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ACTA ZOOLOGICA BULGARICA Acta zool. bulg., 63 (1), 2011: 17-35 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 Sofia, Bulgaria; E-mail: boev@nmnhs.com; zlatozarboev@gmail.com Abstract: A new species of small falcon of the group ‘tinnunculus’ is described on the base of 16 bone finds 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 interarticularis; (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 diversified 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 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 recently only seven of them are considered as valid species (OLSON 1985, BOCHENSKI 1997, MLIKOVSKY 1996, 2002) – (1) F. antiquus MOURER-CHAUVIRÉ (1975) (Middle Pleistocene of Noailles, France); (2) F. medius 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 falcons have been described from the New World localities – (6) F. oregonus BRODKORB, 1946 (Early/Middle Pliocene of Oregon), and (7) F. kurochkini SUFIREZ & OLSON, 2001 (Late Pleistocene/Holocene of Cuba). 17

ACTA ZOOLOGICA BULGARICA

Acta zool. bulg., 63 (1), 2011: 17-35

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 Sofia, Bulgaria;

E-mail: boev@nmnhs.com; zlatozarboev@gmail.com

Abstract:

A new species of small falcon of the group ‘tinnunculus’ is described on the base of 16 bone finds 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 interarticularis;

(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 diversified 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

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 recently

only seven of them are considered as valid species

(OLSON 1985, BOCHENSKI 1997, MLIKOVSKY 1996,

2002) – (1) F. antiquus MOURER-CHAUVIRÉ (1975)

(Middle Pleistocene of Noailles, France); (2) F. medius

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 falcons

have been described from the New World localities

– (6) F. oregonus BRODKORB, 1946 (Early/Middle

Pliocene of Oregon), and (7) F. kurochkini SUFIREZ &

OLSON, 2001 (Late Pleistocene/Holocene of Cuba).

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), Sofia.

Material and Methods

The finds 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 grayyellowish-coloured

sands and clay sands of oblique

and complex inner stratification at a depth of 1.00-

1.50 m (DIMITAR KOVACHEV, NMNHS – pers. comm.).

All the 16 bone finds (NoNo NMNHS 12539-12545;

12559-12567; Fig. 1, 2) represent parts of an unarticulated

skeleton of one adult individual. It was collected

by D. KOVACHEV in 1980s. Material is kept

in the NMNHS. The findings have been identified

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 processus

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 scapulae

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 pectoralis

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 medial

side; b – maximum width of proximal epiphysis;

c – length of depressio m. brachialis; d – thickness

of proximal epiphysis; e – length (transversal diameter)

of cotyla dorsalis; f – longitudinal diameter of

cotyla ventralis (Table 4); radius: a – width of proximal

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 fibularis; b – thickness of the diaphysis

in the distal end of crista fibularis; c – width

of diaphysis in the distal end of crista fibularis; d –

maximum diagonal width of proximal epiphysis; e –

length from the proximal epiphysis to the distal end

of the tibio-fibular symphysis (at the caudal side);

f – length of foramen interosseum distale (Table 7);

tarsometatarsus: a – width of diaphysis in the proximal

end of the fossa metatarsi I; b – thickness of diaphysis

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 trochlea

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 calipers

to 0.05 mm accuracy, but read to the 1st digit

after decimal point. All generic names of the binominals

are given abbreviated in the text and are in full

in the Appendix I. ‘Smaller’, ‘much smaller’, ‘bigger’

or ‘much bigger’ in ‘Comparison and discussion’

section mean that the fossil specimen differs

considerably in size from the specimens of compared

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 tibiofibular

symphysis (at the lateral side) of tibiotarsus

(Table 7 – e) (Fig. 4).

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

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 – dorsal

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).

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).

