Pleistocene avifaunas of Bulgaria: a brief review

Abstract

BOEV Z. 2006. Pleistocene avifaunas of Bulgaria: a brief review. - Historia naturalis bulgarica, 17: 95-107. Abstract. This paper sums up the faunistic results of the exploration of all Pleistocene bird faunas, uncovered so far in Bulgaria. They cover 160 species, 4 Early Pleistocene, 1 Middle Pleistocene, and 13 Late Pleistocene sites.

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Pleistocene avifaunas of Bulgaria: a brief review 95
Historia naturalis bulgarica,
17: 95-107, 2006
Pleistocene avifaunas of Bulgaria: a brief review
1
Zlatozar BOEV
BOEV Z. 2006. Pleistocene avifaunas of Bulgaria: a brief review. – Historia naturalis bulgarica, 17: 95-107.
Abstract. This paper sums up the faunistic results of the exploration of all Pleistocene bird
faunas, uncovered so far in Bulgaria. They cover 160 species, 4 Early Pleistocene, 1 Middle Pleistocene,
and 13 Late Pleistocene sites.
Key words: Paleozoology, Paleofaunas, Quaternary birds, Aves, Bulgaria
Introduction
As a country of southern geographical situation in Europe, Bulgaria has a great significance for
the elucidation of the building of ranges and the former range fluctuations of the recent bird species in
Europe. Bulgaria spans 111,000 km
2
and its territory could be divided in three main climatic regions:
Temperate-continental (N and part of SW Bulgaria), Transitional (most of S and NE Bulgaria) and
Continental-Mediterranean (the most S and the very E parts of the country) (VELEV, 1997).
The Pleistocene avifaunas of Bulgaria are still poorly known. In contrast to Croatia, partly Serbia
and Greece and most of the countries of Central and Western Europe, the Pleistocene bird faunas of
Bulgaria remained completely unstudied until the early 1980-ies. More considerable archaeological
excavations of the Paleolithic sites have been carried out as early as the first decade of last century, but
besides the regular mentioning of bird remains, they contained no taxonomical identifications.
The paleornithology is the youngest field of the ornithological science in Bulgaria. The first paper on fossil
birds in Bulgaria was published in 1982 by the Polish paleornithologist Zygmunt Bocheński, who reports on
23 avian species (two of them, Lagopus mutus and Pyrrhocorax pyrrhocorax disappeared) from the Late Würmian
(50 000-10 000 B.P.) deposits in the Bacho Kiro Cave (BOCHEŃSKI, 1982). It is the first research on the
Pleistocene avifaunas of the country. The second paper, also by a foreigner, presents the results of the
Georgian paleornithologist Prof. Dr. Nikolay Burchak-Abramovich (BURCHAK-ABRAMOVICH &
NIKOLOV, 1984), describing two avian species from two Miocene localities of the country.
In 1997 a Czech paleornithologist published his notes on the Pleistocene findings in one of the
caves in the Karlukovo Karst Area in NW Bulgaria (MLÍKOVSKÝ, 1997).
During the last two decades the author has published a series of studies. Most of the data came
from cave deposits or rock fissures. According to the Bulgarian Speleological Federation at least 5000
caves have been registered throughout the country by 2001. Our data on Pleistocene avifaunas
1
The paper follows the data of a report on the same topic, which was presented on 26 February, 2002 to the
collegiums of the paleontology and zoology sections in the Paleontology Department, the Natural History Museum,
London.

96 Zlatozar Boev
concerns 18 caves, i.e. 0.4 %. The full taxonomic composition of the Bulgarian Pleistocene site has
been reported by BOEV (1999, 2001).
The four Early Pleistocene localities are as follows: Temnata Doupka cave, Cherdzenitsa Cave,
Kunino quarry and Balsha quarry. The Morovitsa Cave is the only Middle Pleistocene locality. Most
of the localities (13) are of the Late Pleistocene. They are all caves: Kozarnika, Cave N 16, Bacho Kiro,
Temnata Doupka, Karlukovo 4, Devetashkata, Filipovska, Mechata Doupka, Mirizlivka, Tsareva
Tsarkva, Kozarskata, Toplya, and Razhishkata (Fig. 1).
