Dimitris S. Kostopoulos sister group of the American on the basis of family. The discovery of the Axios valley fossil sites and their advanced selenodonty and sheathed horns or as the sister exploration during ’70s and ’80 brought to the light several group of Cervidae- or Cervidae clade (JANIS & new bovid taxa, which study strongly influenced the existed SCOTT, 1987; GENTRY &HOOKER, 1988; BEINTEMA et al., evolutionary scenarios. The general interest for the Greek 2003). Early authors already mentioned a basic dichotomy faunas forces the following paleontological research to the of the family in two primary subclades, one with ox-like exploration of several older and younger fossil assemblages, dentition (the so called “Böodontia”) and another one with enlarging the available bovid record and resuscitating the -like dentition (“Aegodontia”). This basic taxonomic discussion on Bovidae evolution and radiation. Today, well division, restored and redefined by many later authors (e.g. known bovid assemblages are recognized from numerous GENTRY, 1992; V RBA &SCHALLER, 2000), largely Greek localities ranging from early Miocene to the late corresponds to the proposal of HASSANIN &DOUZERY Pleistocene. Thus, Greek bovids form one of the best (1999) in classifying living Bovidae in two subfamilies, documented and more complete European mammal records. and . The number and status of supra- generic taxa (mainly tribes) included in each subfamily is Early bovids still open to discussion (e.g. GENTRY, 1992; HASSANIN & DOUZERY, 1999). In a general way, Bovinae group together The current knowledge on earliest bovids is quite limited due the tribes Bovini, Boselaphini and , while mainly to the restricted data sources. E o t r a g u s PI L G R I M , 1939 Antilopinae are divided in two basic tribes Antilopini and is considered by most authors as the first true bovid, appeared Caprini s.l. and their allies. Apart from isolated living and more or less simultaneously in Europe and Asia at about 18 fossil bovid taxa, these classificatory schemes are generally Ma ago (GE N T R Y et al., 1999). Whether the origination of supported by both molecular and paleontological evidences E o t r a g u s predates or postdates the first split into the family and they are widely acceptable (e.g. GA T E Z Y et al., 1997; is, however, a point of scientific controversy; several authors VRBA &SCHALLER, 2000; GENTRY, 2000a). relate the with the basis of Boselaphini lineage(s), while The emergence of the family is placed in Eurasia and it some others still argue about its placing among the Bovinae. is dated by most authors at about 19 Ma ago, a period of In terms of European Mammal Chronology, E o t r a g u s major faunal interchanges with the Afro-Arabian continent appeared in W-C Europe from MN 4 to MN 6, while it is also (GENTRY, 1992, 2000b; VRBA &SCHALLER, 2000 and k n o w n from North and East Africa, Israel, Pakistan and literature therein). As a result of continental separation China. E o t r a g u s was unknown from SE Europe but the during early Miocene (19-17 Ma), the Antilopini (or part of discovery of Agios Antonios fissure-filling faunule, in them) developed in Africa, while the Bovinae remained in Chalkidiki peninsula filled this gap. KO U F O S &SY R I D E S Eurasia, documenting the first major split into the family (1997) refer to this genus several horn-cores from this (GENTRY, 1992; HASSANIN &DOUZERY 1999; VRBA & locality, which is dated in the upper part of MN4/lower part SCHALLER, 2000; ROPIQUET &HASSANIN, 2005). The of MN5. Although the material is still under study, the global warming of the middle Miocene (16-13.5 Ma) presence of the genus is confirmed but the species constrained a subsequent explosive radiation of tribes determination should wait for more precise data. associated by a second important phase of faunal T e t h y t r a g u s AZ A N Z A &MO R A L E S, 1994 definitely interchanges between Eurasia and Africa (VRBA &SCHAL- belongs to the earlier non-boselaphine bovids of the Old LER, 2000). Later on, the expansion of the grassland World, together with the Indian C a p r o t r a g o i d e s and the environments at the end of Miocene (7-5 Ma) assisted the Afro-Arabian G e n t r y t r a g u s.It is known from MN6 to MN8 evolution of open-habitat bovids, marking the next phase on faunas of Western and Central Europe, as well as from the Bovidae evolution, during which several modern groups Turkey. T e t h y t r a g u s horn-core pattern shows advance appeared (e.g. tragelaphines, bovines, reduncines, characters found in several later bovids (GENTRY et al., 1999; aepycerotines). During Plio-Pleistocene most “archaic” GE R A A D S, 2003). GENTRY (2000a, b) detect in T e t h y t r a g u s groups follow a rather in situ evolution, droved by global or in a form close to it the possible forerunner of Caprini but and local environmental changes, while some new bovid other non-boselaphine bovid groups may also originate from clades also appeared (e.g. rupicaprines), allowing to the the same stock. A mandible referred to T. koehleri o r i g i n a l l y mid-late Pleistocene “modernization” of the bovid known from Turkey has been recently found in the early association. Middle Miocene (MN 5) fauna of Thymiana, Chios island (D E BO N I S et al., 1998). Although insufficient for certain The Greek bovid record conclusion, the Chios form fits pretty well with that of T . k o e h l e r i from locality 3 of ²andir (GE R A A D S, 2003). The study of the Greek bovid association goes far back to the The highly specialized H y p s o d o n t u s SO K O L O V, 1949 is late 19th century and it is firmly related to the Pikermi and central to a great discussion and according to several authors Samos faunas. The study of their wealthy bovid association it could belong to a separate bovid clade of tribal or enriches the current knowledge and strongly involves to the subfamily level, which may be biphyletic to other Bovidae reconstruction theories of the evolutionary history of the (GENTRY & HE I Z M A N N, 1996; GENTRY et al., 1999). H y p s o - Hellenic Journal of Geosciences, vol. 41, 141-152 143 144 Dimitris S. Kostopoulos