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

Species a b c d

Fossil – Hadzhidimovo

Falco bulgaricus sp. n. NMNHS 12563 3.4 5.7 3.8 ca. 5.1

Recent

Falco naumanni ISEAK A 4783/90 2.5 5.0 2.6 6.6

Falco naumanni ISEAK A 4958/91 2.8 5.5 2.4 6.6

Falco subbuteo ISEAK A 3196/76 2.8 6.0 2.9 7.2

Falco subbuteo ISEAK A 2071/69 2.9 6.8 3.1 6.6

Falco subbuteo ISEAK A 3620/79 2.8 6.4 3.0 6.8

Falco subbuteo ISEAK 230/41/14 3.1 6.2 3.0 6.8

Falco subbuteo ISEAK 244/38/13 3.0 5.8 2.8 7.3

Falco subbuteo ISEAK 416/62 3.3 6.7 3.0 7.2

Falco subbuteo ISEAK A 1483/64 2.8 5.8 2.8 6.5

Falco tinnunculus ISEAK A 4449/87 2.7 6.0 2.6 5.8

Falco tinnunculus ISEAK A 4734/90 2.7 6.2 2.9 6.5

Falco tinnunculus ISEAK A 4564/88 3.0 6.3 3.9 6.8

Falco tinnunculus ISEAK A 4594/89 2.9 6.7 2.9 6.1

Falco tinnunculus ISEAK A 4509/88 2.8 6.5 3.1 6.1

Falco vespertinus ISEAK A 5056/92 2.7 4.6 2.6 6.1

Falco columbarius NHM 1988.61.1 2.8 6.9 4.0 6.3

Falco columbarius ISEAK A 2532/72 2.6 5.9 2.7 6.9

Falco eleonorae ISEAK A 5093/92 3.8 9.1 3.8 8.8

Falco biarmicus NHM 1976.60.5 ca. 3.8 7.7 5.2 9.4

Falco sparverius ISEAK A 4106/84 2.4 4.0 2.1 5.2

Falco longipennis ISEAK A 5351/94 3.9 6.9 3.1 7.3

Falco cenchroides ISEAK A 4916/91 2.7 4.7 2.7 5.6

Falco rupicoloides ISEAK A 4782/90 2.6 5.8 2.7 6.4

Falco chicquera NHM 1993.2.5 3.1 5.0 3.1 ca.5.3

Falco columbarius NHM 1862.1.18.3 ca. 3.3 - 3.7 5.7

Falco vespertinus NHM 1855.4.4.9 3.1 5.9 3.1 5.6

Falco columbarius NHM 1930.3.24.264 2.7 5.7 3.1 5.9

Falco naumanni NHM 1961.13.4 2.2 5.7 4.0 5.1

Falco naumanni NHM 1955.15.2 2.6 6.3 3.9 6.0

Falco vespertinus NHM 1869.19.12 2.7 4.6 2.5 4.9

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 site

According to the rich associated terrestrial megafauna

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) includes

30 taxa: Mustelidae indet., Hyaenotheriun

gr. wongii (ZDANSKY, 1924), Hyaenictitheriini indet.,

cf. Miohyaenotherium bessarabicum SEMENOV,

1989, Adcrocuta eximia (ROTH AND WAGNER, 1854),

Table 2. Measurements of scapula prox. in fossil and recent

Falco.

Species a b

Fossil – Hadzhidimovo

Falco bulgaricus sp. n. NMNHS 12565 4.0 2.5

Recent

Falco columbarius NMNHS 2/2002 3.8 2.0

Falco columbarius NMNHS 1/1997 3.6 2.2

Falco subbuteo NMNHS 1/1989 5.5 2.8

Falco subbuteo NMNHS 3/1993 3.9 1.9

Falco tinnunculus NMNHS 1/2002 2.8 1.8

Falco tinnunculus NMNHS 7/1989 3.0 1.8

Falco tinnunculus NMNHS 10/1992 4.0 2.4

Falco vespertinus NMNHS 1/1991 2.8 2.0

Boev Z.

