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TABLE 1. Results of absolute dating methods for various layers of Biśnik Cave (Cyrek et al., 2010; Van Asperen and Stefaniak, 2011). EU=Early Uptake model; LU=Linear Uptake model (Pike and Hedges, 2001). (PDF format)

TABLE 2. Bird remains identified in layers 19–8 (fauna complexes IX–V) in Biśnik Cave. Categories of habitat (see Material and Methods): A, amphibious; E, ecotone; F, forest; O, open; T, tundra. To exclude double counting of remains, before computing the minimum number of individuals (MNI) for mixed complexes it was always checked whether the remains from mixed sediments may have belonged to birds from layers with well-defined stratigraphy. (PDF format)

TABLE 3.Not fully ossified specimens of subadult and/or immature birds found in successive layers in Biśnik Cave.

 

Taxa No.

Taxon

Complex IX

Complex VIII

Complex VI

Complex V

Complex IV

Complex III

Complex II

Complex I

Mixed
Complexes

Total

19

18

15

12-13 mixed

10

9

6-7 mixed

4

3-4 mixed

2

1

1-2 mixed

1-12 mixed

?

1

cf. Anas strepera

 

 

 

 

 

 

 

 

 

 

 

 

1

 

1

2

Anas platyrhynchos

 

 

 

 

 

1

 

 

 

 

 

1

 

 

2

 

cf. Anas platyrhynchos

 

 

 

 

 

 

 

 

 

1

 

 

 

 

1

3

Anas querquedula/crecca

 

 

 

 

 

 

 

 

 

 

 

 

 

1

1

4

Melanitta nigra

 

 

 

 

 

 

 

 

 

 

 

 

 

1

1

 

Anatinae indet.

 

 

 

 

 

1

 

 

 

 

 

 

 

 

1

5

Buteo buteo/lagopus

 

 

 

 

 

 

 

 

 

 

 

1

 

2

3

6

Falco tinnunculus

 

1

 

 

 

 

 

 

 

2

1

 

2

9

15

 

Falco cf. tinnunculus

 

 

 

 

 

 

 

 

 

 

 

 

2

2

4

7

Falco peregrinus

 

 

 

 

 

 

 

 

 

 

 

 

 

1

1

 

Falco sp.

 

 

 

 

 

 

1

 

 

 

 

1

2

3

7

 

Falconiformes small size

 

 

 

 

 

 

 

 

 

 

 

1

 

 

1

8

Lagopus lagopus

 

 

 

 

1

 

 

 

 

 

 

 

 

 

1

 

Lagopus sp.

 

 

 

1

 

 

 

 

 

 

 

 

1

 

2

9

Tetrao tetrix

 

 

 

1

 

 

 

 

 

5

 

1

 

2

9

10

Tetrao urogallus

 

 

 

 

 

 

 

 

 

 

 

1

 

1

2

 

Tetrao/Lagopus

 

 

 

 

 

 

 

 

 

1

 

 

 

 

1

11

Gallus gallus

 

 

 

 

 

 

 

 

 

 

1

 

 

10

11

 

Galliformes indet

 

 

 

 

 

 

 

 

 

 

 

1

 

3

4

12

Rallus aquaticus

 

 

 

 

 

 

 

 

 

 

 

1

 

 

1

13

Strix aluco

 

 

 

 

 

 

 

 

 

1

 

 

 

 

1

14

Asio flammeus/Asio otus

 

 

 

 

 

 

 

 

 

 

 

1

 

 

1

 

Strigiformes middle size

 

 

 

 

 

 

 

 

 

 

 

1

 

 

1

15

Hirundo/Cecropis

 

1

 

 

 

 

 

 

 

 

 

 

 

 

1

16

Anthus sp.

1

 

 

 

 

 

 

 

 

 

 

 

 

 

1

17

Turdus viscivorus

 

 

 

 

 

 

 

 

 

 

 

 

1

 

1

 

Turdus viscivorus/pilaris

 

 

 

 

 

 

 

 

 

 

1

 

 

 

1

 

Turdus sp.

