falco bulgaricus sp. n. (aves: falconiformes) from...

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ACTA ZOOLOGICA BULGARICAActa zool. bulg., 63 (1), 2011: 17-35

Introduction Genus Falco comprises 37 recent species, 15 of them from Africa, Madagascar and adjacent islands, 11 – from South-East Asia, Australia and New Zealand, and 7 – from North and South Americas. Ten species occur both in Europe and Asia (WHITE et al., 1994). OLSON (1985) suggests an Afro-South-Asian origin of the genus, in spite of the present-day concentration of the species of family Falconidae in South America.

Few data are known on the fossil history of falcons Falco LINNASEUS, 1758. A summary of the fossil record of the family has been given in other paper (BOEV 1999). A total of 10 fossil species were described in the genus Falco until now, although re-cently only seven of them are considered as valid spe-cies (OLSON 1985, BOCHENSKI 1997, MLIKOVSKY 1996, 2002) – (1) F. antiquus MOURER-CHAUVIRÉ (1975)

(Middle Pleistocene of Noailles, France); (2) F. me-dius UMANSKAYA, 1981 (late Miocene (MN 11-13) from S Ukraine; the only known falcon of Miocene both of Eurasia and Africa; the oldest record of genus Falco at all); (3) Falco umanskajae, SOBOLEV 2003 (late Pliocene (MN 16) from the vicinities of Odessa, Ukraine); (4) F. bakalovi BOEV, 1999 (late Pliocene of Varshets, Bulgaria). MLÍKOVSKÝ (2002) considered F. antiquus as a synonym of F. cherrug GRAY, 1844, and listed only two fossil species of genus Falco: F. medius and F. bakalovi. (5) Falco chowi HOU LIANHAI 1982 is known from Pleistocene of China. Two fal-cons have been described from the New World locali-ties – (6) F. oregonus BRODKORB, 1946 (Early/Middle Pliocene of Oregon), and (7) F. kurochkini SUFIREZ & OLSON, 2001 (Late Pleistocene/Holocene of Cuba).

Falco bulgaricus sp. n. (Aves: Falconiformes) from the Late Miocene of Hadzhidimovo (SW Bulgaria)

Zlatozar Boev

National Museum of Natural History, Bulgarian Academy of Sciences, 1, Tsar Osvoboditel Blvd., 1000 Sofi a, Bulgaria; E-mail: [email protected]; [email protected]

Abstract: A new species of small falcon of the group ‘tinnunculus’ is described on the base of 16 bone fi nds of an unarticulated skeleton of an adult individual, dated late Miocene (Turolian – Maeotian, lower part of the zone MN 11-12). Diagnosis: A medium-sized fossil species in the genus Falco differing from the closest F. tinnunculus by: (1) coracoid – much shorter f. a. clavicularis in cranial view; (2) humerus – relatively longer diaphysis; (3) ulna – wider distal fourth of depressio m. brachialis; (4) tibiotarsus – longer base (more proximally positioned inception) of crista cnemialis lateralis, and less protruding area interarticu-laris; (5) tarsometatarsus – relatively shorter diaphysis. It is suggested that the splitting of the genus Falco into two major groups, ‘tinnunculus’ and ‘cherrug’ occurred at least in Late Pliocene, and probably it may be an indication of the super-genera/genera rank. The ‘tinnunculus’ group is of more ancient origin and it was diversifi ed in late Miocene. The ‘cherrug’ group is not recorded in Neogene and the all Tertiary. Its record appears even in Early Pleistocene, while the ‘tinnunculus’ group is well documented in Neogene of Europe since Late Pliocene.

Key words: Falcons, Fossil birds, Falconiformes, Late Miocene, Bulgaria

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Boev Z.

Abbreviations

Anatomical: dex. – dextra; dig. – digitus, digiti; dist. – distalis; f. a. – facies articularis; max. – maximum; proc. – processus, processi; prox. – proximalis; sin – sinistra; s. a. – sulcus articularis; tbt – tibiotarsus; tmt – tarsometatarsus; Institutional: NHM – Natural History Museum, formerly British Museum (Natural History), Tring; ISEAK – Institute of Systematics and Evolution of Animals (Polish Academy of Sciences), Krakow; NMNHS – National Museum of Natural History (Bulgarian Academy of Sciences), Sofi a.

Material and Methods The fi nds originate from the vicinity of the town of Hadzhidimovo near the town of Gotse Delchev (Blagoevgrad District; SW Bulgaria), Hadzhidimovo-1 locality, 41.30 N, 23.52 E; UTM grid: GM 30; 500 m a. s. l. They come from gray-yellowish-coloured sands and clay sands of oblique and complex inner stratifi cation at a depth of 1.00-1.50 m (DIMITAR KOVACHEV, NMNHS – pers. comm.). All the 16 bone fi nds (NoNo NMNHS 12539-12545; 12559-12567; Fig. 1, 2) represent parts of an unar-ticulated skeleton of one adult individual. It was col-lected by D. KOVACHEV in 1980s. Material is kept in the NMNHS. The fi ndings have been identifi ed through reference to comparative bird collections of the ISEAK, NHM and NMNHS.

Description of the manner of bone measuring in fossil and recent Falco: coracoid: a – thickness of processus acrocoracoideus; b – width of proces-sus acrocoracoideus; c – thickness of diaphysis in the cranial end of impressio m. sternocoracoidei on the cranial side; d – length of f. a. humeralis (Table 1); scapula dex. prox.: a – width of column scapu-lae beyond f. a. humeralis; b – thickness of column scapulae beyond f. a. humeralis (Table 2); humerus: a – maximum length of bone; b – length of crista pec-toralis from tuberculum dorsale; c – length of fossa brachialis; d – thickness of caput humeri; e – height of caput humeri on lateral side; f – thickness of the diaphysis at the foramen nutrinium; g – width of the diaphysis at the foramen nutrinium (Table 3); ulna: a – width in the middle of the diaphysis on the me-dial side; b – maximum width of proximal epiphysis; c – length of depressio m. brachialis; d – thickness

of proximal epiphysis; e – length (transversal diam-eter) of cotyla dorsalis; f – longitudinal diameter of cotyla ventralis (Table 4); radius: a – width of proxi-mal epiphysis; b – thickness of proximal epiphysis; c – minimum width of proximal half of diaphysis; d – thickness of epiphysis in tuberculum bicipitale (Table 5); femur: a – thickness of diaphysis in the end of its 1st (proximal) forth of its length; b – width of diaphysis in the same point (Table 6); tibiotarsus: a – length of crista fi bularis; b – thickness of the dia-physis in the distal end of crista fi bularis; c – width of diaphysis in the distal end of crista fi bularis; d – maximum diagonal width of proximal epiphysis; e – length from the proximal epiphysis to the distal end of the tibio-fi bular symphysis (at the caudal side); f – length of foramen interosseum distale (Table 7); tarsometatarsus: a – width of diaphysis in the proxi-mal end of the fossa metatarsi I; b – thickness of dia-physis in the proximal end of the fossa metatarsi I; c – minimum width of diaphysis (Table 8); phalanx 1 dig. I pedis: a – minimum width of the ‘diaphysis’; b – width of f. a. prox.; c – height of f. a. prox. (Table 9); phalanx 2 dig. II pedis dex.: a – minimum width of the ‘diaphysis’; b – width of f. a. prox.; c – height of f. a. prox. (Table 10); phalanx 2 dig. III pedis dex.: a – minimum width of the ‘diaphysis’; b – width of f. a. prox.; c – height of f. a. prox.; d – diameter of tro-chlea articularis (Table 11); phalanx 3 dig. III pedis dex. a – minimum width of the ‘diaphysis’; b – width of f. a. prox.; c – height of f. a. prox. (Table 12).