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

Species a b c d e f g

Fossil – Hadzhidimovo

Falco 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 4.6

Recent

Falco biarmicus NHM 1976.60.5 75.4 23.9 10.9 5.6 5.6 5.5 6.0

Falco cenchroides ISEAK A 4916/91 50.1 15.3 5.5 3.1 2.9 4.0 4.3

Falco chicquera NHM 1993.2.5 51.2 16.0 5.1 3.4 3.7 3.3 4.2

Falco columbarius NHM 1862.1.18.3 51.8 18.5 ca. 6.4 4.0 3.8 4.0 4.8

Falco columbarius NHM 1930.3.24.264 46.1 15.0 6.6 3.5 3.4 3.9 4.4

Falco columbarius NHM 1988.61.1 51.8 17.1 4.7 3.8 4.2 4.1 4.9

Falco columbarius ISEAK A 2532/72 46.6 14.3 6.7 3.5 3.0 3.7 4.1

Falco eleonorae ISEAK A 5093/92 70.1 19.9 7.9 5.3 5.0 5.4 6.2

Falco longipennis ISEAK A 5351/94 53.8 17.7 6.6 4.1 3.6 4.4 5.0

Falco naumanni NHM 1955.15.2 51.9 16.2 7.8 3.4 3.5 3.7 4.1

Falco naumanni NHM 1961.13.4 48.0 8.9 7.0 2.9 3.3 3.3 3.5

Falco naumanni ISEAK 4958/91 52.0 16.6 7.5 3.1 2.8 3.6 4.1

Falco naumanni ISEAK A 4783/90 51.6 16.8 7.0 3.1 2.7 3.3 4.0

Falco rupicoloides ISEAK A 4782/90 51.3 17.3 6.5 3.3 3.0 3.7 4.1

Falco sparverius ISEAK A 4106/84 42.3 13.2 5.5 2.6 2.5 2.8 3.3

Falco subbuteo ISEAK 230/41/14 53.4 16.5 6.4 3.9 3.7 4.4 4.8

Falco subbuteo ISEAK 244/38/13 54.3 18.1 6.1 3.9 3.7 4.0 4.3

Falco subbuteo ISEAK 416/62 62.3 20.0 6.8 4.3 3.8 4.7 5.2

Falco subbuteo ISEAK A 1483/64 53.7 17.1 7.4 3.6 3.2 4.1 4.6

Falco subbuteo ISEAK A 2071/69 53.2 16.9 6.6 4.0 3.5 4.2 4.8

Falco subbuteo ISEAK A 3196/76 54.7 17.4 6.6 3.9 3.6 4.3 4.8

Falco subbuteo ISEAK A 3620/79 52.9 17.6 7.9 4.3 3.5 4.0 4.6

Falco tinnunculus ISEAK 4509/88 56.0 19.3 6.8 3.4 3.3 4.2 4.7

Falco tinnunculus ISEAK 4594/89 53.6 17.2 7.3 3.3 3.1 4.0 4.1

Falco tinnunculus ISEAK A 4449/87 55.0 19.4 7.6 3.3 2.9 3.8 4.2

Falco tinnunculus ISEAK A 4564/88 53.7 18.4 7.7 3.1 3.0 4.5 4.1

Falco tinnunculus ISEAK A 4734/90 54.9 18.9 7.0 3.4 3.2 4.0 4.5

Falco vespertinus NHM 1855.4.4.9 50.5 18.4 7.0 3.3 3.8 3.3 3.8

Falco vespertinus NHM 1869.19.12 50.1 16.5 6.6 2.9 3.1 3.4 3.8

Falco vespertinus ISEAK A 2371/71 49.9 14.8 7.1 3.1 2.9 3.7 4.0

Falco vespertinus ISEAK A 4737/90 49.6 16.7 6.7 3.1 2.8 3.5 3.8

Falco vespertinus ISEAK A 5056/92 - 17.2 - 3.2 3.1 - -

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.

6

5,5

5

4,5

4

3,5

3

2,5

2

40 45 50 55 60 65 70 75 80

a

Falco bulgaricus sp. n. NMNHS 12567

Falco biarmicus BMNH NHM 1976.60.5

Falco cenchroides ISEAK A 4916/91

Falco chicquera BMNH NHM 1993.2.5

Falco columbarius BMNH NHM 1862.1.18.3

Falco columbarius BMNH NHM 1930.3.24.264

Falco columbarius BMNH NHM 1988.61.1

Falco columbarius ISEAK A 2532/72

Falco eleonorae ISEAK A 5093/92

Falco longipennis ISEAK A 5351/94

Falco naumanni BMNH NHM 1955.15.2

Falco naumanni BMNH NHM 1961.13.4

Falco naumanni ISEAK 4958/91

Falco naumanni ISEAK A 4783/90

Falco rupicoloides ISEAK A 4782/90

Falco sparverius ISEAK A 4106/84

Falco subbuteo ISEAK 230/41/14

Falco subbuteo ISEAK 244/38/13

Falco subbuteo ISEAK 416/62

Falco subbuteo ISEAK A 1483/64

Falco subbuteo ISEAK A 2071/69

Falco subbuteo ISEAK A 3196/76

Falco subbuteo ISEAK A 3620/79

Falco tinnunculus ISEAK 4509/88

Falco tinnunculus ISEAK 4594/89

Falco tinnunculus ISEAK A 4449/87

Falco tinnunculus ISEAK A 4564/88

Falco tinnunculus ISEAK A 4734/90

f

Falco vespertinus BMNH NHM 1855.4.4.9

Falco vespertinus BMNH NHM 1869.19.12

Falco vespertinus ISEAK A 2371/71

Falco vespertinus ISEAK A 4737/90

Fig. 3. Correlation between the maximum length of humerus and the thickness of the diaphysis at the foramen nutrinium [mm] (measurements “a” and “f”, Table 3) in the

species of g. Falco.

Boev Z.