 

 

 

 

 

 

 

 

 

 

1

 

 

 

1

18

cf. Luscinia sp.

 

 

 

 

 

 

 

 

 

 

 

 

1

 

1

 

Turdidae (small) indet.

 

 

 

1

 

 

 

 

1

 

 

 

 

 

2

19

cf. Sylvia sp.

 

 

 

 

 

 

 

 

 

1

 

 

 

 

1

20

Corvus monedula

 

 

1

 

 

1

 

 

 

 

 

 

 

7

9

 

Corvus monedula/Pyrrhocorax

 

 

8

 

 

 

 

 

 

 

 

 

 

 

8

 

Corvidae (small) indet.

 

 

7

 

 

1

1

2

 

2

1

1

3

7

25

21

Corvus corax

 

 

 

 

 

 

 

 

 

 

 

 

 

1

1

 

Aves indet

 

 

 

 

 

 

 

 

 

 

 

 

 

1

1

 

Total

1

2

16

3

1

4

2

2

1

13

5

11

13

51

125

 
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Continuous 300,000-year fossil record: changes in the ornithofauna of Biśnik Cave, Poland

Teresa Tomek, Zbigniew M. Bocheński, Paweł Sochal, and Krzysztof Stefaniak

Plain Language Abstract

Biśnik  Cave is situated in a limestone rock about 50 km northeast of Kraków, southern Poland. Its importance stems from the fact that it is one of the few sites in Europe, and the only one in Poland, with 300,000-year-long sequence of uninterrupted sediments, that cover the time span from before the Saalian (i.e., the Odra and Warta glacial periods) to the Holocene. The excavations yielded about 200,000 animal bones and more than 4,000 stone, bone and antler artifacts. Bird remains from Biśnik  Cave consist of nearly 1,600 skeletal fragments of at least 96 taxa that represent a minimum of 285 individuals. The majority of the remains belong to Galliformes; relatively numerous are also Corvidae, Falconiformes, Anseriformes and species of the genus Turdus. The remains include one extinct falcon (Falco tinnunculus atavus) and four species new for the Polish fossil avifauna: the Eastern Imperial Eagle (Aquila heliaca), Pine Grosbeak (Pinicola enucleator), Parrot Crossbill (Loxia pytyopsittacus) and Common Redpoll (Carduelis flammea). Avian remains indicate a mosaic of various habitats in the surroundings of Biśnik  Cave. Some kind of mature forest or at least sparsely growing trees, as well as water bodies, marshes, wet meadows, steppe and tundra habitats must have been present during the entire time of sedimentation. It is postulated that the Kraków-Częstochowa Upland was a local refugium for the forest fauna during the Saalian and Vistulian glaciations. Due to its geographic location, the region was probable a transition zone between the steppe habitats of Central Asia and the more oceanic environments of Western Europe.

Resumen en Español

Registro fósil continuo a lo largo de 300000 años: cambios en la ornitofauna de la cueva de Biśnik, Polonia

La cueva de Biśnik, excavada en rocas calizas, se sitúa unos 50 kilómetros al noreste de Cracovia, en el sur de Polonia. Su importancia radica en el hecho de que es uno de los pocos yacimientos europeos, y el único en Polonia, con una secuencia ininterrumpida de sedimentos que abarca un intervalo de 300000 años, desde antes del Saaliense hasta el Holoceno. Las excavaciones en dicha cueva han proporcionado cerca de 200000 huesos de animales y más de 4000 utensilios en piedra, hueso y asta. Los restos de pájaros hallados en Biśnik consisten en cerca de 1600 fragmentos esqueléticos de al menos 285 individuos pertenecientes a un mínimo de 96 taxones. La mayoría de los restos son de Galliformes, aunque son también relativamente abundantes los de Corvidae, Falconiformes, Anseriformes y el género Turdus. Los restos incluyen también un taxón extinguido (Falco tinnunculus atavus) y cuatro especies nuevas en la avifauna fósil de Polonia (Aquila heliaca, Pinicola enucleator, Loxia pytyopsittacus y Carduelis flammea). Los restos de aves indican la existencia de un mosaico de hábitats distintos en los alrededores de la cueva de Biśnik. Durante el intervalo de tiempo representado en los sedimentos habría algún tipo de bosque maduro o, al menos, árboles dispersos, así como cuerpos de agua y hábitats de marisma, praderas húmedas, estepa y tundra. La meseta de Cracovia-Częstochowa debió constituir un refugio para la fauna de los bosques durante las glaciaciones del Saaliense y del Vistuliense.

