new fossil record of the late pliocene kestrel (falco ... · to the late pliocene kestrel falco...

13

GEOLOGICA BALCANICA, 40. 1–3, Sofia, Dec. 2011, p. 13–30.

New fossil record of the Late Pliocene kestrel (Falco bakalovi Boev, 1999) from the type locality in Bulgaria

Zlatozar BoevNational Museum of Natural History, Bulgarian Academy of Sciences, 1, Tsar Osvoboditel Blv., 1000 Sofia, Bulgaria; e-mail: [email protected]; [email protected](Accepted in revised form: 10.03.2011)

Abstract. Twenty six new bone finds of 4 individuals (2 adult and 2 juvenile) have been described and referred to the Late Pliocene kestrel Falco bakalovi Boev, 1999. They came from the species type locality near the town of Varshets (northwest Bulgaria) and demonstrate specific distinguishing from all known falcons of the “tinnunculus” group. They enrich the bone morphology of that falcon covering a large variety of 16 different skeletal elements.

Boev, Z. N. 2011. New fossil record of the Late Pliocene kestrel (Falco bakalovi Boev, 1999) from the type locality in Bulgaria. Geologica Balcanica 40(1–3), 13–30.

Key words: Falco bakalovi, Falcons, fossil birds, Late Pliocene, Villafranchian Bulgaria.

IntroductIon

Until now a total of 11 fossil species were described in the genus Falco Linnaeus, 1758. Only two of them were considered valid species from Europe (Mlíkovský, 2002). Until recently, F. medius Umanskaya, 1981 from the Late Miocene (MN 11-13) in south Ukraine was the only known falcon of the Miocene of both Eurasia and Africa and the oldest record of the genus. Recently, F. bul garicus, anoth-er and older Miocene falcon of “tinnunculus” group has been described from SW Bulgaria (Late Miocene; Boev, 2011). The second species is the present F. baka lovi Boev, 1999. Falco umanskajae Sobolev 2003 (Late Pliocene, MN 16) from the vicinities of Odessa in Ukraine, is con-sidered “nomen nudum” (Wikipedia, 2011). Outside Europe only one Neogene species, F. ore gonus Brodkorb, 1946 (Early/Middle Pliocene of Oregon, USA), has been described.

The Pliocene falcon F. bakalovi has been designat-ed through a synsacral fragment (holotype) collected in 1990. Since that time additional excavations for collect-ing of fossil material have been undertaken which lasted several years.

Part of the collected rich material, still unexamined completely, belongs to a small falcon. Boev (1999 a, b; 2002) listed a total of sixteen new finds identified as Falco ex gr. tinnunculus. In addition, some finds col-lected in the last two years were also referred to this fal-con. All this material is the subject of the present paper.

The finds come from the type locality near the town of Varshets in northwest Bulgaria and are associated with F. bakalovi, the only known falcon of the site until now. They are the first confirmation of the existing of this spe-cies and enlarge its morphological characteristics.

MaterIal and Method

Abbreviations: Anatomical: c. – cotyla; cond. – condylus; dex. – dextra; dig. – digitus, digiti; dist. – distalis; f. a. – facies a.; max. – maximum; min. – minimum; pr. – proc-essus, processi; prox. – proximalis; sin – sinistra; s. a. – sulcus articularis; tbt – tibiotarsus; tmt – tarsometatarsus. Institutional: BMNH – Natural History Museum, former-ly British Museum (Natural History), Tring; NMNHS – National Museum of Natural History, Bulgarian Academy of Sciences, Sofia; UCBL – University Claude Bernard, Lyon 1, France.

The material was collected through screening and washing of the sediments. It consists of 26 bones and bone fragments, representing 16 skeletal elements of both fore limbs and the pectoral girdle, and the hind limbs and the pelvis girdle: No NMNHS 122-124; 131-132, 134-136, 143-145, 181, 189, 223, 294, 304-306, 317-321, 364, 14 952 and 14964. The finds belong to 2 adult and 2 juvenile individuals from MNI 4. They have been collected by the author in a 12-year period (1991-2001) from the type locality near the town of Varshets,

14

NW Bulgaria. The material was identified by comparison with the holotype in the reference avian skeleton collec-tion of the NMNHS, as well as the osteological collec-tions of the Centre des Sciences de la Tèrre, Université Claude Bernard – Lyon (UCBL) in 1994–1995 and the Natural History Museum, Tring, a part of the Natural History Museum, London, former British Museum of Natural History (BMNH) in 1999 and 2003. It is kept in the Vertebrate Animal Department of the NMNHS.

The taxonomy follows White et al. (1994). The osteo-logical terminology is after Baumel and Witmer (1993)

and Livezey and Zusi (2006). The chronostratigraphy follows Mein (1990) and Guerin (1990).

All measurements (Tables 1–16, Fig. 1) have been taken using calipers to 0.05 mm accuracy, but read to the 1st digit after decimal point. Description of the meas-uring manner of some bones in fossil and recent Falco: humerus dex. dist.: a – thickness of condylus dorsalis; b – thickness of condylus ventralis; c – total width of c. dorsalis and c. ventralis (Table 4); ulna sin. prox.: a – lon-gitudinal diameter of cotyla ventralis; b – transversal di-ameter (length) of cotyla dorsalis; c – maximum width of

Fig. 1. Manner of measurings of the pelvis in Falconidae (Drawings Vera Hristova)

15

proximal diaphysis; d – length of depressio m. brachialis; e – width of diaphysis at the middle of depressio m. bra-chialis (Table 5); femur sin. dist.: a – width of distal epi-physis; b – diameter of condylus lateralis; c – diameter of condylus medialis; d – diameter (thickness) in sulcus intercondylaris; e – cranio-caudal thickness of the dia-physis above distal epiphysis (Table 10); tibiotarsus dex. dist.: a – width of distal epiphysis; b – diameter of the condylus medialis; c – diameter of the condylus lateralis; d – thickness in the incissura intercondylaris; e – width of diaphysis in the proximal end of sulcus supratendineus (Table 11). The manner of other measuring is given on Fig. 1. All generic names of the binominals are given ab-breviated in the text and are in full in the Appendix 1. “Smaller”, “much smaller”, “larger” or “much larger” in the “Comparison and discussion” section mean that the fossil specimen differs considerably in size from the specimens of the compared species, and thus their taxo-nomic identity is excluded.

SySteMatIc Palaeontology

The general morphology of all 26 finds indicates that they belong to the group of the small falcons (Falco sp. ex gr. tinnunculus). As F. bakalovi is the only known representative of that group in the Varshets locality, we refer all these finds to the same species.