All measurements have been taken using cali-pers to 0.05 mm accuracy, but read to the 1st digit after decimal point. All generic names of the binomi-nals are given abbreviated in the text and are in full in the Appendix I. ‘Smaller’, ‘much smaller’, ‘big-ger’ or ‘much bigger’ in ‘Comparison and discus-sion’ section mean that the fossil specimen differs considerably in size from the specimens of com-pared species, and thus their taxonomic identity is excluded.

Two indices have been calculated with these measurements: (1) max. length of humerus (Table 3 – a): thickness of the diaphysis of humerus at the foramen nutrinium (Table 3 – f) (Fig. 3); (2) max. length of humerus (Table 3 – a): length from the proximal epiphysis to the distal end of the tibio-fi bular symphysis (at the lateral side) of tibiotarsus (Table 7 – e) (Fig. 4).

Falco bulgaricus sp. n. (Aves, Falconiformes) from the Late Miocene of Hadzhidimovo...

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Fig. 1. Falco bulgaricus sp. nov.: Skeletal elements of the fore limbs and the pectoral girdle: coracoid dex. NMNHS 12563 – cranial view (a), and caudal view (b); scapula dex. prox. NMNHS 12565 – ventral view (c); humerus sin. NMNHS 12567 – lateral view (d); humerus dex. prox. NMNHS 12561 – lateral view (e), and dorsal view (f); ulna sin. prox. NMNHS 12560 – medial view (g), lateral view (h), and dorsal view (i); radius dex. prox. NMNHS 12562 – dor-sal view (j); Skeletal elements of the hind limbs and the pelvis girdle: femur dex. prox. NMNHS 12564 – cranio-lateral view (k); tibiotarsus dex. NMNHS 12559 – lateral view (l). Scale bar = 1 cm. (Photos: Assen Ignatov).

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Boev Z.

Fig. 2. Falco bulgaricus sp. nov.: tibiotarsus dex. NMNHS 12559 – caudal view (a), and dorsal view (b); tibiotarsus sin. NMNHS 12539 – cranial view (c); tarsometatarsus sin. dist. NMNHS 12540 – cranial view (d); tarsometatarsus dex. dist. NMNHS 12566 (partly coated) – cranial view – (e); phalanx 1 dig. I pedis sin. NMNHS 12543 (f) – dorsal view; phalanx 2 dig. II pedis dex. NMNHS 12542 – medial view (g), ventral view (h), and caudal view (i); phalanx 2 dig. III pedis dex. NMNHS 12541 – dorsal view (j), and caudal view (k); phalanx 3 dig. III pedis dex. NMNHS 12544 – ventral view (l); Phalanx 3 dig. III pedis sin. NMNHS 12545 ventral view (m). Scale bar = 1 cm. (Photos: Assen Ignatov).


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The taxonomy follows WHITE et al. (1994). The osteological terminology is after BAUMEL & WITMER (1993) and, in some respects, LIVEZEY & ZUSI, 2006. The chronostratigraphy follows MEIN (1990).

Short description of the siteAccording to the rich associated terrestrial mega-fauna this locality is dated back to the Late Miocene (Turolian – Maeotian, lower part of the zone MN 11-12; ca. 7 million years ago). The associated land mammalian megafauna after SPASSOV (2002) in-cludes 30 taxa: Mustelidae indet., Hyaenotheriun gr. wongii (ZDANSKY, 1924), Hyaenictitheriini in-det., cf. Miohyaenotherium bessarabicum SEMENOV, 1989, Adcrocuta eximia (ROTH AND WAGNER, 1854),

Table 1. Measurements of coracoid in fossil and recent Falco.

Species a b c dFossil – HadzhidimovoFalco bulgaricus sp. n. NMNHS 12563 3.4 5.7 3.8 ca. 5.1RecentFalco naumanni ISEAK A 4783/90 2.5 5.0 2.6 6.6Falco naumanni ISEAK A 4958/91 2.8 5.5 2.4 6.6Falco subbuteo ISEAK A 3196/76 2.8 6.0 2.9 7.2Falco subbuteo ISEAK A 2071/69 2.9 6.8 3.1 6.6Falco subbuteo ISEAK A 3620/79 2.8 6.4 3.0 6.8Falco subbuteo ISEAK 230/41/14 3.1 6.2 3.0 6.8Falco subbuteo ISEAK 244/38/13 3.0 5.8 2.8 7.3Falco subbuteo ISEAK 416/62 3.3 6.7 3.0 7.2Falco subbuteo ISEAK A 1483/64 2.8 5.8 2.8 6.5Falco tinnunculus ISEAK A 4449/87 2.7 6.0 2.6 5.8Falco tinnunculus ISEAK A 4734/90 2.7 6.2 2.9 6.5Falco tinnunculus ISEAK A 4564/88 3.0 6.3 3.9 6.8Falco tinnunculus ISEAK A 4594/89 2.9 6.7 2.9 6.1Falco tinnunculus ISEAK A 4509/88 2.8 6.5 3.1 6.1Falco vespertinus ISEAK A 5056/92 2.7 4.6 2.6 6.1Falco columbarius NHM 1988.61.1 2.8 6.9 4.0 6.3Falco columbarius ISEAK A 2532/72 2.6 5.9 2.7 6.9Falco eleonorae ISEAK A 5093/92 3.8 9.1 3.8 8.8Falco biarmicus NHM 1976.60.5 ca. 3.8 7.7 5.2 9.4Falco sparverius ISEAK A 4106/84 2.4 4.0 2.1 5.2Falco longipennis ISEAK A 5351/94 3.9 6.9 3.1 7.3Falco cenchroides ISEAK A 4916/91 2.7 4.7 2.7 5.6Falco rupicoloides ISEAK A 4782/90 2.6 5.8 2.7 6.4Falco chicquera NHM 1993.2.5 3.1 5.0 3.1 ca.5.3Falco columbarius NHM 1862.1.18.3 ca. 3.3 - 3.7 5.7Falco vespertinus NHM 1855.4.4.9 3.1 5.9 3.1 5.6Falco columbarius NHM 1930.3.24.264 2.7 5.7 3.1 5.9Falco naumanni NHM 1961.13.4 2.2 5.7 4.0 5.1Falco naumanni NHM 1955.15.2 2.6 6.3 3.9 6.0Falco vespertinus NHM 1869.19.12 2.7 4.6 2.5 4.9

Table 2. Measurements of scapula prox. in fossil and re-cent Falco.

Species a bFossil – HadzhidimovoFalco bulgaricus sp. n. NMNHS 12565 4.0 2.5RecentFalco columbarius NMNHS 2/2002 3.8 2.0Falco columbarius NMNHS 1/1997 3.6 2.2Falco subbuteo NMNHS 1/1989 5.5 2.8Falco subbuteo NMNHS 3/1993 3.9 1.9Falco tinnunculus NMNHS 1/2002 2.8 1.8Falco tinnunculus NMNHS 7/1989 3.0 1.8Falco tinnunculus NMNHS 10/1992 4.0 2.4Falco vespertinus NMNHS 1/1991 2.8 2.0

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Boev Z.

Paramachairodus cf. orientalis GAUDRY, 1862, Machairodus giganteus (WAGNER, 1848), Metailurus cf. major ZDANSKY 1924, Gomphotheriidae indet., Mammut gr. borsoni, (HAYS, 1834), Ancylotherium pentelicum (GAUDRY & LARTET, 1856), Chalicotheriinae indet., ? Ceratotherium neumayri (OSBORN, 1900), Dicerorhinus pikermiensis GLOGER, 1841, Tapirus aff. jeanpiveteaui BOEUF, 1991, Hipparion gr. mediterraneum ROTH & WAGNER,

1855, Hipparion cf. brachypus HENSEL 1862, Hipparion ? platygenys GROMOVA, 1952, Microstonyx sp., Cervidae indet., Helladotherium duvernoyi (GAUDRY & LARTET, 1856), Palaeotragus rouenii, Gazella (Procapra) sp., Palaeoryx sp., Palaeoreas lindermayeri WAGNER, 1848, Tragoportax sp. – l, Tragoportax sp. – 2, Hystrix primigenia, WAGNER, 1848) and Mesopithecus cf. delsoni BONIS & BOUVRAIN, GERAADS & KOUFOS, 1990.