Fig. 1. Continue Hellenic Journal of Geosciences, vol. 41, 141-152 145 d o n t u s is present in several MN5 to MN 6 Asian faunas of this material is quite questionable. extended westwards to Georgia, Turkey, Slovenia and ex T r a g o p o r t a x appears with two chronologically succeeded Yugoslavia. In Greece, it has been described from the MN 5 species. Tr. rugosifrons ( =Tr. recticornis =? Tr. curvcornis)is Thymiana fauna by D E BO N I S et al. (1998), who ascribe the a form close to the Indian relatives and it appears at the material to H . cf. g a o p e n s e, suggesting interesting beginning of Turolian, predominating in the MN11 faunas. similarities to the Gaopo form from China. Anyway, the Although the chronostratigraphic position of the type material synonymy among several species and genera of this group is of Tr. rugosifrons from Samos is uncertain, it seems more not yet decided. reliable that it comes from the lower fossiliferous levels. During MN12, the species is superseded by Tr. amalthea b u t whether this replacement is a result of evolution or Subfamily BOVINAE immigration is not clear enough. Here again, the Maramena fauna suggests the persistence of Tr. amalthea until the end of Boselaphini MN13 but the data are very scarce for certain conclusion; in Boselaphines were a successful Eurasian bovid group that fact, the Maramena form could be also related to an early appeared during late Middle Miocene and strongly radiated bovine. and expanded during Late Miocene from China to Spain and Africa. Their emergence is dated before 17.5 Ma in Pakistan, Tragelaphini suggesting an Asian origin probably from an E o t r ag u s- l i k e ancestor (GENTRY, 1992 and literature therein). As a result of For a long time several Eurasian spiral-horned of their great diversification, Miocene boselaphines present Miocene and Plio-Pleistocene age have been considered as quite complicate systematic that frequently suffers from belonging to tragelaphines. Nonetheless, GENTRY (1971 and nomenclature problems and controversies both at genus and literature listed) showed than most of these forms are species level. certainly —or more closely related to the Antilopini clade Boselaphines appeared in Central-Western Europe during and have nothing to do with the -so considered— strictly MN7-MN8 but their earliest Greek record is dated quite later, African Tragelaphini . This option has been adopted by coming from the late Vallesian (MN 10) of Macedonia most authors and the recent literature restricts the expansion (BO U V R A I N , 1997). Then after, two genera appear: and evolution of Tragelaphini into Africa (e.g. GE N T RY,1992; M i o t r a g o c e r u s ST R O M E R, 1928 of probably European or VR B A &SC H A L L E R, 2000). In contrast, a newly discovered Eurasian origin, expanded south-eastwards during Turolian, bovid from the late Miocene (early MN 12) of Perivolaki, when boselaphines become much more abundant and Thessaly shows clear relationships to the African tribe T r a g o p o r t a x PI L G R I M, 1939, a probably Asian immigrant. A (KO S T O P O U L O S &KO U F O S, in press). Cladistic analysis great number of species has been ascribed to each of them showed that the Perivolaki form could be considered as a late without however a sharp distinction at generic level; hence, Miocene tragelaphine stem taxon, suggesting a common several species interchange between T r a g o p o r t a x a n d origin from a Eurasian boselaphine ancestor (KO S T O P O U L O S M i o t r a g o c e r u s increasing confusion. The coexistence of these &KO U F O S, in press). This point of view strengthens two genera in most of the Turolian faunas of the Balkans and HAS S A N I N &DO U Z E R Y ’ S (1999) suggestion that “…t r a g e l a - neighboring areas in combination with the insufficiency of the phines migrated in Africa where they have been distinct for at fossil record set off a great discussion about their distinction least 15 Ma…”. (SO L O U N I A S, 1981; SP A S S O V &GE R A A D S, 2004; KO S T OP O U- L O S, 2005). Bovini The undetermined Pentalophos form, documented by a few lower toothrows is quite similar to M i o t r a g o c e r u s c f . At about 6-5 Ma ago the Bovini spit off from a boselaphine p a n n o n i a e from the latest Vallesian (end MN 10) locality of ancestral stock on the Indian subcontinent and then they are Nikiti-1 (KOS T OP O U L O S, 2005). SO L O U N I A S (1981) mention expanded (separately?) to Africa and Eurasia, invading into the presence of monacensis in Samos and N. America too (GE R A A D S, 1992; GENTRY &HE I Z M A N N, Pikermi but the distinction of the referred material