Here we summarize all data on the Pleistocene bird faunas of Bulgaria, based on 5770 avian bone
fossils (BOEV, 1999, 2001а). One site, the Temnata Doupka Cave, contains deposits both of the
Early and the Late Pleistocene. The nesting habitat preferences of the recorded species show a forest-
steppe landscape in the surroundings of the cave. The correlation between the woodland and the
steppe (open land) species is 10:4, indicating the prevailing role of the forest habitats in this part of the
country. The complete taxonomic list includes at least 160 recognized species.
The Balkan Peninsula as a refugium of the arboreal and termophylic avifauna
MOREAU (1954a) and BLONDEL (1984; 1987a,b) in their ornithogeographical analyses state
that during the last glaciation (ca. 70 000 B.P.) the three Mediterranean peninsulas (Pyrenees,
Fig. 1. Pleistocene avian localities in Bulgaria: 1 – Balsha quarry, 2 – Kunino quarry, 3 - Cherdzenitsa
Cave, 4 – Temnata Doupka Cave, 5 – Morovitsa Cave, 6 – Kozarnika Cave, 7 – Cave No 16, 8 –
Bacho Kiro Cave, 9 - Temnata Doupka Cave, 10 - Karlukovo 4 Cave, 11 – Devetashka Cave, 12 –
Filipovska Cave, 13 – Mecha Doupka Cave, 14 – Mirizlivka Cave, 15 - Tsareva Tsarkva Cave, 16 –
Kozarska Cave, 17 – Toplya Cave, 18 – Razhishka Cave.

Pleistocene avifaunas of Bulgaria: a brief review 97
Apennines and Balkans) played a key role in the survival of the wood avifaunistical complex.
BLONDEL (1982) summarizes that the glacial events did not cause extinctions, or speciations in
the Mediterranean region in the Pleistocene. On the contrary, because of the extraordinary
topographic, edaphic and climatic heterogenerosity and the high diversity derived from it, a process
of intensive speciation in the Pyrenees, Apennines and Balkans occurred during the Pleistocene,
but this speciation concerns only a very limited part of the arboreal avifauna of the region
(BLONDEL, 1984; 1985; 1987b).
The southern parts of the Balkan Peninsula, as well as the other two larger peninsulas of Europe
sheltered suitable refugia for the heat-loving fauna in the colder stages during the Middle and the Late
Pleistocene. The occurrence of Testudo sp. in the Early Pleistocene site near Kunino, (possibly) confirms
such a statement. Testudo hermanni is known from Würm-2 in France too (BLONDEL, 1985).
As it is known, the glacial events on the Balkans were considerably poorer, especially in the
southern parts of the peninsula. The main part of the populations of Turdus spp., Sitta spp., Picus spp.,
Picoides spp., Certhia spp., Prunella spp., Alectoris spp., Garrulus glandarius, Hieraeetus fasciatus, Falco
eleonorae, etc. have survived namely in these regions (BLONDEL, 1985). The representatives of most
of these genera have been established in the Pleistocene deposits of the cited localities of Bulgaria also
(BOEV, 2001). They were usually found in several localities. Thus, for the first time the fossil record
of Bulgaria has confirmed the regularities, mentioned above.
According to BLONDEL (1984; 1987a) the evolution of four genera (Alectoris, Sitta, Sylvia and
Monticola) of the Palearctic fauna occurred predominantly in the Mediterranean. The Balkans and
the West Asia Minor form the second (after the Pyrenees-Apennines) center of speciation in warblers
of the genus Sylvia during the Pleistocene. At present this eastern center is inhabited by 11 species.
