GEOLOGICA BALCANICA, 29.3-4, Sofia, December 1999, p. 51-57

Earliest finds of cross bills (genus Loxia) (Aves: Fringillidae) from Varshets (NW Bulgaria)

Zlatozar Boev

National Museum of Natural History, Bulgarian Academy of Sciences, 1000 Sofia

(submitted 26.01. 1999; accepted 02.02. 1999)

Iioes, 3. H. CaMble c}pesHue HaxooKu ICJlecm06 (poo Loxia) (Aves: Fringi//idae) u3 Bblpweqa, Cesepo-3anadHa.R IioAza· pUR. Cn.enaHo o6oJpeHKe 4JocKnbHOii neronKCK po.na Loxia. lionrapCKKe HaXO.IlKK npOKCXO.IlliT K3 4 MeCTOHaxOliC.IleHKH -3 K3 HKX K3 aepxHero nneKCTOileHa (BIOpM) K 1 K3 cpeD.Hero aKnnacjlpaHKa (oKono 2,2 MnH. ner). nocne.nHKH HJ HHX (no­Hop a cKacHCTOM xonMe a OJCPCCTHOCTJIX r. Bwpweua, Ceae­po-3ana,nHall lionrapHll) CO.IleplKHT CaMLie .npeBHHe OCTaTKK KJJeCTOB. 0HH OTKOCJITCJI JC HOBOMy HeHJBeCTKOMy D.O CHX nop BHD.y, Loxia patevi sp. n.

Abstract. A review of the fossil history of g. Loxia is presented. Bulgarian record of crossbills consists of several finds of 4 sites - 3 of Upper Pleistocene (Wurmian) age and one of Middle Villafranchian (ca. 2,2 mya). The last site (a ponor near the town of Varshets, NW Bulgaria) has provided the earliest record of the genus. These finds are described as Loxia patevi sp. n.

Boev, Z. 1999. Earliest finds of crossbills (genus Loxia) (Aves: Fringillidae) from Varshets (NW Bulgar­ia).- Geologica Bale., 29, 3-4; Sl-57.

Key words: fossil birds, passeriformes, Pliocene avifaunas, Bulgaria, Villafranchian.

The common crossbill Loxia curvirostra Linnaeus, 1758-short ecobiogeographical notes

The common crossbill at present is a resident, no­madic and irruptive species in boreal to warm tem­perate zones (Fig. 1 ). It prefers mature conifer for­ests. Northern populations move to south up to 500-1 000 km in the periods of food deficiency (mainly seeds of conifers). In the mountains in southern parts of the range, it occurs up to c. 4500 m a.s.l. (Harri­son, 1982). Inhabits both inner parts of large woods and forest edges, more often of Picea, Pinus and Lar­ix. Its distribution depends on the nearness of water. The present range south of 55° is very broken (Cramp, Perrins, 1994).

Tyrberg (1991) concludes that during the Wurm glaciation the main population of L. curvirostra was confined in Western Europe. The crossbills survived in the Balkans in limited refuges of Picea forests. In the Holocene, following distribution of conifer forest in northern parts of the present

range, the crossbills restored the range northwest­wards, where the subspecies L. c. scotica and L. c. pytyopsittacus appeared. Cramp, Perrins (1994) con­sider them separate species -Loxia scotica (Harter, 1904) and Loxia pytyopsittacus (Borkhausen, 1793).

Fossil records of genus Loxia

No fossil species of g. Loxia have been described up to now (Brodkorb, 1978; Olson, 1985; Tyrberg, 1991, 1998 a; Mlikovsky, 1996; Bochenski, 1997).

Brodkorb (1978) reports on various Pleistocene finds of L. curvirostra from France, former Czech­oslovakia, Hungary, Ukraine, California, New Mexico and Illinois. Tyrberg (1991) lists 32 Pleis­tocene localities from Great Britain, France, Ger­many, Hungary, former Czechoslovakia, Italy, Ru­mania, Austria, Croatia, Malta and Israel. The common crosbill is known also from the following Quaternary sites: Pleistocene of Pusskaporos (Ko­rmos, 1911), Middle Wurm in Istallosko, Remete­hegy and Upper Wurm of Pillisszanto-1 (Janossy,