from 1996). Miotragocerus valenciennesi ( =Graecoryx valenciennesi= Fossil bovines are scarcely known from the Greek fossil T r agoportax gaudryi s e n s u KO S T OPOULOS, in press) is not record. Samokeros minotaurus SO L O U N I A S, 1981 is a clear enough and I suggest that both the Pikermi and Samos fascinating late Miocene bovid from Samos, which already forms belong to the latter species, which is present in a great shows several advanced bovine features and according to number of MN11-MN12 faunas of Greece, Bulgaria, Turkey GE N T R Y &HE I Z M A N N (1996) it could be placed into Bovini. and Ukraine. Later on (MN13) M. macedoniensis replace M . S a m o k e r o s is probably originated from a T r a g o p o r t a x s t o c k v a l e n c i e n n e s i in Northern Greece at least; this species shows and it seems also to be related with some of the early Pliocene some peculiarities that could suggest endemism. M i o t r a - large bovids referred to P a r a b o s AR A M B O U R G &PI V E T E A U, g o c e r u s probably persists in Greece until the end of MN13, 1929 and A l e p h i s GR O M O L A R D, 1980. Although recently referred from the Maramena fauna but the species definition reviewed, the systematic of P a r a b o s and A l e p h i s p r e s e n t s 146 Dimitris S. Kostopoulos Hellenic Journal of Geosciences, vol. 41, 141-152 147 significant incongruities that need further discussion. P a r a b o s type material from Samos. G. capricornis appears perhaps as is also referred from the early Pliocene locality of Megalo early as G. pilgrimi but it prevails during MN 12. Its Emvolo (P. macedoniae) but the identification of the material systematic is not fully defined; several authors consider G . is still open (e.g., SO L O U N I A S, 1981). c a p r i c o r n i s to be synonym of the West European G . The European early Pliocene bovines are rather non- d e p e r d i t a, while there are suspicions that the material of G . c a p r i c o r n i s from Pikermi could include another species close existent. During early Middle Pliocene, the non- or identical to G. mytilinii from Samos (KO S T O P O U L O S, monophyletic L e p t o b o s PI L G R I M, 1939 invades S. Europe 2005). G. deperdita is certainly known from the late Turolian from Asia together with E q u u s and E l e p h a s marking the (MN 13) of Greece and G a z e l l a sp. from Maramena is most beginning of Villafranchian (the so called “E-L-E dispersal likely related to this species. G. deperdita could be also event”). The earliest occurrences of L e p t o b o s in Europe are present in some earlier faunas, while the possibility to be coming from Dusino (MN 16 Italy) and Damatria (MN16 originated from G. capricornis cannot be excluded. Greece; KO U F O S &KO S T O P O U L O S, 1997) but the genus was The climatic changes at the dawn of Pliocene allow never well established in the Balkans and its presence is to decline dramatically in the entire Europe. G a z e l l a rather sporadic. At the end of Pliocene, the widespread West b o r b o n i c a DE P E R E T, 1884 is the single species occurred in S. European species Leptobos etruscus occupies Greece but Europe from Ruscinian to middle Villafranchian. The short later it is replaced by more advanced bovines westward prolongation of the late Pliocene aridification forc- representing early . Whether these primitive bisons ed, however, new Asiatic immigrants to penetrate SE Europe; (“Leptobos” vallisarni, Eobison) are coming from evolution Gazella bouvrainae KO S T O P O U L O S, 1996 a species related to ofL e p t o b o s or by migration is not yet clear. True bisons (e.g. the earlier G. emilii BO U V R A I N, 1998 from Turkey appears at priscus) certainly occur at the Middle Pleistocene that time in Greece and gradually replaces G. borbonica u n t i l together with oxes (e.g primigenius), both representing the beginning of Pleistocene when gazelles definitely northern immigrants escaped southwards probably because disappear from the area. The contemporaneous Gazella aegea of the Pleistocene climatic deterioration. AT H A N A S S I O U, 2002 is more questionable, indicating relations to other genera. Subfamily ANTILOPINAE Heteronymous Spiral horned antelopes Antilopini The origin of this bovid group remains doubtful since In a broad sense, Antilopini could be divided into the gazelles between the definite non-boselaphine ancestral stock of MN and their allies and the heteronymous spiral horned antelopes. 5-6 and the first evidences of anticlockwise spiral horned A third group usually referred to Antilopini includes several antelopes at Middle Sinap (MN 9; GE N T RY, 2003) there is a fossil genera of clockwise (homonymous) spiral-horned time gap of about 3Ma. Nevertheless, from late Vallesian bovids mainly of Miocene age. Nonetheless, it is not clear if onwards, anticlockwise spiral-horned antelopes present a this group certainly originates from the same stock with strong radiation, allowing a great variety of forms, most of anticlockwise forms. In the hypothesis of a separate ancestry them well expressed on the Greek record. between them, Hypsodontinae KO E H L E R, 1987 could have P r o s t r e p s i c e r o s FO R S Y T H- M A J O R, 1891 is unquestionably crucial systematic interest. the most successful and diversified late Miocene genus of this group. The primitive Pr. vallensiensis is the oldest known Gazelles species appeared