Table 3. Measurements of humerus in fossil and recent Falco.

Species a b c d e f gFossil – HadzhidimovoFalco bulgaricus sp. n. NMNHS 12561 - ca. 16.9 - 3.5 3.4 - -Falco bulgaricus sp. n. NMNHS 12567 ca. 58.0 ca. 17.0 6.7 - ca. 3.5 4.3 4.6RecentFalco biarmicus NHM 1976.60.5 75.4 23.9 10.9 5.6 5.6 5.5 6.0Falco cenchroides ISEAK A 4916/91 50.1 15.3 5.5 3.1 2.9 4.0 4.3Falco chicquera NHM 1993.2.5 51.2 16.0 5.1 3.4 3.7 3.3 4.2Falco columbarius NHM 1862.1.18.3 51.8 18.5 ca. 6.4 4.0 3.8 4.0 4.8Falco columbarius NHM 1930.3.24.264 46.1 15.0 6.6 3.5 3.4 3.9 4.4Falco columbarius NHM 1988.61.1 51.8 17.1 4.7 3.8 4.2 4.1 4.9Falco columbarius ISEAK A 2532/72 46.6 14.3 6.7 3.5 3.0 3.7 4.1Falco eleonorae ISEAK A 5093/92 70.1 19.9 7.9 5.3 5.0 5.4 6.2Falco longipennis ISEAK A 5351/94 53.8 17.7 6.6 4.1 3.6 4.4 5.0Falco naumanni NHM 1955.15.2 51.9 16.2 7.8 3.4 3.5 3.7 4.1Falco naumanni NHM 1961.13.4 48.0 8.9 7.0 2.9 3.3 3.3 3.5Falco naumanni ISEAK 4958/91 52.0 16.6 7.5 3.1 2.8 3.6 4.1Falco naumanni ISEAK A 4783/90 51.6 16.8 7.0 3.1 2.7 3.3 4.0Falco rupicoloides ISEAK A 4782/90 51.3 17.3 6.5 3.3 3.0 3.7 4.1Falco sparverius ISEAK A 4106/84 42.3 13.2 5.5 2.6 2.5 2.8 3.3Falco subbuteo ISEAK 230/41/14 53.4 16.5 6.4 3.9 3.7 4.4 4.8Falco subbuteo ISEAK 244/38/13 54.3 18.1 6.1 3.9 3.7 4.0 4.3Falco subbuteo ISEAK 416/62 62.3 20.0 6.8 4.3 3.8 4.7 5.2Falco subbuteo ISEAK A 1483/64 53.7 17.1 7.4 3.6 3.2 4.1 4.6Falco subbuteo ISEAK A 2071/69 53.2 16.9 6.6 4.0 3.5 4.2 4.8Falco subbuteo ISEAK A 3196/76 54.7 17.4 6.6 3.9 3.6 4.3 4.8Falco subbuteo ISEAK A 3620/79 52.9 17.6 7.9 4.3 3.5 4.0 4.6Falco tinnunculus ISEAK 4509/88 56.0 19.3 6.8 3.4 3.3 4.2 4.7Falco tinnunculus ISEAK 4594/89 53.6 17.2 7.3 3.3 3.1 4.0 4.1Falco tinnunculus ISEAK A 4449/87 55.0 19.4 7.6 3.3 2.9 3.8 4.2Falco tinnunculus ISEAK A 4564/88 53.7 18.4 7.7 3.1 3.0 4.5 4.1Falco tinnunculus ISEAK A 4734/90 54.9 18.9 7.0 3.4 3.2 4.0 4.5Falco vespertinus NHM 1855.4.4.9 50.5 18.4 7.0 3.3 3.8 3.3 3.8Falco vespertinus NHM 1869.19.12 50.1 16.5 6.6 2.9 3.1 3.4 3.8Falco vespertinus ISEAK A 2371/71 49.9 14.8 7.1 3.1 2.9 3.7 4.0Falco vespertinus ISEAK A 4737/90 49.6 16.7 6.7 3.1 2.8 3.5 3.8Falco vespertinus ISEAK A 5056/92 - 17.2 - 3.2 3.1 - -


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Fig.

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Falc

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491

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Falc

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993.

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Falc

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532/

72

Falc

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509

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Falc

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535

1/94

Falc

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MN

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955.

15.2

Falc

o na

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MN

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961.

13.4

Falc

o na

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ni IS

EAK

495

8/91

Falc

o na

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A 4

783/

90

Falc

o ru

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EAK

A 4

782/

90

Falc

o sp

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ISEA

K A

410

6/84

Falc

o su

bbut

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EAK

230

/41/

14

Falc

o su

bbut

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EAK

244

/38/

13

Falc

o su

bbut

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EAK

416

/62

Falc

o su

bbut

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EAK

A 1

483/

64

Falc

o su

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EAK

A 2

071/

69

Falc

o su

bbut

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EAK

A 3

196/

76

Falc

o su

bbut

eo IS

EAK

A 3

620/

79

Falc

o tin

nunc

ulus

ISEA

K 4

509/

88

Falc

o tin

nunc

ulus

ISEA

K 4

594/

89

Falc

o tin

nunc

ulus

ISEA

K A

444

9/87

Falc

o tin

nunc

ulus

ISEA

K A

456

4/88

Falc

o tin

nunc

ulus

ISEA

K A

473

4/90

Falc

o ve

sper

tinus

BM

NH

185

5.4.

4.9

Falc

o ve

sper

tinus

BM

NH

186

9.19

.12

Falc

o ve

sper

tinus

ISEA

K A

237

1/71

Falc

o ve

sper

tinus

ISEA

K A

473

7/90

NH

M

NH

M NH

MN

HM

NH

M

NH

MN

HM

NH

M

NH

M

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Boev Z.

The avian fauna includes 3 taxa: an accipitrid, Buteo spassovi BOEV & KOVACHEV 1998, a bucerot-id, Euroceros bulgaricus BOEV & KOVACHEV, 2007 and a struthionid, Struthio karatheodoris (BOEV & SPASSOV, 2009).

Systematic paleontology Order: FALCONIFORMES SHARPE, 1874Family: FALCONIDAE VIGORS, 1824Subfamily: Falconinae (VIGORS, 1824)Genus Falco LINNAEUS, 1758 Falco bulgaricus nov. sp.Holotype: Ulna sin. prox. NMNHS 12560

(Fig. 1 g, h, i); collections of the Vertebrate Animals

Department of the National Museum of Natural History – Sofi a, Bulgarian Academy of Sciences. Collected by D. KOVACHEV in 1980s.

Paratypes: coracoid dex. NMNHS 12563; hu-merus sin. NMNHS 12567; radius sinistra proxima-lis NMNHS 12562; tibiotarsus dex. NMNHS 12559; phalanx 1 dig. I pedis sin. NMNHS 12543; phalanx 2 dig. III pedis dex. NMNHS 12541.

Comparison: For morphological comparison see ‘Comparison and discussion’ section; for osteo-metric comparison see Tables 1-7.

Etymology: The name ‘bulgaricus’ is given af-ter the name of the country, Bulgaria, where the fi nds were discovered.

Table 4. Measurements of ulna prox. in fossil and recent Falco.