51

1986) in Hungary; Upper Pleistocene of Mallorka (McMinn, Alcover, 1992); Middle Pleistocene (150 000 - 125 000 B.P.) from Grotte du Lazaret (Vi­lette, 1993), Wurm (Magdalenian) in Blassac (Bou­chud, Bouchud, 1955), Tardiglacial - Lower Ho­locene in Jean-Pierre I (Mourer-Chauvire, 1994), Wurmian 4 of Massat in Pyrenees (Clot, Mourer­Chauvire, 1986) in France; Riss-Wurm in South Devon (Harrison, 1980), Lower Pleistocene in Wye Valley Cave (Harrison, 1980); Upper Holocene (2000 B.P.) in Ossom's Eyrie Cave (Bramwell et al., 1990) in Great Britain; Lower Paleolithic in Grotta dei Fanciulli in Balzi Rossi (Campana, I 946), Up­per Pleistocene in Riparo di Fumane (Cassoli, Tagliacozzo, 1994), end of Wurm 3 (20 000 B.P.) -middle ofWurm 4 (I2 500 B.P.) in Arene Candide (Cassoli, 1980), Wurmian (Origniac) of Grotte de Fumane (Bartolomei et al., 1992) in Italy; Wurmi­an 2-3 in La Balauziere in Monaco (Bonifay, 1966); Pleistocene of Crimea (Dementiew, 1960); Middle

and Upper Pleistocene of Mall orca (AI cover et al., 1992); Paleolithic in Syuren I in Crimea (Barysh­nikov, Potapova, 1992). Laterly Tyrberg (1998) in his catalogue of the Pleistocene birds of the Pale­arctic summarizes all data and lists 26 sites of Up­per Pleistocene, I site of Middle Pleistocene and I site of unknown Pleistocene age.

Thus, the fossil record of g. Loxia encompasses Lower (? Middle) Pleistocene (Wye Valley Cave (Harrison, 1980)) to Holocene. According to the exhaustive review of Tyrberg (1991 ), the oldest finds of g. Loxia originate from the Stranska Skala Cave in Czech Republic. They are dated initial phases of the Middle Pleistocene (c. 0,2 mya) (Janossy, 1972). According to Tyrberg (1998), the two other Paiearctic crossbills, L. leucoptera, and L. pytyopsittacus are known from the Upper and the Middle Pleistocene respectively. Thus, no evi­dences for the Tertiary history of the genus are available up to now.

Fig. I. Recent range of the genus Loxia in Europe (by Cramp, Perins, 1994) (black) with the location of the Bulgarian sites. Upper Pleistocene (Wurmian): I - Bacho Kiro Cave, 2 - Cave 16, 3- Kozarnika Cave; Upper Pliocene (Middle Villafranchian): 4-Varshets (drawing: Vera Hristova)

52

Fossil data on crossbills from Bulgaria

First data on fossil finds of crossbills (Loxia curvi­rostra) from Bulgaria have been published by Bo­chenski (1982) from the Wurmian deposits of the Bacha Kiro Cave (Central Stara Planina /Cental Balkan Mountains/).

Two other finds of L. curvirostra of Upper Wur­mian (carpometacarpus dex., NMNHS No 794, and carpometacarpus sin., NMNHS No 795) are collected by Dr Vassil Popov in the Cave No 16 (The Temnata Cave system; West Balkan Moun­tains), and other 6 (2 tibiotarsi dex. dist., NoNo 8905, 8906; 3 humeri dex. dist., NoNo 8481-8483; carpometacarpus sin. prox, No 8484) of approxi­mately same age in the Kozarnika Cave (near the town of Belogradchik; West Balkan Mountains) are collected by Dr Nikolay Sirakov and Ms Mar­garita Marinska (Boev, 1998).

The presence of Loxia remains and the first data on the fossil avifauna ofVarshets have been reported by Boev (1995 a, b; 1996; in press). It is evident from the review of fossil data that the Bulgarian finds from Varshets are the oldest record of g. Loxia. They prove the existance of crossbills in the southern parts of the Western Palearctic since the last ca. 2,2 mya.

The material ofVarshets consists of 5 bone frag­ments: humerus dex. dist. - NMNHS 307 (fig. 2 -a), humerus sin. prox. - NMNHS 308 (fig. 2 -b), humerus sin. prox.- NMNHS 309 (fig. 2- c), tibio-

Table I

tarsus dex. dist. - NMNHS 31 0 (fig. 2 - d, e), and ulna dex. prox. - NMNHS 311 (fig. 2 - f, g). Having in mind the rapid Neogene-Quaternary evolution of passerform birds, the established morphological diferences and the considerable age of the finds, we distinguish them into a new separate species.

Taxonomical description

Loxia patevi sp. n.

Ho/otype. Humerus dex. dist. (fig. 2a), Collections of the Recent and Fossil Birds Department of the Na­tional Museum of Natural History, Bulgarian Acad­emy of Sciences, cat. No 307, collected by Z. Boev.

Locality. A ponor in a rocky hill, 6 km NNE of Varshets (43,13 N, 23,17 E). Unconsolidated, un­stratified sediments accumulated in the filling of clay terra-rossa. The fossil bones are broken, sometimes making a kind of bone breccia.