in the MN9-MN10 faunas of Greece and Although the African Antilopini H o m o i o d o r c a s from the late Turkey and it could be phylogenetically related to P r . Miocene of Ngorora (Kenya) could be related to the ancestry v i n a y a k i. Slightly later, the larger but less systematically of gazelles (GE N T RY et al., 1999) the origin of the group and informative Pr. syridisi also occurs in N. Greece. During early its classification are still unresolved. Fossil gazelles are in- Turolian the small Pr . ax i o s i KO S T O P O U L O S, 2004 predo- discriminately referred to the genus G a z e l l a , while several minates associated by a primitive form related to P r . efforts to recognize supra-species taxa did not allow r o t u n d i c o r n i s; the latter species will be fully established generally acceptable results. during the middle Turolian. At that time, in the east margins Earliest G a z e l l a are represented by small forms with of the Aegean region Pr. axiosi seems to give origin to P r . conical horns and they are known from Algeria (G . z i t e l l i . In contrast to the current ideas, it seems quite possible p r a e g a u d r y i; MN9/10) and Turkey (G. ancyrensis;MN that two distinct lineages are included into P r o s t r e p s i c e r o s: 9/10). Late Vallesian gazelles are still very rare and the first originated from Sinapodorcas incarinatus BO U V R A I N insufficiently known. During this time interval the first et al., 1994 and allow to r o t u n d i c o r n i s - z i t e l l i group, while the G a z e l l a-like form also appeared in N. Greece (BO U V R A I N , 1997). From the beginning of Turolian G a z e l l a presents an second arrive from P r o s t r e p s i c e r o s sp. of Sinap and P r. explosive radiation and geographic expansion. Among the v a l l e n s i e n s i s from Greece and let to v i n a y a k i - l y b i c u s group. well established and known species, the long horned G . N i s i d o r c a s BO U V R A I N, 1979 is another important p i l g r i m i seems to predominate in Greece during MN11, but Antilopini genus with apparent primitive morphology there are still questions about the stratigraphic source of the persisting in time. Its origin should be searched among or 148 Dimitris S. Kostopoulos close to the MN9 forms referred to M a j o r e a s KO S T O P O U L O S, the late Turolian (MN 13) of Greece expanding both the 2004. N i s i d o r c a s appears in Greece at the very beginning of chronological and geographical distribution of the genus. Turolian with N. planicornis, which characterizes the MN11 Moreover GENTRY et al. (1999) recognized in several faunas. The species does not reach south-eastern Greece Ukrainian samples of MN 11 age referred to G a z e l l a a where it is probably replaced by Majoreas woodwardi. Both possible relative of H i s p a n o d o r c a s. A form close to H . disappear little after the dawn of MN12. o r i e n t a l i s seems to be also present in the latest Vallesian Similarly to the gazelles, the anticlockwise spiral horned locality of Nikiti-1 (= partimO i o c e r o s cf.a t r o p a t e n e s in KO- antelopes strongly decline during the early Pliocene; none S T O P O U L O S &KO U F O S, 1996), resuscitating the discussion genus is known from Greece during this period. At middle on the genus origin and chronology. Villafranchian the cosmopolitan G a z e l l o s p i r a PI L G R I M & S a m o d o r c a s BO U V R A I N & BO N I S, 1985 is another peculiar SC H A U B, 1939 invade into Greece with G. torticornis,which genus representing medium sized clockwise spiral horned persists until the beginning of Pleistocene. The genus presents of late Miocene age. It is known by a single specimen close affinities with several Asian forms and especially from Samos referred to the species S. kuhlmani (SO L O U N I A S, Parastrepsiceros sokolovi VE K U A, 1968 of MN 16 and its 1981). The morphological characters of this form are quite systematic needs further study. At the end of Pliocene an distinct from all the above mentioned genera and its enigmatic form appears in Northern Greece: Antilope kou - phylogenetic relations are still open. f o s a e, previously ascribed to P a r a s t r e p s i c e r o s (KO S T O P O U L O S During Pliocene clockwise spiral horned antelopes seem to &KO U F O S, in press). This species presents strong similarities disappear from SE Europe. At the beginning of Pleistocene the with the living (Antilope cervicapra) and it probably new genus P o n t o c e r o s VE R E S T C H A G I N et al., 1969 appeared in belongs to its forerunners. On the other hand, the similarities Greece. The genus has been originally referred to tragelaphines between A. koufosae and the earlier P r o s t r e p s i c e r o s could be (CR E G U T- B O N N O U R E &GU E R I N, 1996) but both the skull and of great phylogenetic interest. the limb-bone morphology are opposed to this conclusion, sug- gesting closer affinities to the caprines. The chronological range of the single known species P . amb i g u u s covers the entire early Homonymous Spiral Horned Antelopes Pleistocene of Greece, and somewhat later it still exists in the Clockwise spiral-horned antelopes appear as early as Middle north Black-See coasts, while its presence in Syria and Spain Miocene (e.g. T u r c o c e r u s KO E H L E R, 1987) but their remains speculative. taxonomy is still ambiguous due to their