Order FALCONIFORMES (Sharpe, 1874)Family FALCONIDAE (Vigors, 1824)Subfamily Falconinae (Vigors, 1824)

genus Falco linnaeus, 1758 Falco bakalovi Boev, 1999

Holotype: Postacetabular part of the left half of pelvis, NMNHS 1642. Collected on July 25th, 1990 by Z. Boev.Paratypes: humerus dex. dist. NMNHS 135; ulna sin. prox. NMNHS 131; ulna dex. dist. NMNHS 14 964; fe-mur sin. dist. NMNHS 223; tbt dex. dist. NMNHS 136.Comparison: See Tables 1–16; Fig. 1 and “Description and comparison” section. Measurements of the paratypes: Tables 1–16; Fig. 1. Neodiagnosis: A small-sized (between F. tinnunculus and F. subbuteo) fossil species of genus Falco, differing by the sharp transition (turn) of crista iliaca dorsolateralis over the ala ischii and the rounded (oval), but not angular shape of the caudal edge of foramen ilioischiadicum. Distinguishing differences from: (A) the modern F. tin-nunculus: (1) distal humerus: blunter (rounder) epicondy-lus ventralis; (2) рroximal ulna: shallower fossa under c. dorsalis, and almost twice narrower depressio m. brachi-alis; (3) distal ulna: thicker cond. ventralis ulnae in caudal view; (4) distal femur: much higher crista tibiofibularis in caudal view; (5) distal tbt: much thicker wall of the medial tendineal fossa above c. medialis; from (B) the Miocene F. bul garicus: (1) proximal ulna, (2) distal humerus and (3) distal tibiotarsus much stouter and robust.Locality: A ponor in a rocky hill, 6 km NNE of the town of Varshets (43.13° N, 23.17° E). Unconsolidated, un-stratified sediments accumulated as terra-rossa clay. The fossil bones are broken, locally forming a bone breccia. Chronology: Villafranchian. The associated mam mal fauna (Spassov 1997 a, b, 2000; Popov, 2001) gave

Taxa a b c d eFossil – Varshets

Falco bakalovi NMNHS 304 2.4 2.8 2.1 10.2 4.6Recent

Falco chicquera BMNH 1965.18.1 3.5 3.5 2.0 11.7 ca. 7.1Falco chicquera BMNH 1993.2.5 2.6 2.8 1.7 10.0 6.1Falco columbarius BMNH 1930.3.24.264 2.4 2.8 1.9 10.0 5.8Falco columbarius BMNH 1930.3.24.267 2.7 2.6 1.8 10.5 6.6Falco columbarius BMNH 1988.61.1 2.7 3.1 1.7 10.6 7.0Falco naumanni BMNH 1955.15.2 2.6 2.9 1.6 8.9 6.2Falco naumanni BMNH 1961.13.4 2.2 2.8 1.6 3.8 5.5Falco newtoni BMNH 1897.5.10.29 2.2 2.6 1.5 7.4 ca. 4.7Falco sparverius BMNH 1954.3.2 2.0 2.4 1.7 7.2 5.2Falco subbuteo BMNH 1985.76.1 2.8 3.2 2.3 11.1 7.3Falco subbuteo BMNH 1994.39.1 3.0 3.3 1.8 11.5 7.3Falco subbuteo UCBL 111/2 2.9 2.7 1.8 11.4 5.2Falco tinnunculus BMNH 1930.3.24.275 2.9 3.1 1.8 10.3 6.9Falco tinnunculus UCBL 119/2 2.6 2.4 1.6 9.2 4.6Falco tinnunculus UCBL 119/3 2.8 2.6 1.7 9.3 4.6Falco vespertinus BMNH 1855.4.4.9 2.3 2.9 2.0 9.4 6.0Falco vespertinus BMNH 1869.10.19.12 2.3 2.4 1.8 8.7 5.2Falco vespertinus BMNH 1952.3.122 2.5 3.2 1.8 9.7 6.3

Table 1The measurements of coracoid sin. in some fossil and recent Falco (ref. to Fig. 1 A)

17

the site a MN17/MNQ17 zone attribution (Mein, 1990; Guerin, 1990). Spassov (2000, 2003) attributed the site in the St. Vallier unit between the levels of the localities Roccaneyra and St. Vallier.

deScrIPtIon and coMParISon

Skeletal elements of the fore limbs and the pectoral girdle

Coracoid sin. NMNHS 304 (Figs 2 a, b, Table 1). The specimen belongs to a juvenile/subadult individual. It differs from: F. chicquera: similar in size and general morphology, but differs by the thinner shaft of coracoid; F. columbarius: sharper pr. acrocoracoideus; F. nau-manni: by the larger size, and the relatively narrower

f. a. humeralis; F. newtoni: the same way as F. sparverius; F. sparverius: by the larger size, and the thicker shaft in c. scapularis; F. subbuteo: morphologically similar, but smaller and differs by the narrower f. a. humera-lis, and narrower pr. acrocoracoideus; F. tinnunculus: high similarity in size, general shape and proportions, but differs by the almost twice narrower f. a. humeralis; F. vespertinus: sharper pr. acrocoracoideus, narrower f. a. humeralis.

Coracoid sin. NMNHS 305 (Fig. 2 c, Table 2). The specimen lacks caudal part of the sternal end and all humeral end. Its preserved part (mainly the bone shaft) completely fits to No 304 in both morphology and size.

Clavicula sin. prox. NMNHS 306 (Figs 2 d, e, Table 3). The specimen preserves the distal (humeral) half of the bone and differs from: F. chicquera: by the less rounded pr. acromialis; F. columbarius: shallower relief of f. a. ac-

Fig. 2. Falco bakalovi Boev (Photographs: Assen Ignatov)a, b, coracoid sin. NMNHS 304c, coracoid sin. NMNHS 305d, e, clavicula sin. prox. NMNHS 306 f, humerus dex. dist. NMNHS 14 952g, h, humerus dex. dist. NMNHS 134 i, j, k, humerus dex. dist. NMNHS 135 l, ulna sin. prox. NMNHS 132

←



Falco chicquera BMNH 1965.18.1 – – 3.2 – –Falco chicquera BMNH 1993.2.5 ca. 9.2 ca. 18.9 2.9 – 6.1Falco columbarius 1930.3.24.267 9.6 20.2 2.8 2.3 5.3Falco columbarius BMNH 1930.3.24.264 8.2 17.8 2.7 2.3 5.5Falco columbarius BMNH 1988.61.1 8.7 20.0 3.1 2.3 6.3Falco naumanni BMNH 1955.15.2 6.9 14.8 2.9 2.2 5.4Falco naumanni BMNH 1961.13.4 6.3 14.0 2.8 2.0 5.3Falco newtoni BMNH 1897.5.10.29 7.5 13.7 2.7 2.2 5.0Falco sparverius BMNH 1954.3.2 8.1 15.4 2.3 1.9 4.8Falco subbuteo BMNH 1985.76.1 9.0 18.0 3.0 2.4 6.2Falco subbuteo BMNH 1994.39.1 8.5 18.1 3.2 2.5 6.5Falco subbuteo UCBL 111/2 10.2 17.6 3.1 2.4 6.6Falco tinnunculus BMNH 1930.3.24.275 9.3 17.9 3.3 2.4 5.7Falco tinnunculus UCBL 119/2 9.0 16.4 2.8 2.1 5.2Falco tinnunculus UCBL 119/3 8.2 16.0 3.4 2.2 5.2Falco vespertinus BMNH 1855.4.4.9 7.6 16.1 2.2 1.7 5.7Falco vespertinus BMNH 1869.10.19.12 8.7 16.4 2.5 2.3 5.1Falco vespertinus UCBL 117/1 10.0 18.9 3.1 2.0 5.8

Table 2The measurements of coracoid sin. in some fossil and recent Falco (ref. to Fig. 1 B)

18

Table 3The measurements of clavicula sin. prox. in some fossil and recent Falco (ref. to Fig. 1 C, D, E)