Blondel calls it “eastern center of endemism and dispersion of the “melanocephala” group” in contrast
to the “western center of “undata”. Sylvia communis has been established in Devetashkata Cave, while
the remains of S. cf. atricapilla occurred in the Cherdzenitsa Cave – 1. These finds mark the genus’s
distribution during the Pleistocene in Bulgaria: the former one from the Middle Pleistocene and the
latter from the Late Pleistocene.
As BLONDEL (1987b) summarizes, the oldest record of Alectoris, Sitta and Sylvia comes from
the Mediterranean and dates back to the Middle to Late Pleistocene. Here the record of the Late
Pliocene Sitta sp. n. from W Bulgaria, the oldest record of this genus in the world, deserves special
attention, but the Late Pliocene and all the Tertiary avian records are beyond the scope of the present
report.
The fossil record of Bulgaria shows that in the Pleistocene at least 50 recent avian genera were
spread: Accipiter, Falco, Buteo, Aquila, Aegypius, Circus, Falco, Alectoris, Perdix, Coturnix, Phasianus,
Lagopus, Tetrao (incl. Lyrurus), Crex, Porzana, Rallus, Gallinula, Otis, Columba, Streptopelia, Otus, Bubo,
Nyctea, Strix, Apus, Dendrocopus, Lullula, Alauda, Riparia, Ptyonoprogne, Anthus, Lanius, Oenanthe,
Turdus, Monticola, Erithacus, Cettia, Sitta, Bombycilla, Fringilla, Loxia, Carduelis, Pyrrhula, Passer,
Sturnus, Garrulus, Pica, Nucifraga, Pyrrhocorax, Corvus, etc. Most of them could be unquestionably
referred to the group of the dendrophylous, or arboreal birds. A smaller part (Alectoris, Apus, Riparia,
etc.) are obligatory termophylous, chiefly petrophylous elements.
The representatives of the arboreal avifauna have been localized in the glacials south of the
Balkan Range, where besides coniferous, there were vast deciduous forests in the southern parts. It is
believed that in Bulgaria the most important were the broad-leaved forests in the SE parts of the
country (Strandza and Sakar mountains). These regions as a whole, are determined by MOREAU
(1954a,b) as refugia for the survival of Honey Buzzard, Tawny Owl, Middle Spotted Woodpecker,

98 Zlatozar Boev
Green Woodpecker, Golden Oriole, Nuthatch, Icterine Warbler, Blackcap, Whitethroat, Wood Warbler,
Red-breasted Flycatcher, Hawfinch, etc.
Tetraonids are determined as a specific element of the Pleistocene avifaunas of Europe by
MOURER-CHAUVIRÉ (1993). This regularity is present in the Pleistocene localities in the Balkans
too. All the five tetraonids spread in Europe were widely spread during the Pleistocene in Bulgaria too.
The steppe landscapes prevailed in N Bulgaria in the Pleistocene. In the steppes of W Europe
(France, Spain) by the Middle and the early Late Pleistocene Perdix palaeoperdix, the ancestor of the
modern gray partridge was widely spread (MOURER-CHAUVIRÉ, 1993; SANCHÉZ, 1996b). This
species was established for the first time on the Balkan Peninsula in four Bulgarian caves – Kozarnika,
Cave No 16, Devetashka and Razhishka.
The following bird species of the termophylic fauna have been established in Bulgaria: Honey
Buzzard, Swift, Alpine Swift, Nightjar, Bee-eater, Sand Martin, Northern Wheatear (entomophagous),
Quail, Corncrake, Baillon’s Crake, Turtle Dove (phytophagous), Short-toed Eagle (herpetophagous),
etc. All of them are now represented in Bulgaria and the Balkans by their breeding populations. Some
heat-loving species of Micromammalia (Apodemus mystacinus, Jordanomys major) have been recorded
even in the Pleistocene localities of N Bulgaria (POPOV & DELCHEV, 1997). The range of A.
mystacinus at present is confined to the E Mediterranean.
