ffistorianaturalis bulgarica, 14, 2002: 109-115 Fossil record and disappearance of peafowl (Pavo Linnaeus) from the Balkan Peninsula and Europe (Aves: Phasianidae) BOEV Zlatozar Introduction The Balkan Tertiary avifauna is still poorly studied. A total of21 Tertiary sites, containing fossils of birds are known from all 10 Balkan countries (including the Balkan parts of Romania and Turkey), 10 of them from Bulgaria (Mlikovsky, 1996a), Because oftheir unique geographical position, the Balkans served as a bridge for the terrestrial faunal migrations during the whole Tertiary period and later as well, a fact that was well documented by the fossil mammalian fauna (Spassov, 2000). This determines their significance as a region of considerable palaeozoogeographical interest for the whole European continent. Some of the latest (end of 1999) palaeontological discoveries contributeto the evaluation ofthe importance of the Balkan Peninsula as one of the regions of Europe of the last "tropical" elements among the terrestrial fossil bird fauna as well. The fossil record ofpeafowl from the Balkan Peninsula Recently five bones, four ofthem originating from Bulgaria and one ft-om Greece, have contributedto the so far scantyfossil record ofthe genus Pavoon the Balkans. The Bulgarian finds represent four pedal phalanxes of an adult individual, kept in the National Museum ofNatural History (Sofia): phalanx 2 dig. II pedis; phalanx 2 dig. Ill pedis; phalanx 3 dig. IV pedis; phalanx distalis dig. I pedis. The phalanx 2 dig. Ill pedis is broken and the proximal articular part ofthe bone is not preserved (BoEV, 2001). All other finds are of excellent preservation and even the tinyvascular pores on the bone surfaces £ire clearly seen. The find fi'om Greece represents a distal halfofthe left tibiotarsus ofan adult individual. The surface of the bone is damaged, but the distal end is preserved in good condition. The find is kept in the Laboratory ofGeology and Palaeontology of the University of Thessaloniki (BoEV & KouFOS, 2000). All these finds are referred to the fossil Bravard's Peafowl Pavo bravardi(Gervais, 1849), known until now from Western Europe only (BoEV, 2001). A detailed 109 \ morphological description of the Bulgarian and Greek fossil remains of P. bravardiis given in BoEV (2001) and BoEV & KOUFOS (2000). Short data on the Balkan locaUties ofpeafowl m Muselievo locality (Bulgaria). The site is a rock shelter ofup to 1,5 depth in a destroyed Pliocene cave in ? Sarmatian limestones, about 1 km SE ofthe Muselievo Village (Lovech District, N Bulgaria) and 350 m offthe right m bank ofthe Ossam River; 150 a. s. 1. (43.36 N, 24.50 E). Associated fauna: Reptnia: Chelonia fam. indet.; Aves: Accipiter sp., Falco sp. ex gr. cherrug, Lagopus aff. atavus, Pavo bravardi, Passeres fam. indet. and Aves indet. (Boev, 2001); Mammalia: Mimomys occitanus, Pseudomeriones abbreviatus, Rhagapodemus hautimagnensis, Myomimus dehmi, Sus minor, Sciurus sp., GLissp., Apodemussp., Rhinolophussp., Miniopterussp., Dolichopithecussp. (Popov & Delchev, 1997), Talpa cf. csarnotana, BLarinoides mariae, Deinsdorfia kordosi, Mafia csarnotensis, Episoriculusgibberodon, Hypolagus brachignatus, Trischizolagus cf. dimitrescuae, PliopentaLagits cf. dietrichi, Pratilepus kutschurganicus, Ochotonoides csarnotanus, Dolomys odessanus, Pliomys hungaricus, Dryomimus cf. eliomyoides, Glis minor, Apodemus cf. dominnans, Allocricetus cf. bursae, A. ehki, Trilophomys pyrenaicus, Prospalaxpriscus, Pliospalaxcompositodontus, Myotissp., Pliopetauristasp. and Sciurotamias sp. (Popov, in press), Vulpes sp., aff. Nyctereutes sp., Felidae gen.