Palabras clave: aves fósiles, Pleistoceno, paleoambiente, paleoclima.

Traducción: Miguel Company

Résumé en Français

Un enregistrement continu de 300.000 ans: changements dans l’ornithofaune de la grotte de Biśnik, Pologne

La grotte de Biśnik est située dans des roches calcaires à environ 50 km au nord-est de Cracovie, sud de la Pologne. Sont importance repose sur la fait qu’elle est un des rares sites en Europe, et le seul en Pologne, avec une séquence ininterrompue de sédiments longue de 30.000 ans, couvrant une période allant du Saalien à l’Holocène. Les fouilles ont livré environ 200.000 os d’animaux et plus de 4.000 pierres, et artefacts d’os et de bois. Les restes d’oiseaux de la grotte de Biśnik se composent de près de 1.600 fragments de squelettes provenant d’au moins 96 taxa qui représentent au minimum 285 individus. La majorité des restes appartiennent aux Galliformes ; les Corvidae, Falconiformes, Anseriformes et le genre Turdus sont aussi relativement nombreux. Les restes incluent un taxon éteint (Falco tinnunculus atavus) et quatre nouvelles espèces pour les avifaunes fossiles polonaises (Aquila heliaca, Pinicola enucleator, Loxia pytyopsittacus and Carduelis flammea). Les restes d’oiseaux indiquent l’existence d’une mosaïque d’habitats variés dans les alentours de la grotte de Biśnik. Un certain type de forêt mature ou au moins d’arbres peu développés, de même que d’étendue d’eau, marais, prairies humides, et d’habitat de steppe et toundra ont dû être présents pendant toute la période de sédimentation. L’hypothèse est faite que les hautes terres de Cracovie-Częstochowa constituaient un refuge local pour les faunes forestières pendant le Saalien et les glaciations Vistuliennes.

Mots clés: oiseaux fossiles, Pléistocène, paléoenvironnent, paléoclimat

Translator: Olivier Maridet

Deutsche Zusammenfassung

Ein durchgehender Fossilrekord von 300 000 Jahren: Änderungen in der Ornithofauna der Biśnik Höhle von Polen

Die Biśnik Höhle befindet sich in einem Kalksteingebirge ungefähr 50 km nordöstlich von Krakau, Südpolen. Ihre Bedeutung liegt darin, dass sie eine der wenigen Höhlen in Europa ist, und die einzige in Polen, die mit einer 300 000 Jahre andauernden ununterbrochenen Sedimentation eine Zeitspanne vom Saalium bis zum Holozän abdeckt. Die Ausgrabung hat mehr als 200 000 Tierknochen und mehr als 4 000 Stein-, Knochen-und Geweihartefakte hervorgebracht. Vogelreste aus der Biśnik Höhle bestehen aus beinahe 1 600 Skelettfragmenten von mindestens 96 Taxa, die ein Minimum an 285 Individuen repräsentieren. Der Großteil der Funde gehört zu den Galliformes. Relativ häufig sind auch Corvidae, Falconiformes, Anseriformes und die Gattung Turdus. Die Reste beinhalten ein ausgestorbenes Taxon (Falco tinnunculus atavus) und vier neue Arten der fossilen polnischen Avifauna (Aquila heliaca, Pinicola enucleator, Loxia pytyopsittacus and Carduelis flammea). Die Vogelreste deuten auf ein Mosaik von verschiedenen Habitaten in der Umgebung der Biśnik Höhle hin. Während der gesamten Sedimentationsperiode müssen sowohl eine Art von Waldbestand oder zumindest niedrig wachsende Bäume vorhanden gewesen sein, als auch Gewässer, Sümpfe, Feuchtwiesen, Steppe und Tundra Habitate. Es ist postuliert, dass das Kraków-Częstochowa Hochland während der Saale -und Vistula - Vereisungen ein lokales Refugium für die Waldfauna war.