Falco chicquera BMNH 1965.18.1 – 3.5 1.5 7.2 –Falco chicquera BMNH 1993.2.5 6.6 2.9 1.1 6.6 3.9Falco columbarius BMNH 1930.3.24.264 1.7 2.1 0.9 5.0 3.8Falco columbarius BMNH 1930.3.24.267 1.8 3.2 0.7 5.5 6.0Falco columbarius BMNH 1988.61.1 1.7 3.0 0.8 6.1 4.9Falco naumanni BMNH 1955.15.2 1.5 2.8 0.9 4.9 4.4Falco newtoni BMNH 1897.5.10.29 1.4 2.6 0.9 4.0 3.0Falco sparverius BMNH 1954.3.2 1.7 2.4 1.0 3.9 3.4Falco subbuteo BMNH 1994.39.1 1.9 3.8 1.3 6.5 4.2Falco subbuteo UCBL 111/3 1.8 3.4 1.0 6.8 5.5Falco subbuteo UCBL 111/4 1.5 2.8 1.1 6.0 5.3Falco tinnunculus BMNH 1930.3.24.275 1.5 2.8 1.8 4.7 5.2Falco tinnunculus BMNH 1930.3.24.278 1.2 2.8 1.7 5.2 4.4Falco tinnunculus UCBL 119/7 1.5 2.7 1.0 5.1 5.0Falco tinnunculus UCBL 119/8 1.2 2.6 0.9 5.1 4.1Falco vespertinus BMNH 1855.4.4.9 1.8 2.8 1.1 5.2 4.2Falco vespertinus BMNH 1869.10.19.12 1.8 2.8 0.9 4.3 3.6Falco vespertinus BMNH 1952.3.122 1.7 2.8 0.9 4.3 3.8

Table 4The measurements of humerus dex. dist. in some fossil and recent Falco

Taxa a b cFossil – Varshets

Falco bakalovi NMNHS 134 4.7 2.2 ca. 7.4Falco bakalovi NMNHS 135 4.8 2.4 ca. 7.6Falco bakalovi NMNHS 14 952 5.8 ca. 2.8 ca. 8.5

RecentFalco chicquera BMNH 1965.18.1 6.3 3.2 7.8Falco chicquera BMNH 1993.2.5 5.0 2.7 6.5Falco columbarius BMNH 1930.3.24.264 5.3 2.6 6.9Falco columbarius BMNH 1930.3.24.267 5.6 2.7 7.4Falco columbarius BMNH 1988.61.1 5.7 2.9 7.4Falco naumanni BMNH 1955.15.2 5.5 2.6 6.9Falco naumanni BMNH 1961.13.4 4.9 2.7 5.8Falco newtoni BMNH 1897.5.10.29 4.6 2.3 5.1Falco sparverius BMNH 1954.3.2 4.8 2.2 5.7Falco subbuteo BMNH 1974.16.1 6.3 3.0 7.9Falco subbuteo BMNH 1985.76.1 6.2 3.0 8.4Falco subbuteo BMNH 1994.39.1 6.5 3.0 8.5Falco tinnunculus BMNH 1930.3.24.275 4.2 2.9 7.8Falco tinnunculus BMNH 1930.3.24.276 6.1 2.9 8.2Falco tinnunculus BMNH 1930.3.24.278 5.3 2.7 6.8Falco vespertinus BMNH 1855.4.4.9 5.2 2.7 6.7Falco vespertinus BMNH 1869.10.19.12 5.1 2.7 6.7Falco vespertinus BMNH 1952.3.122 5.2 2.6 7.4

19

rocoracoidea, larger distance between the f. a. acrocora-coidea; F. naumanni: by the wider f. a. acrocoracoidea; F. newtoni: by the larger size, and the deeper relief of the acrocoracoidal articular end; F. sparverius: by the larger size; F. subbuteo: by the shorter f. a. acrocoracoidea; F. subbuteo: smaller, in cranio-lateral view the vow-like profile between the f. a. acrocoracoidaea is narrower and round than open and elliptic; F. tinnunculus: wider f. a. acrocoracoidea and sharper pr. acrocoracoideus; F. vespertinus: larger size and the wider humeral part in lateral view.

Humerus dex. dist. NMNHS 14 952 (Fig. 2 f, Table 4). The specimen preserves only fragment of dorsal (lateral) part of the distal epiphysis. It differs from: F. chicquera: by the deeper relief on the dorsal side of distal epiphysis. F. columbarius: less developed pr. supracondylaris dor-salis, smaller diameter of cond. dorsalis; F. naumanni: mainly by the larger size. F. newtoni: the same way as F. sparverius; F. sparveriusi: by the larger size. F. sub-buteo: smaller, in caudal view cond. dorsalis more short-ened; F. tinnunculus: similar, but slightly larger, differs by the wider facies a. acrocoracoidea, and narrower cond. ventralis humeralis; F. vespertinus: similar, but cond. dorsalis slightly bigger in caudal view.

Humerus dex. dist. NMNHS 134 (Figs 2 g, h, Table 4). The specimen belongs to a juvenile/subadult individual. F. tinnunculus: dimensionally completely fits.

Humerus dex. dist. NMNHS 135 (Figs 2 i, j, k, Table 4). F. columbarius: more straight distal diaphysis, shorter epicondylus dorsalis; F. subbuteo: smaller size;

shallower sulcus m. humerotricipitis, shorter pr. flexo-rius in medial view; F. tinnunculus: blunter (rounder) epicondylus ventralis; F. vespertinus: the same way as F. tinnunculus.

Ulna sin. prox. NMNHS 131 (Table 5). The speci-men differs from: F. chicquera: by the more straight, instead slightly curved shaft of the proximal part of dia-physis; F. chicquera: by the relatively slightly narrower c. dorsalis; F. columbarius: by the wider diaphysis, and the wider depressio m. brachialis; F. naumanni: by the larger size, and the less oval, instead angular, in dorsal edge, c. dorsalis; F. subbuteo: smaller, c. ventralis is more angular, instead round; F. tinnunculus: similar, but differs by the thicker diaphysis, shallower fossa under c. dorsalis, and almost twice narrower depressio m. brachialis; F. vespertinus: more robust, longer ole-cranon, c. dorsalis more angular than round.

Ulna sin. prox. NMNHS 132 (Fig. 2 l, Table 5). The specimen differs the same way as No 131.

Ulna dex. dist. NMNHS 294 (Fig. 3 a, Table 6). The specimen belongs to a juvenile/subadult individual, reached its definitive size. The dorsal part of c. dorsalis ulnaris had been broken. Beside the incompletely de-veloped articular surfaces, the distal epiphysis bears the characteristic features of Falconiformes and Falconidae, especially the widely open depressio radialis, the wider (in comparison to Accipitridae) s. intercondylicus, and the more protuberant in ventral direction tuber carpale. Metrically and morphologically completely fits to small-er Palearctic falcons of genus Falco. It differs from:

Table 5The measurements of ulna sin. prox. in some fossil and recent Falco


Falco bakalovi NMNHS 131 4.4 4.0 8.3 13.5 4.7Falco bakalovi NMNHS 132 4.5 4.0 7.9 10.6 4.9