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

Species a b c d e f

Fossil – Hadzhidimovo

Falco bulgaricus sp. n. NMNHS 12560 3.6 ca. 7.8 12.3 ca. 5.5 3.8 ca. 4.9

Recent

Falco naumanni NHM 1955.15.2 3.6 7.6 12.0 5.2 3.4 3.2

Falco naumanni NHM 1961.13.4 3.0 5.7 9.8 4.9 3.0 3.9

Falco naumanni ISEAK A 4783/90 3.3 7.5 11.8 5.0 3.2 3.3

Falco naumanni ISEAK 4958/91 3.6 7.5 12.0 5.3 3.2 3.7

Falco subbuteo ISEAK A 3196/76 3.9 8.5 14.2 5.7 3.9 4.5

Falco 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 4.1

Falco subbuteo ISEAK 230/41/14 3.7 8.9 11.2 5.8 3.8 4.1

Falco subbuteo ISEAK 244/38/13 3.6 8.5 12.8 5.6 3.6 4.3

Falco subbuteo ISEAK 416/62 4.2 9.3 13.9 5.9 4.0 4.3

Falco tinnunculus ISEAK A 4564/88 3.6 7.8 11.7 5.3 2.9 3.8

Falco tinnunculus ISEAK 4509/88 3.7 8.1 12.7 6.5 3.7 4.6

Falco tinnunculus ISEAK 4594/89 3.5 7.8 13.0 5.5 3.0 4.2

Falco tinnunculus ISEAK A 4449/87 3.1 7.6 11.5 6.1 3.1 4.3

Falco tinnunculus ISEAK A 4734/90 3.5 7.9 11.4 5.3 3.7 4.3

Falco vespertinus NHM 1869.19.12 3.2 7.2 12.0 4.8 3.1 3.8

Falco vespertinus NHM 1855.4.4.9 3.3 7.8 10.8 5.2 3.3 4.2

Falco vespertinus ISEAK A 2371/71 3.4 7.5 11.7 6.0 3.1 4.0

Falco vespertinus ISEAK A 4737/90 3.2 6.7 12.3 4.9 3.0 4.2

Falco 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 3.9

Falco 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 4.5

Falco eleonorae ISEAK A 5093/92 4.9 9.7 17.9 7.3 7.1 5.2

Falco biarmicus NHM 1976.60.5 5.0 11.4 16.7 7.2 4.3 6.7

Falco sparverius ISEAK A 4106/84 2.6 6.1 7.7 4.1 3.2 2.4

Falco longipennis ISEAK A 5351/94 3.7 8.5 11.9 6.8 4.0 3.6

Falco cenchroides ISEAK A 4916/91 3.4 7.3 10.8 5.8 3.2 3.4

Falco chicquera NHM 1993.2.5 3.3 7.8 ca. 8.9 4.9 3.4 4.0

Falco rupicoloides ISEAK A 4782/90 3.4 7.9 11.9 5.1 2.8 4.3

The avian fauna includes 3 taxa: an accipitrid,

Buteo spassovi BOEV & KOVACHEV 1998, a bucerotid,

Euroceros bulgaricus BOEV & KOVACHEV, 2007

and a struthionid, Struthio karatheodoris (BOEV &

SPASSOV, 2009).

Systematic paleontology

Order: FALCONIFORMES SHARPE, 1874

Family: FALCONIDAE VIGORS, 1824

Subfamily: 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 – Sofia, Bulgarian Academy of Sciences.

Collected by D. KOVACHEV in 1980s.

Paratypes: coracoid dex. NMNHS 12563; humerus

sin. NMNHS 12567; radius sinistra proximalis

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 osteometric

comparison see Tables 1-7.

Etymology: The name ‘bulgaricus’ is given after

the name of the country, Bulgaria, where the finds

were discovered.

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

Species a b c d

Fossil – Hadzhidimovo

Falco bulgaricus sp. n. NMNHS 12562 3.9 2.3 2.0 2.1

Recent

Falco naumanni ISEAK A 4783/90 3.3 2.4 2.0 2.1

Falco naumanni ISEAK 4958/91 3.5 2.6 2.2 2.4

Falco subbuteo ISEAK A 3196/76 3.9 3.0 2.0 2.6

Falco subbuteo ISEAK A 2071/69 3.5 2.8 2.1 2.3

Falco subbuteo ISEAK 230/41/14 3.9 2.8 2.8 2.8

Falco subbuteo ISEAK 244/38/13 3.9 2.8 2.8 2.8

Falco subbuteo ISEAK A 3620/79 3.8 2.9 2.4 2.5

Falco subbuteo ISEAK 416/62 4.3 3.0 3.0 3.2

Falco tinnunculus ISEAK A 4449/87 3.1 2.5 1.8 2.2

Falco tinnunculus ISEAK A 4734/90 3.6 2.7 2.7 2.4

Falco tinnunculus ISEAK A 4564/88 3.5 2.7 2.7 2.4

Falco tinnunculus ISEAK 4509/88 3.6 2.9 3.0 3.0

Falco tinnunculus ISEAK 4594/89 3.7 2.7 2.4 2.4

Falco vespertinus ISEAK A 4737/90 3.3 2.3 2.0 1.9

Falco vespertinus ISEAK A 2371/71 3.0 2.3 1.9 2.1

Falco columbarius ISEAK A 2532/72 3.5 2.4 2.0 2.0

Falco columbarius NHM 1988.61.1 3.8 2.8 2.0 2.5

Falco eleonorae ISEAK A 5093/92 4.8 3.4 2.9 3.2

Falco biarmicus NHM 1976.60.5 5.0 4.0 3.0 3.6

Falco sparverius ISEAK A 4106/84 2.8 2.0 1.8 1.8

Falco longipennis ISEAK A 5351/94 3.5 2.8 2.3 2.5

Falco cenchroides ISEAK A 4916/91 3.4 2.5 2.4 2.4

Falco rupicoloides ISEAK A 4782/90 3.5 2.4 2.3 2.3

Falco chicquera NHM 1993.2.5 3.4 ca. 2.5 ca. 1.9 1.8

Falco vespertinus NHM 1855.4.4.9 3.5 2.5 1.7 2.4

Falco columbarius NHM 1930.3.24.264 3.4 2.5 1.9 2.1

Falco naumanni NHM 1961.13.4 3.0 2.2 1.5 2.2

Falco naumanni NHM 1955.15.2 3.1 2.4 1.7 1.8

Falco vespertinus NHM 1869.19.12 3.1 2.4 2.8 2.4

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 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 interarticularis; (5) tarsometatarsus

– relatively shorter diaphysis smaller.