Species a b c d e fFossil – HadzhidimovoFalco bulgaricus sp. n. NMNHS 12560 3.6 ca. 7.8 12.3 ca. 5.5 3.8 ca. 4.9RecentFalco naumanni NHM 1955.15.2 3.6 7.6 12.0 5.2 3.4 3.2Falco naumanni NHM 1961.13.4 3.0 5.7 9.8 4.9 3.0 3.9Falco naumanni ISEAK A 4783/90 3.3 7.5 11.8 5.0 3.2 3.3Falco naumanni ISEAK 4958/91 3.6 7.5 12.0 5.3 3.2 3.7Falco subbuteo ISEAK A 3196/76 3.9 8.5 14.2 5.7 3.9 4.5Falco subbuteo ISEAK A 2071/69 3.8 8.0 11.8 5.4 3.3 -Falco subbuteo ISEAK A 3620/79 3.6 8.4 12.1 6.7 3.1 4.1Falco subbuteo ISEAK 230/41/14 3.7 8.9 11.2 5.8 3.8 4.1Falco subbuteo ISEAK 244/38/13 3.6 8.5 12.8 5.6 3.6 4.3Falco subbuteo ISEAK 416/62 4.2 9.3 13.9 5.9 4.0 4.3Falco tinnunculus ISEAK A 4564/88 3.6 7.8 11.7 5.3 2.9 3.8Falco tinnunculus ISEAK 4509/88 3.7 8.1 12.7 6.5 3.7 4.6Falco tinnunculus ISEAK 4594/89 3.5 7.8 13.0 5.5 3.0 4.2Falco tinnunculus ISEAK A 4449/87 3.1 7.6 11.5 6.1 3.1 4.3Falco tinnunculus ISEAK A 4734/90 3.5 7.9 11.4 5.3 3.7 4.3Falco vespertinus NHM 1869.19.12 3.2 7.2 12.0 4.8 3.1 3.8Falco vespertinus NHM 1855.4.4.9 3.3 7.8 10.8 5.2 3.3 4.2Falco vespertinus ISEAK A 2371/71 3.4 7.5 11.7 6.0 3.1 4.0Falco vespertinus ISEAK A 4737/90 3.2 6.7 12.3 4.9 3.0 4.2Falco columbarius NHM 1930.3.24.264 3.4 - ca. 11.5 - - -Falco columbarius ISEAK A 2532/72 3.3 6.4 8.0 5.8 3.1 3.9Falco columbarius NHM 1862.1.18.3 3.8 7.8 12.1 - - -Falco columbarius NHM 1988.61.1 3.6 8.0 11.8 5.3 3.8 4.5Falco eleonorae ISEAK A 5093/92 4.9 9.7 17.9 7.3 7.1 5.2Falco biarmicus NHM 1976.60.5 5.0 11.4 16.7 7.2 4.3 6.7Falco sparverius ISEAK A 4106/84 2.6 6.1 7.7 4.1 3.2 2.4Falco longipennis ISEAK A 5351/94 3.7 8.5 11.9 6.8 4.0 3.6Falco cenchroides ISEAK A 4916/91 3.4 7.3 10.8 5.8 3.2 3.4Falco chicquera NHM 1993.2.5 3.3 7.8 ca. 8.9 4.9 3.4 4.0Falco rupicoloides ISEAK A 4782/90 3.4 7.9 11.9 5.1 2.8 4.3


25

Measurements of the holotype: Table 4.Measurements of the paratypes: Tables 1, 3,

5, 7, 9, and 11. Diagnosis: A medium-sized fossil species in the

genus Falco differing from the closest F. tinnunculus: (1) coracoid – much shorter f. a. clavicularis in cra-nial view; (2) humerus – relatively longer diaphysis; (3) ulna – wider distal fourth of depressio m. brachi-alis; (4) tibiotarsus – longer base (more proximally positioned inception) of crista cnemialis lateralis, and less protruding area interarticularis; (5) tarsometatar-sus – relatively shorter diaphysis smaller.

Locality: Hadzhidimovo-1, near the town of Gotse Delchev (Blagoevgrad District; SW Bulgaria).

Table 5. Measurements of radius prox. in fossil and recent Falco.

Species a b c dFossil – HadzhidimovoFalco bulgaricus sp. n. NMNHS 12562 3.9 2.3 2.0 2.1RecentFalco naumanni ISEAK A 4783/90 3.3 2.4 2.0 2.1Falco naumanni ISEAK 4958/91 3.5 2.6 2.2 2.4Falco subbuteo ISEAK A 3196/76 3.9 3.0 2.0 2.6Falco subbuteo ISEAK A 2071/69 3.5 2.8 2.1 2.3Falco subbuteo ISEAK 230/41/14 3.9 2.8 2.8 2.8Falco subbuteo ISEAK 244/38/13 3.9 2.8 2.8 2.8Falco subbuteo ISEAK A 3620/79 3.8 2.9 2.4 2.5Falco subbuteo ISEAK 416/62 4.3 3.0 3.0 3.2Falco tinnunculus ISEAK A 4449/87 3.1 2.5 1.8 2.2Falco tinnunculus ISEAK A 4734/90 3.6 2.7 2.7 2.4Falco tinnunculus ISEAK A 4564/88 3.5 2.7 2.7 2.4Falco tinnunculus ISEAK 4509/88 3.6 2.9 3.0 3.0Falco tinnunculus ISEAK 4594/89 3.7 2.7 2.4 2.4Falco vespertinus ISEAK A 4737/90 3.3 2.3 2.0 1.9Falco vespertinus ISEAK A 2371/71 3.0 2.3 1.9 2.1Falco columbarius ISEAK A 2532/72 3.5 2.4 2.0 2.0Falco columbarius NHM 1988.61.1 3.8 2.8 2.0 2.5Falco eleonorae ISEAK A 5093/92 4.8 3.4 2.9 3.2Falco biarmicus NHM 1976.60.5 5.0 4.0 3.0 3.6Falco sparverius ISEAK A 4106/84 2.8 2.0 1.8 1.8Falco longipennis ISEAK A 5351/94 3.5 2.8 2.3 2.5Falco cenchroides ISEAK A 4916/91 3.4 2.5 2.4 2.4Falco rupicoloides ISEAK A 4782/90 3.5 2.4 2.3 2.3Falco chicquera NHM 1993.2.5 3.4 ca. 2.5 ca. 1.9 1.8Falco vespertinus NHM 1855.4.4.9 3.5 2.5 1.7 2.4Falco columbarius NHM 1930.3.24.264 3.4 2.5 1.9 2.1Falco naumanni NHM 1961.13.4 3.0 2.2 1.5 2.2Falco naumanni NHM 1955.15.2 3.1 2.4 1.7 1.8Falco vespertinus NHM 1869.19.12 3.1 2.4 2.8 2.4

Table 6. Measurements of femur dex. in fossil and recent Falco.

Species a bFossil – HadzhidimovoFalco bulgaricus sp. n. NMNHS 12564 3.7 4.0RecentFalco columbarius NMNHS 2/2002 4.0 4.5Falco columbarius NMNHS 1/1997 4.0 4.6Falco subbuteo NMNHS 1/1989 4.4 5.4Falco subbuteo NMNHS 3/1993 4.1 5.0Falco tinnunculus NMNHS 1/2002 3.3 3.7Falco tinnunculus NMNHS 7/1989 3.5 3.8Falco tinnunculus NMNHS 10/1992 4.2 4.5Falco vespertinus NMNHS 1/1991 3.6 2.4

26

Boev Z.

Fig.

4.

Cor

rela

tion

betw

een

the

max

imum

leng

th o

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erus

and

the

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th fr

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ysis

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ular

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the

spec

ies o

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.

1012141618202224

4045

5055

6065

7075

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a

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Falc

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icus

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. NM

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930.

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Falc

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61.1

Falc

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532/

72

Falc

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rae

ISEA

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509

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Falc

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535

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Falc

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MN

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961.

13.4

Falc

o na

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495

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Falc

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783/

90

Falc

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782/

90Fa

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106/

84

Falc

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230

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14

Falc

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244

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416

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Falc

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319

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Falc

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620/

79

Falc

o tin

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ulus

ISEA

K 4

509/

88

Falc

o tin

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ulus

ISEA

K 4

594/

89Fa

lco

tinnu

ncul

us IS

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A 4

449/

87

Falc

o tin

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ulus

ISEA

K A

456

4/88

Falc

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ISEA

K A

473

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Falc

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BM

NH

185

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4.9

Falc

o ve

sper

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186

9.19

.12

Falc

o ve

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ISEA

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237

1/71

NH

M NH

MN

HM

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M

NH

MN

HM

NH

M

NH

MN

HM


27

Chronology: Late Miocene (Turolian – Maeotian, lower part of the zone MN 11-12).