RecentFalco chicquera BMNH 1965.18.1 5.1 4.8 8.9 11.2 5.1Falco chicquera BMNH 1993.2.5 4.5 3.8 7.8 ca. 9.4 4.1Falco columbarius BMNH 1930.3.24.264 4.0 4.2 7.4 ca. 10.2 ca. 4.2Falco columbarius BMNH 1930.3.24.267 4.3 3.4 8.0 9.5 4.3Falco columbarius BMNH 1988.61.1 4.5 3.9 8.1 9.7 4.4Falco naumanni BMNH 1955.15.2 4.2 3.7 7.7 12.3 4.1Falco naumanni BMNH 1961.13.4 3.9 3.5 6.9 10.5 3.9Falco newtoni BMNH 1897.5.10.29 3.7 3.0 6.2 7.6 3.6Falco sparverius BMNH 1954.3.2 3.6 2.8 6.5 8.2 4.1Falco subbuteo BMNH 1974.16.1 4.7 4.0 9.3 11.0 5.2Falco subbuteo BMNH 1985.76.1 4.4 4.3 8.5 ca. 10.9 4.9Falco subbuteo BMNH 1994.39.1 4.9 3.9 8.9 11.2 4.7Falco tinnunculus BMNH 1930.3.24.275 4.1 3.8 8.0 11.4 4.5Falco tinnunculus BMNH 1930.3.24.276 4.8 3.9 8.3 11.3 4.9Falco tinnunculus BMNH 1930.3.24.278 4.4 3.6 7.8 10.8 4.3Falco vespertinus BMNH 1855.4.4.9 4.8 3.8 7.8 11.7 4.0Falco vespertinus BMNH 1869.10.19.12 3.9 3.2 7.3 11.7 4.1Falco vespertinus BMNH 1952.3.122 4.2 3.3 7.4 9.0 4.6

21

F. columbarius: larger and more robust, more angular c. ventralis and c. dorsalis; F. subbuteo: smaller size, shallower s. radialis; F. tinnunculus: shallower s. tend-ineus; F. vespertinus: lower cond. dorsalis.

Ulna dex. dist. NMNHS 14 964 (Fig. 3 b, Table 6). The specimen differs from: F. chicquera: by the absence of clear constriction between the diaphysis and cond. ventralis in lateral view; F. columbarius: slightly bigger, wider s. radialis, higher c. ventralis; F. naumanni: by the larger size, and the shallower s. radialis; F. newtoni: by the larger size, and by the straight, instead slightly curved distal fourth of diaphysis in lateral view; F. sparverius: by the larger size; F. subbuteo: by the deeper s. radialis; F. tinnunculus: very similar, but slightly larger; thicker

cond. ventralis ulnae in caudal view; F. vespertinus: thicker diaphysis, deeper s. radialis.

Ulna sin. dist. NMNHS 364 (Fig. 3 c, Table 6). The specimen differs the same way as No 294.

Carpometacarpus dex. prox. NMNHS 124 (Figs 3 d, e, Table 7). The specimen belongs to a juvenile/subadult in-dividual. It differs from: F. columbarius: of similar size; F. subbuteo: smaller, less steep caudal profile of tr. car-palis; F. tinnunculus: by the narrower f. a. radiocarpalis; F. vespertinus: more angular cranial edge of the dorsal condyle of trochlea carpalis.

Carpometacarpus sin. prox. NMNHS 189 (Figs 3 f, g, h, Table 8). The specimen differs from: F. chicquera: by the larger size and the slightly concaved caudal pro-

←Fig. 3. Falco bakalovi Boev (Photographs: Assen Ignatov)

a, ulna dex. dist. NMNHS 294 b, ulna dex. dist. NMNHS 14 964 c, ulna sin. dist. NMNHS 364 d, e, carpometacarpus dex. prox. NMNHS 124 f, g, h, carpometacarpus sin. prox. NMNHS 189 i, carpometacarpus sin. dist. NMNHS 122 j, k, l, femur sin. dist. NMNHS 223

Taxa a b c dFossil – Varshets

Falco bakalovi NMNHS 294 4.5 6.3 4.1 3.8Falco bakalovi NMNHS 14 964 4.7 6.8 4.2 3.7

RecentFalco chicquera BMNH 1965.18.1 – 7.0 4.9 4.4Falco chicquera BMNH 1993.2.5 4.2 6.0 3.5 3.3Falco columbarius BMNH 1930.3.24.264 4.1 6.2 3.8 3.4Falco columbarius BMNH 1930.3.24.267 4.5 6.7 3.5 3.5Falco columbarius BMNH 1988.61.1 4.6 6.6 3.5 3.6Falco naumanni BMNH 1955.15.2 4.5 6.0 4.0 3.4Falco naumanni BMNH 1961.13.4 4.1 5.9 3.4 3.1Falco newtoni BMNH 1897.5.10.29 3.6 5.1 3.1 2.9Falco sparverius BMNH 1954.3.2 3.6 5.4 3.4 3.0Falco subboteo UCBL 111/4 5.4 7.4 4.2 3.8Falco subbuteo UCBL 111/2 4.8 7.2 4.3 3.7Falco subbuteo UCBL 111/3 4.8 6.9 4.4 3.7Falco tinnunculus UCBL 119/1 4.6 7.1 4.6 3.8Falco tinnunculus UCBL 119/2 4.2 5.9 3.7 3.3Falco tinnunculus UCBL 119/3 4.4 6.2 4.0 3.4Falco tinnunculus UCBL 119/7 4.6 6.7 4.2 3.9Falco vespertinus BMNH 1855.4.4.9 4.2 6.3 4.2 3.8Falco vespertinus UCBL 117/1 4.4 6.1 3.8 3.4Falco vespertinus UCBL 117/2 4.2 5.7 3.6 3.0

Table 6The measurements of ulna dex. dist. in some fossil and recent Falco (ref. to Fig. 1 F)

22

Table 7The measurements of carpometacarpus dex. prox. in some fossil and recent Falco (ref. to Fig. 1 G)

Taxa a b cFossil – Varshets

Falco bakalovi NMNHS 124 4.8 5.1 2.6Recent

Falco chicquera BMNH 1965.18.1 5.2 5.9 3.1Falco chicquera BMNH 1993.2.5 3.7 5.3 2.8Falco columbarius BMNH 1930.3.24.264 4.5 5.4 2.7Falco columbarius BMNH 1930.3.24.267 5.2 6.8 2.7Falco columbarius BMNH 1988.61.1 4.4 5.7 2.9Falco naumanni BMNH 1955.15.2 4.5 4.8 2.6Falco naumanni BMNH 1961.13.4 4.3 5.0 2.4Falco newtoni BMNH 1897.5.10.29 4.0 5.1 2.6Falco sparverius BMNH 1954.3.2 3.8 4.5 2.5Falco subbuteo BMNH 1974.16.1 5.6 6.6 3.6Falco subbuteo BMNH 1985.76.1 5.0 6.5 4.3Falco subbuteo BMNH 1994.39.1 5.5 7.2 3.6Falco tinnunculus UCBL 119/2 4.5 5.6 2.7Falco tinnunculus UCBL 119/3 4.2 5.6 2.6Falco tinnunculus UCBL 119/4 4.4 5.7 2.7Falco vespertinus BMNH 1855.4.4.9 4.6 5.2 2.9Falco vespertinus BMNH 1869.10.19.12 4.3 5.3 2.8Falco vespertinus BMNH 1952.3.122 4.9 5.4 2.6

Table 8The measurements of carpometacarpus sin. prox. in some fossil and recent Falco (ref. to Fig. 1 H)



