Bulletin of the Natural History Museum - Plovdiv Bull. Nat. Hist. Mus. Plovdiv, 2020, vol. 5: 1-14

Late Pleistocene Avifauna of the Pešturina (Nišava District, SE ) and its Implications for Late Pleistocene Refugia in the Central

1*, 2 Zlatozar N. Boev Stefan Milošević

1 - National Museum of Natural History, Bulgarian Academy of Sciences, 1 Blvd. Tsar Osvoboditel, 1000 Sofia, 2 - University of Belgrade, Faculty of Philosophy, Laboratory for Bioarchaeology, Čika Ljubina 18-20, 11000 Belgrade, SERBIA *Corresponding author: [email protected]; [email protected]

Abstract. Avian remains presented here were collected from the deposits spanning between late MIS 5 to 3, and give a rare opportunity to observe Last Interglacial/Last Glacial avian succession in the Central Balkans. The Late Pleistocene avifauna from Pešturina cave (Niš District, SE Serbia), comprises 26 taxa from 18 families and 10 orders, 16 of which are reported for the first time from Pleistocene deposits of the Central Balkans. Today, these species live in a variety of habitats – open grassland, forest, rocky, and aquatic habitats. The species from forest habitat were the most abundant. All species (еxcept black grouse) are present in the country’s modern avifauna. The cave provides some examples of “mixed” Pleistocene avifaunas, including species both of Euro-Siberian and Mediterranean/temperate present day distribution. The occurrence of rock sparrow marks one of the northernmost Pleistocene ones in so far, while presence of other species is suggestive for complex mosaic ecosystems, which probably characterized Last Glacial landscapes in the Central Balkans, at least until the LGM.

Key words: Quaternary/Late Pleistocene , Late Pleistocene refugia, Central Balkans, avian paleoecology, avian taphonomy, avian zoogeography.

Introduction present the uncovered avifauna and its nature Paleontological studies, based on avifaunal of accumulation. Since studied material comes remains, are important for understanding the from an archaeological site, the paleoecological changes in paleoecological settings of the reconstruction is important in assessing the archaeological/paleontological sites. range of habitats also available to Paleolithic Quaternary record of birds on the Balkan – late and modern Peninsula is still insufficiently studied. About 20 humans of the culture. The results Pleistocene cave localities so far revealed are compared with other Late Pleistocene exhaustive avifaunal complexes (Fig. 1). avifaunas from the Balkans, Aegean, and the Pešturina cave (Niš District, SE Serbia) northern Mediterranean arch. Paleoecological represents a previously unexamined Late studies of the Late Pleistocene avifaunas Pleistocene site in the region, located in the throughout Balkans and the Aegean are central part of the Balkan Peninsula. The paper numerous (MOURER-CHAUVIRÉ, 1981; presents results of paleoavifaunal study, with an MLÍKOVSKÝ, 1995A; SÁNCHEZ MARCO, 2004; aim to fill in the Last Interglacial/Early Glacial MICHAILIDIS, 2013; WILCZYNSKI et al., 2016), ecological gap in the Central Balkans, and but do not allow precise ecological division

© Bull. Nat. Hist. Mus. Plovdiv Regional Natural History Museum – Plovdiv http://rnhm.org/en/ University of Plovdiv Publishing House Late Pleistocene Avifauna of the Pešturina Cave (Nišava District, SE Serbia)... between various warm/cold stages of the Late knot (MICHAILIDIS, 2013), redwing (MOURER- Pleistocene, mostly due to lack of absolute CHAIVIRÉ, 1981). At present, the main parts of dating. Based on them, it can be established their summer (breeding) range lie out of the that some taxa occur quite frequently Balkan region, or northwards from the region throughout the peninsula – grey partridge, of the Pešturina cave. In the same time, a large brambling, Eurasian bullfinch and jay, peregrine number of species preferring warm climate falcon, and various corvids, while other were so (swifts, swallows, larks, pipits, orioles, shrikes, far often encountered in the interior/montane, warblers, etc.) have been also found at these or Mediterranean (Aegean/Ionian/Adriatic) localities. parts of Balkans (black grouse, Eurasian Further importance of this material is that woodcock and skylark, griffon vulture, the majority of remains come from Last common quail). A number of species of the Interglacial (MIS 5) deposits of the site, which “boreal” avian complex have been found at are so far rarely observed and studied on many of the Late Pleistocene localities in the Central Balkans, and the Balkans in general. Balkans: short-eared owl, whimbrel, white- Also, the occurrences of several bird taxa were winged snowfinch, horned lark (WILCZYŃSKI et identified for the first time in the Late al., 2016), red-breasted goose, boreal owl, red Pleistocene record of this region.

Fig. 1. Sites of the major Late Pleistocene avifaunas of Balkan Peninsula: Pešturina (*); Tylos (1); Mavromori (2); Liko Gumbes C and Gerani complex (3); Kitsos (4); Vraona (5); Elaichoria 3 (6); Mecha Dupka (7); Bacho Kiro (8); Devetashka (9); Temnata and Cave 16 (10); Kozarska (11); Filipovska and Tsareva Tsarkva (12); Razhishkata (13); (14); Smolućka (15); Trebački krš (16); Mališina stijena (17); Crvena stijena; (18) Bijambarska (19); Zelena (20); Velika; (21) Veternica (22); Krapina (24); Šandalja I and II.