It is usually believed that the recent avifauna of the Temperate zone of the Palearcic is a result of
the Holocene migration of the faunistic complexes. For the bird fauna of the neighboring Romania
for example, DRUGESCU (1987) states that its genesis is due to the migration of the elements of the
Mediterranean, Caucasian, Syrian, Indian and Mongolian refugia after the end of Pleistocene.
Our data for Bulgaria, at least for S Bulgaria, do not support this hypothesis. The Early Holocene
faunal changes were actually the last considerable events, which led to the modern state of the recent
Balkan avifauna. But it should be evaluated in reference to the faunistic complexes of the terrestrial
fauna of the vertebrates that had been formed earlier, i. e. in the Pleistocene. As a whole, the avifauna
of Bulgaria had almost a modern definite character.
Seasonal migrants in the Pleistocene localities
Now it is well known that most of the modern species of birds appeared in the Late Pliocene and
the Pleistocene. Many relict routes of migrations have survived until the present, as stated by KUMARI
(1959). Hence, the seasonal migrations of birds are a paleogeographic heritage.
On the other hand, migrations as an adaptation of birds to evade the unfavorable environmental
conditions are very dynamic and “plastic”. Many examples refute the hypothesis of their
conservativeness. Data of modern ornithology on the neo-migrations show that the routs and the
winter areas (winter ranges) could be considerably changed in very short terms.
One of the most evident proofs of the completely changed migration is the Red-breasted Goose.
In two decades only (1960-1980) this species has drastically changed its winter range from the Kazil-
Agach steppe along the Caspian Sea coast in Azerbaijan with arable lands in South Dobrudzha (the
Shabla-Ezerets Lake complex) along the Black Sea coast in Bulgaria. Almost the whole winter range
has moved here and during the last several years almost the whole species’ population of this Siberian
species (56 000 birds) spent the winter in the NE of Bulgaria (GEORGIEV, 1995).
VILETTE (1983) divides the European species according to their breeding migratory populations
into two main types: migrants breeding in the northern latitudes, and migrants breeding in the

Pleistocene avifaunas of Bulgaria: a brief review 99
temperate latitudes. As we know, during the Pleistocene, only a small limited part of the S European
territory preserved suitable habitats for wintering of the European migratory populations. As MOREAU
(1954a,b) states, relatively warmer open grounds that are now common in many regions of the
Mediterranean peninsulas, were absent in the cold phases of the Pleistocene. This reflected the
duration, distance and direction of the migrations. Most of the populations made trans-Mediterranean
migrations to N Africa or SW Asia. The water birds, however, were not forced to leave their shelters
in the S-European limits along the coasts of the Tyrrenian, Adriatic, Aegean, Black, Asov and Caspian
seas, where they had suitable wintering grounds.
We have established the occurrence of some aquatic species of a more “northern” for the region
distribution in the Early to Late Holocene too. Some of the localities of the S Black Sea Coast and of
N Bulgaria: Oxyura leucocephala, Aythya nyroca, A. ferina, A. fuligula, and Phalacrocorax aristotelis (BOEV,
1995; unpubl. data). Most of them are now only winter visitors of the country.
Some migrants that now visit the country only in the winter season have been established by their
numerous fossil (and subfossil) remains. These species are: Gavia arctica, G. immer, G. stellata, Anser
fabalis, A. albifrons, A. erythropus, Anas penelope, A. crecca, Bucephala clangula, Melanita nigra, Buteo
lagopus, Cyrcus cyaneus, Cygnus olor, Tringa nebularia, Tr. glareola, etc. During the Middle and Late
Pleistocene they were visiting the country, but we do not know their status of sojourn. Their breeding
range now lie further N of the territory of Bulgaria.
Fig. 2. Present day range of Lagopus mutus (dotted) in Europe (after JONSSON, 1994) and the location
of the Pleistocene sites of the species in Bulgaria: 1 – Bacho Kiro Cave (after BOCHEŃSKI, 1982), 2
– Devetashka Cave (incl. finds of L. mutus/lagopus).