relative rareness (e.g. BO U V R A I N &BO N I S, 1985, GENTRY et al., 1999, Caprini and relatives RO U S S I A K I S, 2003) and I would like to keep at the moment the classification proposed by BO U V R A I N &BO N I S (1985) as Caprini (= s e n s u GENTRY, 1992) and their relatives more reliable. The earliest evidence of such a bovid in Greece (hippotragines, alcelaphines) is an extremely diversified is S a m o t r a g u s BO U V R A I N &BO N I S, 1985 known from the group including several bovid branches of probably different late Vallesian (MN 10) localities of Axios valley, where it is origin that makes their supra-generic classification quite deli- represented by the small and quite primitive form S . cate. The origin of the tribe is dated back to the early-middle p r a e c u r s o r. The chronological range of this species is very Miocene and although Caprini and their allies could descend short; its final occurrence is rather coming from the latest from the T e t h y t r a g u s - G e n t r y t r a g u s - C a p r o t r a g o i d e s c o m p l e x Vallesian locality of Nikiti-1 (=partim O i o c e r o s c f . (GENTRY,1992;GENTRY et al., 1999; VRBA & SC H A L L E R , a t r o p a t e n e s in KO S T O P O U L O S &KO U F O S, 1996). The genus 2000) alternative solutions can be also possible for several appears later in Samos with the large S. crassicornis but the members of this group. Recent molecular analyses time gap between the two species and their phyletic relations (RO P I Q U E T &HA S S A N I N, 2005) showed that the sister-group are not easy to define. of Caprini s . l . is the clade uniting alcelaphines and O i o c e r o s GA I L L A R D, 1902 is a less well known genus of hippotragines and that apart from Caprini s . s . n e i t h e r this Late Miocene group. Although several earlier forms are Ovibovini nor Rupicaprini are monophyletic. ascribed to it, the genus appears in Greece as soon as middle Turolian (MN 12) with two more or less contemporaneous -like forms species: Oioceros rothi mainly from Pikermi and O i o c e r o s w e g n e r i from Samos; the relations between them as well as From middle-late Miocene onwards goat-like bovids p r e s e n t with S a m o t r a g u s are still open to discussion (e.g. GENTRY e t strong radiation and during late Miocene they effectively a l., 1999; RO U S S I A K I S, 2003) but I think more possible that occupy the entire eastern Mediterranean. The earliest O i o c e r o s and S a m o t r a g u s belong to quite distinct lineages evidences are coming from the MN7-9 faunas of Turkey and with O. wegneri more closely related to the latter genus. Tunisia (Protoryx enanus,“Pachytragus” solignaci)butthe Until recently H i s p a n o d o r c a s TH O M A S et al., 1982 was relations of the middle-late Miocene forms with the living considered to be a west European genus known from the ones are still debatable, see-sawing between the Eurasian Spanish middle Turolian. Later on, a new species H . C a p r a and the African and probably repre- o r i e n t a l i s BO U V R A I N &BO N I S , 1988 has been discovered in senting the ancestral stock of both lineages. The apomorphy Hellenic Journal of Geosciences, vol. 41, 141-152 149 of short metapodials shown in living Caprini s . s. is already twisted horn-cores is also central to a great debate (e.g. attested in the MN 10 Aragoral mudejar AL C A L A & SO L O U N I A S, 1981; GENTRY &HE I Z M A N N, 1996; GENTRY, MO R A L E S, 1997 from Spain giving rise to sev e r a l 2000a, b). The only known species C. argalioides is mainly evolutionary scenarios (e.g., GE N T R Y, 2000b; RO P I Q U E T & known from Samos old levels and it probably disappears at HA S S A N I N, 2005 and literature therein). the beginning of MN 12. More or less at the same time the Among the different genera referred from the late smaller Parurmiatherium rugosifrons occurs in the Greek Miocene of Greece, three seem to persist in time: P r o t o r y x faunas, persisting until the end of middle Turolian. The genus FO R S Y T H- M A J O R, 1891, Pachytragus SC H L O S S E R,1904and is considered by GENTRY (2000a, b) to be a synomym of P s e u d o t r a g u s SC H L O S S E R, 1904. An intermediate form U r m i a t h e r i u m . between the earlier P . s o l i g n a c i and the Turolian P r o t o r y x- P a l a e o r y x GA U D R Y, 1861 is probably the most successful P a c h y t r a g u s species firstly occurred in Greece during late late Miocene ovibovine-like bovid. Its earliest documentation Vallesian (BO U V R A I N, 1997). The presence of P a c h y t r a g u s is dated to the late Vallesian of Greece and the genus l a t i c e p s in the early Turolian faunas of N. Greece is still occurrences remain limited during early Turolian. P a l a e o r y x uncertain, but the species occurs at the end of this time presents a larger expansion at the middle Turolian, known by interval at Perivolaki-Thessaly as well as in Samos and two species: P. majori and P. pallasi (KO S T O P O U L O S, 2005). neighboring areas, where it is replaced during middle-late Its attribution to the ovibovines is not yet profoundly Turolian by the more advanced P a c h y t r a g u s c r a s s i c o r n i s established but it seems to be quite possible (see also (GENTRY, 1971). At the same time Protoryx carolinae GENTRY,2000b). appears in Pikermi. The closely related genus P s e u d o t r a g u s, During Pliocene