2 Zlatozar N. Boev, Stefan Milošević Short description of the site Materials and methods Pešturina cave (Niš District, SE Serbia; N Presented material was collected in course 43° 17’58’’, E 22° 3’7’’, 305 m a.s.l.; Fig. 1) is of systematic archaeological excavations during located on the rim of the Nišava river basin, at 2010 – 2013 campaigns from trench spanning the transition from mountainous-broken terrain over 16 m2, and up to the depth of around 2,5 and a canyon entrance to lowlands. It is a small m. The material was collected through the cave, 22 m deep, and 15 m wide. The cave washing and sieving of the excavated sediments entrance is 3.5 m high. on 4 and 2 mm mesh. The main stratigraphic units consist of 3 A total of 43 bones and bone fragments of Pleistocene (layers 2-4) layers, which contained birds were collected. All finds are kept in the the assemblage for this study, and one Laboratory for Bioarchaeology (University of Holocene layer (layer 1). Late Pleistocene age is Belgrade, Serbia). Remains were determined confirmed by C14 dating in cal. BP (ALEX & using comparative osteological collection from BOARETTO, 2014; ALEX et al., 2019), and ESR the Vertebrate Animals Department of the (Blackwell et al., 2014): National Museum of Natural History 1. Layer 2: 15 960 – 16 380 (RTD-7148), 28 (Bulgarian Academy of Sciences, Bulgaria). 990 – 31 270 (RTK-6446), >37 800 (RTK- Avian taxonomy follows del HOYO & COLLAR 6445); (2014, 2016). Species’ habitat preferences are 2. Layer 3: 32 067 – 33 400 (RTD-7231B), after Harrison (1982). >40 660 (RTK-6449); 36 400 – 41 400 ESR Abbreviations: (1) anatomical: cmc. – 3. Layer 3/4: >41 100 (RTK-6447), 36 340- carpometacarpus; dex. – dextra; dist. – distalis; 46 330 (RTK-6450); juv. – juvenis; ph. – phalanx; prox. – 4. Layer 4: 43 080 – 45 170 (RTD-7149; 81 proximalis; sin. – sinistra; tbt. – tibiotarsus; tmt. 600 – 105 700 ESR. – tarsometatarsus; (2) ecological: GS - open The uncovered mammalian is grassland; WS – woodland; RS – rocks and rock abundant (34 766 specimens) and is massifs; AS - aquatic. characterized by large diversity of species: Mammuthus sp., Coelodonta antiquitatis, Results Bison priscus, Equus ferus germanicus, Equus Species composition and its hydruntinus, giganteus, Cervus paleoenvironmental implications elaphus, dama, Capreolus capreolus, The established avifauna is rich and varied. Capra ibex, Rupicapra rupicapra, Sus scrofa, The 26 taxa represent 13% of contemporary Lepus europaeus, Castor fiber, Hystrix avifauna in the Balkans part of Serbia (SCIBAN vinogradovi, Ursus spelaeus, Ursus arctos, et al., 2015). Out of them, more than one half Panthera leo, Panthera pardus, Crocuta of the identified taxa (16) are recorded for the crocuta, Canis lupus, vulpes, Felis first time in the Late Pleistocene of Central silvestris, Meles meles (Milošević, 2016; Balkans. Most of the taxa (10) identified at Dimitrijević, in prep.). The site also yielded first Pešturina cave is stenotypic for forests, remains in Serbia found in secure followed by those inhabiting grassland and stratigraphic context (RADOVIĆ et al., 2019). rocky habitats (6), while one is from the aquatic Nearly a thousand chipped stone artefacts were habitat. collected in course of excavations. These are mostly Avian remains were at least partly quartzite backed and naturally backed knives, accumulated by large raptor, most probably retouched flakes, denticulates, and notches, atypical Eurasian eagle owl (Bubo bubo). Bone scrapers, with small proportion of flint products fragmentation and beak punctures observed on and low levallois index, giving the Mousterian some specimens fully correspond to that species industries from layers 4 and 3 Charentian character (BOCHENSKI et al., 1993; LAROULANDIE, 2002). rich in denticulates. Layer 2 contained Gravettian Similar avian accumulations have been studied in artifacts, mostly finished tools among which the region at several Pleistocene sites, both in microgravettes were most numerous (MIHAILOVIĆ Serbia and Bulgaria – Smolućka cave (MALEZ & & MILOŠEVIĆ, 2013). Description and examination DIMITRIJEVIĆ, 1990) (in Serbia), and Cave 16 of all artefacts found in the cave remain out of the (BOEV, 1999), Razhiska cave (BOEV, 2000a), scope of present paper. Kozarnika cave (BOEV, 2001a) (in Bulgaria).