Falco chicquera BMNH 1965.18.1 6.5 – 3.3 12.0 –Falco chicquera BMNH 1993.2.5 5.5 7.2 3.1 10.2 3.9Falco columbarius BMNH 1930.3.24.264 5.5 8.8 2.0 9.8 3.5Falco columbarius BMNH 1930.3.24.267 6.8 9.3 2.6 11.8 3.7Falco columbarius BMNH 1988.61.1 6.1 9.2 3.0 11.7 3.8Falco naumanni BMNH 1955.15.2 5.2 8.8 3.8 10.4 3.6Falco naumanni BMNH 1961.13.4 5.2 8.4 2.7 10.4 3.5Falco newtoni BMNH 1897.5.10.29 5.1 6.3 2.7 ca. 8.6 3.2Falco sparverius BMNH 1954.3.2 4.8 6.3 2.6 8.8 3.3Falco subbuteo BMNH 1974.16.1 6.8 8.8 3.8 12.4 4.6Falco subbuteo BMNH 1985.76.1 6.3 10.7 3.6 12.0 4.1Falco subbuteo BMNH 1994.39.1 6.5 9.0 3.7 11.2 4.4Falco tinnunculus BMNH 1930.3.24.275 5.8 8.4 3.0 10.7 4.0Falco tinnunculus BMNH 1930.3.24.278 6.1 9.0 3.0 10.3 3.6Falco tinnunculus UCBL 119/7 5.5 9.4 3.2 10.2 3.8Falco vespertinus BMNH 1855.4.4.9 5.5 9.2 3.0 11.6 4.0Falco vespertinus BMNH 1869.10.19.12 5.2 7.2 3.0 10.8 3.4Falco vespertinus BMNH 1952.3.122 5.7 9.1 3.1 9.2 3.2

23

Table 9The measurements of carpometacarpus sin. dist. in some fossil and recent Falco (ref. to Fig. 1 I)


Falco bakalovi NMNHS 122 ca. 3.3 6.5 3.1 4.4Recent

Falco chicquera BMNH 1965.18.1 3.0 6.8 3.1 4.6Falco chicquera BMNH 1993.2.5 3.0 6.3 2.2 3.7Falco columbarius BMNH 1930.3.24.264 3.9 6.0 2.7 3.7Falco columbarius BMNH 1930.3.24.267 4.2 6.2 2.4 3.8Falco columbarius BMNH 1988.61.1 4.1 5.6 2.5 4.2Falco naumanni BMNH 1955.15.2 3.7 6.2 2.5 4.0Falco naumanni BMNH 1961.13.4 3.2 5.4 2.3 3.6Falco newtoni BMNH 1897.5.10.29 2.7 5.0 2.2 3.3Falco sparverius BMNH 1954.3.2 3.3 4.8 2.2 3.0Falco subbuteo BMNH 1985.76.1 4.5 6.8 2.7 4.3Falco subbuteo BMNH 1994.39.1 4.6 6.8 2.8 4.4Falco subbuteo UCBL 111/3 3.8 7.0 3.2 4.0Falco tinnunculus UCBL 119/1 3.8 7.3 2.9 4.2Falco tinnunculus UCBL 119/2 3.7 6.7 2.8 4.0Falco tinnunculus UCBL 119/6 3.6 7.0 2.9 4.0Falco vespertinus BMNH 1855.4.4.9 4.4 5.9 2.5 3.9Falco vespertinus UCBL 117/2 3.5 6.1 2.6 3.6Falco vespertinus UCBL 117/4 3.8 6.4 2.6 3.7

file of trochlea carpalis; F. columbarius: slightly larger in size, deeper s. between pr. extesorius and the lateral con-dyle of dorso-lateral view, longer pr. extensorius; F. nau-manni: by the larger size and the bigger pr. pisiformis; F. new toni: the same way as F. sparverius; F. sparverius: by the larger size, and the much deeper constriction be-tween pr. extensorius and the lateral condyle of the troch-lea carpalis in lateral view; F. subbuteo: more proximal position of the synostosis metacarpalis proximalis, more reversed (upright) inception of the os metacarpale minus; F. tinnunculus: by the slightly larger size, shorter incis-sura tendineus on the os metacarpalis majoris, and the shorter and rounder pr. pisiformis; F. vespertinus: larger size, and by the bigger pr. pisisformis.

Carpometacarpus sin. dist. NMNHS 122 (Fig. 3 i, Table 9). The specimen differs from: F. columbarius: slightly bigger; F. subbuteo: smaller, by the round, in-stead of pear-shaped protuberance of the middle part of f. a. digitalis major, by its stronger protruding, by its vertical orientation towards the joint axis, by its more ventral position (reaching the edge), by the shallower s. tendineus in its distal end; F. tinnunculus: f. a. digiti mi-noris narrower and all differences listen for F. subbuteo; F. vespertinus: bigger, thicker f. a. digiti majoris.

Skeletal elements of the hind limbs and the pelvis girdle

Femur sin. dist. NMNHS 223 (Figs 3 j, k, l, Table 10). The specimen differs from: F. chicquera: by the clearly

wider cond. lateralis in ventral view; F. columbarius: wider impressio ansae m. iliofibularis, shorter crista su-pracondylaris medialis, wider cond. medialis in cranial view; F. naumanni: by the larger size, and the relatively larger crista supracondylaris medialis; F. newtoni: by the larger size; F. sparverius: by the larger size, and the more asymmetrical s. patellaris in ventral view; F. subbuteo: smaller, more developed and sharper tuberculum muscu-li gastrocnemialis lateralis; F. tinnunculus: much higher crista tibiofibularis in caudal view, by the wider crista supracondylaris medialis; F. vespertinus: bigger in size, and the smaller tuberculum m. gastrocnemialis lateralis.

Tbt dex. dist. NMNHS 136 (Fig. 4 a, Table 11). The specimen differs from: F. chicquera: the same way as F. subbuteo; F. columbarius: slightly larger general size, larger distal part of diaphysis, wider pons supratendineus; F. naumanni: the same way as F. vespertinus; F. new-toni: the same way as F. sparverius; F. subbuteo: more robust and larger distal part of diaphysis, deeper depres-sio epicondylaris lateralis; F. sparverius and F. newtoni: by the larger size, and the relatively wider distal part of diaphysis; F. tinnunculus: by the much thicker wall of the medial tendineal fossa above c. medialis, by the thinner cond. medialis in ventral view and by the wider (more ro-bust) distal third of diaphysis; F. vespertinus: larger size, and the relatively wider distal part of diaphysis.

Tbt dex. dist. NMNHS 145 (Fig. 4 b, Table 11). The specimen differs the same way as No 136.

Tmt dex. prox. NMNHS 123 (Figs 4 c, d, Table 13). The specimen differs from: F. columbarius: bigger in

24



Falco chicquera BMNH 1965.18.1 ca. 7.3 6.9 7.5 – 5.6Falco chicquera BMNH 1993.2.5 7.4 6.9 6.5 4.4 4.6Falco columbarius BMNH 1930.3.24.264 7.0 6.2 6.0 4.4 4.0Falco columbarius BMNH 1930.3.24.267 7.5 6.5 6.3 4.3 4.4Falco columbarius BMNH 1988.61.1 7.9 6.9 6.8 4.4 4.3Falco naumanni BMNH 1955.15.2 6.6 5.8 5.3 4.0 3.8Falco naumanni BMNH 1961.13.4 6.1 5.8 5.2 4.0 3.5Falco newtoni BMNH 1897.5.10.29 6.4 5.0 5.0 ca. 2.6 3.7Falco sparverius BMNH 1954.3.2 6.3 5.7 5.5 3.7 4.4Falco subbuteo BMNH 1974.16.1 8.3 7.4 7.0 5.0 4.8Falco subbuteo BMNH 1985.76.1 7.7 6.6 6.6 4.1 4.0Falco subbuteo BMNH 1994.39.1 8.5 7.2 6.8 4.6 4.7Falco tinnunculus BMNH 1930.3.24.275 8.2 7.1 6.2 5.6 4.5Falco tinnunculus BMNH 1930.3.24.276 8.2 7.2 6.5 4.6 5.0Falco tinnunculus BMNH 1930.3.24.278 7.7 6.8 6.0 4.1 4.0Falco vespertinus BMNH 1855.4.4.9 6.4 5.9 5.8 4.6 4.2Falco vespertinus BMNH 1869.10.19.12 6.2 5.5 5.4 4.5 4.9Falco vespertinus BMNH 1952.3.122 6.8 5.9 6.0 4.2 4.2