ovibovines almost disappear from representing a smaller form with longer horns, associates Europe but they still exist in Asia. At the beginning of P r o t o r y x-P a c h y t r a g u s faunas in both the east and the west Pleistocene new forms invade into north Mediterranean coasts of the Aegean sea. coasts. This is the time of the first appearance of S o e r g e l i a After the middle Turolian explosion, the group shows a SC H A U B,1951andPraeovibos ST A U D I N G E R, 1908, two strong reduction. The single known Greek taxon of latest genera that will later characterize the middle Pleistocene of Miocene age which presents clear relations with P r o t o r y x i s C. Europe. A largeS o e r g e l i a species that could be forerunner N o r b e r t i a KO E H L E R et al., 1995, while the middle of the middle Pleistocene S. elisabethae occurs in Greece Villafranchian Eythyceros AT H A N A S S I O U, 2002 could be also during late early Pleistocene, associated by a primitive allied to the same bovid stock. At the end of Pliocene the new Praeovibos. Both genera show a wide expansion on the Asiatic immigrant H e m i t r a g u s invades into the Balkans from whole S. Europe but from middle Pleistocene onwards they Asia. The genus is barely known from Greece but it seems to are restricted above the Alpine chain. persist until the end of early Pleistocene. True of the Rupicaprines include today three Eurasian and one North genus C a p r a certainly occur during late middle Pleistocene, American genus. The earliest evidence of rupicaprine-like documented is several cave deposits of Greece mainly by the forms cannot be defined with accuracy but it seems quite species Capra ibex. During Holocene Capra aegagrus a l s o possible that this group appears as early as latest Miocene/ appears in Greece, being today the single living earliest Pliocene. K o u f o t r a g u s BO U V R A I N, 1996 is probably representative of this genus. the earliest record of this bovid group in Greece. It is about a medium sized form, originally referred by AR A M B O U R G & PI V E T E A U (1929) to the new species Gazella bailloudi a n d Ovibos- and Rurpicapra-like forms coming from the early Pliocene deposits of Megalo Emvolo The classification of living ovibovines and rupicaprines is (MN15). Years after, the type skull has been re-discovered in quite complicate and becomes much worse for the earlier the collections of the Natural History Museum of Paris and forms of these groups. Recent molecular data group together re-described as a new taxon allied to rupicaprines. O v i b o s with N a e m o r h e d u s (RO P I Q U E T &HA S S A N I N, 2 0 0 5 ) P r o c a m p t o c e r a s SC H A U B , 1923 is phenetically close to the d i s p r o v ing the monophyly of the relevant tribes and thus the living R u p i c a p r a, and it appears more or less simultaneously in phylogenetic displacement of several of the late Miocene S. Europe at the beginning of Villafranchian. It is poorly genera remains open to discussion. documented in Greece from some late Pliocene faunas from M e s e m b r i a c e r u s BO U V R A I N , 1975 is considered to be one the North. of the earliest ovibovines mainly because of its basicranial During middle-late Villafranchian (MN 17) G a l l o g o r a l and atlas morphology, reminding living members of this GU E R I N, 1965 invades into S. Europe probably from North- group. The genus is known by a single species M. melentisi, East. The genus could be originated from a P a l a e o r y x coming from the late Vallesian (MN 10) faunas of N. Greece ancestor. The earliest occurrence of this large bovid is dated and at the moment it does not appear to have direct relatives at about 2.5-2.0 Ma ago, when it appears to inhabit the peri- in earlier or later faunas. Pontic area. In Greece the sub-species G. meneghinii During early Turolian the genus C r i o t h e r i u m FO R S Y T H- s i c k e n b e r g i i KO S T O P O U L O S, 1998 characterizes the middle MA J O R, 1891 appears in Greece. This large sized and Villafranchian (MN 17) faunas and it seems to extend extremely specialized bovid with robust anticlockwise eastwards until the east coasts of the Black Sea. Somewhat 150 Dimitris S. Kostopoulos later a more advanced race appears in SW Europe (France, to be present until the end of Turolian but unpublished data Italy). Some authors suggest that the dwarf antelopes of the indicate that the latest forms referred to it may be significantly West Mediterranean islands may originate from this genus. different at species level. P l i o t r a g u s KR E T Z O I, 1941 is a quite problematic genus Tragoreas oryxoides SC H L O S S E R, 1904 is a very showingaffinitiestoboththelivingrupicaprinesand questionable taxon, known by a few specimens from Samos ovibovines. Although Pliotragus ardeus is rather well known but also referred from some other late Miocene localities of from the middle-late Villafranchian of W. Europe and it Greece. SO L O U N I A S (1981) extensively discuss the probably extends eastwards to the Balkans, it was never morphological relationships of this form, and although he mentioned from Greece. However, it seems that a more ad- retains its taxonomic validity, his comparisons let me suggest vanced form penetrates (?) Greece during middle Pleistocene, that it could be a fictional form based on undetermined occurring in the old horizons of the Petralona