Late Pleistocene and Earliest Holocene avifauna from the Loutra Almopias Cave (Macedonia, Greece)

The bird fauna of the Loutra Almopias Cave comprised eight orders, 22 families, 41 genera and at least 47 species (68 different taxa) established based on 551 avian bone finds. The Loutra Almopias Cave is a new (26 th) Pleistocene avian locality in Greece. About 92.5% of the material was dated to the Earliest Holocene (11.230 ± 110 y. BP), coming from the upper chamber of the cave, whereas an older assemblage was dated to the Late Pleistocene (37,880 y. ± 370/360 BP), coming from the floor chambers of the cave. Here is provided the first fossil record of 13 species in Greece (Buteo lagopus, Francolinus francolinus, Lagopus lagopus, Bonasa bonasia, Dryocopus martius, Lulula arborea, Bombicylla garrulus, Cinclus cinclus, Sylvia borin, Carduelis carduelis, Loxia curvirostra, Plectrophenax nivalis and Emberiza cirlus). Three genera (Francolinus, Anthus and Riparia/Ptyonoprogne) were established for the first time in Pleistocene deposits in the country. The record of Fr. francolinus was the first one for the Balkans and Eastern Europe. At both sites inside the cave, regardless of their different age, the habitat preferences of the identified bird species indicated that the surrounding paleoenvironment included both grassy openlands (dominated by gramineans) and woodland (coniferous or mixed woods). In addition, rocky habitats were represented in the surroundings of the locality. The presence of small species of Perdicinae (26%) and corvids (34%) indicated the former existence and prevelance of open grassy fields and rocky environments. The Alpine chough was the most abundant species at both localities. In the Earliest Holocene it comprised 29.5%, whereas in the Late Pleistocene it reached 25.6% of the material. Such a representation suggests a natural non-human accumulation of the material, probably due to the feeding behavior of Bubo bubo. The Late Pleistocene record (19 taxa; 41 finds) included Falco sp. cf. F. peregrinus, Perdix perdix, Perdix sp., Alectoris graeca, Alectoris sp., Lagopus cf. lagopus, cf. Bonasa bonasia, Columba livia, Columba palumbus, Bubo bubo, Melanocorhypha calandra, Anthus sp., Turdus sp., Pica pica, Pyrrhocorax graculus, Loxia curvirostra, Coccothraustes coccothraustes, Pyrrhula pyrrhula, Fringillidae gen. indet., non-Passeriformes indet. and also suggests the site has been used by the eagle owl. The climate probably used to be drier and cooler than in the Earliest Holocene. The record of 17 woodland species from the Earliest Holocene that were absent in the Late Pleistocene could be explained by the more humid and moderate climate at the very end of the Pleistocene.


Introduction
The bear cave site (Loutra Almopias Cave: LAC) is located in northern Greece, NW of and about 120 km from Thessaloniki (Fig. 1). The investigations of the area have lasted more than 20 years and the site has yielded abundant cave bear remains among large mammals, as well as micromammals. All these consist Phasianidae Vigors, 1825 Perdix perdix (Linnaeus, 1758). (Pl. 1, Fig. 4) Known from the EP to LP from numerous Eurasian localities from France to Syria and China (Tyrberg, 1998;2008). The locality falls within the recent breeding range of the species (Hagemeijer & Blair, 1997).
Occurs mainly in moister grasslands, grass moors, heathlands, borders of bog and marshland, steppes, alpine and lowland meadows and open scrubs (Harrison, 1982).
Perdix sp. Finds of Perdix sp. are known from the MP to LP from numerous localities in China, f. Czechoslovakia, France, Israel and Portugal (Tyrberg, 1998;2008). Perdix cf. jurcsaki has been established in the Early Middle and MP of Petralona (Chalkidiki), while Perdix palaeoperdix -from the EP of Tourkobounia 2 (Attiki) (Tyrberg, 1998). The material from LAC did not allow identification to species level.
B 150: The morphology fully agreed with the coracoid of F. francolinus and clearly distinguished it from all other listed genera/species. Dimensionally it also was closer to the black francolin. It could be distinguished clearly from L. muta and B. bonasia based on the much shorter processus lateralis coracoidei and the much deeper bow of crista articularis sternalis. B 163: In cranial view the find clearly differed from L. muta because of the considerably thicker labrum internum coracoidei. In Zlatozar Boev, Evangelia Tsoukala dorsal view it had two well-developed fossa separated by a protruding ridge, completely absent in L. muta and B. bonasia. Previous fossil records of F. francolinus originate only from the LP of Spain, Israel, Syria, Iraq and Iran (Tyrberg, 1998). So far, Fr. francolinus has not been established from Greece and the Balkans nor from Eastern Europe at all (Tyrberg, 1998;2008). Francolinus capeki Lambrecht, 1933 is known from the MP of Croatia, Francolinus sp. -from the EP of Croatia. Until the 19 th century, Fr. francolinus was spread in the W Mediterranean, but the present range approaches westwards up to S Turkey and Cyprus (Babayev et al., 1997). The locality lies outside of the recent breeding range of the species (Hagemeijer & Blair 1997). Occurs mainly in moister areas with good growth of vegetation, shrubby thickets bordering watercourses and lakes, extensive areas of scrub or shrubby herbage, lush grasses, marshy areas with tall herbage and grasses, forest edges, forest clearings within the 26ºC July isotherm (Harrison, 1982). Its historical range includes SE Spain and Sicily and probably some Aegean Islands (until the middle of 19 th century; Harrison, 1982). In general, all the francolins' records in the Pleistocene deposits are scant. According to Tyrberg (1998), five species of Francolinus Stephens, 1819 have been recorded in the Palearctic, four of them in Europe. Mourer-Chauviré (1993) considers Francolinus spp. as Tertiary relicts, which have progressively vanished in the Pleistocene.
Francolinus sp. The finds clearly differed dimensionally from Perdix and Alectoris (and Lagopus and Bonasa) ( Table 2). Some measurements differed by ca. 30% of the measurement value (for example measurement "d" of A. graeca, "b" of P. perdix, "c" of L. lagopus, etc.).
Alectoris/Francolinus. Both phalangeal finds were thinner than A. graeca. Based on their size and overall morphology, they could be attributed to a phasianid species, close to Alectoris or Francolinus, but precise identification was not possible.
Perdix/Francolinus. The five phalangeal finds were smaller, shorter and thicker than P. perdix. Based on their size and overall morphology, they could be attributed to a phasianid species, close to Perdix or Francolinus, but precise identification was not possible.
Perdicinae gen. indet. This material (seven pedal phalanges) needs further detailed comparisons of wider series of specimens (mainly francolins, Ammoperdix spp.), but it could be attributed to Perdicinae, based on the size and morphological similarity (round profile of the distal end and compact phalangeal body of the ph. 1 and 2).