Locality: Hadzhidimovo-1, near the town of

Gotse Delchev (Blagoevgrad District; SW Bulgaria).

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

Falco.

Species a b

Fossil – Hadzhidimovo

Falco bulgaricus sp. n. NMNHS 12564 3.7 4.0

Recent

Falco columbarius NMNHS 2/2002 4.0 4.5

Falco columbarius NMNHS 1/1997 4.0 4.6

Falco subbuteo NMNHS 1/1989 4.4 5.4

Falco subbuteo NMNHS 3/1993 4.1 5.0

Falco tinnunculus NMNHS 1/2002 3.3 3.7

Falco tinnunculus NMNHS 7/1989 3.5 3.8

Falco tinnunculus NMNHS 10/1992 4.2 4.5

Falco vespertinus NMNHS 1/1991 3.6 2.4

Boev Z.

24

22

20

18

e

16

14

12

10

40 45 50 55 60 65 70 75 80

a

Falco bulgaricus sp. n. NMNHS 12567 NMNHS 12559

Falco biarmicus BMNH NHM 1976.60.5

Falco cenchroides ISEAK A 4916/91

Falco chicquera BMNH NHM 1993.2.5

Falco columbarius BMNH NHM 1862.1.18.3

Falco columbarius BMNH NHM 1930.3.24.264

Falco columbarius NHM BMNH 1988.61.1

Falco columbarius ISEAK A 2532/72

Falco eleonorae ISEAK A 5093/92

Falco longipennis ISEAK A 5351/94

Falco naumanni BMNH NHM 1955.15.2

Falco naumanni BMNH NHM 1961.13.4

Falco naumanni ISEAK 4958/91

Falco naumanni ISEAK A 4783/90

Falco rupicoloides ISEAK A 4782/90

Falco sparverius ISEAK A 4106/84

Falco subbuteo ISEAK 230/41/14

Falco subbuteo ISEAK 244/38/13

Falco subbuteo ISEAK 416/62

Falco subbuteo ISEAK A 2071/69

Falco subbuteo ISEAK A 3196/76

Falco subbuteo ISEAK A 3620/79

Falco tinnunculus ISEAK 4509/88

Falco tinnunculus ISEAK 4594/89

Falco tinnunculus ISEAK A 4449/87

Falco tinnunculus ISEAK A 4564/88

Falco tinnunculus ISEAK A 4734/90

Falco vespertinus BMNH NHM 1855.4.4.9

Falco vespertinus BMNH NHM 1869.19.12

Falco vespertinus ISEAK A 2371/71

Fig. 4. Correlation between the maximum length of humerus and the length from the proximal epiphysis to the distal end of the tibio-fibular symphysis (at the lateral side) of

tibiotarsus [mm] (measurement “a”, Table 3 and measurement “e”, Table 7) in the species of g. Falco.