cave (CR E G U T - material of other taxa. BO N N O U R E &TS O U K A L A,2005) . R u p i c a p r a is the single living representative of this group CONCLUSIONS survived in Greece in some restricted and isolated herds of P indos and Rhodope mountain c h a i n s . The chrono-spacial distribution of the referred bovid genera and species is given in Figs. 1 and 2. Statistical elaboration of Incertae sedis forms the available data allow to some interesting but not surprising conclusions. Thus the total number of Bovidae species ever recorded in Greece is >70: 20% represents Bovinae, 35% H e l l a d o r c a s BO U V R A I N, 1997 is a late Vallesian (MN 10) Antilopini and 32% Caprini s . l . , while a 13% rests incertae genus exclusively known from Greece. The single species H . sedis. g e r a a d s i presents similarities with both P r o t r a g e l a p h u s ( s e e Bovinae appear to present a general increasing tendency below) and P r o s t r e p s i c e r o s and its systematic relationships from Miocene to Pleistocene (Fig. 3), which is mainly due to are not yet clear enough but I think it more closely related to the emergence and radiation of Bovini. Boselaphini increase the Protragelaphus lineage. toward middle Turolian and then gradually decrease until In the MN 10 faunas of N. Greece H e l l a d o r c a s i s their disappearance after MN 14 (Fig. 4). Bovini emerge associated by the genus O u z o c e r u s BO U V R A I N &BO N I S,1986 during MN12 and increase rather gradually toward MN19 known by two species, O. gracilis and O. pentalophosi, (the observed pits are certainly fictitious due to the merge of which show non-spiralled but moderately-strongly twisted MN14-15-16 and MN18-19 faunas), while from the horn-cores. The primitive characters of the O u z o c e r u s s k u l l beginning of Pleistocene they seem to rise more rapidly and horn-cores might be also seen in some MN 9 specimens toward the middle-late Pleistocene (Fig. 4). The participation of carpines is always important (Fig. 3) from Middle Sinap (MN 9, Turkey) ascribed to P a l a e o r e a s but they generally increase from Miocene to Pleistocene. This and they could satisfy the conditions for the ancestry of this general trend is interrupted by a decreasing phase at the end of genus but certain conclusions are not possible at the moment. Miocene (which could be however related to the scarcity of According to the available data O u z o c e r u s is also referred data) and a second decline at the end of Pliocene which is from the latest Miocene locality of Maramena but a fresh look probably related to climatic factors. From the beginning of on this material shows that most probably it represents a Pleistocene the tribe increases rapidly because of the arrival of P r o t r a g e l a p h u s DA M E S, 1883. BO U V R A I N (1992) states t h a t new immigrants occupying the available niches (Fig. 4). most of the apomorphic features observed in P r o t r a g e l a p h u s Antilopini predominate during Late Miocene and suggest affinities with alcelaphines or Caprini in general. P r o t r a g e l a p h u s appears in Greece at the beginning of Middle Turolian with the widespread species Pr. skouzesi,whichis replaced in the late Turolian by the larger Pr. theodori. The genus presence in the Maramena fauna prolongs its chronological distribution to the end of Miocene. P a l a e o r e a s GA U D R Y, 1861 belongs to the anticlockwise spiral-horned antelopes with tightly twisted and straight horns. Its systematic position remains unclear, referred either to Antilopini or to Caprini and therefore we shall follow GE R A A D S et al. (2003) in suggesting that it cannot be ascribed to any modern tribe. Early forms related to this genus are coming from the MN 9 faunas of Turkey, while during early Turolian (MN 11) the species Palaeoreas zouavei appears in N. Greece. Slightly later (end of MN 11) another species, Palaeoreas lindermayeri makes its first appearance and expanded rapidly, participating in most of the middle Fig. 3. % partition of Greek bovid tribes through time. Turolian mammal faunas of the area. The species is supposed Hellenic Journal of Geosciences, vol. 41, 141-152 151

Fig. 4. % partition of Greek bovid tribes per MN zone.

Pliocene but strongly decline from the beginning of SC H A L L E R ( E d s )Antelopes, and relatives, 11-25, Yale Univ. Pleistocene until their disappearance after MN20 (Figs. 3, 4). Press. Into this general trend a secondary reduction phase occurs at G E N T R Y , A. W. (2000b). Caprinae and Hippotragini (Bovidae, the beginning of Pliocene, which is more probably related to Mammalia) in the upper Miocene. I n : E. VBRA & G. the global climatic changes. S C H A L L E R ( E d s ), Antelopes, deer and relatives,65-83.Yale Univ. Press. GENTRY, A. W. (2003). Ruminantia (Artiodactyla). I n : M . REFERENCES FORTELIUS, J. KAPPELMAN, S. SEN, AND R. L. BERNOR ( E d s . ) Geology and Paleontology of the Miocene Sinap Formation, ARA M BOU RG, C. & J. P I V E T E A U ( 1929) . Les vértebr és du Pontien T u r k e y, 332-379. Columbia Univ. Press. de Salonique. Ann. Paléontologie, 18, 1-82. GENTRY, A.W. & P. HOOKER (1988). The phylogeny of the BEINTEMA, J. J., H. BREUKELMAN, J.