Tetraonidae Vigors, 1825
Tetrao tetrix Linnaeus, 1758. (Pl. 1, Fig. 11) The find (left ulnare bone) completely fitted the size and shape of the compared black grouses. Known from the EP to LP from Spain to Ukraine (Tyrberg, 1998;2008). In Greece, the species is known from the LP of Elaiochoria 3 (Chalkidiki) (Tyrberg, 1998). The locality lies outside of the recent breeding range of the species. Until 1990 s , the species still has been breeding in the north-westernmost parts of the neighboring North Macedonia (Hagemeijer & Blair 1997). It occurs mainly in conifer and birch forests along forest edges, in open forests and areas of rock and broken woodland scrubs, swampy heathlands, moorlands and bogs within the 11º C and 21-24º C July isotherms (Harrison, 1982).
Lagopus cf. lagopus (Linnaeus, 1758). (Pl. 1, Fig. 12; Pl. 2, Figs. 1-2) The find (distal/claw/pedal phalanx and a symphysal half of a furcula) fitted the size and shape of the compared willow ptarmigan. Known from the EP to LP from numerous Eurasian localities from Spain to Ukraine (Tyrberg, 1998;2008). Recently, also found in the LP in Mališina Stijena (Montenegro; Bogoćević & Dimitriević, 2004). Until now, the species has not been established in fossil records from Greece. The locality lies outside of the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in swampy scrub tundra, marshy heathlands, bogs in conifer forests, birch and willow forests, riverine scrubs of steppes (Harrison, 1982).
aff. Lagopus sp. With typical shape and proportions of a gallinaceous bird. Dimensionally close to the genus Alectoris, but the general proportions were similar to Tetraonidae, especially its size, which indicated that it should be considered as a member of Lagopus. Two species of Lagopus used to be widely distributed in the Pleistocene of Europe, established in numerous localities far south of the recent breeding range. All of them were dated to the Late Pleistocene (Tyrberg, 1998;2008 (Tyrberg, 1998). So far, the species has not been established in the fossil record of Greece (Tyrberg, 1998;2008). The locality falls within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in thick forests with undergrowth, mountain conifer forests, spruce forests with bilberry layer, mixed forests with undershrubs, beech forests and riverine alder forests Late Pleistocene and Earliest Holocene Avifauna from the Loutra Almopias Cave (Macedonia, Greece) within the 13º C and 21-22º C July isotherm (Harrison, 1982).
Bonasa bonasia/Lagopus muta. The size and morphology of the three pedal phalanges suggested a small tetraonid. Further identification was not possible.
Columbiformes (Latham, 1790) Columbidae (Illiger, 1811) Columba livia Gmelin, 1789. (Pl. 2, Fig. 7) Known from EP to LP from numerous Eurasian localities from Spain to Jordan and China (Tyrberg, 1998;2008). In Greece, the species is known from the LP of Elaiochoria 3 (Chalkidiki) and Vraóna (Attiki) (Tyrberg, 1998); from the ?LP of the Liko Cave (Crete); the LP of the Kalamakia Cave (Lakonia) (Tyrberg, 2008) and from the LP of the Klissoura Cave 1 (Bochenski & Tomek, 2010). The locality falls within the recent breeding range of the species (feral pigeon) (Hagemeijer & Blair, 1997). Occurs mainly in rocky areas with nearby open spaces, rocky coasts and islands, rocky outcrops in mountains, in open grasslands or steppe (Harrison, 1982). Theoretically, a small part of the identified material could belong to Columba oenas Linnaeus, 1758, a species often undistinguishable osteomorphologically from the rock pigeon. On the other hand, the rock pigeon lives in flocks and usually is much more numerous. The only collected long bone (a distal fragment of a femur) did not allow firm determination, although Tomek & Bochenski (2005) found that the two species differed in size to some extent.
Columba palumbus Linnaeus, 1758 (Pl. 2, Figs. 5-6). The distal pedal phalanx completely corresponded in terms of its bend and higher articular end to that of the wood pigeon. The bows of the symphysal portion of furcular also fitted that of C. palumbus. The wood pigeon is the only columbid species of Europe, which is much larger than the other pigeons and doves. Known from the EP to LP of numerous Eurasian localities from Spain to Georgia (Tyrberg, 1998;2008). In Greece, the species is known from the MP of Petralona (Tyrberg, 1998) and the Liko Cave (Crete) (Tyrberg, 2008). The locality falls within the recent breeding range of the species (Hagemeijer & Blair, 1997). It occurs mainly in broadleaf forests and forest edges (Harrison, 1982).