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

Species a b c d e f

Fossil – Hadzhidimovo

Falco 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 23.8

Recent

Falco biarmicus NHM 1976.60.5 13.2 4.3 6.4 12.0 24.0 22.0

Falco cenchroides ISEAK A 4916/91 8.7 3.1 4.7 7.7 16.9 17.2

Falco chicquera NHM 1993.2.5 9.1 2.8 3.9 8.6 18.2 17.2

Falco columbarius NHM 1862.1.18.3 8.0 3.4 5.0 9.2 19.9 18.8

Falco columbarius NHM 1930.3.24.264 6.8 2.8 4.4 7.9 15.8 17.0

Falco columbarius NHM 1988.61.1 9.6 3.0 4.8 9.1 17.0 17.6

Falco columbarius ISEAK A 2532/72 6.7 2.9 4.4 7.6 14.2 19.3

Falco eleonorae ISEAK A 5093/92 9.0 3.7 4.3 10.4 19.5 16.3

Falco longipennis ISEAK A 5351/94 8.4 3.5 4.6 8.8 18.6 17.4

Falco naumanni NHM 1955.15.2 8.4 2.8 3.0 7.2 16.8 14.9

Falco naumanni NHM 1961.13.4 6.8 2.8 3.1 6.7 14.8 15.0

Falco naumanni ISEAK 4958/91 7.7 2.8 3.6 7.4 15.9 15.9

Falco naumanni ISEAK A 4783/90 8.1 2.8 4.0 7.4 15.0 15.4

Falco rupicoloides ISEAK A 4782/90 8.0 2.9 4.4 7.7 14.1 13.7

Falco sparverius ISEAK A 4106/84 7.1 2.6 3.1 6.7 14.1 13.8

Falco subbuteo ISEAK 230/41/14 7.6 3.2 4.5 8.4 15.8 17.7

Falco subbuteo ISEAK 244/38/13 7.7 2.9 4.2 8.1 15.6 18.4

Falco subbuteo ISEAK 416/62 9.8 3.2 4.2 8.8 17.6 15.3

Falco subbuteo ISEAK A 2071/69 7.0 2.8 4.4 8.3 15.6 17.3

Falco subbuteo ISEAK A 3196/76 7.8 3.0 3.9 8.4 11.8 16.7

Falco subbuteo ISEAK A 3620/79 7.3 3.0 4.3 8.7 16.0 17.6

Falco tinnunculus ISEAK 4509/88 10.7 3.3 4.5 8.4 18.7 18.0

Falco tinnunculus ISEAK 4594/89 12.2 3.4 4.8 9.5 20.3 15.8

Falco tinnunculus ISEAK A 4449/87 10.3 3.0 4.8 8.2 17.6 19.9

Falco tinnunculus ISEAK A 4564/88 10.1 3.2 4.8 8.6 17.4 19.7

Falco tinnunculus ISEAK A 4734/90 10.1 3.4 4.6 8.4 17.6 18.4

Falco vespertinus NHM 1855.4.4.9 7.4 2.7 3.3 7.1 16.1 14.4

Falco vespertinus NHM 1869.19.12 7.1 2.5 2.4 7.1 13.9 14.8

Falco vespertinus ISEAK A 2371/71 7.1 2.7 4.0 ca. 7.0 13.8 19.8

Chronology: Late Miocene (Turolian –

Maeotian, lower part of the zone MN 11-12).

Comparison and Discussion

The general morphology of all skeletal elements unequivocally

indicates that the findings 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 firm affiliation to that genus.

According to general dimensions, the specimen

of Hadzhidimovo is referred to the s. c. ‘tinnunculus’

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 concentrated

on the comparison mainly with the species of

‘tinnunculus’ group. On the other hand, the smallest

Old World falcons (Microhierax SHARPE, 1874

and Polihierax KAUP, 1847) are excluded from our

comparison, because of their evident strong and significant

metrical differences. Their skeletal elements

are more of twice smaller (WHITE et al. 199) than the

compared specimen. The late Pliocene F. bakalovi

Boev Z.

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

Species a b c

Fossil – Hadzhidimovo

Falco bulgaricus sp. n. NMNHS 12540 3.5 1.9 -

Falco bulgaricus sp. n. NMNHS 12566 - - 2.2

Recent

Falco 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 -

differs by the smaller size (comparable to modern F.

columbarius and F. tinnunculus), while F. bulgaricus

sp. n. dimensionally stands between F. tinnunculus

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 carpometacarpus

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 relatively

longer wings and shorter legs indicate a specific

morphologic adaptation. Anyway, thus we exclude

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

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 medius

using the mean proportion (1.271) for ‘tinnunculus’

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. rupicoloides:

larger size; F. chicquera: similar in size

and general morphology; F. columbarius: thicker pr.

acrocoracoideus (measurement ‘c’), thinner diaphysis

(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. acrocoracoideus

(measurement ‘a’) and almost twice

wider impressio ligamenti acrocoracohumeralis; F.

tinnunculus: similar both in general morphology and

size, but sharper proc. acrocoracoideus, better developed

longitudinal groove on the supracoracoidal

surface, and much shorter f. a. clavicularis in cranial

view; F. vespertinus: bigger, and the thicker proc. acrocoracoideus.

Scapula dex. prox. NMNHS 12565 (Fig. 1 –

c; Table 2). F. columbarius: bigger and much thicker;

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 parallel

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. columbarius:

considerably longer humeral bone and more

proximally, instead distally, positioned foramen nutritium

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. eleonorae:

much smaller; F. longipennis: longer and thinner

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 proportions

as its diaphysis is relatively longer. Foramen nutritium

is situated much more distally. Nevertheless

most of the details are similar, confirming its referring

to ‘tinnunculus’ group of genus Falco; F. vespertinus:

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 thinner

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 depressio

m. brachialis; F. cenchroides: larger, wider cotyla

dorsalis; F. chicquera: larger, and deeper depressio

m. brachialis; F. columbarius: more proximally positioned

foramen nutritium is the only difference;

F. eleonorae: much smaller; F. longipennis: smaller;

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 intermuscularis;

Falco naumanni: larger, deeper fossa m.

brachialis (esp. in its proximal end), and better developed

tuberculum lig. collateralis ventralis.

Radius dex. prox. NMNHS 12562 (Fig. 1 –

j; Table 5). F. biarmicus: much smaller, better developed

margo interosseus; F. chicquera: slightly

larger; F. tinnunculus: shallower f. a. ulnaris; F. vespertinus:

larger and bigger tuberculum bicipitale;

Falco columbarius: bigger tuberculum bicipitale; F.

subbuteo: slightly smaller; F. tinnunculus: more protruding

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 intermuscularis

cranialis; F. subbuteo: slightly smaller

and less developed impressiones iliotrochantericae;

Boev Z.