3 Late Pleistocene Avifauna of the Pešturina Cave (Nišava District, SE Serbia)... Galliformes from Serbian and Bulgarian contemporary avifaunas. The species was found in: Velika Phasianidae Pećina, (MALEZ, 1984), Smolućka cave, Serbia (MALEZ & DIMITRIJEVIĆ, 1990; Perdix perdix (Linnaeus, 1758) DIMITRIJEVIĆ, 1998), Zelena cave, Bosnia and Material: tbt. dex. dist.; femur dex. dist. ? Herzegovina (MALEZ, 1973, 1983), Trebjest juv.; tmt. sin. juv.; humerus sin. prox.; ph. 1 dig. cave, Montenegro (LENARDIC, 1990), Visoka 3 pedis sin.; cmc. dex. dist.; cmc. sin.; cmc. sin. Pećina, Croatia (LENARDIC, 1990), Vindija, prox.; humerus sin. prox. (Layers 2, 4). MNI 3. Velika Pećina, Croatia (MALEZ, 1988a, 1997; Remarks: Second /subfossil record for MALEZ-BAČIĆ, 1975), and Veternica, Croatia Serbia. The species was found in: Smolućka Cave, (MALEZ, 1963; MALEZ-BAČIĆ, 1975), Filipovska cave, Bulgaria (BOEV, 2001b), Kozarnika cave, Serbia (MALEZ & DIMITRIJEVIĆ, 1990; Dimitrijević, 1998), Crvena stijena rockshelter, Montenegro Bulgaria (BOEV, 2001a), Devetashka cave, Bulgaria (BOEV, 2001d), and Cave 16, Bulgaria (MALEZ & MALEZ-BAČIĆ, 1974c), Gornja Bijambarska Pećina and Zelena pećina, Bosnia and (Boev, 1999); and Elaichoria 3 cave, , (MLÍKOVSKÝ, 1995a). Herzegovina (MALEZ, 1970a, 1973), Velika Pećina, Habitat: WS. Croatia (MALEZ, 1975, 1984a), Šandalja I cave, Croatia (MALEZ & MALEZ-BAČIĆ, 1974a, 1974b), Krapina, Croatia (MALEZ & MALEZ, 1988; MALEZ, Anseriformes 1970b), and Veternica (Medvednica) cave, Croatia (/ „Perdix sp.“/; MALEZ, 1963). Anatidae The species was also found in: Razhishka cave, Bulgaria (BOEV, 2000a), Filipovska cave, Anas crecca Linnaeus, 1758 Bulgaria (BOEV, 2001b), Kozarska cave, Bulgaria Material: cmc. sin. prox. (Layer 4). MNI 1 (BOEV, 2001c), Kozarnika cave, Bulgaria (BOEV, Remarks: Second record in the 2001a), Temnata cave, Bulgaria (BOEV, 1994); fossil/subfossil record for Serbia. The species Devetashka cave, Bulgaria (BOEV, 2001d), Bacho was found in: Smolućka cave, Serbia (MALEZ & Kiro cave, Bulgaria (BOCHENSKI, 1982), Cave 16, DIMITRIJEVIĆ, 1990; DIMITRIJEVIĆ, 1998); Bulgaria (BOEV, 1999), and Mecha Dupka cave, Zelena cave, (MALEZ, Bulgaria (BOEV, 1999, 2001d). 1973), Gornja Bijambarska cave, Bosnia and Habitat: GS. Herzegovina (MALEZ, 1970a, 1973), Vindija and Velika Pećina, Croatia (MALEZ, 1975, 1984, Coturnix coturnix (Linnaeus, 1758) 1997; MALEZ-BAČIĆ, 1975); Vlasac, Serbia Material: cmc. sin.; cmc. sin. (Layer 4). MNI 2. (DIMITRIJEVIC et al., 2016), Kozarnika Cave, Remarks: First fossil/subfossil record for Bulgaria (BOEV, 2001a), Devetashka Cave, Serbia. The species was found in: Bacho Kiro Bulgaria (BOEV, 2001d), Vraona, Attika, Greece cave, Bulgaria (BOCHENSKI, 1982), Tsareva (BACHMAYER et al., 1989; Mlíkovský, 1995b; Tsarkva cave, Bulgaria (BOEV, 2001b), SYMEONIDIS et al., 1973), Filipovska cave, Bulgaria (BOEV, 2001b), Habitat: AS. Kozarnika cave, Bulgaria (BOEV, 2001a), Temnata cave, Bulgaria (BOEV, 1994), Gruiformes Devetashka cave, Bulgaria (BOEV, 2001d); Cave 16, Bulgaria (BOEV, 1999), Kitsos cave, Attika, Rallidae Greece (MOURER-CHAUVIRÉ, 1971, 1981). Habitat: GS. Crex crex (Linnaeus, 1758) Material: coracoid sin. (Layer 4). MNI 1. Tetrao tetrix Linnaeus, 1758 Remarks: First fossil/subfossil record for Material: humerus dex. dist.; ph. prox. dig. Serbia. The species was found in: Kozarnika maj. sin.; ph. 1 dig. 1 pedis sin. (Layer 4). MNI Cave, Bulgaria (BOEV, 2001a), Temnata Cave, 1. Bulgaria (BOEV, 1994), Devetashka Cave, Remarks: Third record in the Bulgaria (BOEV, 2001d), Cave 16, Bulgaria fossil/subfossil record for Serbia. The black (BOEV, 1999). grouse is species that recently disappeared, both Habitat: GS.