SCHLÜSSELWÖRTER: fossile Vögel, Pleistozän, Paläoumwelt, Paläoklima

Translator: Eva Gebauer

Arabic

277 arab

Translator: Ashraf M.T. Elewa

Polski Abstrakt

Ciągły, 300.000-letni, zapis kopalny: zmiany ornitofauny jaskini Biśnik, Polska

Jaskinia Biśnik jest usytuowana w wapiennych skałach południowej Polski, ok. 50 km na północny-wschód od Krakowa. Jest to jedno z niewielu stanowisk w Europie, a jedyne w Polsce, posiadające nieprzerwaną sekwencję osadów gromadzących się 300,000 lat i obejmujących okres od zlodowaceń Odry i Warty (Saalian) aż do holocenu. Wykopaliska prowadzone w tej jaskini dostarczyły około 200 000 szczątków zwierzęcych i ponad 4 000 zabytków z kamienia, kości i rogu. Szczątki ptasie z Jaskini Biśnik zawierały około 1 600 fragmentów kostnych należących do 96 taksonów i łącznie reprezentujących nie mniej niż 285 osobników. Większość szczątków ptasich należała do przedstawicieli grzebiących Galliformes; stosunkowo liczne były również kości reprezentujące krukowate Corvidae, szponiaste Falconiformes, blaszkodziobe Anseriformes oraz drozdy (rodzaj Turdus). Zachowane szczątki zawierały jednego wymarłego sokoła (Falco tinnunculus atavus) i cztery gatunki nowe dla kopalnej fauny Polski: orła cesarskiego (Aquila heliaca), łuskowca (Pinicola enucleator), krzyżodzioba sosnowego (Loxia pytyopsittacus) i czeczotkę (Carduelis flammea). Szczątki ptasie wskazują na obecność mozaiki biotopów w okolicy jaskini Biśnik. Dojrzały las, rzadko rosnące kępy drzew, otwarte wody, bagna, podmokłe łąki, step i tundra musiały być obecne w sąsiedztwie jaskini podczas całego okresu sedymentacji. To sugeruje, że Wyżyna Krakowsko-Częstochowska była lokalnym refugium dla fauny leśnej podczas zlodowaceń Odry, Warty i Wisły. Z powodu geograficznego położenia Wyżyny region ten był prawdopodobnie strefą przejściową między stepami centralnej Azji a strefą lasów liściastych Europy zachodniej.

Słowa kluczowe: kopalne ptaki, plejstocen, paleośrodowisko, paleoklimat

Translation: supplied by authors

Riassunto in Italiano

Un record fossile continuo lungo 300,000 anni: variazioni nell’ornitofauna di Biśnik Cave, Polonia

La Biśnik Cave è impostata in rocce calcaree a 50 km a nordest di Cracovia, Polonia meridionale. La sua importanza risiede nel fatto che è uno dei pochi siti in Europa e l’unico in Polonia che registra una sequenza ininterrotta di 300,000 anni, ovvero da epoche antecedenti al Saaliano all’Olocene. I lavori di scavo hanno riportato alla luce circa 200,000 ossa di animali e più di 4,000 manufatti di pietra, osso e corno. I resti di uccelli della Biśnik Cave consistono in quasi 1,600 frammenti scheletrici di almeno 96 taxa per un numero minimo di 285 individui. La maggior parte dei resti appartiene all’ordine Galliformes, relativamente numerosi sono Corvidae, Falconiformes, Anseriformes e il genere Turdus. Annoveriamo fra l’altro un taxon estinto (Falco tinnunculus atavus) e quattro specie nuove nel record fossile dell’avifauna polacca (Aquila heliaca, Pinicola enucleator, Loxia pytyopsittacus e Carduelis flammea). I resti di uccelli indicano un mosaico di diversi habitat nei dintorni della Biśnik Cave: foreste, o quantomeno alberi sparsi, specchi o corsi d’acqua, paludi, wet meadows, steppe e tundra sembrano essere stati presente durante l’intero intervallo di sedimentazione. Ipotizziamo che la Kraków-Częstochowa Upland costituisse un rifugio per la fauna di foresta durante le glaciazioni Saaliana e Vistuliana.