Table 10The measurements of femur sin. dist. in some fossil and recent Falco

Fig. 4. Falco bakalovi Boev (Photographs: Assen Ignatov)a, tbt dex. dist. NMNHS 136 b, tbt dex. dist. NMNHS 145 c, d, tmt dex. prox. NMNHS 123 e, tmt dex. prox. NMNHS 143 f, tmt sin. NMNHS 144 g, phalanx 1 dig. I pedis dex. NMNHS 319 h, phalanx 1 dig. I pedis dex. NMNHS 320 i, phalanx 1 dig. I pedis dex. NMNHS 321 j, phalanx 2 dig. II pedis dex. NMNHS 317 k, phalanx 2 dig. II pedis sin. NMNHS 318 l, phalanx dist. dig. 1 pedis dex. NMNHS 181 – lateral view (above) and caudal view (bellow)

←

size, less prominent lateral edge of c. lateralis; F. sub-buteo: smaller size, probably lower crista medialis hy-potarsi (partly broken); F. tinnunculus: by the shallower c. medialis in cranial view; F. vespertinus: bigger in size.

Tmt dex. prox. NMNHS 143 (Fig. 4 e, Table 12). The specimen differs from: F. columbarius: more angu-lar instead round, c. medialis; F. subbuteo: higher facies subcutanea medialis and facies subcutanea lateralis; F. tinnunculus: lower eminentia intercondylaris; F. colum-barius: bigger, wider proximal third of diaphysis. F. ves-pertinus: bigger in size.

Tmt sin. NMNHS 144 (Fig. 4 f; Table 12). The speci-men belongs to a juvenile individual of almost definite size. The specimen differs the same way as No 123.

Phalanx 1 dig. I pedis dex. NMNHS 319 (Fig. 4 g, Table 14). The specimen differs from: F. columbarius: thicker phalangeal body; F. subbuteo: smaller, slight-ly more asymmetrical trochlea a. in cranial view; F. tinnunculus: by the smaller dimensions, and particu-larly lesser diameter of the condyles of the trochlea; F. vespertinus: bigger, both condyles more parallel in dorsal view.

26

Phalanx 1 dig. I pedis dex. NMNHS 320 (Fig. 4 h, Table 14). The specimen differs the same way as No 319.

Phalanx 1 dig. I pedis dex. NMNHS 321 (Fig. 4 i, Table 14). The specimen belongs to a juvenile/sub-adult individual of almost definite size. It differs from: F. columbarius: more concave ventral side in the proximal end; F. subbuteo: smaller; F. tinnun-culus: by the smaller dimensions, and by the higher

Table 12The measurements of tarsometatarsus sin. in some fossil and recent Falco (ref. to Fig. 1 J, K)


Falco bakalovi NMNHS 143 7.1 – – 2.9 –Falco bakalovi NMNHS 144 7.6 39.5 ca. 2.2 2.8 3.5

RecentFalco columbarius NMNHS 1/1997 6.9 35.5 2.5 3.3 4.6Falco columbarius NMNHS 2/2002 6.5 36.3 2.2 2.8 4.3Falco subbuteo NMNHS 1/1989 8.4 35.7 3.4 3.4 5.1Falco subbuteo NMNHS 3/1993 6.9 35.4 3.2 3.1 4.6Falco tinnunculus UCBL 119/7 7.4 40.7 2.6 3.2 4.3Falco tinnunculus NMNHS 10/1991 7.4 41.4 2.7 3.0 4.4Falco vespertinus NMNHS 1/1991 6.2 29.3 2.3 2.7 3.6

Table 11The measurements of tibiotarsus dex. dist. in some fossil and recent Falco


Falco bakalovi NMNHS 136 6.8 5.5 ca. 5.1 4.3 6.0Falco bakalovi NMNHS 145 ca. 7.2 5.5 – 4.0 5.0

RecentFalco chicquera BMNH 1965.18.1 7.8 6.0 6.3 4.6 5.4Falco chicquera BMNH 1993.2.5 6.7 5.0 5.4 3.9 4.3Falco columbarius BMNH 1930.3.24.264 6.0 5.0 4.7 3.6 4.0Falco columbarius BMNH 1930.3.24.267 6.7 5.3 5.0 3.5 4.3Falco columbarius BMNH 1988.61.1 6.9 5.4 5.0 3.8 4.4Falco naumanni BMNH 1955.15.2 6.0 4.6 4.8 3.2 4.0Falco naumanni BMNH 1961.13.4 5.6 4.7 4.6 3.3 3.5Falco newtoni BMNH 1897.5.10.29 5.6 4.5 4.5 3.1 4.3Falco sparverius BMNH 1954.3.2 5.6 4.5 4.4 3.5 4.0Falco subbuteo BMNH 1974.16.1 7.1 5.3 5.8 3.7 4.4Falco subbuteo BMNH 1985.76.1 6.7 5.1 5.2 3.3 4.1Falco subbuteo BMNH 1994.39.1 7.2 5.6 5.9 4.2 4.3Falco tinnunculus BMNH 1930.3.24.275 7.1 5.6 5.7 4.4 4.8Falco tinnunculus BMNH 1930.3.24.276 6.8 5.7 5.6 4.1 4.6Falco tinnunculus BMNH 1930.3.24.278 6.6 5.4 5.3 3.7 4.2Falco vespertinus BMNH 1855.4.4.9 5.7 4.6 4.7 3.4 4.1Falco vespertinus BMNH 1869.10.19.12 5.2 4.7 4.5 3.3 3.5Falco vespertinus BMNH 1952.3.122 6.2 4.9 4.6 3.6 4.4

“parallelity” of the surfaces of both condyles of the trochlea; F. vespertinus: biger, both condyles more parallel in dorsal view.

Phalanx 2 dig. II pedis dex. NMNHS 317 (Fig. 4 j, Table 15). The specimen differs from: F. columbarius: bigger and more robust; F. subbuteo: smaller, by the wid-er f. a. (measurement “c”); F. tinnunculus: by the smaller diameter of the phalangeal trochlea, narrower trochlea

27

Table 13The measurements of tarsometatarsus dex. prox. in some fossil and recent Falco (ref. to Fig. 1 L)