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

Species a b c

Fossil – Hadzhidimovo

Falco bulgaricus sp. n. NMNHS 12543 1.9 3.6 2.4

Recent

Falco naumanni NHM 1955.15.2 1.6 3.0 2.0

Falco naumanni ISEAK 4958/91 1.7 3.0 2.2

Falco naumanni ISEAK A 4783/90 1.3 2.9 2.4

Falco subbuteo ISEAK A 3196/76 1.8 3.3 2.2

Falco subbuteo ISEAK A 3620/79 1.8 3.3 2.4

Falco subbuteo ISEAK 416/62 1.9 3.5 2.5

Falco tinnunculus ISEAK 4509/88 2.2 3.7 2.5

Falco tinnunculus ISEAK 4594/89 2.2 3.7 2.3

Falco tinnunculus ISEAK A 4449/87 1.9 3.2 2.2

Falco tinnunculus ISEAK A 4734/90 2.1 3.3 2.4

Falco tinnunculus ISEAK A 4564/88 2.3 3.6 2.3

Falco vespertinus NHM 1855.4.4.9 1.7 3.2 ca. 2.0

Falco vespertinus NHM 1869.19.12 1.8 2.6 2.0

Falco vespertinus ISEAK A 2371/71 1.7 2.8 1.9

Falco columbarius NHM 1930.3.24.264 1.6 3.0 2.2

Falco columbarius ISEAK A 2532/72 1.8 3.2 2.2

Falco columbarius NHM 1988.61.1 1.7 2.7 2.4

Falco columbarius NHM 1862.1.18.3 1.8 3.4 2.6

Falco eleonorae ISEAK A 5093/92 2.4 4.4 3.0

Falco biarmicus NHM 1976.60.5 3.2 5.6 3.8

Falco sparverius ISEAK A 4106/84 1.7 2.7 1.9

Falco longipennis ISEAK A 5351/94 1.8 3.2 2.3

Falco cenchroides ISEAK A 4916/91 2.0 3.4 2.2

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

Species a b c

Fossil – Hadzhidimovo

Falco bulgaricus sp. n. NMNHS 12542 2.0 2.7 2.5

Recent

Falco columbarius NMNHS 2/2002 1.6 1.9 2.0

Falco subbuteo NMNHS 1/1989 2.0 3.1 2.9

Falco subbuteo NMNHS 3/1993 1.6 2.5 2.3

Falco tinnunculus NMNHS 1/2002 2.2 2.7 2.6

Falco tinnunculus NMNHS 7/1989 2.0 2.8 2.7

Falco tinnunculus NMNHS 10/1992 2.1 2.9 2.7

Falco vespertinus NMNHS 1/1991 1.4 1.9 1.9

F. tinnununculus: more gracile and slender; F. vespertinus:

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 laterally,

positioned base of the crista cnemialis lateralis;

F. chicquera: more medially, but not laterally, positioned

base of crista cnemialis lateralis; F. columbarius:

similar in size, but more distal position of crista

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

Species a b c d

Fossil – Hadzhidimovo

Falco bulgaricus sp. n. NMNHS 12541 1.7 2.5 2.3 1.9

Recent

Falco columbarius NMNHS 2/2002 1.4 2.3 2.1 1.6

Falco subbuteo NMNHS 1/1989 1.7 3.0 2.5 2.0

Falco subbuteo NMNHS 3/1993 1.3 2.2 1.9 1.5

Falco tinnunculus NMNHS1/2002 1.7 2.2 2.2 1.9

Falco tinnunculus NMNHS 7/1989 1.6 2.6 2.6 1.9

Falco tinnunculus NMNHS 10/1992 1.7 2.7 2.5 2.0

Falco 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 c

Fossil – Hadzhidimovo

Falco bulgaricus sp. n. NMNHS 12544 ca. 1.6 ca. 2.2 ca. 1.8

Falco bulgaricus sp. n. NMNHS 12545 ca. 1.6 - -

Recent

Falco columbarius NMNHS 2/2002 1.3 2.4 1.6

Falco subbuteo NMNHS 3/1993 1.4 2.0 1.9

Falco tinnunculus NMNHS 1/2002 1.6 2.4 2.2

Falco tinnunculus NMNHS 7/1989 1.7 2.4 2.2

Falco tinnunculus NMNHS 10/1992 1.8 2.4 2.3

Falco 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 distal

epiphysis

Length of spatium

intermeta-carpalis

Fossil – Cherevichniy Hutor - Ukraine 1

Falco medius 45-4033 39.8 10.4 7.4 26.2

Recent - Bulgaria

‘tinnunculus’ group

Falco vespertinus NMNHS 1/1991 34.1 8.4 6.1 22.2

Falco tinnunculus NMNHS 8/1989 44.6 19.4 16.3 31.5

Falco tinnunculus NMNHS 7/1989 44.6 19.3 17.4 32.4

Falco tinnunculus NMNHS 19/2005 45.5 19.4 17.0 33.1

Falco columbarius NMNHS 2/2002 37.0 14.1 10.6 25.1

Falco subbuteo NMNHS 1/1989 53.0 17.5 18.8 40.7

Falco 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 43.5

Falco cherrug NMNHS 1/1990 71.9 26.5 22.4 50.8

After Umanskaya (1981).