Parole chiave: uccelli fossili, Pleistocene, paleoambiente, paleoclima

Translator: Chiara Angelone

 
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FIGURE 1. Location of Biśnik Cave in southern Poland.

fig1

FIGURE 2. Left ulna in cranial view and left tarsometatarsus in dorsal view. Falco tinnunculus atavus (1, 3) and modern female of Falco tinnunculus (2, 4). The fossil specimen is much larger and more robust than a large modern female.

fig2

FIGURE 3. Percentage share of bird taxa (numbers of taxa) breeding in five groups of habitats, excavated in particular layers in Biśnik Cave (based on data from Table 2). The correlation between particular fauna complexes and their geological age is explained in Table 1.

fig3

 
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tomekThis email address is being protected from spambots. You need JavaScript enabled to view it.
Institute of Systematics and Evolution of Animals
Polish Academy of Sciences
Slawkowska 17
31-016 Krakow
Poland

Teresa Tomek graduated from the Jagiellonian University, Kraków, Poland in 1967 and received her PhD in 1976. She has been working at the Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków since 1971. She published more than 60 scientific papers on various aspects of ornithology, paleontology and comparative osteology of birds. Teresa took part in eight ornithological expeditions to the Democratic People's Republic of Korea.

bochenskiThis email address is being protected from spambots. You need JavaScript enabled to view it.
Institute of Systematics and Evolution of Animals
Polish Academy of Sciences
Slawkowska 17
31-016 Krakow
Poland (correspondance author)

Zbigniew M. Bocheński graduated from the Jagiellonian University, Kraków, Poland in 1987 and received his PhD in 1994. He has been working at the Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland since 1987. He divides his research activity between fossil and modern bird remains. He has published a number of papers on avian taphonomy, fossil and archaeological bird remains, and several works on the comparative osteology of modern birds. He was the recipient of three major one-year each scholarships, which helped him gain professional experience in other academic centers, including Germany, Switzerland and the USA.

sochaThis email address is being protected from spambots. You need JavaScript enabled to view it.
Department of Palaeozoology
Zoological Institute
University of Wroclaw
Sienkiewicza 21
50-335 Wrocław
Poland

Paweł Socha graduated from the Wroclaw University, Poland in 1992 and received his PhD in 2010. He has been working at the Department of Palaeozoology, Zoological Institute, University of Wroclaw, Poland since 1991. His interests are focused on fossil mammals, especially Neogene and Quaternary rodents and palaeoecology. Further areas of research include palaeoclimatology, caves and fossil DNA. He has participated in excavations in karst areas of Poland and the Ukraine for 25 years. He lectures palaeontology, archaeozoology and the history of the formation of the modern fauna and flora during the Quaternary at the University of Wroclaw.

stefThis email address is being protected from spambots. You need JavaScript enabled to view it.
Department of Palaeozoology
Zoological Institute
University of Wroclaw
Sienkiewicza 21
50-335 Wrocław
Poland

Krzysztof Stefaniak graduated from the Wroclaw University, Poland in 1986 and received his PhD in 2001. He has been working at the Department of Palaeozoology, Zoological Institute, University of Wroclaw, Poland since 1984. His interests are focused on fossil mammals, especially ungulates, including Neogene and Quaternary deer. His research activity includes also archaeozoology, caves and fossil DNA. He has participated in excavations in karst areas of Poland and the Ukraine for 30 years. He lectures palaeontology, archaeozoology and the history of the formation of the modern fauna and flora during the Quaternary at the University of Wroclaw.