100 Zlatozar Boev
Boreal elements in the Pleistocene avifaunas
Now for the first time the whole “boreo-mountain”, “glacial”, “alpine”, “Pleistocene” avifaunistical
complex, characteristic for the glacial stages of the Pleistocene in the Temperate Zone of Europe, has been
established in Bulgaria too. At present the arcto-alpine fauna has a very limited distribution in Bulgaria.
Our data show that the avian boreal complex numbers at least 15 woodland, steppe and
hydrophilous species. These species are: Willow Grouse (Fig. 2), Ptarmigan (Fig. 3), Snowy Owl,
Short-eared Owl, Pygmy Owl, Tengmalm’s Owl, Greate Grey Owl (identified as Strix aff. nebulosa),
Rough-legged Buzzard, Whimbrel or Slender-billed Curlew (Numenius phaeopus/tenuirostris), Red-
breasted Goose, Goldeneye, Common Scotter, Shore Lark, Chough and Alpine Chough. This list
completes the information available so far from some other Balkan countries (mainly Croatia,
Slovenia, Serbia and Romania).
The boreal avifauna is best established in the Late Pleistocene deposits of the Devetashkata Cave
in the Central Stara Planina Mountain (Central part of the Balkan Range). Some specific faunistic
elements have been established also in other caves. The presence of the Snowy Owl for example has
been proved in the Kozarnika Cave in the Western Stara Planina Mountain (BOEV, 1998b). All these
records also confirm for the first time in Bulgaria the southward extension of the ranges of boreal
elements during the cold phases of the Pleistocene.
Fig. 3. Present day range of Lagopus lagopus (dotted) in Europe (after JONSSON, 1994) and the
location of the Pleistocene sites of the species in Bulgaria: 1 – Devetashka Cave, 2 – Kozarnika cave,
3 – Filipovska cave.

Pleistocene avifaunas of Bulgaria: a brief review 101
“Mixed” avifaunas
According to NAZARENKO (1990) a large meridional “forest” bridge between the southern,
temperate and northern regions had a key role in the interchange of the terrestrial faunas in the
Palearctic during the last 35 000 years. It was used by the woodland birds for the extension of their
ranges from South to North in the Eastern Palearctic. It is very likely that the mountain chains of the
Apennines and the Balkans in the southern parts of the Western Palearctic had a similar role. That is
why, because of the high diversity of the relief and the mosaic character of the landscape, in the
neighboring parts of a territory birds so different in type such as the termophilic “Mediterranean”
Sylvia atricapilla, and the cold-loving “boreal” Aegolius funereus occurred together. The occurrence of
S. atricapilla during the Early Pleistocene in the Cherdzenitsa Cave was proven, while S. communis
was spread in the vicinity of the Devetashkata Cave in the Late Pleistocene. In addition, a find of
Sylvia sp. has been established in the Razhishkata Cave dating back to the Late Pleistocene too
(BOEV, 2000).
S. atricapilla coexisted in the Early Pleistocene with Dendrocopos minor and Pyrrhula pyrrhula, while
in the Late Pleistocene S. communis coexisted with Tetrao tetrix, Lagopus lagopus, Tetrastes bonasia,
Glaucidium passerinum and Aegolius funereus. Such “mixed” avifaunas were discovered in the Pleistocene
Fig. 4. Present day range of Lagopus lagopus (dotted) and Pyrrocorax graculus (black) in Europe (after
JONSSON, 1994) and the location of the Kozarnika cave (triangle), where both species coexisted in
the Late Pleistocene.