Caprimulgiformes Ridgway, 1881
Caprimulgidae Vigors, 1825 ?Caprimulgus sp. (Pl. 2,Fig. 12). The find was tentatively referred to nightjars. Young individual with still distally infused metarsalia. General morphology suggested Caprimulgus. Known from the MP sites of France, Italy and from the LP of Croatia and Romania. Caprimulgus europaeus is known from the LP of the Liko Cave (Tyrberg, 1998;2008). The locality lies within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in open or broken broadleaf, mixed or conifer forests, on forest edges and in clearings, in birch and poplar scrubland grass and scrub steppes and groves (Harrison, 1982 (Tyrberg, 1998;2008). So far, the species has not been established in the fossil record of Greece. The locality lies outside of the recent breeding range of the species but falls within its present wintering range (Hagemeijer & Blair, 1997), although on the Balkans it is a rare winter visitor (Cramp & Simmons, 1980). It occurs mainly in open country, on bare moist tundra and shrub to forest tundra, bare upland moors and heathlands, mountains (Harrison, 1982 (Tyrberg, 1998). In Greece, the species is known from the ?LP of the Liko Cave (Crete) (Tyrberg, 1998;2008). Recently found in the LP in Mališina Stijena (Montenegro; Bogoćević & Dimitriević 2004). The locality falls within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in open habitats (from tundra and moorlands to steppes and scrubs) and rarely in forested areas. Widespread, but of very sporadic distribution (Harrison, 1982).   (Tyrberg, 1998;2008). So far, the species has not been established in the fossil record of Greece (Tyrberg, 1998). The locality lies outside of the recent breeding range of the species (Hagemeijer & Blair, 1997) and probably it has survived in the neighboring regions in the country. Occurs mainly in open areas, areas with scattered trees, riverine areas, open forests and plaines, grass and tree steppes and mountain plateaus (Harrison, 1982). The find could be firmly determined only as "large falcon", i. e. Falco sp. ex gr. F. cherrug.
Falco sp. cf. tinnunculus (Linnaeus, 1758) (Pl. 1, Fig. 3). All finds were pedal phalanges of a small falcon of the "tinnunculus" group. The morphology and size excluded F. naumanni and F. vespertinus and inclined to the common kestrel. Known from numerous Eurasian localities of the EP to LP from Spain to Syria and China (Tyrberg, 1998;2008). In Greece, the species is known from the LP of Kitsos (Atika), the Liko Cave (Crete), the Tylos, Vraóna (Attiki) (Tyrberg, 1998). The locality falls within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in open country, grassy or partly forested areas (Harrison, 1982).
Falco sp. cf. vespertinus Linnaeus, 1766. The four pedal phalanges of a small falcon resembled most to the ones of the red-footed falcon. Known from the EP of Hungary and Romania; the MP of f. Czechoslovakia, France and Hungary and from the LP of Bulgaria (cf.), Switzerland, Croatia, France, Georgia, Hungary, Italy, Moldavia, Romania, Spain and Ukraine (Tyrberg, 1998;2008). Recently found in the LP of Mališina Stijena (Montenegro; Bogoćević & Dimitriević, 2004). The locality lies outside of the recent breeding range of the species, but falls within its present wintering range (Hagemeijer & Blair, 1997). Occurs mainly in drier open areas with scattered trees, woodlands, forest steppes, riverine forests and forest edges (Harrison, 1982).
Falco sp. ex gr. tinnunculus. The size and morphology of this pedal phalange firmly determines it only as a "small falcon", i. e. Falco sp. ex gr. tinnunculus.

Strigidae (Vigors, 1825)
Bubo bubo (Linnaeus, 1758) (Pl. 2, Fig. 10). All finds (a tmt and seven pedal phalanges) of an owl fitted dimensionally only to the largest Eurasian owl, the eagle owl. Known from the EP to LP of numerous Eurasian localities from Spain to Jordan, Iraq, Syria and China (Tyrberg, 1998;2008). So far, the species has been established in Greece in the Upper Palaeolithic deposits of the Klissoura Cave 1 (Bochenski & Tomek, 2010). The Loutra Almopias Cave provides the second species record in Greece. The locality falls within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in rocky or wooded areas, from mountains and forests to low rocky outcrops in grasslands or semideserts, from dense forests to bare areas with groups of trees (Harrison, 1982).
Athene noctua (Scopoli, 1769). (Pl. 2, Fig. 9) The only find (ph. 2 d. III p. dex. dist.) completely corresponded to that of the little owl. Known from EP to LP of numerous Eurasian localities from Spain to Israel, Syria and China (Tyrberg, 1998;2008). In Greece, the species is known from the LP/MP of the SE Pigadia Cave (Karpathos), the LP of the Tylos and Vraóna (Attiki) (Tyrberg, 1998). The locality falls within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in more open country, open and broken forests and groves and areas with scattered trees or scrubs, rocky arid areas, steppes, semidesert and dry rocky mountains (Harrison, 1982).
cf. Otus scops (Linnaeus, 1758). (Pl. 2, Fig. 11) The sharp distal end of the phalanx and the sculpture of the articular surface, as well as its size, corresponded to the Eurasian scops owl. Known from the LP sites of Switzerland, Croatia, f. Czechoslovakia, Hungary, Italy, Moldavia, Portugal, Spain, Ukraine and Uzbekistan. In Greece, the species has been known from the LP of the Gumbes C and the Liko Cave (Crete) and the Tylos Island (Tyrberg, 1998;2008). The locality lies within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in lightly forested regions, open or broken forests, riverine forests, areas with scattered trees (Harrison, 1982).