Falco chicquera BMNH 1965.18.1 4.2 3.6 5.4 8.9 7.2Falco chicquera BMNH 1993.2.5 3.3 3.2 4.7 7.8 7.0Falco columbarius BMNH 1930.3.24.264 2.9 2.8 4.0 6.5 6.4Falco columbarius BMNH 1930.3.24.267 3.9 2.9 4.5 7.6 7.0Falco columbarius BMNH 1988.61.1 3.4 3.1 4.4 7.5 6.9Falco naumanni BMNH 1955.15.2 3.0 2.7 3.8 6.5 6.2Falco naumanni BMNH 1961.13.4 3.1 2.7 3.7 6.3 5.7Falco newtoni BMNH 1897.5.10.29 3.0 2.6 3.7 5.2 5.8Falco sparverius BMNH 1954.3.2 3.1 2.7 3.6 6.2 5.7Falco subbuteo BMNH 1974.16.1 3.6 3.2 4.8 7.9 7.4Falco subbuteo BMNH 1985.76.1 3.6 3.4 4.8 7.6 7.0Falco subbuteo BMNH 1994.39.1 3.6 3.4 5.2 8.3 7.2Falco tinnunculus BMNH 1930.3.24.275 3.7 3.3 4.5 7.7 7.0Falco tinnunculus BMNH 1930.3.24.278 3.6 2.8 4.5 7.3 6.9Falco tinnunculus UCBL 119/7 3.4 2.8 3.8 7.7 7.0Falco vespertinus BMNH 1855.4.4.9 3.1 3.0 3.7 6.5 6.2Falco vespertinus BMNH 1869.10.19.12 2.8 2.7 3.6 6.0 5.4Falco vespertinus BMNH 1952.3.122 3.0 2.6 3.9 6.9 6.7

Table 14The measurements of phalanx 1 dig. I pedis in some fossil and recent Falco (ref. to Fig. 1 M)

Taxa a b c d e fFossil – Varshets

Falco bakalovi NMNHS 319 2.4 2.2 3.2 2.2 9.9 1.4Falco bakalovi NMNHS 320 2.6 2.6 3.3 2.3 10.3 1.3Falco bakalovi NMNHS 321 2.2 2.0 3.3 2.2 10.4 1.3

RecentFalco columbarius NMNHS 2/2002 2.2 2.4 3.3 2.2 9.7 1.2Falco subbuteo NMNHS 1/1989 2.6 2.5 4.1 2.9 12.6 1.6Falco subbuteo NMNHS 3/1993 2.2 2.0 3.2 2.2 10.3 1.2Falco tinnunculus NMNHS 10/1991 2.6 2.3 3.8 2.6 11.0 1.4Falco tinnunculus UCBL 119/1 2.5 2.4 3.9 2.5 11.4 1.5Falco tinnunculus UCBL 119/2 2.4 2.2 3.6 2.5 10.3 1.4Falco tinnunculus UCBL 119/3 2.2 2.1 3.3 2.3 9.8 1.3Falco tinnunculus UCBL 119/7 2.4 2.2 3.7 2.3 10.7 1.4Falco vespertinus NMNHS 1/1991 2.0 2.0 3.0 2.0 8.5 1.1

(measurement “a”), possibly indication of more primitive raptority, i. e. narrower and less strong claws; F. vesper-tinus: bigger and more robust.

Phalanx 2 dig. II pedis sin. NMNHS 318 (Fig. 4 k, Table 15). The specimen belongs to a juvenile/subadult individual of almost definite size. The specimen differs from: F. columbarius: narrower distal end; F. subbuteo: by the wider f. a. (measurement “c”); F. subbuteo: small-

er; F. tinnunculus: The same way as No 317; F. vesperti-nus: bigger and more robust.

Phalanx dist. dig. 1 pedis dex. NMNHS 181 (Fig.4 l, Table 16). The specimen differs from: F. columbarius: more robust; F. subbuteo: wider articular surfaces (meas-urement “d”); F. vespertinus: bigger in size and wider f. a.; F. vespertinus: bigger; Falco tinnunculus: by the shorter base of the attachment of the muscle.

28

dIScuSSIon

The comparative analysis of the examined finds un-equivocally shows that they could only be referred to the so-called “tinnunculus” group (Falco sp. ex gr. tin-nunculus) of the smaller falcons in the genus Falco, well separated from the “cherrug” group of the larger falcons. Some finds, however, because of the worse degree of their preservation, or being specimens of immature indi-viduals, could not be accepted as reliable proofs for the former existing of a concrete definite species of Falco sp. ex gr. tinnunculus. Having in mind the common col-lecting of all material (common sample and horizon), and their dimensional characteristics, we refer all finds to a single species. Its skeletal elements clearly differ from all compared species of the recent fauna. They could not be referred also to the hitherto known fossil taxa of genus Falco. Considerations for that have been given in

other paper (Boev, 1999a), describing the fossil species Falco bakalovi. The abundant excavated avian material (1589 finds, Boev, 2002; over 1700 finds, Boev, inpubl. data) collected so far, and its examination shows that no taxa are represented by now by a single find, and most of them are represented by several (even several dozens of bones).

The smallest Old World falcons (Microhierax Sharpe, 1874 and Polihierax Kaup, 1847) are excluded from our comparison, because of their evident strong metrical dif-ferences. Their skeletal elements are more than twice smaller compared to specimens of these genera.

The sexual dimorphism in size should also be taken into account. Hill (1944) examining Falconiformes in general, stated that falcons of genus Falco constitute a group of species demonstrating the most expressed sexual dimorphism (concerning length of the wing, tail, tarsus, and bill), reaching up to 37.8 % (male to female meas-

Table 15The measurements of phalanx 2 dig. II pedis in some fossil and recent Falco (ref. to Fig. 1 M)

Taxa a b c d e fFossil – Varshets

Falco bakalovi NMNHS 317 1.9 1.8 2.6 2.2 9.7 1.4Falco bakalovi NMNHS 318 1.9 1.9 2.6 2.2 9.8 1.3

RecentFalco columbarius NMNHS 2/2002 2.0 1.9 2.5 2.5 9.3 1.1Falco subbuteo NMNHS 1/1989 2.5 2.6 3.1 2.9 12.5 1.6Falco subbuteo NMNHS 3/1993 1.8 1.8 2.0 1.9 9.8 1.0Falco subbuteo UCBL 111/1 1.6 1.7 1.9 1.8 9.0 1.2Falco tinnunculus NMNHS 10/1991 2.1 2.0 2.4 2.4 9.8 1.3Falco tinnunculus UCBL 119/2 1.9 1.9 2.2 2.2 9.1 1.2Falco tinnunculus UCBL 119/3 2.0 2.0 2.5 2.4 9.7 1.3Falco tinnunculus UCBL 119/7 2.1 2.1 2.4 2.3 9.9 1.2Falco vespertinus NMNHS 1/1991 1.7 1.8 2.2 2.0 8.7 1.1

Table 16The measurements of phalanx dist. dig.I pedis in some fossil and recent Falco (ref. to Fig. 1 N)


Falco bakalovi NMNHS 181 3.2 2.8 ca. 10.2 2.6Recent

Falco columbarius NMNHS 2/2002 2.8 3.1 9.5 2.4Falco subbuteo NMNHS 1/1989 3.2 3.2 10.1 2.7Falco subbuteo NMNHS 3/1993 3.4 4.7 9.5 2.1Falco subbuteo UCBL 111/1 3.0 2.7 9.1 2.0Falco tinnunculus NMNHS 10/1991 3.8 2.9 10.5 2.6Falco tinnunculus UCBL 119/1 3.2 3.0 10.1 2.6Falco tinnunculus UCBL 119/2 3.1 2.7 10.2 2.4Falco tinnunculus UCBL 119/3 3.0 2.9 9.5 2.4Falco tinnunculus UCBL 119/7 3.5 2.9 10.2 2.8Falco vespertinus NMNHS 1/1991 4.0 2.8 8.0 1.8

29

urements). The only feature (tarsus length) where the bone structure determines much the size varies between 2.0 and 20.5 %. On the other hand, Cramp and Simmons (1980) noted that in F. t. tinnunculus, for example, the “sex differences [are] significant for adult and juvenile, except for tarsus”. According to them these differences in the wing length reach 6.30 to 8.48 %. Obviously the skeletal elements differ less.

