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 Bovidae through time is given, focus on the taxonomic, phylogenetic and chronological aspects and questions that arise from their study. Bovini 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. Antilopini 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 mammal 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 Antilocapridae 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-Giraffidae 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 goat-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. Bovinae and Antilopinae. 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 Tragelaphini, 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 genus 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.