Piciformes (Meyer et Wolf, 1810)
Picidae Vigors, 1825 cf. Dryocopus martius (Linnaeus, 1758) (Pl. 3, Fig. 1). The large size and general shape (stumpy and strongly curved phal. body) suggested the find belonged to the black woodpecker. All other woodpeckers in the Western Palearctic have much smaller pedal ph. Known only from the LP of France, Georgia and Poland (Tyrberg, 1998;2008). In Greece, the species has not been established in the fossil record (Tyrberg, 1998;2008). The locality falls within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs Late Pleistocene and Earliest Holocene Avifauna from the Loutra Almopias Cave (Macedonia, Greece) mainly in climax forests, conifer forests with pines, mixed forests, climax beech forests, more open forests or partial clearings with dead timber and mountain forests (Harrison, 1982).
cf. Picidae. The second specimen of Piciformes from LAC Ia chamber was smaller than Dryocopus martius, but its morphology suggested that it could be referred to Picidae.
Passeriformes (Linnaeus, 1758) Alaudidae (Vigors, 1825) Galerida sp. cf. G. cristata Linnaeus, 1758 (Pl. 3, Fig.  3). The fragments of humerus and tarsometatarsi were characteristic of species of Alaudidae. The deep fossa pneumotricipitalis, the round caput humeri and the projected inception of crista pectoralis of an alaudid fitted to the ones of the crested lark. The shape of the almost parallel trochleae of the three distal portions of tmt also corresponded to the same species. The size, proportions and the shape of the osteological details pointed to G. cristata. Galerida cristata is known from the EP to LP from numerous Eurasian localities from Spain to Jordan and China (Tyrberg, 1998;2008). The locality falls within the recent breeding range of the species (Hagemeijer & Blair, 1997). It occurs mainly in lowlands, drier less vegetated areas, sandy areas of steppes, semideserts or deserts, sandy coasts and dunes, areas of sparse tufted vegetation, bare sandy or stony soils with scattered scrubs, dry hillsides, river banks, desert edges with scrubs (Harrison, 1982).
Melanocorhypha calandra Linnaeus, 1766 (Pl. 3, Fig. 5). Lark-like morphology of distal tmt was dimensionally closer to the largest European lark, calandra lark. Melanocorhypha calandra is known from the EP to LP of numerous Eurasian localities from Spain to Russia and Israel (Tyrberg, 1998;2008). The locality falls within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in drier warmer grass steppes in lowland areas, grass steppes with some tall herbage, shrubby growths, feathergrass and short Artemisia and arable grounds (Harrison, 1982).
Alauda sp. (Pl.3, Fig. 2). The two distal humeri were bigger than these of Lullula and Calandrella and smaller than the ones of Melanocorypha and Galerida and resembled most to Alauda in size and shape. Finds of Alauda sp. are known from the LP from France, Germany, f. Czechoslovakia and Ukraine (Tyrberg, 1998;2008). In Greece, the genus Alauda has not yet been established in the fossil records (Tyrberg, 1998).
Lulula arborea (Linnaeus, 1758) (Pl. 3, Fig. 4). The distal end of tmt showed the features (dimensions and structure) of a woodlark. The specimen was compared with all European larks. Known from the EP of Spain; the MP of France, Italy, Romania, Spain and the LP of Belarus, Bulgaria (cf.), f. Czechoslovakia, France, Italy, Poland, Romania, Spain, the U.K. and Ukraine (Tyrberg, 1998;2008). Recently found in the LP in Mališina Stijena (Montenegro; Bogoćević & Dimitriević, 2004). The species has not been previously established in the fossil records of Greece. The locality lies within the breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in thin or open birch or oak forests, heathlands with scattered trees, or bordering forests, dry hill slopes (Harrison, 1982). The last author notes that the species is sensitive to small climatic changes, which determines the northern limits of the range.
Hirundinidae Vigors, 1825 cf. Hirundo sp. (Pl. 3, Fig. 6). The only find (distal ulna fragment) was compared with all European swallows and resembled most to Hirundo/Cercopis. Its fragmentarity excluded further identification. Finds of g. Hirundo are widely known from the EP to LP from Belarus to Spain and from the United Kigdom to Kirgizia (Tyrberg, 1998;2008). In Greece, the Hirundo swallows have been established in Vraóna (H. rustica) and in the Liko Cave (H. daurica).
Riparia/ Ptyonoprogne (Pl. 3., Fig. 7) cf. Riparia riparia/ Ptyonoprogne rupestris. The only find (a distal end of tarsometatarsus) was compared with all European swallows and resembles most to Riparia riparia and Ptyonoprogne rupestris. Its fragmentarity excluded further identification. Finds of Riparia and Ptyonoprogne are known from the MP from France to China, from the LP of Belarus to Egypt and from Spain to Georgia (Tyrberg, 1998;2008). These genera have not been previously established in the fossil records of Greece. The locality lies within the recent breeding range of Riparia riparia and Ptyonoprogne rupestris (Hagemeijer & Blair, 1997). It occurs in a Historia naturalis bulgarica 40 (2019) Zlatozar Boev, Evangelia Tsoukala broad range of habitats: from sandy estuaries and seacoast banks to rocky outcrops and cliffs and drier mountain slopes (Harrison, 1982).

Motacillidae Vigors, 1825
Anthus sp. (cf. A. trivialis) (Pl. 3, Fig. 8). The only find (a distal end of tarsometatarsus) was compared with most of the European wagtails and pipits and resembles most to the tree pipit, but further determination would have been speculative. The find was strongly elongated and typical for g. Anthus.
Remarks: Anthus sp. is known from the MP to LP of f. Czechoslovakia, Georgia, Italy, Belgium, Switzerland, Hungary and Uzbekistan (Tyrberg, 1998;2008). The water/rock pipit complex (A. spinoleta/petrosus) is considered "montane" element in the LP deposits throughoout Europe (Tyrberg, 1991). Recently, A. cf. campestris has been found in LP of Mališina Stijena (Montenegro; Bogoćević & Dimitriević, 2004). In Greece, the genus Anthus has not been established in the fossil record so far (Tyrberg, 1998). Pipits occur both in areas of open and scattered trees and shrubs and/or open treeless habitats (Harrison, 1982).

Bombycillidae Swainson, 1831
Bombycilla garrulus (Linnaeus, 1758) (Pl. 3, Fig. 9). The only find (a distal end of tarsometatarsus) showed clear similarity to the waxwing. The shape and position of foramen vascular distale and the extensor groove, as well as the deeper relief of the three trochleae completely fitted the description of B. garrulus. Known from the MP of France and from the LP of Bosnia-Herzegovina, Croatia, f. Czechoslovakia, France, Hungary, Italy, Poland, Romania and the United Kingdom (Tyrberg, 1998;2008). So far, the species has not been established in the fossil records of Greece. The locality lies outside of the recent breeding range of the species, but falls within its present wintering range (Hagemeijer & Blair, 1997). It occurs mainly in thick conifer or mixed forests with a berry-bearing shrub layers, riverbanks, forest tundra. In the wintertime accomplishes irruptive movements (Harrison, 1982).

Cinclidae Sundevall, 1836
Cinclus cinclus (Linnaeus, 1758) (Pl. 3, Fig. 10). All pedal phalanges were distal (claw) phalanges and bore specific features for the dipper, i. e. high and compressed phalangeal body, deep lateral grove and deep relief of the articular facies. The similarity with C. cinclus was unequivocal. Known from the MP of Germany and France and from the LP from the U.K.
to Belarus and from Germany to Romania (Tyrberg, 1998;2008). So far, the species has not been established in the fossil records of Greece. The locality lies within the recent breeding range of the species (Hagemeijer & Blair, 1997). It occurs near clear running waters, mainly with a stony or gravelly bed and usually around rocky mountain streams and small rivers (Harrison, 1982).
Turdidae Bonaparte, 1838 (Pl. 4, Fig. 1). This find was heavily damaged. Morphologically resembled to g. Turdus and dimensionally was closer to T. viscivorus. In all remaining European thrushes tmt is smaller. Known from the EP to LP from numerous Eurasian localities from Spain to Russia and Iraq (Tyrberg, 1998;2008). In Greece, the species is known from the LP of the Kitsos Cave (Attiki) (Tyrberg, 1998). Remains of Turdus pilaris/viscivorus have been found in the Liko Cave (Crete) (Tyrberg, 2008). The locality falls within the recent breeding range of the species (Hagemeijer & Blair, 1997). Turdus viscivorus occurs mainly in mature open forests with little undergrowth, in mountain forests and in winter more frequently in open grasslands (Harrison, 1982).
Turdus sp. The bone fragment was so damaged, that even generic identification was difficult. Although it was heavily damaged, dist. epiphysis of the tmt was typical for g. Turdus. In Greece, remains of Turdus sp. are known from the LP of the Klissoura Cave I (Argolis) (Tyrberg, 2008;Bochenski & Tomek, 2010); the ?LP of Gumbes B, the Liko Cave (both in Crete) and the LP of the Tylos Island (Tyrberg, 1998;2008).