4 Zlatozar N. Boev, Stefan Milošević Otidiformes Bulgaria (/“?Aegolius”/; BOEV, 1994); Kitsos cave, Attika, Greece (MOURER-CHAUVIRÉ, Otididae 1971, 1981), Liko cave, Crete, Greece (WEESIE, 1982, 1987a, 1987b). Otis tarda/Tetrax tetrax Habitat: WS. Material: (cf.) vertebra cervicalis 9 (Layer 4). MNI 1. Accipitriformes Remarks: As the size of female O. tarda overlaps with that of males of T. tetrax we Accipitridae prefer to leave identification open. First record in the fossil/subfossil record for Serbia. O. Gyps fulvus (Hablizl, 1783) tarda and T. tetrax were found in the Material: (cf.) vertebra cervicalis 5 (Layer 4). Pleistocene of Šandalja I and Šandalja II, MNI 1. Croatia (MALEZ & MALEZ-BAČIĆ, 1974a,b; Remarks: Second record in the MALEZ, 1993). “Otis tarda/Tetrax tetrax” was fossil/subfossil record for Serbia. The species found in the Devetashka cave, Bulgaria (BOEV, was found in: Smolućka cave, Serbia (MALEZ & 2001d), and Vraona, Attika , Greece; (O. tarda; DIMITRIJEVIĆ, 1990; DIMITRIJEVIĆ, 1998), Šandalja, Croatia (MALEZ, 1988b), Krapina, BACHMAYER et al., 1989; MLÍKOVSKÝ, 1995b; Croatia (MALEZ & MALEZ, 1988; MALEZ SYMEONIDIS et al., 1978). 1970b; MALEZ-BAČIĆ, 1975), Liko cave, Crete, Habitat: GS. Greece (WEESIE, 1982, 1987a, 1987b; WEESIE, 1982, 1987a, 1987b), Rethymnon fissure, Crete, Charadriiformes Greece (WEESIE, 1982, 1987a, 1987b); while all Quaternary records (5 sites) of G. fulvus in Scolopacidae Bulgaria came from late Holocene archaeological localities (BOEV, 1996, 1999). Scolopax rusticola Linnaeus, 1758 Habitat: RS. Material: scapula dex. prox. (Layer 4). MNI 1. Piciformes Remarks: First fossil/subfossil record for Serbia. The species was found in: Zelena cave, Picidae Bosnia and Herzegovina (MALEZ, 1973, 1975; MALEZ-BAČIĆ, 1975); Vindija, Croatia (MALEZ- Picus canus Gmelin, 1788 BAČIĆ, 1975); Gornja Bijambarska cave, Bosnia Material: humerus sin. (Layer 2). MNI 1. and Herzegovina (/„Scolopax sp.“/; MALEZ, Remarks: First fossil/subfossil record for 1970a); Vindija and Velika Pećina, Croatia Serbia. The Grey-headed woodpecker has not (MALEZ, 1984, 1988a, 1997); Devetashka cave, been encountered before in the fossil/subfossil Bulgaria (BOEV, 1999, 2001d) Gerani IV cave, record in the Balkans. Pešturina cave is its first Crete, Greece (WEESIE, 1987a, 1987b). Liko Quaternary site. cave, Crete, Greece (WEESIE, 1982, 1987a, Habitat: WS. 1987b; WEESIE, 1982, 1987a, 1987b). Habitat: WS. Falconiformes Strigiformes Falconidae

Strigidae Falco tinnunculus (Linnaeus, 1758) Material: coracoid sin. prox. (Layer 2). MNI 1 Aegolius funereus (Linnaeus, 1758) Remarks: Second record in the fossil/subfossil Material: (cf. Aegolius funereus) coracoid record for Serbia. The species was found in: sin. prox. (Layer 4). MNI 1. Smolućka cave, Serbia (MALEZ & DIMITRIJEVIĆ, Remarks: First fossil/subfossil record for 1990; DIMITRIJEVIĆ, 1998), Šandalja I, Croatia Serbia. The species was found in: Devetashka (MALEZ, 1988b; MALEZ & MALEZ-BAČIĆ, 1974a, cave, Bulgaria (BOEV, 2001d), Temnata cave, 1974b), Crvena Stijena rockshelter, Montenegro