102 Zlatozar Boev
for different parts of the Palearctic: Transcaucasia (Gallus gallus and Lagopus lagopus) of the Paleolithic
in Georgia (BURCHAK-ABRAMOVICH, 1986), western Mediterranean (Lagopus sp. and Gyps fulvus,
Tetrao tetrix and Alectoris sp.) of Würm 2 in Monaco (MOURER-CHAUVIRÉ, 1975a), Falco naumanni
and Lagopus mutus and Monticola saxatilis and Nyctea scandiaca) of Riss in S France, etc. Similar are the
associations of “Nyctea scandiaca - Pyrrhocorax graculus” from the Kozarnika Cave in W Bulgaria (Late
Pleistocene; BOEV, 2001b) and “Nyctea scandiaca - Cervus dama” in the Middle Pleistocene in Italy
(BARBATO et al., 1992).
“Mixed” Pleistocene faunas are known also from Middle Europe. Some boreal forms as Tetrao
conjugens, T. macropus and Lagopus atavus coexisted with the Mediterranean Francolinus capeki and Fr.
minor (JANOSSY, unpubl. data; BOCHEŃSKI, 1985) in the Plio-Pleistocene of Hungary. Similar
coexistence is known from the Late Pleistocene in Poland, where BOCHEŃSKI (1974) established
the boreal Falco columbarius, Eudromias morinellus and Numenius phaeopus and the Mediterranean
Aegypius monachus and Apus melba. The summarized data of TYRBERG (1991a) show that in the 850
Würmian localities of W and C Europe L. lagopus and L. mutus coexisted with C. coturnix and T.
tetrax. He proved that very often some arcto-alpine forms such as Aquila chrysaetos, Falco tinnunculus,
Nyctea scandiaca, Alauda arvensis, Corvus corax, P. pyrrhocorax and P. graculus coexisted with steppe and
Mediterranean ones such as Buteo rufinus, Circus macrourus, Falco vespertinus, Alectoris spp., Melanocorypha
spp. Calandrella spp. and Sturnus roseus). BARBATO et al. (1992) reported on common occurrence of
Fig. 5. Present day range of the genus Nyctea (black) and the genus Alectoris (dotted) in Europe (after
JONSSON, 1994) and the location of the Kozarnika Cave (triangle), where species of both genera
coexisted in the Late Pleistocene.

Pleistocene avifaunas of Bulgaria: a brief review 103
Alectoris graeca and Nyctea scandiaca and of Pyrrhocorax graculus and Perdix perdix. This overlapping of
the ranges of the “northern” and “southern” forms is one of the most specific features of the Pleistocene
avifaunas of Europe. P. graculus and L. lagopus also coexisted during the Late Pleistocene in Bulgaria
(Fig. 4).
Almost all the rest of the records of the Snowy Owl and Rock Partridges are among the best
examples of the “mixed” avifaunas. This association is also established in Bulgaria (the Kozarnika
Cave and the Devetashkata Cave; BOEV, 1998a,b; 2001b; Fig. 5). A similar example from the Kozarnika
Cave is the coexistence of the Willow Grouse and the Red-rumped Martin.
The summarizing data for Europe of BOCHEŃSKI (1985) outline the Würmian coexistence of
L. lagopus and Surnia ulula and Melanocorypha calandra and Pyrrhocorax graculus on the other hand.
Such an association (Melanocorypha sp. - Pyrrhocorax pyrrhocorax/ P. graculus) we have established for
the first time in Bulgaria in the Razhishkata Cave (BOEV, 1998a,b; 2000). Other examples of the
“mixed” avifaunas are also known from Bulgaria. BOCHEŃSKI (1982) reports on remains of
Lagopus mutus and Alectoris graeca in the Bacho Kiro Cave that have coexisted. Our data (BOEV,
1994) proved that in the surroundings of the Temnata Doupka Cave ca. 28 900 B. C. some
Mediterranean (A. monachus and Al. graeca) and some boreo-mountain (N. caryocatactes) species also
coexisted. In the Kozarnika Cave the Chough coexisted with the Willow Grouse (BOEV, 2001b).