Sylviidae Vigors, 1825
Sylvia sp. cf. S. borin (Boddaert, 1783) (Pl. 4, Fig. 2). The only and fragmentary find of a tiny distal tmt was compared with most of European Sylvia warblers and the kinglets; the preserved features allowed referring it only to g. Sylvia. Known from the MP of the U.K. and from the LP of Croatia and Israel (Tyrberg, 1998;2008). The species has not been previously established in the fossil records of Greece. The locality lies within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in the shrubby undergrowth in mixed and broadleaf forests, clearings, birch scrubs; in the southern parts of its range occurs mainly in mountain forests (Harrison, 1982).
Erithacus/ Luscinia (Pl. 3, Fig. 12). The only find was a distal tmt which was compared with species of all European genera of smaller muscicapids. The distal epiphysis was much similar than that of Erithacus/Luscinia. Erithacus rubecula has been found in some ?LP layers of the deposits in the Liko Cave (Crete) (Tyrberg, 2008). Erithacus and Luscinia are known from the EP of Spain and f. Czechoslovakia and from the LP from the U.K. to Georgia and Israel. Erithacus rubecula is known from the LP of the Liko Cave (Tyrberg, 1998;2008). The locality lies within the resent breeding range of the E. rubecula (Hagemeijer & Blair, 1997). The species of the Erithacus-Luscinia complex occur in thick forests with shrubby undergrowth, marshy areas, clearings, in stunted conifers and birch zones, alpine meadows and lowland areas and on hills usually in broadleaf forests (Harrison, 1982).

Paridae Vigors, 1825
Parus major Linnaeus, 1758 (Pl. 4, Fig. 3). The only find was a distal tmt which was compared with species of all European tits. Dimensionally and morphologically completely fitted to the great tit. Known from the EP of Romania, Spain and Ukraine; the MP of Austria and China and from the LP between the U.K. and Ukraine (Tyrberg, 1998;2008). The locality lies within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in the broadleaf forests, but also in conifer or mixed forests and groves (Harrison, 1982).

Sittidae Lesson, 1828
Sitta cf. europaea Linnaeus, 1758 (Pl. 4, Fig. 4). The only find was a complete claw phalanx of the third (largest) toe. It was thinner and longer and all other features entirely matched those of the corresponding skeletal element of the Eurasian nuthatch. Known from the EP of Austria; the MP of f. Czechoslovakia and the U.K. and from the LP of Austria, Bulgaria, Croatia, Switzerland, France, Poland, f. Czechoslovakia and Romania (Tyrberg, 1998;2008). The locality lies within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in broadleaved, mixed or conifer forests; often in well-grown more open forests with tall trees, mainly in the mountains but also in the open hillside forests (Harrison, 1982).
Sitta sp. The lateral condyle of tr. m. III wider than in S. europaea. Nuthatches are common in many Late Pleistocene localities in Europe, including neighbouring Bulgaria (Boev, 2001b).
Pica sp. The find (Fig. 3) is dimensionally smaller than C. monedula Linnaeus, 1758, P. graculus, P. pyrrhocorax and bigger than N. caryocatactes, P. pica, C. cooki and G. glandarius (Table 4). According dos Anjos (2009), P. infaustus is the smallest European corvid in terms of body length and mass, except for C. cooki (Table 5), but this species has not been compared. Pica pica is the most similar species but LAC B13 differed from it by its larger size. From N. caryocatactes it differed by the better developed narrowing ("neck") of prox. end and the much shallower longitudinal concavity on the dorsal surface of the bone. LAC B13 differed from P. pica by the relatively longer "neck" of prox. end.