5 Late Pleistocene Avifauna of the Pešturina Cave (Nišava District, SE Serbia)...

(MALEZ & MALEZ-BAČIĆ, 1974c), Trebjest cave, Habitat: WS. Montenegro (LENARDIC, 1990), Gornja Bijambarska cave, Bosnia and Herzegovina Sittidae (MALEZ, 1973); Kozarnika cave, Bulgaria (BOEV, 2001a), Temnata cave, Bulgaria (BOEV, 1994), Sitta europaea Linnaeus, 1758 Devetashka cave, Bulgaria (BOEV, 2001d), Cave 16, Material: humerus sin. dist. (Layer 4). MNI 1. Bulgaria (BOEV, 1999), Gumbes B cave, Crete, Remarks: First fossil/subfossil record for Greece (WEESIE 1982, 1987a, 1987b), Kitsos cave, Serbia. The species was found in: Gornja Attika, Greece (MOURER-CHAUVIRÉ, 1971, 1981), Bijambarska cave, Bosnia and Herzegovina Liko cave, Crete, Greece (WEESIE 1982, 1987a, (MALEZ, 1970a), Vindija and Velika Pećina, 1987B; WEESIE 1982,1987a,1987b), , Croatia (MALEZ, 1975, 1997; MALEZ-BAČIĆ, Chalkidiki, Greece (KRETZOI 1977; KRETZOI & 1975); Filipovska cave, Bulgaria (BOEV, 2001b), POULIANOS, 1981), Tylos cave, Greece (ALCOVER Cave 16, Bulgaria (BOEV, 1999). et al., 1992), Vraona, Attika, Greece (BACHMAYER Habitat: WS. et al., 1989; Mlíkovský, 1995b; SYMEONIDIS et al., 1978). Fringillidae Habitat: RS. Fringilla coelebs Linnaeus, 1758 Passeriformes Material: (cf.) humerus sin. juv. (Layer 4). MNI 1. Alaudidae Remarks: First fossil/subfossil record for Serbia. The species was found in: Šandalja I Alauda arvensis Linnaeus, 1758 cave, Croatia (MALEZ & MALEZ-BAČIĆ, 1974a, Material: humerus dex. (Layer 2). MNI 1. 1974b), Vindija and Velika Pećina, Croatia Remarks: First fossil/subfossil record for (MALEZ, 1975, 1988A, 1997; MALEZ-BAČIĆ, Serbia. The species was found in: Bacho Kiro 1975), Kozarnika cave, Bulgaria (BOEV, 2001a), cave, Bulgaria (BOCHENSKI, 1982), Temnata Cave 16, Bulgaria (BOEV, 1999), Liko cave, cave, Bulgaria (BOEV, 1994), Cave 16, Bulgaria Crete, Greece (Fr. coelebs/montifringilla; (BOEV, 1999); Vraona, Attika, Greece WEESIE 1982, 1987a, 1978b). (MLÍKOVSKÝ, 1995b). Habitat: WS. Habitat: GS. Pyrrhula pyrrhula (Linnaeus, 1758) Galerida cristata Linnaeus, 1758 Material: humerus sin. (Layer 2). MNI 1. Remarks: First fossil/subfossil record for Material: humerus dex. (Layer 3). MNI 1. Serbia. The species was found in: Šandalja I Remarks: First fossil/subfossil record for cave, Croatia (MALEZ & MALEZ-BAČIĆ, 1974a, Serbia. The species was found in: Gornja 1974b), Vindija and Velika Pećina, Croatia Bijambarska cave, Bosnia and Herzegovina (MALEZ, 1975, 1997; MALEZ-BAČIĆ, 1975), (MALEZ, 1970a), Vindija and Velika Pećina, Filipovska cave, Bulgaria (BOEV, 2001b), Croatia (MALEZ, 1975, 1997). Kozarnika cave, Bulgaria (Boev, 2001a), Habitat: GS. Temnata cave, Bulgaria (Boev, 1994), Devetashka cave, Bulgaria (BOEV, 2001d), Cave Motacillidae 16, Bulgaria (BOEV, 1999), Liko cave, Crete, Greece (WEESIE, 1982, 1987a, 1987b). Anthus trivialis (Linnaeus, 1758) Habitat: WS. Material: (cf.) humerus sin. prox. (Layer 3). MNI 1. Remarks: First fossil/subfossil record for Hirundinidae Serbia. The species was found in: Velika Pećina, Croatia (MALEZ, 1975, 1984; MALEZ-BAČIĆ, Ptyonoprogne rupestris (Scopoli, 1769) 1975), and , Croatia (MALEZ, Material: (cf.) humerus sin. (Layer 4). 1988a, 1997), Kozarnika cave, Bulgaria (BOEV, MNI 1. 2001a), Devetashka cave, Bulgaria (BOEV, Remarks: First fossil/subfossil record for 2001d); Cave 16, Bulgaria (BOEV, 1999). Serbia. The species was found in: Bacho Kiro

6 Zlatozar N. Boev, Stefan Milošević Cave, Bulgaria (BOCHENSKI, 1982), and Cave, Chalkidiki, Greece (KRETZOI 1977; Kozarnika Cave, Bulgaria (BOEV, 2001a). KRETZOI & POULIANOS, 1981). Habitat: RS. Habitat: RS.