Chronostratigraphy. Middle Villafranchian. The associated fauna of mammals (Spassov, 1995; V. Popov- pers. comm.) attributes the site to the MN 17 zone according to chronostratigraphical system of Mein (1990).

Etymology. The name "patevi" is given after the name of the eminent Bulgarian ornithologist, Dr Pavel Patev (13.1 0.1 889 - 22.03.1950), the author of the monograph "The Birds of Bulgaria" (1950).

Measurements. See tables 1 - 4.

Measurements of the proximal humerus of fossil and recent Loxia (ref to fig. 3)

Species a b c d

Fossil Loxia patevi sp.n. - NMNHS 308 6,9 2.2 3,4 2.1 Loxia patevi sp.n. - NMNHS 309 6,7 2,4 3,4 2,1 Recent Loxia curvirostra - UCBL 430/1 7,3 2,3 3,4 2,2 Loxia curvirostra - UCBL 430/2 7,1 2,5 3,1 2,0 Loxia curvirostra - UCBL 430/3 7.9 2.7 3,7 2,2 Loxia cun1irostra - NMNHS 3/1992 7,4 2.45 3,3 2,1 Pyrrhula pyrrhula · UCBL 427/5 5,9 1,9 2,6 1.7 Pinicola enuclearor- UCBL 429/1 7,9 2,6 4,0 2,4

Table 2 Measurements of the distal humerus offossil and recent Loxia (ref to fig . 3)

Species I a b c d

Fossil Loxia patevi sp.n. - NMNHS 307 2,7 5,6 1,6 1,05 Recent Loxia C'.4rvirostra - UCBL 430/1 2,9 5,9 1,6 1,2 Loxia curvirostra - UCBL 430/2 2,75 5,7 1,6 1,0 Loxia curvirostra - UCBL 430/3 3,2 6,2 1,8 1,2 Loxia curvirostra- NMNHS 3/1992 2,9 6,1 1,7 1.4 Pyrrhu/a pyrrhula- UCBL 427/3 2,7 5,9 1,5 1,3 Pinicola enuc/eator- UCBL 429/1 3,1 6,9 1,8 1,1

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Table 3 Measurements of the proximal ulna of fossil and recent Loxia (ref to fig . 3)

Species I a b c d

Fossil Loxia patevi sp. n . - NMNHS 311 3,6 2,0 3,9 3,7 Recent Loxia curvirostra- UCBL 430/1 3,6 2,0 3,8 3,8 Loxia curvirostra - UCBL 430/2 3,6 2,0 3,7 3,7 Loxia curvirostra - UCBL 430/3 3,9 2,2 4,0 4,0 Loxia curvirostra - NMNHS 3/1992 3,8 2,2 3,8 4,0 Pyrrlwla pyrrhula- UCBL 427/3 3,7 1,9 3,5 3,8 Pinicola enucleator- UCBL 429/1 4,2 2,3 4,2 3.4

Table 4 Measurements of the distal tibiotarsus of fossil and recent Loxia (ref to fig. 3)

Species a

Fossil Loxia patevi sp.n. - NMNHS 310 2,1 Recent Loxia curvirostra- UCBL 430/1 2,2 Loxia curvirostra - UCBL 430/2 2,2 Loxia curvirostra - UCBL 430/3 2,0 Loxia curvirostra- NMNHS 3/1992 2,2 Pyrrhula pyrrhula UCBL 427/3 Pinicola enucleator UCBL 429/1

Comparison. The general shape of the bone sug­gests a Fringillidae humerus. It defers from Pyr­rhula by the shallower and more unclearly formed fossa olecrani. The differences from Pinicola con­sist of the less prominent laterally processus su­pracondylaris dorsalis. Both, dimensionally and morphologically Loxia patevi sp. n. is very close to the recent crossbill L. curvirostra, but some differ­ences are clearly represented.

The proximal end of humerus is close to Loxia and defers from Pinicola by: 1) the absence of fos­sa pneumotricipitalis; 2) the longer crista del­topectoralis and 3) more distally positioned linea intermuscularis on the crista deltopectoralis.

The proximal ulna of L. patevi sp. n. defers from that of L. curvirostra by the narrower base of ole­cranon ulnae.

1,8 1,9

The distal tibiotarsus has narrower trochlea of distal epiphysys and narrower condylus medialis and condylus lateralis and less developed sulcus extensorius.