Zlatozar Boev, Evangelia Tsoukala
On the other hand, morphologically it significantly resembled P. pica but was much bigger in size. From C. monedula it differed by the steeper (sharper) widening after "neck" and the shallower relief of the dorsal surface. LAC B13 specimen was compared with the measurements of the European corvids (Tomek & Bochenski, 2000). It did not fit to any of the 662 European specimens of corvids, measured by Tomek & Bochenski (2000 : Table XII, p. 50), including P. infaustus. As seen in Table 4, the measurement "a" much approached only to the maximum value of N. caryocatactes (13.9), the measurement "b" lied within the ranges of P. pica (3.2-4.9), P. graculus (3.7-5.1) and C. monedula (4.2-5.1), the measurement "c" lied within the ranges of G. glandarius (2.2-3.0), P. pica (2.3-3.0), P. graculus (2.7-3.7) and C. monedula (2.9-3.6), whereas the measurement "d" lied within the ranges of P. graculus (2.1-2.2) and C. monedula (1.8-2.4). In comparison with these species, LAC B13 specimen differed from: C. monedula by the much smaller (shorter) phalanx (measurement "a") and the more angular ventral end of distal f. a.; P. pica by the bigger size, thinner ventral edge on the medial side and the better developed pila cranialis; P. graculus by the much smaller size and the relatively thicker "neck" in the proximal end; G. glandarius and C. cooki by the much bigger size. Index "total length: proximal width" (Table 4) of B13 was 3.404, whereas for P. pica it was 3.378. This was the smallest difference when compared to the values of this index among other small corvids in Europe: C. cooki -3.076; G. glandarius -3.171, N. caryocatactes -3.500, P. infaustus -3.520, P. pyrrhocorax -3.627, C. monedula -3.630 and Pyrrhocorax graculus -3.638. Mlíkovský (2002) synonymised Pica pica major Janossy 1972 from Stránská Skála I (Czechia) with C. monedula. After the original diagnosis of D. Janossy, skeletal elements of this subspecies were bigger than those of the modern European magpie. Pica p. major originates from Southern Czechia (Stránská skála) and is dated to EP (MQ 1b) (Mlíkovský, 2002). Although the total length of the humerus and femur of P. p. major exceeds these measurements in modern P. pica (by 13.52 % and 13.83 %, respectively; calculations after measurements of Janossy, 1972), because of the lack of analogous skeletal elements for comparison and the considerable chronostratigraphical difference (ca. 0.6 Ma), we prefer to designate the EEH magpie from the Loutra Almopias Cave to Pica sp. and not to refer it tentatively to the P. p. major (even less so to C. monedula after Mlíkovský, 2002).
Pyrrhocorax graculus (Linnaeus, 1766) (Pl. 4, Fig. 10). The Alpine chough was the most numerous species among the material. All skeletal elements bore the features of a large passeriform and both morphologically and dimensionally resembled completely P. graculus. Known from the EP to LP of numerous Eurasian localities from Spain to Israel, Georgia and Azerbaijan (Tyrberg, 1998;2008). Recently found in the LP of Mališina Stijena (Montenegro; Bogoćević & Dimitriević, 2004). In Greece, the species is known from the LP of Elaiochoria (Chalkidiki); the ?LP of Gerani IV (Crete) and of the Liko Cave (Crete) and the LP of Vraóna (Attiki) (Tyrberg, 1998;2008). The locality falls within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in areas where steep rocky slopes adjoin open grasslands in the mountains (Harrison, 1982).
Pyrrhocorax cf. pyrrhocorax (Linnaeus, 1758) (Pl. 4, Fig. 9). All finds (distal tmt and pedal phalanges) represented large passerines (corvids). The size and robustness of the tmt-fragment distinguished it from P. graculus and completely fitted to the redbilled chough. Widely known from the EP to LP of numerous Eurasian localities from Portugal to China (Tyrberg, 1998;2008). In Greece, the species is known from the LP of Vraóna (Attiki) and of the Tylos; the?LP of the Liko Cave (Crete) (Tyrberg, 1998;2008), (cf.) from the LP of the Klissoura Cave 1 (Bochenski & Tomek 2010). The locality lies outside (but not very remote; ca. 150 km away) the recent breeding range of the species (Hagemeijer & Blair, 1997). It occurs mainly in areas where steep cliffs and crags occur close to open grassy areas in the mountains (Harrison, 1982).
Corvidae gen. indet. -2. Dimensionally close to C. monedula and P. graculus, but the shape of cotyla humeralis was elliptical instead round. Dimensions of the specimen B 342 lied within the ranges of three species: C. monedula, P. graculus and P. pyrrhocorax ( Table 6) after data of Tomek & Bochenski (2000). On the other hand, morphologically this specimen differed from P. pyrrhocorax by the elliptical shape of cotyla humeralis and the slender shaft (diaphysis). It differed from P. pica by the elliptical shape of cotyla humeralis and the rounder instead angular section of the proximal diaphysis. From C. monedula it also differed by the elliptical shape of cotyla humeralis and the more uprised tuberculum bicipitale. From N. caryocatactes it differed by the elliptical shape of cotyla humeralis, more uprised tuberculum bicipitale and the rounder instead angular section of the proximal diaphysis.

Late Pleistocene and Earliest Holocene Avifauna from the Loutra Almopias Cave (Macedonia, Greece)
cf. Corvidae gen. indet. Only the dist. fourth of the wing ph. prox. d. majoris dex. was preserved. Size of G. glandarius but further identification was not possible.

Sturnidae Vigors, 1825
Sturnus sp. cf. S. vulgaris Linnaeus, 1758 (Pl. 4, Figs. 5-6). Known from the EP to LP of numerous Eurasian localities from Spain to Russia, Israel and Iraq (Tyrberg, 1998;2008). In Greece, the species is known from the LP of the Kitsos (Attiki) (Tyrberg, 1998). Remains of Sturnus sp. are known from the ? LP of the Liko Cave (Crete) (Tyrberg, 2008). Recently found in the LP of Mališina Stijena (Montenegro; Bogoćević & Dimitriević, 2004). The locality falls within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs in open montane forests with grassy clearings, riverine forests in meadows and groves (Harrison, 1982). Probably most of the material belonged to S. vulgaris, but due to the higher morphological resemblance of both species and the preservation of the finds, it could be further identified. Theoretically, part of the identified material could belong to S. roseus (Linnaeus, 1758), a species hardly distinguishable osteomorphologically from the common starling. The rosy starling was also regularly found in the Würmian deposits of Europe (Tyrberg, 1991).

Fringillidae Vigors, 1825
Fringilla sp. cf. F. coelebs Linnaeus, 1758 (Pl. 5, Fig.  2). The find (proximal part of a humerus) closely resembled, in terms of the shape of crista pectoralis, tuberculum ventral and caput humeri, to the common chaffinch. Known from numerous Eurasian localities from Spain to China (Tyrberg, 1998;2008). Remains of Fringilla coelebs/montefringila have been reported from the ?LP of the Liko Cave (Crete) (Tyrberg, 2008). The locality falls within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in the more open and marginal areas of all types of forest and small woodlands (Harrison, 1982).
Carduelis sp. cf. C. carduelis (Linnaeus, 1758) (Pl. 4. Fig. 12). Carduelis carduelis is known from the EP to LP of a number of Eurasian localities from Spain to Ukraine and Israel (Tyrberg, 1998;2008). The species has not been previously established in the fossil records of Greece (Tyrberg, 1998). The locality falls within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in the open-foliaged broadleaved trees, weed-grown grounds or mixed herbages, edges and clearings of forests, scattered groups of trees, steppes, riverine forests, shrubs and groves (Harrison, 1982).
cf. Chloris chloris (Linnaeus, 1758) (Pl. 5, Fig.  1). The similarity of these fragmentary find of a proximal cmc (synostosis metacarpalis prox.) was highest to the European creenfinch. Known from the EP and MP of France and Israel, the MP of Malta and from the LP from France to Israel and from the U.K. to Ukraine. Recently found in the LP of Mališina Stijena (Montenegro; Bogoćević & Dimitriević, 2004). In Greece, the species has been found in the LP deposits of the Liko Cave (Tyrberg, 1998;2008). The locality lies within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in the open-foliaged broadleaved forests. This species prefers tall and well-grown trees (Harrison, 1982).
Carduelis sp. The specimen (tmt dex. dist.) was slightly bigger than C. spinus but identification to the species level was not possible.
Loxia curvirostra Linnaeus, 1758 (Pl. 5., Fig. 5). Known from the MP to LP of a number of Eurasian localities from France to Russia and Israel (Tyrberg, 1998;2008). The species has not been previously established in the fossil records of Greece (Tyrberg, 1998). The locality lies outside of the recent breeding range of the species, but it frequently inhabits the Pindos and Rhodopes Mountains in N Greece (Hagemeijer & Blair, 1997). Occurs mainly in conifer (spruce) forests, mixed forests and mountain forests (Harrison, 1982).
Pyrrhula pyrrhula (Linnaeus, 1758) (Pl. 5, Fig.  3). Various skeletal elements were found among the material: mandibulae, humeri, os quadratum and tarsometatarsi. The proximal (articular) fragment of the mandible for example was rather diagnostic with its high ramus mandibulae (in the region of os praearticulare). Known from the MP to LP of a number of Eurasian localities from Spain to Greece (Tyrberg, 1998;2008). Recently found in the LP of Mališina Stijena (Montenegro; Bogoćević & Dimitriević, 2004). In Greece, the species is known from the ?LP of the Liko Cave (Crete) (Tyrberg, 1998;2008). The locality falls within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in conifer and mixed forests and Zlatozar Boev, Evangelia Tsoukala broadleaf forests with thick undergrowth (Harrison, 1982).
Fringillidae gen. indet. Similar to Fringilla coelebs size but further identification was not possible.