There are also a number of other examples from Bulgaria. According to BLONDEL (1985) this
combination of birds of different biotas could be explained by the uniqueness of the physico-
geographical characteristics of the N Mediterranean being a kaleidoscope of landscapes.
Thus, during all phases of the Pleistocene, due to the topographic diversification of the region,
various species of the humid (i. e. mesophilous) forests, as well as the inhabitants of the drier
xerophitic cold steppes coexisted in the mosaic landscapes of the Mediterranean. As a whole, the
“mixed” avifaunas in Europe had survived until ca. 50 000 B.P., when the distribution of birds
acquired its modern aspect. “Mixed” faunas in mammals are much better known from most of the
Pleistocene localities. According to POPOV (1990a,b) the Late Pleistocene fossil assemblages from
the Temnata Doupka Cave contained European-West-Siberian (wood and forest-steppe) and East-
European-Kazakhstan (arid) elements. These “mixed” assemblages consisted of species that at present
never occur together and POPOV (1990a,b) concludes that they are younger and more dynamic
formations - a result of the local peculiarities of the fauna development.
All these data on the Pleistocene record of birds from Bulgaria allow to draw the following:
Conclusions
The avian Pleistocene record of Bulgaria is still poorly known although it includes 160 species of
13 orders – 40.0 % of the recent country’s bird fauna. Most of the material originates from Würmian
(Late Würmian) deposits, making two thirds of the Pleistocene sites.
The Pleistocene avifauna of Bulgaria contained 8 species that disappeared from the recent
country’s fauna: Lagopus mutus, Lagopus lagopus, Tetrao tetrix, Phasianus colchicus (native subspecies
colchicus form), Glaucidium passerinum, Nyctea scandiaca, Strix aff. nebulosa and Pyrrhocorax pyrrhocorax.
In addition another 4 species established in the Pleistocene deposits (Black Vulture, Snipe, Great
Bustard and Little Bustards) disappeared from the Late Holocene breeding bird fauna of the country.
The Late Pleistocene avifaunas included fossil taxa – one fossil species (Perdix palaeoperdix) and
one fossil subspecies (Pyrrhocorax graculus vetus). The Late Pleistocene localities of Perdix palaeoperdix

104 Zlatozar Boev
probably prove its (?) coexistence with the modern Grey Partridge (Perdix perdix). They are the youngest
records of that extinct species so far.
The Early Pleistocene avifaunas of Bulgaria are known from 4 localities with 26 taxa of 5 orders.
The most numerous are the Passeriformes. One species (the Chough) has disappeared. The
Cherdzenitsa Cave is the fourth Early Pleistocene locality of that species in the Palearctic (TYRBERG,
1998). The records of the Lesser Spotted Woodpecker, Red-rumped Swallow, Sand Martin, Rock
Thrush and the House Sparrow are the oldest ones, while the finds of the Woodlark are the second
known from the Early Pleistocene in the Palearctic.
The Middle Pleistocene avifaunas are least known among the Pleistocene faunas of Bulgaria.
Only 1 site with 7 taxa of 2 orders has been established so far.
The Late Pleistocene avifaunas are studied best. They are known from 13 sites with 134 taxa of 13
orders.
In the interglacials some Mediterranean species (Alpine Swift, Red-rumped Swallow) extended
their ranges northwards in the Late Pleistocene, reaching the northern parts of the country.
The Pleistocene avifaunas of Bulgaria developed in close connection with the remaining bird
faunas of the neighbour regions of SE Europe. For some groups they show a wider distribution of
some recent species. The country’s Pleistocene (at least Würmian) avifauna showed the whole
boreo-mountain complex, consisting of 15 species at least, specific for the Temperate Zone of Europe
in the Middle and Late Pleistocene.
The Colchic Pheasant in Bulgaria is a species of autochtonous origin, recorded from 3 Late
Pleistocene sites. Phasianus colchicus (native subspecies colchicus) disappeared from the recent bird
fauna of the country and the whole of Europe. On the other hand the Black Grouse was actually
spread, even widely spread, throughout the whole country. Its remains were found in 7 Middle and
Late Pleistocene localities and they sufficiently document its distribution during the last 80 000 years.