(? Dinofelis sp.), Sus arvernensis minor, Cervus cf. pardinensis, Procapreolus sp., aff. Gazella sp., Tapirus arvernensis, cf. Stephanorhinus jeanvireti, Lynx sp., aff. Macaca sp. (Spassov, 2000). Taphonomy: Most probablythe accumulation ofthe avian remains is a result ofthe life activity at a feeding place of large owls (Strigiformes) and carnivore mammals (Carnivora) that shared the cave. Age: The site dates backto the second half of the Middle Ruscinian, MN 15; 3,3-3,1 MA, the Early Pliocene (PoPOV & Delchev, 1997). The chronostratigraphyfollows (1990). Latterlythe age MN has been determined the second halfof 15 (Spassov, 2000). Megalo Emvolon locality (Greece). The site is situated 20 km SW of Thessaloniki, Macedonia, N Greece (40:35 N; 22:46 E). It is the best known Ruscinian locality containing fossils of Tertiary vertebrate fauna in Greece. Associated fauna: Reptilia: Testudo cf. graeca, Testudo sp.; Mammalia: Oryctolagus odessanus, Trischizolagus dumitrescuae, Trischizolagus cf. maritsae, Microspalax odessanus, Dolichopithecus ruscinensis, Nyctereutes tingi, Hipparion longipes, Parabos macedoniae, Koufotragus bailloudi, Gazella borbonica and Sus minor(BoEV & KouFOS, 2000). Taphonomy: The deposits are fluvial and consist mainlyofcross-bedded sands, gravel, silts and sand-silts. The character ofthe sediments indicates a very rapid deposition, while the faunas from the different horizons do not differ (BOEV & KouFOS, 2000). For this reason all the material is referred to as Megalo Emvolon fauna. Age: The associated megafauna dates the site back to the Early Pliocene (Ruscinian, MN 15 zone) (BoEV & KouFOS, 2000). 110 Review of the fossil record ofpeafowl in Europe The fossil record ofthe genus Pavois based on two species - Pavo bravardi and P. aesculapi (Gaudry, 1862). Adding the new discoveries of Pliocene peafowl in the Balkans, the summary data on the history ofthese birds will look as follows: . The Bravard's Peafowl isknownfromseven sites(Fig. 1)infour countries, all in Europe (France, Moldova, Bulgaria and Greece): Serrat-d'en-Vacquer (MN 15), Arde (MN 16), Saint-VaUier (MN 17), Seneze (MN 17) in France (Mourer- Chauvire, 1990, 1993, 1996), Lucheshti (MN 16 /or ?15/) in Moldova (Bochenski & KUROCimiN, 1987; Mlikovsi^, 1996b), Muselievo (MN 15) in Bulgaria (BOEV, 1996, 2001) and Megalo Emvolon (MN 15) in Greece (BoEV & KOUFOS, 2000). The other species, Pavo aesculapi, is known from five sites in four countries, all European as well (Greece, Moldova, Ukraine and Hungary): Pikermi (MN 12-13) in Greece (Janossy, 1991; Mlikovsky, 1996c), Kolkotova Balka (MN 9-10) in Moldova (Janossy, 1991; Mlikovsky, 1996b), Odessa catacombs (MN 15) (Janossy, 1991; Mlikovsky, 1996d) and Novoelisavetovka (MN 11) in Ukraine (Janossy, 1991; Mlikovsky, 1996d) and Polgardi (MN 13) in Hungary (Janossy, 1991; Mlikovsky, 1996e). Thus, the fossil record of peafowl in Europe proves their distribution in twelve sites on the continent, three ofthem on the Balkan Peninsula. Greece is the only country where both Tertisiry peafowl were spread, but their occurrence there was not contemporaneous. The Late Miocene record of P. aesculapi from Pikermi antedates the new find of P. bravardi from Megalo Emvolon by about 3 million years. ItisworthtomentionthatinSWFrance(Sansan, MN6, 15-12 Ma) remains of Miophasianus alius have been found (Cheneval, 1996). Later this species has been considered to be much more closely related to peafowl (Cheneval, 2000). That fact could indicate the much older (MN 6) "European" history of the lineage ofthe peafowl than what has been considered until now (MN 9). The habitats of the fossil peafowl The palaeobotanical analysis of the localities where P. bravardi was established shows the presence ofsome exotic thermophilous floral elements. Their analogues now are spread out oftheWestern Palearctic - the Caucasus, the Himalayas, Japan, Southeast Asia or Central America (Mourer- Chauvire, 1989). At all these sites the remains of deers are numerous. They are considered indicators for woodland habitats with scattered trees. At the new Balkan site of Muselievo an abundance of finds of Cervidae (Cervus cf. pardinensis, Procapreolus sp.) is also well established (Spassov, 2000). The Macromammalian faunaindicates presence offorested savanna, light forests to open lands with scattered bush (Spassov, 2000). The same habitats are recognized at the other Balkan site, Megalo Emvolon (BoEV & KouFOS, 2000). In the Western Europe (France) mixed broad-leafed woods were the preferred habitats of the Bravard's Peafowl and their remains have been 111 Fig. 