Comparison and Discussion The general morphology of all skeletal elements un-equivocally indicates that the fi ndings belong to a medium-sized falconiform of Falconidae. The size of all skeletal elements and the general comparison with genus Falco show considerable morphological similarity and suggest fi rm affi liation to that genus.

According to general dimensions, the speci-men of Hadzhidimovo is referred to the s. c. ‘tin-

nunculus’ group (Falco sp. ex gr. tinnunculus) of the smaller falcons in the genus Falco, well separated from ‘cherrug’ group of the larger falcons. That is why the morphological comparison will be concen-trated on the comparison mainly with the species of ‘tinnunculus’ group. On the other hand, the small-est Old World falcons (Microhierax SHARPE, 1874 and Polihierax KAUP, 1847) are excluded from our comparison, because of their evident strong and sig-nifi cant metrical differences. Their skeletal elements are more of twice smaller (WHITE et al. 199) than the compared specimen. The late Pliocene F. bakalovi

Table 7. Measurements of tibiotarsus dist. in fossil and recent Falco.

Species a b c d e fFossil – HadzhidimovoFalco bulgaricus sp. n. NMNHS 12539 7.5 3.4 4.2 - - -Falco bulgaricus sp. n. NMNHS 12559 7.0 3.6 4.4 ca. 8.6 ca. 17.3 23.8RecentFalco biarmicus NHM 1976.60.5 13.2 4.3 6.4 12.0 24.0 22.0Falco cenchroides ISEAK A 4916/91 8.7 3.1 4.7 7.7 16.9 17.2Falco chicquera NHM 1993.2.5 9.1 2.8 3.9 8.6 18.2 17.2Falco columbarius NHM 1862.1.18.3 8.0 3.4 5.0 9.2 19.9 18.8Falco columbarius NHM 1930.3.24.264 6.8 2.8 4.4 7.9 15.8 17.0Falco columbarius NHM 1988.61.1 9.6 3.0 4.8 9.1 17.0 17.6Falco columbarius ISEAK A 2532/72 6.7 2.9 4.4 7.6 14.2 19.3Falco eleonorae ISEAK A 5093/92 9.0 3.7 4.3 10.4 19.5 16.3Falco longipennis ISEAK A 5351/94 8.4 3.5 4.6 8.8 18.6 17.4Falco naumanni NHM 1955.15.2 8.4 2.8 3.0 7.2 16.8 14.9Falco naumanni NHM 1961.13.4 6.8 2.8 3.1 6.7 14.8 15.0Falco naumanni ISEAK 4958/91 7.7 2.8 3.6 7.4 15.9 15.9Falco naumanni ISEAK A 4783/90 8.1 2.8 4.0 7.4 15.0 15.4Falco rupicoloides ISEAK A 4782/90 8.0 2.9 4.4 7.7 14.1 13.7Falco sparverius ISEAK A 4106/84 7.1 2.6 3.1 6.7 14.1 13.8Falco subbuteo ISEAK 230/41/14 7.6 3.2 4.5 8.4 15.8 17.7Falco subbuteo ISEAK 244/38/13 7.7 2.9 4.2 8.1 15.6 18.4Falco subbuteo ISEAK 416/62 9.8 3.2 4.2 8.8 17.6 15.3Falco subbuteo ISEAK A 2071/69 7.0 2.8 4.4 8.3 15.6 17.3Falco subbuteo ISEAK A 3196/76 7.8 3.0 3.9 8.4 11.8 16.7Falco subbuteo ISEAK A 3620/79 7.3 3.0 4.3 8.7 16.0 17.6Falco tinnunculus ISEAK 4509/88 10.7 3.3 4.5 8.4 18.7 18.0Falco tinnunculus ISEAK 4594/89 12.2 3.4 4.8 9.5 20.3 15.8Falco tinnunculus ISEAK A 4449/87 10.3 3.0 4.8 8.2 17.6 19.9Falco tinnunculus ISEAK A 4564/88 10.1 3.2 4.8 8.6 17.4 19.7Falco tinnunculus ISEAK A 4734/90 10.1 3.4 4.6 8.4 17.6 18.4Falco vespertinus NHM 1855.4.4.9 7.4 2.7 3.3 7.1 16.1 14.4Falco vespertinus NHM 1869.19.12 7.1 2.5 2.4 7.1 13.9 14.8Falco vespertinus ISEAK A 2371/71 7.1 2.7 4.0 ca. 7.0 13.8 19.8

28

Boev Z.

differs by the smaller size (comparable to modern F. columbarius and F. tinnunculus), while F. bulgari-cus sp. n. dimensionally stands between F. tinnun-culus and F. subbuteo. UMANSKAJA (1981) stated that (in terms of the carpometacarpus length) F. medius stands closer to F. tinnunculus, F. vespertinus and F. naumanni (Table 13). Its holotype is a left carpomet-acarpus and no other materials have been collected. The type locality is Cherevichniy Hutor (Belyaevskiy District of the Odessa Region), which is about 670 km from the Hadzhidimovo locality, and the age is almost the same as of F. bulgaricus sp. n.

Figs 3 and 4 clearly show that F. bulgaricus sp. n. was a larger ‘small’ falcon, standing between F. tinnunculus and F. subbuteo. Even more its rela-tively longer wings and shorter legs indicate a spe-cifi c morphologic adaptation. Anyway, thus we ex-clude any taxonomical identity between F. medius and the falcon of late Miocene from Hadzhidimovo. Although we have much more skeletal elements of F. bulgaricus sp. n., the carpometacarpus is the only large bone of ‘ossa longa tubulosa’, unrepresented in the collected material. Thus it remains incomparable. For the comparison purpose we calculated the mean

Table 8. Measurements of tarsometatarsus in fossil and recent Falco.

Species a b cFossil – HadzhidimovoFalco bulgaricus sp. n. NMNHS 12540 3.5 1.9 -Falco bulgaricus sp. n. NMNHS 12566 - - 2.2RecentFalco naumanni ISEAK 4958/91 3.0 2.0 -Falco naumanni ISEAK A 4783/90 2.7 1.9 -Falco subbuteo ISEAK A 3196/76 2.8 2.5 -Falco subbuteo ISEAK A 2071/69 2.7 2.0 -Falco subbuteo ISEAK A 3620/79 2.8 2.0 -Falco subbuteo ISEAK 230/41/14 2.9 2.0 -Falco subbuteo ISEAK 244/38/13 2.7 1.8 -Falco subbuteo ISEAK 416/62 3.2 2.3 -Falco tinnunculus ISEAK 4509/88 3.6 2.3 -Falco tinnunculus ISEAK 4594/89 3.4 2.4 -Falco tinnunculus ISEAK A 4449/87 3.0 2.0 -Falco tinnunculus ISEAK A 4734/90 3.3 2.0 -Falco tinnunculus ISEAK A 4564/88 3.4 2.0 -Falco vespertinus ISEAK A 2371/71 2.6 1.8 -Falco columbarius ISEAK A 2532/72 2.8 2.0 -Falco columbarius NHM 1988.61.1 3.0 2.5 -Falco eleonorae ISEAK A 5093/92 3.8 2.5 -Falco biarmicus NHM 1976.60.5 5.4 3.3 -Falco sparverius ISEAK A 4106/84 2.5 1.6 -Falco longipennis ISEAK A 5351/94 3.3 2.2 -Falco cenchroides ISEAK A 4916/91 3.3 2.0 -Falco rupicoloides ISEAK A 4782/90 2.8 1.9 -Falco chicquera NHM 1993.2.5 3.0 2.2 -Falco columbarius NHM 1862.1.18.3 3.2 2.2 -Falco vespertinus NHM 1855.4.4.9 3.0 1.8 -Falco columbarius NHM 1930.3.24.264 2.8 2.0 -Falco naumanni NHM 1961.13.4 2.6 1.8 -Falco naumanni NHM 1955.15.2 2.7 2.0 -Falco vespertinus NHM 1869.19.12 2.6 1.8 -


29

values of both carpometacarpus and humerus for the falcons (Table 14) and the ratio between the humerus length and the carpometacarpus length (Table 15). Thus the extraplolated humerus length of Falco me-dius using the mean proportion (1.271) for ‘tinnun-culus’ group is ca. 50.6 mm, while the extrapolated carpometacarpus length of Falco bulgaricus sp. n. using the same proportion is ca. 45.6 mm.