Emberizidae Vigors, 1831
Emberiza calandra Linnaeus, 1758 (Pl. 5, Figs. 8-9). The finds were compared with most of the European buntings. The corn bunting is the largest among them and the distal tmt and clow phalanx fully fitted in size and shape of that species. Known from the LP of Israel, Spain, Framce, Germany and Greece (Liko Cave, Crete) (Tyrberg, 2008). The locality falls within the recent breeding range of the species (Hagemeijer & Blair, 1997). In Greece, this species has been established in the LP deposits of the Liko Cave.
Occurs mainly in open grassy areas with some tall herbage or low shrubs, in the vegetation of dry and semiarid regions, grass steppes with sparse shrub, stony steppes with sparse herbage, grassy areas of sandy desert, hilly steppe areas (Harrison, 1982).
cf. Plectrophenax nivalis (Linnaeus, 1758) (Pl. 5, Fig. 7). Seven distal tmt were identified. Known from the MP of France and Ukraine, from the LP from the U.K. and France to Italy and Ukraine (Tyrberg, 1998;2008). The snow bunting has not been previously established in the fossil records of Greece. The locality lies outside both of the recent breeding and wintering range of the species. On the Balkans, the species has been recorded as an accidental winter visitor in Bulgaria, f. Yugoslavia (Cramp & Perrins, 1994) and Greece (Anonym, 2018). Breeds on bare rocky tundra but winters on the stony or sandy shores and dunes of low-laying coastal areas, grass moorlands and grass steppes (Harrison, 1982).
Emberiza sp. cf. E. cirlus Linnaeus, 1766 (Pl. 5, Fig. 6). Known from the LP sites of Croatia, Italy and Spain (Tyrberg, 1998;2008). Previously, the species has not been established in the fossil records of Greece. The locality lies within the recent breeding range of the species (Hagemeijer & Blair, 1997). Occurs mainly in grassy hillsides with scattered trees, open broadleaf forests and groves (Harrison, 1982).
Emberizidae gen. indet. (size of Plectrophenax nivalis). The LAC B 173 bone fragment was so damaged that even generic identification would be impossible, although it demonstrated some resemblance in terms of size and morphology with the buntings (and especially with Plectrophenax nivalis).
Non-Passeriformes indet. The LAC B 55 specimen represented an articular (prox.) half of a large claw ph. with extremely deep grows on its lateral sides. Further identification was not possible.    Fig. 2-B).

Taphonomy
The majority of avian remains recovered in the Loutra Almopias Cave were heavily damaged, usually fragmented. Almost all of the ossa longa tubulosa were broken and mainly epiphyses, or even separate bone splinters (of diaphyses) were present. Sometimes, only fragments of the bone endings were preserved, for example caput humeri, trochlaeae mt., even endings (only f. a.) of the pedal phalanges. In general, for all the collected avian material was characteristic the very high level of fragmentation and complete absence of the traces of erosion. Most numerous were the small/tiny bones as the limb phalanges (mainly pedal), representing 2/3 of the collected material (67%). All avian remains (probably with two exceptions) belonged to mature (adult) individuals. Only several juveniles have been recorded among them. On the other hand, over 2/3 of the finds (73%) were preserved only partially by their prox./dist. half or epiphyses or the phal. bodies. Over 1/4 of the finds (26%) belonged to small Perdicinae species (Table 1). Similar was the share of the corvids, representing 34% of the material. The Alpine chough was the most abundant species in both localities, representing 25.6% and 29.5%, respectively in the LP and EEH localities. Such a representation suggested a natural non-human accumulation of the material, probably indicating a former feeding place of a large raptor (eagle owl) in the cave. The collected material corresponded to the taphonomic conclusions of Bochenski (2005). Some bones (LAC B37 from LAC Ia) demonstrated features of digestion: their bone surface was not smooth and it was covered by a number of tiny pores. Both finds belonged to Perdix perdix from the EEH. On the other hand, the dominance of large to medium-sized preys (grey partridge, rock partridge, rock pigeon, common wood pigeon, yellow-billed Alpine chough), reaching 56.5% of the finds, explained the accumulation of the faunal remains by a large raptor, most probably eagle owl (Andrews, 1990), which was also represented in the collected bone material in the EEH cave locality. A detailed research on the metric characteristics of the recent eagle owl's preys in the region (Balkans; S Bulgaria), has found that the common bird preys of B. bubo are bird species with body mass between 100 and 500 grams (Boev, 1993). Namely, the five species listed above fall within this metric range. Although suggesting the same taphonomy, the comparison of both avifaunas of the Loutra Almopias Cave showed that the LP avian locality was over five times poorer and provided 2.2 finds/taxa as compared to 10.8 finds/taxa from the EEH locality.
The anatomical representation of the avian skeletal elements was a rather unusual specific feature of both avian localities in the Loutra Almopias Cave. Over 68.4% (377 of the total of 551 finds) were phalanges and only three of them were wing phalanges. Another interesting feature was that 99.0% of the finds belonged to adult individuals and only six finds could be aged as pullets, juveniles or sub-adults.  A. graeca, C. livia, B. bubo, P. graculus), openlands with scattered trees and shrubs (Anthus sp., P. pica). The complete absence of aquatic birds might be a result of the appearance of thermal springs in the area that influenced the hydrobionts, essential prey and food resource for many hydrophylous birds. Such thermal springs exist at present, but they could change their temperatures and appear and disappear during the millennia. The eagle owl (present in the fossil material) was the most probable accumulator of the faunal remains, which was suggested by the record of medium to large-sized birds, such as buzzards and black grouse, the high fragmentation of bones and lacking of skulls and the rare mandibles in the material (Bochenski et al., 1993).