Diagnosis. An Upper PlioceneLoxia of the same size of L. curvirostra, whose distal humerus has less constricted shape of condylus ventralis and less outlined sulcus scapulotricipitalis.

b c d I e

2,8 3,9 2,7 0,9

3,1 2,8 2,7 0,9 3,0 2,7 2,6 1,0 3,3 2,9 2,9 1,1 3,1 3,0 2,9 0,9 2,7 2,6 2,6 0,8 3,3 3,2 3,1 1,0

Collections acronyms. NMNHS- National Mu­seum of Natural History - Sofia; UCBL - Centre des Sciences de Ia Terre at the Universite Claude Bernard - Lyon 1.

Comparative material examined Fossils from Varshets were compared with skeletons of the fol­lowing species: UCBL: Loxia curvirostra - UCBL -430/1; 430/2; 430/3; Pyrrhula pyrrhula UCBL 427/ 3; Pinicola enucleator UCBL 429/1; NMNHS: 3/ 1992.

Palaeofaunistical and palaecological notes

The fauna recovered in that site is very rich and in­dicates a forest-steppe landscape. Over 140 verte­brate species have been established so far, at least 50 of them are birds (Boev, 1995 a, b; 1996): Lago­pus balcanicus Boev, 1995 b, Apus baranensis Janossy, 1977, Gallinula balcanica (Boev, in press), Chauvireria balcanica (Boev, 1998 b), and there­maining bird taxa, still determined up to genus lev­el Gyps sp. n., Gypaetinae gen. indet., Aquila sp., Hieraaetus sp. n., Falco sp. (2 species), Circaetus sp., Accipiter sp., Tetrao cf. partium, Otis cf. kho-

Fig. 2. Loxia patevi sp. n.: a- medial view of humerus dex. dist.- NMNHS 307-holotype, b- medial view of humerus sin. prox. -NMNHS 308, c- medial view of humerus sin. prox.- NMNHS 309, d -lateral view of of tibiotarsus dex. dist. - NMNHS 310, e­cranial view oftibiotarsus dex. dist., f -medial view ofulnadex. prox. -NMNHS 311, g-lateral view (Photographs: Boris Andreev)

54

llllljlllllllll 1''''11111111 1''''1'''' em __ 1 em 1 em 1

I'J~fi"H" -~-

11 11 "It 1111 rrr, I If: 1 11 11111t" 11 trr d

55

d a

II

III

IV 45Jc

b

Fig. 3. Manner of measurings of the bone finds: I - proximal humerus; II -distal humerus; III - proximal ulna; IV -distal tibiotarsus (drawing: Vera Hristova)

satzkii, Porzana sp. n., Columba sp., Actitis sp. n., Alauda sp., Lulu/a arborea sp., Galerida sp., Anthus sp., Motacilla sp., Sitta sp. n., Turdus cf. philomelos, Turdus cf. merula, Turdus sp., Luscinia sp., Musci­capa sp.,Regulus sp. n.,Parus sp. (3 species), Ember­iza sp., Lanius sp. n., Fringilla cf. coelebs, Fringilla sp., Carduelis cf. carduelis, Carduelis sp., Cocco­thraustes sp. n., Pyrrhula sp., Sturnus sp., Pyrrhoco­rax graculus cf. vetus, Corvus cf. monedula, Corvus sp., Pica sp. n., Passeres fam. (6 species), etc.

Five species of tree hemixerophytes have been established by their fossilized seeds in the deposits of Varshets: Celtis praebalcanica sp. n., Prunus fruticosa Pall. f. fossilis, Crataegus pentagyna Waldst. & Kit. ex Willdenow f.jossilis, Pyracantha coccinea Roem., f.fossilis H Swida sanguinea (L.) f. fossilis (Palamarev, in press). All of them are indi­cators of dry and warm climate and xeric or semi­arid landscape, similar to present-day African sa-

56

vanah-steppe. The fossil material of seeds of Celtis is rich. It contains over 500 seeds, almost all of them broken by the beacks of seed-eating avian species. These data confirm the type of the palae­oenvironment restored through the palaeoavifau­nistical analysis, indicating a subxerophylous sub­arid climate.

Possibly, the Middle Villafranchian crossbills of Balkan Mountains have inhabited the mixed open deciduous forest of park type, which have grown on the lots alternating by the steppe grassland lots.

Acknowledgements. I am very grateful to Dr cecile Mourer­Chauvire for the opportunity to work on fossil birds of Bulgar­ia at the Earth Sciences Department of the Universi~ Claude Bernard - Lyon 1. I also thank Prof. Emanuil Palamarev for the identification of plant fossils from the site. This work was partially sponsored by the Fondation Scientifique de Lyon et du Sud-Est (Lyon), the National Science Fund (Sofia) (grant No B-202/1992) and the Society of Avian Paleontology and Evolution.,

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