Skeletal elements of fore limbs and pectoral girdle

Coracoid dex. NMNHS 12563 (Fig. 1 – a, b; Table 1). F. biarmicus: much smaller size, and the relatively narrower humeral part of the bone; F. cenchroides: considerably larger size; F. rupi-coloides: larger size; F. chicquera: similar in size and general morphology; F. columbarius: thicker pr. acrocoracoideus (measurement ‘c’), thinner diaphy-sis (measurement ‘d’); F. longipennis: thicker proc. acrocoracoideus (measurement ‘a’); F. naumanni: larger, wider s. m. supracoracoidei, and less rounded proc. acrocoracoideus; F. subbuteo: thicker pr. ac-rocoracoideus (measurement ‘a’) and almost twice wider impressio ligamenti acrocoracohumeralis; F. tinnunculus: similar both in general morphology and size, but sharper proc. acrocoracoideus, better de-veloped longitudinal groove on the supracoracoidal surface, and much shorter f. a. clavicularis in cranial view; F. vespertinus: bigger, and the thicker proc. ac-rocoracoideus.

Scapula dex. prox. NMNHS 12565 (Fig. 1 – c; Table 2). F. columbarius: bigger and much thick-er; F. subbuteo: thicker margo dorsalis at the area of collum scapulae; F. tinnununculus: more robust; F. vespertinus: bigger and more robust.

Humerus sin. NMNHS 12567 (Fig. 1 – d; Table 3) and humerus dex. prox. NMNHS 12561 (Fig. 1 – e, f; Table 3). F. biarmicus: resembles by the position of the foramen nutritium, and by the presence of two well developed transversal and par-allel grooves on the lateral side of the bone between the intumescentia, but differs by the much smaller size; F. chicquera: longer humerus, and the more proximally positioned foramen nutritium; F. colum-barius: considerably longer humeral bone and more proximally, instead distally, positioned foramen nu-tritium on the diaphysis. (The correlation between the measurements ‘a’ and ‘b’ in the fossil specimen

is 12.8, against 14.1 in F. columbarius); F. eleono-rae: much smaller; F. longipennis: longer and thin-ner diaphysis – the clearest difference; F. naumanni: larger, deeper foramen nutritium, deeper fossa m. brachialis, and longer humeral bone; F. sparverius: too smaller; F. tinnunculus: clearly differs in propor-tions as its diaphysis is relatively longer. Foramen nu-tritium is situated much more distally. Nevertheless most of the details are similar, confi rming its refer-ring to ‘tinnunculus’ group of genus Falco; F. ves-pertinus: considerably larger size, the presence of two well developed transversal and parallel grooves (instead one hardly seen) on the lateral side of the bone between the intumescentia and the neighboring narrower diaphysal parts (they are not represented in all recent small falcons /!/), and the longer and thin-ner diaphysis; F. subbuteo: smaller, more S-shape bent instead straight in dorsal view.

Ulna sin. prox. NMNHS 12560 (Fig. 1 – g, h, i; Table 4). F. biarmicus: smaller, and deeper depres-sio m. brachialis; F. cenchroides: larger, wider cotyla dorsalis; F. chicquera: larger, and deeper depressio m. brachialis; F. columbarius: more proximally po-sitioned foramen nutritium is the only difference; F. eleonorae: much smaller; F. longipennis: small-er; F. sparverius: considerably larger; F. subbuteo: more proximally situated tuberculum bicipitale, and sharper linea intermuscularis; F. tinnunculus: similar, but wider distal fourth of depressio m. brachialis; F. vespertinus: better marked longitudinal linea inter-muscularis; Falco naumanni: larger, deeper fossa m. brachialis (esp. in its proximal end), and better de-veloped tuberculum lig. collateralis ventralis.

Radius dex. prox. NMNHS 12562 (Fig. 1 – j; Table 5). F. biarmicus: much smaller, better de-veloped margo interosseus; F. chicquera: slightly larger; F. tinnunculus: shallower f. a. ulnaris; F. ves-pertinus: larger and bigger tuberculum bicipitale; Falco columbarius: bigger tuberculum bicipitale; F. subbuteo: slightly smaller; F. tinnunculus: more pro-truding f. a. ulnaris.

Skeletal elements of the hind limbs and the pelvis girdle

Femur dex. prox. NMNHS 12564 (Fig. 1 – k; Table 6). F. columbarius: better developed linea inter-muscularis cranialis; F. subbuteo: slightly smaller and less developed impressiones iliotrochantericae;

30

Boev Z.

F. tinnununculus: more gracile and slender; F. ves-pertinus: slightly bigger.

Tibiotarsus dex. NMNHS 12559 (Fig. 1 – l and Fig. 2 – a, b; Table 7) and tibiotarsus sin. prox. NMNHS 12539 (Fig. 2 – c; Table 7). F. biarmicus:

much smaller, and the more medially, but not later-ally, positioned base of the crista cnemialis lateralis; F. chicquera: more medially, but not laterally, posi-tioned base of crista cnemialis lateralis; F. columbar-ius: similar in size, but more distal position of crista

Table 10. Measurements of phalanx 2 dig. II pedis in fossil and recent Falco.

Species a b cFossil – HadzhidimovoFalco bulgaricus sp. n. NMNHS 12542 2.0 2.7 2.5RecentFalco columbarius NMNHS 2/2002 1.6 1.9 2.0Falco subbuteo NMNHS 1/1989 2.0 3.1 2.9Falco subbuteo NMNHS 3/1993 1.6 2.5 2.3Falco tinnunculus NMNHS 1/2002 2.2 2.7 2.6Falco tinnunculus NMNHS 7/1989 2.0 2.8 2.7Falco tinnunculus NMNHS 10/1992 2.1 2.9 2.7Falco vespertinus NMNHS 1/1991 1.4 1.9 1.9

Table 9. Measurements of phalanx 1 dig. I pedis in fossil and recent Falco.

Species a b cFossil – HadzhidimovoFalco bulgaricus sp. n. NMNHS 12543 1.9 3.6 2.4RecentFalco naumanni NHM 1955.15.2 1.6 3.0 2.0Falco naumanni ISEAK 4958/91 1.7 3.0 2.2Falco naumanni ISEAK A 4783/90 1.3 2.9 2.4Falco subbuteo ISEAK A 3196/76 1.8 3.3 2.2Falco subbuteo ISEAK A 3620/79 1.8 3.3 2.4Falco subbuteo ISEAK 416/62 1.9 3.5 2.5Falco tinnunculus ISEAK 4509/88 2.2 3.7 2.5Falco tinnunculus ISEAK 4594/89 2.2 3.7 2.3Falco tinnunculus ISEAK A 4449/87 1.9 3.2 2.2Falco tinnunculus ISEAK A 4734/90 2.1 3.3 2.4Falco tinnunculus ISEAK A 4564/88 2.3 3.6 2.3Falco vespertinus NHM 1855.4.4.9 1.7 3.2 ca. 2.0Falco vespertinus NHM 1869.19.12 1.8 2.6 2.0Falco vespertinus ISEAK A 2371/71 1.7 2.8 1.9Falco columbarius NHM 1930.3.24.264 1.6 3.0 2.2Falco columbarius ISEAK A 2532/72 1.8 3.2 2.2Falco columbarius NHM 1988.61.1 1.7 2.7 2.4Falco columbarius NHM 1862.1.18.3 1.8 3.4 2.6Falco eleonorae ISEAK A 5093/92 2.4 4.4 3.0Falco biarmicus NHM 1976.60.5 3.2 5.6 3.8Falco sparverius ISEAK A 4106/84 1.7 2.7 1.9Falco longipennis ISEAK A 5351/94 1.8 3.2 2.3Falco cenchroides ISEAK A 4916/91 2.0 3.4 2.2


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Table 11. Measurements of phalanx 2 dig. III pedis in fossil and recent Falco.