Paleoenvironment
Earliest Holocene (upper chamber LAC Ia): EEH record (48 species/genera; 510 finds). The uncovered fauna included elements of the "boreal complex" (7 species), inhabiting mainly coniferous forests (T. represented by a relatively large number of finds (11 finds; 2.2 % of the LAC assemblage). The whitethroated dipper is an indicator of the fast-running waters, i.e., rocky mountain streams and small rivers in the wooded mountains (Harrison, 1982 Francolinus francolinus could also be referred to this group but this species usually prefers moister areas, shrubby thickets bordering water bodies and areas of scrubby herbage. Such a wide range of habitats may be explained by a mosaic-type distribution in the very end of the Pleistocene. It is well known from other LP localities in Europe and the continental parts of the Balkans (Bulgaria -Kozarnika Cave, Boev, 2001a, Cherdzhenitsa Cave, Boev, 2000a, Razhishka Cave, Boev, 2000b; and many others).
F. francolinus, B. lagopus, T. tetrix, L. lagopus and P. nivalis have disappeared from the recent bird fauna of Greece. The last four species have Arctic-Alpine ("boreal"), i.e., Arctic/Subarctic distribution and are presently found in high mountains during the summer, while the black francolin gradually reduced its breeding/resident range and disappeared from Europe (Harrison, 1982;Hagemeijer & Blair, 1997).
The occurrence of Lagopus, a typical representative of the modern Arctic/Boreal fauna (and partly of T. tetrix), is among the southernmost Pleistocene records for this genus. This record (in both localities, LP and EEH) likely suggests that the genus Lagopus (as noticed by Bochenski, 2007) did not use to be strictly associated with a cold climate as it is today. It must be noted, that overall the origin of all the members of the family Tetraonidae (including genera Tetrao and Lagopus) was not associated with the cold climate (Boev, 1999;. Present-day climatic (temperature) preferences of some species (T. tetrix, B. bonasia, F. francolinus, found in both cave localities, LP and EEH) indicate that the July isotherm (Harrison, 1982) of 21-26º C encompassed the region of the cave and the climate probably used to be similar but slightly colder than the present one.
Faunal comparison of both avian localities in the Loutra Almopias Cave showed another clear difference: a number of species/taxa, inhabiting mainly woodland habitats, established in the EEH locality were completely absent in the older LP locality. These 15 (17) species/taxa were: L. arborea, B. garrulus, S. borin, S. europaea, T. viscivorus, Erithacus/Luscinia, G. glandarius, F. coelebs, C. chloris, S. vulgaris, P. major, D. martius, A. noctua, O. scops and Caprimulgus sp. Such a difference could be explained probably with the comparatively more humid (and relatively moderate) climate, allowing wide landscape forestation of the region surrounding the cave during the EEH. The large wooded areas probably used to be dominated by various broad-leaved species of trees (at least the distribution of the Eurasian jay follows the oak forests). In addition, the numerous records of the whitethroated dipper indicated the presence of running nonfreezing waters (woodland streams and small rivers) in the mountains. C. cinclus and Ph. ochruros are more closely related to rocky riverine habitats; nevertheless often they lie within the woodland ones.
In both localities the great majority of species represented in the fossil material belonged to the modern recent Balkan avifauna. The great majority of them were represented by their present-day breeding populations: 85.7% of species/taxa for the LP and 92.6% for the EEH.
It is interesting to note, that more of the boreal species have been found in the EEH locality. The paleoenvironment probably used to be more open than today and the mosaic woodland-open grassland landscape dominated. Forty-seven of the recorded species/taxa (69.1%) from the Pleistocene deposits of the Loutra Almopias Cave are still spread in the region of the cave in the Voras Mts. (Table 1; Hagemeijer & Blair, 1997). Among the disappeared species, the "boreal" and open-land steppe birds prevailed, indicating once again the drier and cooler former climate in the region in comparison to the local modern climate.
It is notable, that the fossil record of the Loutra Almopias Cave documented the continuous distribution of only eight species (11.8%) -Falco sp. cf. F. peregrinus, A. graeca, C. palumbus, B. bubo, Anthus sp., P. graculus, C. coccothraustes and P. pyrrhula, that have been spread both in the LP and EEH and at present in the vicinity of the cave. On the other hand, ten species/genera (14.7 %) -Falco sp. cf. F. peregrinus, A. graeca, cf. B. bonasia, C. palumbus, B. bubo, Anthus sp., Turdus sp., P. graculus, C. coccothraustes, P. pyrrhula, have survived since the LP to the present in the region of the cave and 30 (44.1 %) of the species that inhabit the cave vicinity at present, used to existen-Zlatozar Boev, Evangelia Tsoukala ce in the region even in the EEH, but were not recorded in the LP layers (see Table 1). Most of them are typical woodland birds, e.g. cf. Dryocopus martius, Lulula arborea, Bombicylla garrulus, Turdus sp. cf. T. vuscivorus, Erithacus/ Luscinia, Parus major, Sitta cf. europaea, Garrulus glandarius and Fringilla sp. cf. F. coelebs. One of them (Cinclus cinclus) is also a specific aquatic inhabitant.

Concluding Remarks
The fossil avian record of the Loutra Almopias Cave, from both sites in the cave of different age, corresponds to 15.4% of the recent avifauna of Greece, numbering 443 species (Lepage, 2017). In general, the fossil material proved that during the Late Pleistocene -Earliest Holocene at least 73.3% (33 species/taxa) of the total of 45 species/taxa at present used to be represented by their resident populations, seven are summer migrants and four are winter migrants.
Six species of the uncovered fauna disappeared in the Holocene from the modern avifauna of Greece: B. lagopus, F. francolinus, T. tetrix, L. lagopus, F. vespertinus, and P. nivalis. The present breeding range of B. lagopus and P. nivalis is retreating northwards. Only one species (Fr. francolinus) disappeared from the modern bird fauna of Europe.
The range of F. francolinus retreated southeastwards in the Late Holocene, whereas T. tetrix and F. vespertinus still are breeding species in the neighbouring Bulgaria (Hagemeijer & Blair 1997). Sometimes B. lagopus, B. garrulus and even P. nivalis make irregular irruptive migration to the Balkan region in the coldest winter periods (Cramp & Simmons, 1980;Cramp & Perrins 1994).