1. Geographical distribution of the genus Pavo in Europe: Pavo bravardi (1-7): 1 - Serrat-d'en-Vacquer; 2 - Arde; 3 - Saint-Vallier; 4 - Seneze (1-4: France); 5 - Lucheshti (Moldova); 6 - MuseHevo (Bulgaria); 7 - Megalo Emvolon (Greece); Pavoaesculapi(8-11): 8 - Pikermi (Greece); 9 - Odessa; 10 - Novoelisavetovka (Ukraine); 11 - Kolkotova Balka (Moldova) (Drawing: Vera Hristova) found in fluviatile, loessic, or lacustrine sites (Mourer-Chauvire, 1990). Actually, the two recent species oftheAsian peafowl are spread mainlyin the open forest with undergrowth along streams, open woodlands, forest-edge and riverine forest, a peculiarity that could explain the occurrence of the Pavoremains in the European localities. Disappearance of peafowl from Europe The Bravard's Peafowl roamed South-European forests, both in the West and the East of the continent. This species must have appeared there not later than the Late Miocene - Early Pliocene. Possibly, P. aesculapiwas of a more eastern distribution, where it had spread much earlierthan P. bravardi. It is considered that the European peafowl disappeared soon after the first cold phases ofthe Pretegelen (Pretiglian) (Mourer-Chauvire, 1990), i. e. over 112 1 million years ago, together with the disappearance ofthe whole "tropical" complex. Muselievo and Megalo Emvolon are the 6''' and the 7"" site ofPavo bravardi up to now. These sites mark its distribution on the Balkans, both in the extreme North ofthe Peninsula andin its southern regions. Togetherwiththe third site ofPikermi, providing finds ofPavo aesculapi, they proved the Late Miocene - Early Pliocene distribution of peafowl in the South-east corner of the continent. Being on the cross-road of the terrestrial zoogeographical influences, the Balkans were one of the European territories where the last heat-loving elements disappeared. The Bravard's Peafowl survived in the western parts ofthe continent (S France) muchlonger, i.e. bythe very end of MN the Tertiary (the Late Pliocene - Early Pleistocene border) in 18. Summarizing the data available so far, it is clear that the stratigraphic MN range ofPavo bravardiis 15-17 (Earlyto Late Pliocene). P. aesculapiwas MN MN in existence between 9 and 15 (Late Miocene to Early Pliocene). Pavo aesculapi was spread in Eastern and Central Europe, while P. bravardi roamed the whole Southern Europe, both Western and Eastern Europe. In Western Europe the latter survived up to the very end of the Pliocene (Mourer-Chauvire, 1996). It is likely that in SE Europe this peafowl had survived in the relatively warmer habitats until more recent time, i.e. by the beginning ofthe Pleistocene. Acknowledgments The author is very grateful to Dr. Cecile Mourer-Chauvire (Universite Claude Bernard - Lyon), Dr. Evgeny Kurochkin (PaleontologicEil Institute - Moscow) and two anonymous reviewers for their helpful comments and additions to the earlier version ofthe manuscript ofthe paper. References BocHENSKi Z., E. Kurochkin. 1987. New data onPliocene phasianids(Aves: Phasianidae) of Moldaviaand S. Ukraine. - Actazool. cracov., 30 (7): 81-96. BOEV Z. 1996. Tertiary avian localities of Bulgaria. - Acta Univ. Carolinae Geologica, 39 [1995]: 541-545. BOEV Z. 2001. Early Pliocene avifauna of MuseUevo (C Northern Bulgaria). - Acta zool. cracov., 44 (1): 37-52. BOEVZ., G.KouFOS. 2000. ThepresenceofPavobravardi{Gervsds, 1849)(Aves, Phasianidae) intheRuscinianlocalityofMegaloEmvolon,Macedonia,Greece. -GeologicaBalcanica, 30 (1-2): 69-74. ChenevalJ. 1996. MioceneAvianLocalities ofFrance. -ActaUniv. Carolinae Geologica, 39 [1995]: 599-611. Cheneval J. 2000. L'Avifaune de Sansan. - Mem. 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Receivedon 12.12.2001 Author's address: Dr Zlatozeu*Boev National Museum ofNaturalHistory Tsar OsvoboditelBlvd. 1 -: 1000 Sofia, Bulgaria [email protected] 114 u {Pavo Linnaeus) u (Aves: Phasianidae) () . . . . . 6 . (MN 15 ( 3,3 - .) - u . Haxogkimie bravardi) - -egpama 6 - £1 , . , Pavo go ( ) ( . ). - ,{Pavo aesculapi) u P. b.ravardi. MN Pavo bra,vardi 15-17 - MN .u 6 u 9-15 . bravardi -- - ,""u , (). 6 u go u go 115