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Received: 14.01.2005
Author’s address:
Dr. Zlatozar Boev,
National Museum of Natural History,
Tsar Osvoboditel Blvd. 1
1000 Sofia, Bulgaria
E-mail: boev@nmnh.bas.bg;
boevzaro@yahoo.co.uk

Pleistocene avifaunas of Bulgaria: a brief review 107
Плейстоценските орнитофауни в България – кратък преглед
Златозар БОЕВ
(Р е з ю м е)
Обобщени са фаунистичните резултати от проучването на плейстоценските
находища на птичи останки, известни досега в България. Те обхващат 160 вида, 4
ранноплейстоценски, 1 средноплейстоценско и 13 късноплейстоценски находища. Въпреки
това, все още плейстоценската авифауна на България е слабо проучена. От почти 5000
пещери и над 400 рецентни вида птици досега са проучени под 0,4 % от пещерите и са
установени 40,0 % от състава на птиците. Най-многобройни са материалите с вюрмска
възраст, съставляващи 2/3 от всички плейстоценски находки.
Плейстоценската авифауна включвала 8 изчезнали от съвременната фауна на страната
видове – Lagopus mutus, Lagopus lagopus, Tetrao tetrix, Phasianus colchicus colchicus, Glaucidium
passerinum, Nyctea scandiaca, ?Strix aff. nebulosa и Pyrrhocorax pyrrhocorax. Освен тях, 4 други вида
(Aegypius monachus, Gallinago gallinago, Otis tarda и Tetrax tetrax) са изчезнали в късния холоцен (20
век) като гнездещи птици в страната.
Късноплейстоценската орнитофауна включвала 2 фосилни таксона – полската
праяребица (Perdix palaeoperdix) и древната жълтоклюна гарга (Pyrrhocorax graculus vetus).
Плейстоценските находища на Perdix palaeoperdix вероятно доказват съвместното
съществуване на вида с полската яребица (Perdix perdix). Те са най-младите досега известни
находки на този вид.
Ранноплейстоценските авифауни са известни от 4 находища с 26 таксона и 5 разреда,
най-многобройни от които са Passeriformes. Един вид от тях (Pyrrhocorax pyrrhocorax) е изчезнал
от страната. Пещерата Чердженица е 4-то ранноплейстоценско находище на
червеноклюната хайдушка гарга в Палеарктика. Находките на Dendrocopos minor, Hirundo daurica,
Riparia riparia, Monticola saxatilis и Passer domesticus са най-древните находки в Палеарктика, а
находката на Lullula arborea е втора за ранния плейстоцен в Палеарктика.
Средноплейстоценските авифауни са най-слабо проучени. За тях свидетелства има само
от 1 находище със 7 таксона и 2 разреда.
Късноплейстоценските птици засега са изследвани най-пълно. Техни останки са намерени
в 13 находища със 134 таксона от 13 разреда. През интерглациалите някои видове с
медитеранско разпространение като Apus melba и Hirundo daurica са разширявали ареалите
си на север, достигайки дори до северните части на страната.
Плейстоценските авифауни в България се развивали в непосредствена връзка с фауните
на съседните райони в Югоизточна Европа. За някои групи птици те показват по-широки
ареали от съвременните. Плейстоценската (главно вюрмска) орнитофауна на страната
включвала целият борео-монтанен комплекс от над 15 вида, специфични за умерения пояс в
Европа през средния и късния плейстоцен.
Phasianus colchicus colchicus е с автохтонен произход, доказан от находките му в три
късно-плейстоценски находища. Tetrao tetrix действително е бил разпространен широко из
цялата страна. Останките му са намерени в 7 средно- и късноплейстоценски находища,
документирайки по такъв начин непрекъснатото му разпространение в България през
последните 80 000 г.