Passeridae Corvus monedula (Linnaeus, 1758) Material: humerus sin. dist.; (cf.) tbt. sin. Petronia petronia (Linnaeus, 1758) dist. juv. (Layer 2). MNI 1. Material: coracoid sin. (Layer 4). MNI 1. Remarks: Third record in the Remarks: First fossil/subfossil record for fossil/subfossil record for Serbia. The species Serbia. The species was found in: Razhishka was found in: Smolućka cave, Serbia (MALEZ & Cave, Bulgaria (BOEV, 2000). DIMITRIJEVIĆ, 1990; DIMITRIJEVIĆ, 1998), Habitat: RS. Visoka Pećina, Croatia (LENARDIC, 1990), Zelena cave, Bosnia and Herzegovina (MALEZ, Oriolidae 1973), Crvena Stijena rockshelter, Montenegro (MALEZ & MALEZ-BAČIĆ, 1974c), Šandalja I, Oriolus oriolus (Linnaeus, 1758) Croatia (MALEZ & MALEZ-BAČIĆ, 1974a, Material: cmc. dex. prox. (Layer 2). MNI 1. 1974b), Maliśina stijena, Montenegro Remarks: First fossil/subfossil record for (LENARDIC, 1990); Vindija and Velika Pećina, Croatia (MALEZ, 1975, 1988a, 1997; MALEZ- Serbia. The find of “Oriolus sp.” species was BAČIĆ, 1975), , Bulgaria found in: Zelena cave, Bosnia and (BOCHENSKI, 1982), Razhishka cave, Bulgaria Herzegovina (MALEZ, 1973), Šandalja, (BOEV, 2000), Tsareva Tsarkva cave, Bulgaria Croatia (MALEZ, 1986), and Vindija cave, (BOEV, 2001b), Filipovska cave, Bulgaria Croatia (Malez, 1988a). (BOEV, 2001b), Kozarska cave, Bulgaria (BOEV, Habitat: WS. 2001c), Kozarnika cave, Bulgaria (BOEV, 2001a), Temnata cave, Bulgaria (BOEV, 1994), Corvidae Devetashka cave, Bulgaria (BOEV, 2001d), Cave 16, Bulgaria (BOEV, 1999), Gerani IV cave, Pyrrhocorax graculus (Linnaeus, 1766) Crete, Greece (WEESIE, 1987a, 1987b), Material: tbt. sin. dist.; ulna sin. dist.; Gumbes B cave, Crete, Greece (WEESIE, 1982, humerus sin. (diaphysis part); (cf): tmt. dex. 1987a, 1987b), Gumbes C cave, Crete, Greece dist. juv. (Layers 2, 3, 4). MNI 3. (WEESIE 1982, 1987a, 1987b), Liko cave, Crete, Remarks: First fossil/subfossil record for Greece (WEESIE 1982, 1987a, 1987b;), Mavro Serbia. The species was found in: Crvena Mouri cave, Crete, Greece (WEESIE 1987a, Stijena rockshelter, Montenegro (MALEZ & 1987b), Sourida fissure, Crete, Greece (WEESIE MALEZ-BAČIĆ, 1974c), Šandalja I, Croatia 1987a, 1987b), Tylos Cave, Greece (ALCOVER (MALEZ & MALEZ-BAČIĆ, 1974a, 1974b), et al., 1992). Vindija, Croatia (MALEZ, 1988a); Bacho Kiro, Habitat: RS. Bulgaria (BOCHENSKI, 1982), Razhishka, Bulgaria (BOEV, 2000), Tsareva Tsarkva, Garrulus glandarius (Linnaeus, 1758) Bulgaria (BOEV, 2001b), Filipovska, Bulgaria Material: humerus dex. dist.; cmc. sin. (BOEV, 2001b); Kozarnika, Bulgaria (BOEV, (Layer 4). MNI 1. 2001a), Temnata Dupka, Bulgaria (BOEV, Remarks: First fossil/subfossil record for 1994), Devetashka, Bulgaria (BOEV, 2001d), Serbia. The species was found in: Zelena cave, Cave 16, Bulgaria (BOEV, 1999); Elaichoria 3 Bosnia and Herzegovina (MALEZ, 1973), cave, Chalkidiki, Greece (MLÍKOVSKÝ, 1995a), Gornja Bijambarska cave, Bosnia and Gerani IV cave, Crete, Greece (WEESIE, 1987a, Herzegovina (MALEZ, 1970a), Veternica 1987b), Gumbes B cave, Crete, Greece (Medvednica) cave, Croatia (MALEZ, 1963), (WEESIE, 1982, 1987a, 1987b), Liko cave, Krapina cave, Croatia (MALEZ & MALEZ, 1988; Crete, Greece (WEESIE, 1982, 1987a, 1987b, MALEZ 1970b), Vindija and Velika Pećina, WEESIE, 1982,1987a,1987b), Vraona, Attika, Croatia (MALEZ, 1975; MALEZ-BAČIĆ, 1975; Greece (BACHMAYER et al., 1989; MLÍKOVSKÝ, MALEZ, 1988a, 1997), Vlasac, Serbia (BORIĆ et 1995b; SYMEONIDIS et al., 1978), and Petralona al., 2014); Kozarnika cave, Bulgaria (BOEV,