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

carpometacarpus

Fossil – Cherevichniy Hutor - Ukraine 2

Falco medius 45-4033 1 39.8

Recent – Bulgaria

‘tinnunculus’ group

Falco columbarius 1 37.0

Falco subbuteo 2 50.5

Falco tinnunculus 3 44.9

Falco vespertinus NMNHS 1/1991 1 34.1

‘cherrug’ group

Falco cherrug NMNHS 1/1990 1 71.9

Falco peregrinus NMNHS 2/1989 1 64.2

humerus

Fossil – Hadzhidimovo

Falco bulgaricus sp. n. NMNHS 12567 1 ca. 58.0

Recent

‘tinnunculus’ group

Falco columbarius 4 49.0

Falco subbuteo 7 53.9

Falco tinnunculus 5 54.6

Falco vespertinus 4 50.5

Falco cherrug NMNHS 1/1990 1 97.3

Falco 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

Fossil

Falco medius 45-4033 1 -

Falco bulgaricus sp. n. 1 -

Recent

‘tinnunculus’ group

Falco columbarius NMNHS 2/2002 1 1.324

Falco subbuteo 2 1.067

Falco tinnunculus 3 1.216

Falco vespertinus NMNHS 1/1991 1 1.480

Falco spp. ex gr. tinnunculus (mean) 7 1.271

‘cherrug’ group

Falco cherrug NMNHS 1/1990 1 1.269

Falco peregrinus NMNHS 2/1989 1 1.309

After Umanskaya (1981).

fibularis, and more medial, than lateral, inception of

distal end of crista cnemialis cranialis; F. longipennis:

dimensionally and morphologically similar, but

sharper bend of crista cnemialis cranialis; F. naumanni:

bigger, relatively longer crista fibularis, uneven

crista fibularis, and smaller general size of bone; F.

sparverius: much larger; F. subbuteo: more medially,

but not laterally, positioned base of the crista cnemialis

lateralis, longer base of crista cnemialis lateralis,

deeper fossa flexoria and longer foramen interosseum

distale, nevertheless general dimensional similarity;

F. tinnuncculus: longer base (more proximally positioned

inception) of crista cnemialis lateralis, and less

protruding area interarticularis; F. vespertinus: considerably

larger, and presence of a shallow concavity

on the cranial surface of crista fibularis. 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 flat

diaphysis; F. chicquera: lower crista plantaris medialis;

F. columbarius: wider distal part of diaphysis

at foramen vasculare distale, considerably less developed

crista plantaris mediana in plantar aspect,

and less developed edge between the f. subcutanea

lateralis and the f. dorsalis in dorsal aspect; F. naumanni:

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 concave

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 proximal

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 profile of trochlea

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. columbarius:

slightly bigger; F. subbuteo: smaller; F.

tinnununculus: smaller; F. vespertinus: bigger and

more robust.

General comparison. The comparison of skeletal

elements of the fore-limbs/pectoral girdle and

the hind-limbs/pelvic girdle clearly demonstrate

more developed flight capability, i. e. F. bulgaricus

sp. n. to a certain extent was more aerial than terrestrial,

in comparison to its modern closer relatives.

Its wings were relatively more developed, while the

legs were less developed in comparison to the closest

dimensionally and in many respects, morphologically,

species. The compared specimen has relatively

longer wings (longer proximal part, i. e. humeri) and

shorter legs (shorter distal part, i.e. thinner, and possibly

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 specific both for the species

of ‘tinnunculus’ group, and for the specimen of

Hadzhidimovo and they are lacking in all the compared

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.

Boev Z.

Probably these major groups deserve to be

ranked as separate super-genera/genera. According

to the fossil record the large falcons (of ‘cherrug’

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 confirmed

by NITTINGER et al. (2005).

Acknowledgments: The Short-term Visits Program of the Royal

Society (London) and the National Museum of Natural History

(Sofia) 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

BAUMEL J. J., L. M. WITMER 1993. 4 Osteologia, in: Baumel, J.,

King, A., Breazile, J., Evans, H., Vanden Berge, J., (Eds.): –

In: Handbook of Avian Anatomy, Nomina Anatomica

Avium. Nutall Ornithological Club, 23:45-132.

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). – Geologica

Balcanica, 29 (1-2): 131-135.

BOEV Z., D. KOVACHEV 1998. Buteo spassovi sp. n. – a Late Miocene

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 Miocene)

– the first European record of Hornbills (Aves:

Coraciiformes). – Geobios, 40: 39-49.

BOEV Z., N. SPASSOV 2009. First record of ostriches (Aves, Struthioniformes,

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., Fahlbusch

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 Falconidae).

– 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 (Falconiformes,

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 falcon

from the Quaternary of Cuba (Aves: Falconidae: Falco).

– Proceed. Biological Soc. Washington, 114 (2): 34-41.

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, English summary).

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

APPENDIX 1

Examined specimens belonging to recent species

in the Falconidae

Lanner 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 columbarius

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 tinnunculus

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 5056/92

Collared Falconet Microhierax caerulescens

(LINNAEUS, 1758): NHM 2002.41.2; African Pygmyfalcon

Polihierax semitorqutus (SMITH, 1846): NHM

2002.41.1.

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