Species a b c d

Fossil – HadzhidimovoFalco bulgaricus sp. n. NMNHS 12541 1.7 2.5 2.3 1.9RecentFalco columbarius NMNHS 2/2002 1.4 2.3 2.1 1.6Falco subbuteo NMNHS 1/1989 1.7 3.0 2.5 2.0Falco subbuteo NMNHS 3/1993 1.3 2.2 1.9 1.5Falco tinnunculus NMNHS1/2002 1.7 2.2 2.2 1.9Falco tinnunculus NMNHS 7/1989 1.6 2.6 2.6 1.9Falco tinnunculus NMNHS 10/1992 1.7 2.7 2.5 2.0Falco vespertinus NMNHS 1/1991 1.4 2.2 2.1 1.7

Table 12. Measurements of phalanx 3 dig. III pedis in fossil and recent Falco.

Species a b cFossil – HadzhidimovoFalco bulgaricus sp. n. NMNHS 12544 ca. 1.6 ca. 2.2 ca. 1.8Falco bulgaricus sp. n. NMNHS 12545 ca. 1.6 - -RecentFalco columbarius NMNHS 2/2002 1.3 2.4 1.6Falco subbuteo NMNHS 3/1993 1.4 2.0 1.9Falco tinnunculus NMNHS 1/2002 1.6 2.4 2.2Falco tinnunculus NMNHS 7/1989 1.7 2.4 2.2Falco tinnunculus NMNHS 10/1992 1.8 2.4 2.3Falco vespertinus NMNHS 1/1991 1.4 1.9 2.1

Table 13. Measurements of the carpometacarpus in fossil and recent Falco.

Species Maximal length

Maximal width of proximal epiphysis

Width of dis-tal epiphysis

Length of spa-tium interme-

ta-carpalis

Fossil – Cherevichniy Hutor - Ukraine1

Falco medius 45-4033 39.8 10.4 7.4 26.2Recent - Bulgaria‘tinnunculus’ group Falco vespertinus NMNHS 1/1991 34.1 8.4 6.1 22.2Falco tinnunculus NMNHS 8/1989 44.6 19.4 16.3 31.5Falco tinnunculus NMNHS 7/1989 44.6 19.3 17.4 32.4Falco tinnunculus NMNHS 19/2005 45.5 19.4 17.0 33.1Falco columbarius NMNHS 2/2002 37.0 14.1 10.6 25.1Falco subbuteo NMNHS 1/1989 53.0 17.5 18.8 40.7Falco subbuteo NMNHS 3/1993 49.7 20.4 17.0 35.0‘cherrug’ group Falco peregrinus NMNHS 2/1989 64.2 21.8 15.8 43.5Falco cherrug NMNHS 1/1990 71.9 26.5 22.4 50.8

After Umanskaya (1981).

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Boev Z.

Table 14. Mean values (for n >1) of the total length of carpometacarpus and humerus of some fossil and recent Falco.

Species n Total length of the bone

carpometacarpusFossil – Cherevichniy Hutor - Ukraine2

Falco medius 45-4033 1 39.8Recent – Bulgaria‘tinnunculus’ group Falco columbarius 1 37.0Falco subbuteo 2 50.5Falco tinnunculus 3 44.9Falco vespertinus NMNHS 1/1991 1 34.1‘cherrug’ group Falco cherrug NMNHS 1/1990 1 71.9Falco peregrinus NMNHS 2/1989 1 64.2humerusFossil – HadzhidimovoFalco bulgaricus sp. n. NMNHS 12567 1 ca. 58.0Recent‘tinnunculus’ group Falco columbarius 4 49.0Falco subbuteo 7 53.9Falco tinnunculus 5 54.6Falco vespertinus 4 50.5Falco cherrug NMNHS 1/1990 1 97.3Falco peregrinus NMNHS 2/1989 1 84.1

Table 15. Humerus length to carpometacarpus length in some fossil and recent Falco.

Species n Humerus length: carpometacarpus length

FossilFalco medius 45-4033 1 -Falco bulgaricus sp. n. 1 -Recent‘tinnunculus’ group Falco columbarius NMNHS 2/2002 1 1.324Falco subbuteo 2 1.067Falco tinnunculus 3 1.216Falco vespertinus NMNHS 1/1991 1 1.480Falco spp. ex gr. tinnunculus (mean) 7 1.271‘cherrug’ group Falco cherrug NMNHS 1/1990 1 1.269Falco peregrinus NMNHS 2/1989 1 1.309

After Umanskaya (1981).


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fi bularis, and more medial, than lateral, inception of distal end of crista cnemialis cranialis; F. longipen-nis: dimensionally and morphologically similar, but sharper bend of crista cnemialis cranialis; F. nauman-ni: bigger, relatively longer crista fi bularis, uneven crista fi bularis, and smaller general size of bone; F. sparverius: much larger; F. subbuteo: more medially, but not laterally, positioned base of the crista cnemia-lis lateralis, longer base of crista cnemialis lateralis, deeper fossa fl exoria and longer foramen interosseum distale, nevertheless general dimensional similarity; F. tinnuncculus: longer base (more proximally posi-tioned inception) of crista cnemialis lateralis, and less protruding area interarticularis; F. vespertinus: con-siderably larger, and presence of a shallow concavity on the cranial surface of crista fi bularis. F. subbuteo: slightly smaller, longer foramen interosseum distale.

Tarsometatarsus sin. dist. NMNHS 12540 (Fig. 2 – d; Table 8) and tarsometatarsus dex. dist. NMNHS 12566 (Fig. 2 – e; Table 8). F. biarmicus: much smaller size, and more cranio-caudally fl at diaphysis; F. chicquera: lower crista plantaris me-dialis; F. columbarius: wider distal part of diaphysis at foramen vasculare distale, considerably less de-veloped crista plantaris mediana in plantar aspect, and less developed edge between the f. subcutanea lateralis and the f. dorsalis in dorsal aspect; F. nau-manni: larger size, shallower relief on distal part of dorsal surface of tmt, and deeper groove of foramen vasculare distale on the cranial surface; F. subbuteo: the same way as F. columbarius; F. tinnununculus: smaller dimensions of tmt; F. vespertinus: bigger size, and lower crista plantaris medialis.

Phalanx 1 dig. I pedis sin. NMNHS 12543 (Fig. 2 – f; Table 9). F. columbarius: larger size and bigger asymmetry in proximal half of the phalanx; F. naumanni: bigger size, deeper relief on ventral (plantar) surface, and straighter shaft; F. subbuteo: smaller and more concave in proximal end on facies plantaris; F. tinnununculus: slightly smaller, less con-cave f. plantaris in prox. end; F. vespertinus: larger.

Phalanx 2 dig. II pedis dex. NMNHS 12542 (Fig. 2 – g, h, i; Table 10). F. columbarius: bigger; F. subbuteo: slightly smaller, and more concave in prox-imal end of facies plantaris; F. tinnununculus: slightly smaller; F. vespertinus: bigger and more robust.

Phalanx 2 dig. III pedis dex. NMNHS 12541 (Fig. 2 – j, k; Table 11). F. columbarius: bigger,

plantar edge of f. a. prox. less concave; F. subbuteo: more concave in proximal end of facies plataris; F. tinnununculus: smaller, less concave profi le of troch-lea articularis in dorsal view; F. vespertinus: thicker phalangeal body.