-Y. DUBOIS & H. Artiodactyla. I n : M.J. BENTON ( E d . ) T he ph yl oge ny an d W A R M E L S (2003). Phylogeny of r uminants s ecr etory classification of the Tetrapods, v.2: M a m m a l s: Systematics ribonuclease gene sequences of (A n t i l o c a p r a Association Sp. 35 B, 235-272. a m e r i c a n a). Mol. Phyl. Evol. 26, 18-25. GENTRY, A.W. & P . H E I N Z M A N N (1996) . M iocene of B O U V R A I N,G. (1992). Antilopes à chevilles spiralées du Miocène de the Central and Eastern Tethys and Paratethys, I n : R. BERNOR, la province Grèco-Iranienne. Nouvelles diagnoses. A n n. V. FAHLBUSCH & H.-W MITTMANN (Eds) The Evolution of P a l é o n t o l o g i e,74, 43-63. western Eurasian Neogene mammal faunas, 378-391, Columbia B O U V R A I N, G. (1997). Les bovidés du Miocène supérieur de Univ. Press. Pentalophos (Macèdoine, Grèce). Muench. Geowiss. Abh. (A) 34, GENTRY, A. W., G. RÖSSNER & E .H E I N Z M A N N (1999). Suborder 5 - 2 2 . Ruminantia. I n : RÖSSNER G. & HEISSIG K. (Eds) The Miocene BOUVRAIN, G. & L. DE BONIS (1985). Le genre S a m o t r a g u s Land of Europe, 225-258, Verlag, Dr F. Pfeil. (Artiodactyla, Bovidae), une antilope du Miocène supérieur de G E R A A D S, D. (1992). Phylogenetic analysis of the tribe Bovini Grèce. Ann. Paléontologie 71(4), 257-299. (Mammalia: Artiodactyla). Zool. J. Linnean Soc., 102, 193-207. DE BONIS, L., G.D. KOUFOS & S. SEN (1998). Ruminants (Bovidae G E R A A D S, D . (2003). Ruminants, other than G ir aff idae fr om the and Tragulidae) from the middle Miocene (MN5) of the island of middle Miocene hominoid locality of Candir (Turkey). C o u r . Chios, Aegean sea (Greece). N. Jb. Geol. Palaeont. Abh.210(3), Forsch.-Inst. Senckenberg, 240, 181-199. 3s 99-420. GERAADS, D., N. SPASSOV & D.KOVACHEV (2003). P a l a e o r e a s CREGUT-BONNOURE, E. & C. G U E R I N (1996). Famille des l i n d e r m a y e r i (W A G N E R, 1848 ) ( M amma li a, Bo vid ae) f rom th e Bovidae. I n : C. GU E R I N &M.PA T O U- M A T H I S (E d s) Les grands upper Miocene of Bulgaria, and a revision of the species. mammifères Plio-Pleistocenes d’Europe, 62-106. Masson Paris G e o d i v e r s i t a s,25 (2), 405-415. CREGUT-BONNOURE, E. & E. T S O U K A L A ( 2 0 0 5) . Th e P lei st oc ene JANIS CH. & K. S C O T T 1987. The interrelationships of higher Bovids from the Petralona Cave (Macedonia, Greece): new families with special emphasis on the members of the interpretations and biogeographical implications. Q u a t e r n a i r e, Cervoidea. Am.Mus.Nov., 2893, 1-85. hors serie 2005, 2 (in press) HASSANIN, A., & E. J. P. DOUZERY (1999). The tribal radiation of GATESY, J., G. AMATO, E. VRBA, G. SCHALLER, & R. DESALLE the family Bovidae (Artiodactyla) and the evolution of the (1 9 9 7 ) . A cladistic analysis of the mitochondrial DNA from the mitochondrial cytochrome b gene. Mol. Phyl. Evol., 13, 227-243. Bovidae. Molecular Phylogenetics and Evolution, 7, 303-319. KOSTOPOULOS, D.S. (2004). – Revision of some late Miocene spiral GENTRY, A. W. (1971). The earliest goats and other antelopes from horned antelopes (Bovidae, Mammalia). N. Jb. Geol. Palaeont. Samos H i p p a r i o n Fauna. Bulletin British Museum (Natural A b h., 231(2), 167-190. History) Geology, 20, 229-296. KOSTOPOULOS, D.S. (2005). The Late Miocene mammal locality of G E N T R Y, A. W. (1992). The subfamilies and tribes of the family A k k a s d a πi (Central Anatolia, Turkey). Bovidae (Artiodactyla, Bovidae. Mammal Review, 22, 1-32. Mammalia). G e od i v e r s i t a s,27(4), in press. G E N T R Y , A. W. (2000a). The Ruminant r adiation. I n : E. VR B A &G. KOSTOPOU LOS, D.S. & G.D. K O U F O S (1996) . Late M iocene bovids 152 Dimitris S. Kostopoulos

(mammalia, Artiodactyla) from the locality “Nikiti-1” (NKT), 4 9 - 6 0 . Macedonia, Greece. Ann. Paléont., 81, 251-300. R O U S S I A K I S , S. J. (2003). Oioceros rothii (W A G N E R, 1 857) from the KOSTOPOULOS, D.S. & G.D. KOUFOS (in press). – P h e r a i o s late Miocene of Pikermi (Greece): cranial and dental morphology, chryssomallos n.g. n.sp. (Mammalia, Bovidae, Tragelaphini) from the late Miocene of Thessaly (Greece); contribution to the trage- comparison with related forms. G e o d i v e r s i t a s , 25 (4), 717-735. laphine biogeography. J. Vert. Paleont. in press. SOLOUNIAS, N. (1981). The Turolian fauna from the island of Samos, KOUFOS, G.D. & D.S. K O S T O P O U L O S (1997). Ne w ex ca vat io ns i n Greece. Contrib. Vert. Evol., 6, 1-232. the Neogene mammalian localities of Mytilinii, Samos island, SP ASSO V, N . & D . G E R A A D S ( 2 0 0 4 ). T r a g o p o r t a x P I L G R I M, 1937 Greece. G e o d i v e r s i t a s,19 (4), 877-885. and M i o t r a g o c e r u s S T R O M E R, 1928 (Mammalia, Bovidae) from KOUFOS, G.D. & G . S Y R I D E S (1997). A new early-middle Miocene mammal locality from Macedonia, Greece. C. R. Acad. Sc. Paris, the Turolian of Hadjidimovo, Bulgaria, and a revision of the late 325, 511-516 Miocene Mediterranean Boselaphini. G e o d i v e r s i t a s, 26 (2), 339- R E I F, W.-E. (2003). – Problematic issues of cladistics: 1.ancestor 3 7 0 . recognition and phylogenetic classification. N. Jb. Geol. Palaeont. VRBA, E. S. & G. B. SCHALLER (2000). Phylogeny of Bovidae based A b h., 230 (1), 97-143. on behavior, glands, skulls, and postcrania; I n : E. S. VBRA & G. ROPIQUET, A. & A. H A S S A N I N (2005). Molecular phylogeny of caprines (Bovidae, Antilopinae): the question of their origin and B. S C H A L L E R ( E d s ), Antelopes, deer, and relatives, 203-222. diversification during the Miocene. J. Zool. Syst. Evol. Res., 43 (1), Yale University Press.