Two measurements of the proximal ulna show that the length of depressio m. brachialis in F. bulgaricus is 12.3 mm, while in F. bakalovi it is 7.9 to 8.3. On the other hand, the longitudinal diameter of cotyla ventralis in F. bulgaricus is ca. 4.9 mm while in F. bakalovi it is 4.4 to 4.5 mm (Boev, 2011; Table 4). The general ap-pearance of the proximal ulna, distal humerus and dis-tal tibiotarsus of F. bakalovi considerably differs from F. bulgaricus. All these skeletal elements are much stouter and robust.

The morphology and size of F. bakalovi show that it was a close late Pliocene relative to recent F. tinnun-culus, having similar ecological features and biology. Until now it is the most abundant raptorial bird, found in the Varshets locality and in Bulgaria at all (26 finds; 4 individuals).

Acknowledgments

The Foundation Scientifique de Lyon et du Sud-Est (Lyon), the Short-term Visits Program of the Royal Society (London), the National Science Fund, Sofia, grant No B-202/1992), and the National Museum of Natural History (Sofia) have supported the study. The author is very grateful to Dr. Cécile Mourer-Chauviré (UCBL), Dr. Robert Prys-Jones and Dr. Joanne Cooper (BMNH)

for their hospitality and help during the research visits at their institutions. Special thanks to Dr. Nikolay Spassov (NMNHS) and an anonymous reviewer who improved the former version of the manuscript.

aPPendIX 1

examined specimens belonging to recent species of Falconiformes

Accipiter nisus (Linnaeus, 1758); UCBL 96/6; UCBL 96/10; Falco chicquera Daudin, 1800: BMNH 1965.18.1, BMNH 1993.2.5;Falco columbarius Linnaeus, 1758: BMNH 1930.3.24. 264, 1930.3.24.267, BMNH 1988.61.1; Falco naumanni Fleischer, 1818: BMNH 1955.15.2, BMNH 1961.13.4; Falco newtoni Gurney, 1863: BMNH 1897.5.10.29;Falco sparverius Linnaeus, 1758: BMNH 1954.3.2; Falco subboteo Linnaeus, 1758: BMNH 1974.16.1, BMNH 1994.39.1, UCBL 111/1, UCBL 111/2, UCBL 111/3, UCBL 111/4; Falco tinnunculus Linnaeus, 1758: BMNH 1930.3.24.275, BMNH 1930.3.24.276, BMNH 1930.3.24.278, UCBL 119/1, UCBL 119/2, UCBL 119/3, UCBL 119/4, UCBL 119/6, UCBL 119/7, UCBL 119/8; Falco vespertinus Linnaeus, 1766: BMNH 1855.4.4.9, BMNH 1869.10.19.12, BMNH 1952.3.122, UCBL 117/1, UCBL 117/2, UCBL 117/4; Microhierax caerulescens (Linnaeus, 1758): BMNH 2002.41.2; Polihierax semitorqutus (Smith, 1846): BMNH 2002.41.1.

reFerenceS

Baumel, J. J., Witmer, L. M. 1993. 4. Osteologia. In: Baumel, J., King, A., Breazile, J., Evans, H., Van den Berge, J., (Eds.). Handbook of Avian Anatomy, Nomina Anatomica Avium 23, 45–132. Nutall Ornithological Club.

Boev, Z. 1999a. Falco bakalovi sp. n. – a Late Pliocene falcon (Falconidae, Aves) from Varshets (W Bulgaria). Geologica Balcanica 29(1–2), 131–135.

Boev, Z. 1999b. Neogenski i kvaternerni ptitsi (Aves) ot Balgariya. [Neogene and Quaternary birds (Aves) from Bulgaria]. DSci. Thesis, National Museum of Natural History, Bulgarian Academy of Sciences, Sofia, 243 pp. supplements (in Bulgarian, unpublished).

Boev, Z. 2002. Neogene avifauna of Bulgaria. In: Zhou, Z., Zhang, F. (Eds.) Proceedings of the 5th Symposium of the Society of Avian Paleontology and Evolution, Beijing, 29–40.

Boev, Z. 2011. Falco bulgaricus sp. n. (Aves, Falconiformes) from the Middle Miocene of Hadzhidimovo (SW Bulgaria). Acta zoologica bulgarica 63(1), 17–35.

Cramp, S., K., Simmons, E. L. (Eds.). 1980. Handbook of the Birds of Europe the Middle East and North Africa. The

Birds of Western Palearctic, Vol. II. Hawks to Bustards. Oxford University Press, 695 pp.

Guerin, C. 1990. Biozones or Mammal Units? Methods and limits in Biochronology. In: Linday, E.H., Fahlbusch, V., Mein, P. (Eds.). European Neogene mammal Chronology. Plenum Press, New York, 116–130.

Hill, N. P. 1944. Sexual dimorphism in the Falconiformes. The Auk, 61, 228–234.

Livezey, B. C., Zusi, R. L. 2006. Higher-order phylogeny of modern birds (Theropoda, Aves: Neornithes) based on comparative anatomy: I. Methods and Characters. Bulletin of Carnegie Museum of Natural History, Pittsburgh, 37, 502–544.

Mein, P. 1990. Updating of MN zones. In: Lindsay, E.H., Fahlbusch, V., Mein, P. (Eds.) European Neogene mammal chronology. Plenum Press, New York, 73-90.

Mlíkovský, J., 2002. Cenozoic Birds of the World. Part 1, Europe. Praha, Ninox Press. 406 pp.

Popov, V. 2001. Late Pliocene voles (Mammalia, Arvicolidae) from Varshets (North Bulgaria). Acta zoologica cracovien-sia 44(2), 143–172.

30

Spassov, N. 1997a. Varshets and Slivnitsa – new localities of Villafranchian vertebrate fauna from Bulgaria (taxonomic composition, biostratigraphy and climatochronology). Geologica Balcanica 27(1–2), 83–90.

Spassov, N., 1997b. Villafranchian succession of mammalian megafaunas from Bulgaria and the biozonation of South-East Europe. In: Aguilar, J.-P., Legendre, S., Michaux, J. (Eds.), Actes du Congres Biochrom’97. Mémoires et travaux de l’Institut de Montpellier de l’École Pratique des Hautes Études 21, 669–676.

Spassov, N. 2000. Biochronology and zoogeographic affinities of the Villafranchian faunas of Bulgaria and South Europe. Historia naturalis bulgarica 12, 89–128.

Spassov, N. 2003. The Plio-Pleistocene vertebrate fauna in South-Eastern Europe and the megafaunal migratory waves from the east to Europe. Revue de Paléobiologie. 22(1), 197–229.

Umanskaya, A. S. 1981. The Miocene birds of the western Black Sea coasts of the Ukrainian SSR. Vestnik Zoologii 17(3), 17–21 (in Russian with English abstract).

White, C. M., Olsen, P. F., Kiff, L. F. 1994. Family Falconidae (Falcons and Caracaras). In: del Hoyo, J., Elliot, A., Sargatal, J. (Eds.). Handbook of the Birds of the World. Vol. 2. New World Vultures to Guineafowl. Lynx Edicions, Barcelona, 216–275.

Wikipedia 2011. Falcon. At: http://en.wikipedia.org/wiki/Falcon.

new fossil record of the late pliocene kestrel (falco ... · to the late pliocene kestrel falco...

Documents