Phalanx 3 dig. III pedis dex. NMNHS 12544 (Fig. 2 – l; Table 12). and Phalanx 3 dig. III pedis sin. NMNHS 12545 (Fig. 2 – m; Table 12). F. co-lumbarius: slightly bigger; F. subbuteo: smaller; F. tinnununculus: smaller; F. vespertinus: bigger and more robust.

General comparison. The comparison of skel-etal elements of the fore-limbs/pectoral girdle and the hind-limbs/pelvic girdle clearly demonstrate more developed fl ight capability, i. e. F. bulgaricus sp. n. to a certain extent was more aerial than ter-restrial, in comparison to its modern closer relatives. Its wings were relatively more developed, while the legs were less developed in comparison to the clos-est dimensionally and in many respects, morphologi-cally, species. The compared specimen has relatively longer wings (longer proximal part, i. e. humeri) and shorter legs (shorter distal part, i.e. thinner, and pos-sibly shorter, tarsometatarsi).

The two parallel transversal grooves (lineae intermusculares) on the lateral side of the humeral bone bellow the intumescentia, marking the end of crista bicipitalis, are specifi c both for the spe-cies of ‘tinnunculus’ group, and for the specimen of Hadzhidimovo and they are lacking in all the com-pared species of ‘cherrug’ group. Thus they may be a plesiomorphic ferature for the smaller falcons.

Conclusions

The presence of parallel transversal grooves on humerus could be considered as one of the slight osteological distinguishing features, suggesting an ancient, at least at Late Pliocene separation of these groups of falcons – (1) ‘tinnunculus’, kestrels (including hobbies, g. Hypotriorchis BOIE, 1826), and (2) ‘cherrug’, hierofalcons Hierofalco CUVER, 1817 (including Peregrine Falcon Falco peregrinus TUNSTALL, 1771). On the other hand, the position of the foramen nutritium of humeral bone (NMNHS 12567) is more distal, approaching to F. vesperinus, instead to F. tinnunculus.

34

Boev Z.

Probably these major groups deserve to be ranked as separate super-genera/genera. According to the fossil record the large falcons (of ‘cher-rug’ group) appeared in the Early Pleistocene and they are not represented in the Tertiary record (MLÍKOVSKÝ, 2002). On the other side, the Neogene record of small falcons (of ‘tinnunculus’ group) in Europe is well documented (Bulgaria, Hungary and Ukraine, all of Late Pliocene) (MLÍKOVSKÝ, 2002). The relatively younger origin of ‘cherrug’ complex

in comparison to smaller falcons is also confi rmed by NITTINGER et al. (2005).

Acknowledgments: The Short-term Visits Program of the Royal Society (London) and the National Museum of Natural History (Sofi a) have supported the study. The author is very grateful to Mr Dimitar Kovachev for handing the material for examination and Dr. Robert Prys-Jones and Dr. Joanne Cooper (NHM) and Dr. Tereza Tomek and Dr. Zbigniew Bochenski (ISEAK) for providing excellent working conditions, which facilitated much of this study. Special thanks to Dr. Nikolay Spassov (NMNHS) for the helpful review of an earlier version of the manuscript.

References

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BOCHEŃSKI Z., 1997. List of European fossil bird species. – Acta zoologica cracoviensia 40 (2): 293-33.

BOEV, Z. 1999. Falco bakalovi sp. n. – a Late Pliocene falcon (Falconidae, Aves) from Varshets (W Bulgaria). – Geo-logica Balcanica, 29 (1-2): 131-135.

BOEV Z., D. KOVACHEV 1998. Buteo spassovi sp. n. – a Late Mio-cene Buzzard (Accipitridae, Aves) from SW Bulgaria. – Geologica Balcanica, 29 (1-2): 125-129.

BOEV Z., D. KOVACHEV 2007. Euroceros bulgaricus gen. nov., sp. nov. from Hadzhidimovo (SW Bulgaria) (Late Mio-cene) – the fi rst European record of Hornbills (Aves: Coraciiformes). – Geobios, 40: 39-49.

BOEV Z., N. SPASSOV 2009. First record of ostriches (Aves, Stru-thioniformes, Struthionidae) from the late Miocene of Bulgaria with taxonomic and zoogeographic discussion. – Geodiversitas, 31 (3): 493-507.

BOEV Z. 2011. New fossil record of the Late Pliocene falcon (Falco bakalovi Boev, 1999) from type locality in Bulgaria. – Geologica Balcanica. (In press).

LIVEZEY B. C., R. L. ZUSI. 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., Fahl-busch V., Mein P. (Eds.): – In: European Neogene mammal chronology. New York. Plenum Press, 73-90.

MLIKOVSKY J. 1996 (Ed.): Tertiary avian localities of Europe. Acta universitatis Carolinae Geologica. Univerzita Karlova. Praha, 39 (1995): 519-852.

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

NITTINGER F., HARING, E., PINSKER, W., WINK, MICHAEL & GAMAUF, A. 2005. Out of Africa? Phylogenetic relationships between Falco biarmicus and other hierofalcons (Aves Falconi-dae). – Journal of Zoological Systematics and Evolutionary Research 43 (4): 321-331.

OLSON S. L. 1985. The fossil record of birds. – In: King, J. R., D. C. Parker, Eds. Avian Biology, Vol. VIII, Academic Press, New York, 79-252.

SOBOLEV D. V. 2003. New species of Pliocene falcon (Falconi-formes, Falconidae). – Vestnik zoologii, 37 (6): 85-87.

SPASSOV N. 2002. The Turolian Megafauna of West Bulgaria and the character of the Late Miocene ‘Pikermian biome’. – Bollettino della Società Paleontologica Italiana, 41 (1): 69-81.

SUAREZ W., S. OLSON. 2001. A remarkable new species of small fal-con from the Quaternary of Cuba (Aves: Falconidae: Falco). – Proceed. Biological Soc. Washington, 114 (2): 34-41.

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

Received: 29.10.2010 Accepted: 13.01.2011


35

APPENDIX 1Examined specimens belonging to recent species in the FalconidaeLanner Falcon Falco biarmicus NHM 1976.60.5; Nankeen Kestrel Falco cenchroides ISEAK A 4916/91; Red-nacked Falcon Falco chicquera DAUDIN, 1800: NHM 1993.2.5; Merlin Falco co-lumbarius LINNAEUS, 1758: NHM 1862.1.18.3, NHM 1930.3.24.264, NHM 1988.61.1, ISEAK A 2532/72; Eleonora’s Falcon Falco eleonorae ISEAK A 5093/92; Australian Hobby Falco longipennis ISEAK A 5351/94; Lesser Kestrel Falco naumanni FLEISCHER, 1818: NHM 1955.15.2, NHM 1961.13.4, ISEAK 4958/91, ISEAK A 4783/90, ISEAK A 4958/91; Greater Kestrel Falco rupicoloides ISEAK A 4782/90; American Kestrel Falco sparverius LINNAEUS, 1758: ISEAK A 4106/84; Eurasian Hobby

Falco subboteo LINNAEUS, 1758: ISEAK 230/41/14, ISEAK 244/38/13, ISEAK 416/62, ISEAK A 1483/64, ISEAK A 2071/69, ISEAK A 3196/76, ISEAK A 3620/79; Common Kestrel Falco tin-nunculus LINNAEUS, 1758: ISEAK 4509/88, ISEAK 4594/89, ISEAK A 4449/87, ISEAK A 4509/88, ISEAK A 4594/89, ISEAK A 4734/90; Red-footed Falcon Falco vespertinus LINNAEUS, 1766: NHM 1855.4.4.9, NHM 1869.19.12, ISEAK A 2371/71, ISEAK A 4737/90, ISEAK A 5056/92Collared Falconet Microhierax caerulescens (LINNAEUS, 1758): NHM 2002.41.2; African Pygmy-falcon Polihierax semitorqutus (SMITH, 1846): NHM 2002.41.1.

falco bulgaricus sp. n. (aves: falconiformes) from...

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