7 Late Pleistocene Avifauna of the Pešturina Cave (Nišava District, SE Serbia)...

2001a), Devetashka cave, Bulgaria (BOEV, 2001d), accuracy of ESR for layer 4 – especially since Cave 16, Bulgaria (BOEV, 1999); Gerani II cave, the minimum obtained age ranges from two Crete, Greece (WEESIE, 1982, 1987a, 1987b), different samples are 57 500 to 65 600 Gumbes C cave, Crete, Greece (WEESIE, 1982, (BLACKWELL et al., 2012). 1987a, 1987b), Liko cave, Crete, Greece (WEESIE, In general, the material from layers 2 and 3 1982,1987a, 1987b), Mavro Mouri cave, Crete, refers to Last Glacial, but that of layer 4 reveals Greece (WEESIE, 1987a, 1987b), Tylos cave, avifauna of the Last Interglacial. Greece (ALCOVER et al., 1992). Habitat: WS. The avifauna of Pešturina Cave among other late Pleistocene avifaunas on the Balkan Discussion Peninsula and northern Mediterranean The avifauna of the MIS 3 environment (layers Layer and habitat species distribution 2 and 3) is represented by some steppe (grey The species numbers according to main partridge, skylark), and some forest taxa (grey- habitats is as follows: GS – 4(5) species, WS – headed woodpecker, bullfinch, tree pipit). An 10, RS – 6, and AS – 2. The number of GS and interesting record is the golden oriole, a species that WS is almost equal in layers 2 and 3, while in “is typically a bird of open broadleaf forest”, but layer 4 GS taxa show the greatest taxonomic also lives in forest steppes (HARRISON, 1982). In diversity. context of layer 2 (maximum age of 34 kyA BP), it Layer 2 (15 960 – 37 800 cal. BP) – 11 taxa, is interesting to observe a cohabitation species that 9 species – GS: Falco tinnunculus, Perdix prefer to feed in broadleaf (golden oriole), perdix, Alauda arvensis, Pyrrhocorax graculus, coniferous (grey-headed woodpecker), and mixed Corvus monedula. WS: Picus canus, Pyrrhula (bullfinch) forests. Such coexistence probably pyrrhula, Oriolus oriolus. This layer proves that Balkans were a suitable refugium even corresponds to the MIS 3. in the cool stages of the Late Pleistocene that Layer 3 (32 067 – 46 330 cal. BP) – 6 taxa, 5 directly preceded the LGM. species – GS: Perdicinae gen. indet., BLONDEL et al. (2010) states that Eurasian jay Pyrrhocorax graculus, Corvus cf. monedula. and common chaffinch (species with very large WS: Anthus trivialis. This layer corresponds to modern-day distributional ranges) survived in the the MIS 3. Balkans during the glacial periods. Records of Layer 4 (105 700 – 81 600) – 17 taxa, 14 Pešturina (and other sites such as Krapina, Vindija, and Velika Pećina) fully support the continuity of species – GS obviously prevail: Gyps fulvus, their persistence in the Central Balkans since the Perdix perdix, Coturnix coturnix, Crex crex, Last Interglacial. Otis tarda/Tetrax tetrax, Ptyonoprogne rupestris, These authors also argue that contemporaneous Petronia petronia, Pyrrhocorax graculus, Corvus colonies of Eurasian crag martins “occur near the monedula. WS: Tetrao tetrix, Scolopax rusticola, coast” in the Mediterranean region (BLONDEL et al., Aegolius funereus, Sitta europaea, Fringilla 2010: p. 127), but numerous evidences from coelebs, Garrulus glandarius. This layer Pleistocene deposits, including Pešturina, prove its corresponds to the late MIS 5 and perhaps MIS 4 former wider inland distribution. Tree pipit is at (ca. 110 000 – 70 000 BP). present more common in the northern European According to C14 dates, it is apparent that forests than in the Mediterranean basin, but it was stratigraphic mixing occurred at the site, since widely spread throughout the Balkans during the dates from layer 3 already present infinite C14 temperate/warm stages of Pleistocene. ages, as well as earlier dates in upper portions As stated, at the height of the cool stages, “no of layer 4 compared to some samples from arboreal vegetation persisted north of … the layer 3. However, according to Bayesian model Carpathians” in Southeastern Europe (BLONDEL, (ALEX et al., 2019), layer 2 is not older than 34 1997: cxxiv), the broad-leaved forests in the Balkans kyA cal. BP, while layer 3 is not older than 45 (including those in the region of Pešturina) were kyA cal. BP. Also, the dates are not consistent refugia for Eurasian jay, tree pipit, grey-headed at layer 3 and 4 boundary. Since there are woodpecker, golden oriole, common chaffinch, infinite C14 ages both from layer 3 and 3-4 layer woodcock, bullfinch, etc. Data from layer 4 in boundary, it is reasonable to assume the Pešturina show that: (1) Typical steppe species occur

8 Zlatozar N. Boev, Stefan Milošević in layer 4: Perdix perdix, Coturnix coturnix, Crex In comparison to other contemporaneous crex, Otis tarda/Tetrax tetrax; (2) typical boreal Late Pleistocene avifaunas in the Balkans forest species occur in layer 4: Tetrao tetrix, Scolopax (Krapina cave, NW Croatia, around 580 km rusticola, Aegolius funereus, Sitta europaea, Fringilla away from the Pešturina cave), we also found coelebs, Garrulus glandarius; (3) typical broad-leaf common species. The 3 common bird species from Pešturina cave (total taxa/species – 26) forest occur in layers 3 and 2: Picus canus, Pyrrhula and Krapina cave (total taxa/species – 10; pyrrhula, Oriolus oriolus, Anthus trivialis. MALEZ & MALEZ, 1988; MALEZ, 1970b) The “steppe” taxa most widespread in the represent 11.1 % and 30.0 % respectively. interglacial environment (after HARRISON, These are: Gyps fulvus, Perdix perdix, and 1982) were grey partridge, common quail, corncrake, bustards, rock sparrow, or “forest” Garrulus glandarius. The refugia character of Central Balkans, as (HARRISON, 1982) taxa - black grouse, woodcock, boreal (Tengmalm's) owl, Eurasian suggested by avifaunal remains discussed here, nuthatch, common chaffinch, and Eurasian jay. could also had impact of other faunal The presence of the crag martin suggests communities, as well as Paleolithic humans. relatively high summer temperatures, allowing it Various warm adapted terrestrial fauna may to hunt abundant small flying dipteran insects. have found favourable paleoecological Well-represented steppe and woodland species conditions here, both from Mediterranean in confirm the mosaic forest-steppe landscape in course of Last Interglacial, and Danube this region of the Central Balkans, similarly to corridor and Central Europe during the Last those already observed in the eastern part of Glacial. These may have given the rise of specific mosaic environments rich primarily in the Balkan Peninsula, i. g. Bulgaria (BOEV, 1999, 2001d, 2006). species diversity, and perhaps shifts in their All species, except black grouse, found in population density at various stages of the last the Quaternary deposits of the Pešturina cave Ice Age as well. This may opened up the are present in the recent avifauna of the possibility for Paleolithic humans to choose from a wider array of habitats in which they country (SCIBAN et al., 2015). could establish successful subsistence. At least In general, the species composition partially when the site of Pešturina is concerned, there corresponds to the Late Pleistocene avifaunas are no marks of sharp changes in the habitats from some of the neighbouring regions in the presented by avifauna, and no true boreal Balkans – Croatia, Bosnia, Montenegro, and species, although the abrupt paucity in Western Bulgaria where most of the species taxonomic representation from the Last Glacial encountered at Pešturina have been registered deposits is evident in comparison with the so far (BOEV, 1994, 1999, 2000 a, b, 2001 a, b, Interglacial assemblage. c, d, 2006 a, b, c). All of them confirm their As many other Pleistocene localities on the wider distribution throughout the Central Balkans, Pešturina cave provides interesting Balkans during the Late Pleistocene. This is best illustrated over the example of Late examples of the s. c. „mixed” avifaunas: Tetrao Pleistocene avifauna from Kozarnika cave (NW tetrix and Oriolus oriolus (Fig. 2), Bulgaria, around 70 km away from Pešturina; Ptyonoprogne rupestris and Scolopax rusticola Gravettian, 26000-19 000 B.P), which show (Fig. 3). At the present, breeding ranges of striking similarity (BOEV, 2001a). The 14 these species are more or less widely separated, common bird species from Pešturina cave (total but in the mosaic forest-steppe landscapes of taxa/species – 26) and Kozarnika cave (total Pleistocene they coexisted in the vicinities of taxa/species – 43; BOEV, 2001a) represent 51.8 the cave. % in the Pešturina cave and 32.6 % in the As FINLAYSON (2011) writes, the glaciations Kozarnika cave respectively. These are: Anas cold did not affect the Palearctic’s birds crecca, Falco tinnunculus, Tetrao tetrix, Perdix [dramatically], because the cold seasonality could be dealt with through migration, while perdix, Coturnix coturnix, Crex crex, Aegolius sedentary species simply shifted geographical funereus, Anthus trivialis, Ptionoprogne position. Among those that found refuge on rupestris, Fringilla coelebs, Pyrrhula pyrrhuia, the southern peninsulas, and other cryptic refugia, Garrulus glandarius, Pyrrhocorax graculus, were the sedentary and partially migratory and Corvus monedula.

9 Late Pleistocene Avifauna of the Pešturina Cave (Nišava District, SE Serbia)... bioclimatic tolerants. The boreal birds returned (MINGOZZI & ONRUBIA, 1997). This is the one of north too, but also left prisoners, this time trapped the northernmost occurrences of the rock sparrow around the mountains of the mid-latitude belt (s. c. in Pleistocene Europe. glacial relicts). (p. 236). This author accepts the Established avifauna includes 16 species western Balkans, and to a lesser degree the Alps previously unknown in the Pleistocene record of and the Italian mountains, as “the major European Serbia (TYRBERG, 1998; 2008): C. coturnix, C. refugia for broadleaved trees during the last crex, S. rusticola, A. funereus, P. canus, A. arvensis, glaciation”. (p. 29). G. cristata, A. trivialis, S. europaea, F. coelebs, P. pyrrhula, O. oriolus, G. glandarius, P. rupestris, P. petronia, and P. graculus.

Acknowledgements Excavations at Pešturina cave are funded thanks to the Serbian Ministry of Culture grant No 177023, as is the study of presented Pešturina osteological material for one of the authors PhD (SM). We thank National Museum of Natural History in Sofia, Bulgaria, for avifauna comparative material, and Vesna Dimitrijević and Laboratory for Bioarchaeology at Faculty of Philosophy in Belgrade and its study collection. Special thanks to Fig. 2. Present day ranges of Tetrao tetrix two anonymous reviewers of the manuscript for (after Schmitz, 1997) and Oriolus oriolus (after their helpful suggestions and remarks. WASSMANN, 1997) in Europe, with the position of Pešturina indicated. The areas of overlapped References ranges are given in grey color. ALCOVER J. A., FLORIT F., MOURER-CHAUVIRÉ C., WEESIE P.D.M. 1992. The avifaunas of the isolated Mediterranean islands during the Middle and Late Pleistocene. In: Campbell, K. E. (Еd.), Papers in avian honoring Pierce Brodkorb. Science Series. Natural History Museum of Los Angeles County, Los Angeles 36: 273-283. ALEX B., BOARETTO E. 2014. Radiocarbon Chronology of Pešturina Cave. In: Mihailović D. (Еd.), Palaeolithic and Mesolithic Research in the Central Balkans. Serbian Archaeological Society, Belgrade 39-48. Fig. 3. Present day ranges of Ptyonoprogne ALEX B. MIHAILOVIĆ D. MILOŠEVIĆ S. BOARETTO rupestris (after Sackl & Sere, 1997) and E. 2019. Radiocarbon chronology of Middle Scolopax rusticola (after HOODLESS & SAARI, and sites in Serbia, Central 1997) in Europe, with the position of Pešturina Balkans. Journal of Archaeological Science: indicated. The areas of overlapped ranges are Reports, 25: 266-279. given in grey color. BACHMAYER F., MALEZ V., SYMEONIDIS M., THEODOROU G., ZAPFE H. 1989. Die Pešturina cave reveals distribution of 5 present Ausgrabung in der Hohle von Vraona (Attika) long-distance migratory birds, which are summer im Jahre 1985. Sitzungsberichte Osterreichische visitors in the Balkans. These belong to 3 orders – Akademie der Wissenschaften. Mathematisch- C. coturnix, A. trivialis, O. oriolus, P. rupestris, Naturwissenschaftliche Klasse, Abtteilung 1, and C. crex. One species (P. petronia) deserves 197: 287-307. special attention, as its present day (summer) range BLACKWELL A. B., CHU S., CHAITY I., HUANG Y. lies far south of the location of the Pešturina cave E. W., MIHAILOVIĆ D., ROKSANDIĆ M.,

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