The trophic ecology of wolves and their predatory role in ungulate communities of forest ecosystems in Europe

• Autorzy: Okarma H.
• Języki publikacji: EN

Abstrakty

EN

Predation by wolves Canis lupus Linnaeus, 1758 in ungulate communities in Europe, with special reference to the multi-species system of Białowieża Primeval Forest (Poland/Belarus), was assessed on the basis results of original research and literature. In historical times (post-glacial period), the geographical range of the wolf and most ungulate species in Europe decreased considerably. Community richness of ungulates and potential prey for wolves, decreased over most of the continent from 5-6 species to 2-3 species. The wolf is typically an opportunistic predator with a highly diverse diet; however, cervids are its preferred prey. Red deer Ceruus eiaphus are positively selected from ungulate communities in all localities, moose Alces alces are the major prey only where middle-sized species are scarce. Roe deer Capreolus capreolus are locally preyed on intensively, especially where they have high density, co-exist mainly with moose or wild boar Sus scrofa, and red deer is scarce or absent. Wild boar are generally avoided, except in a few locations; and European bison Bison bonasus are not preyed upon by wolves. Wolf predation contributes substantially to the total natural mortality of ungulates in Europe: 42.5% for red deer, 34.5% for moose, 25.7% for roe der, and only 16% for wild boar. Food niche breadth (B) of wolves in Europe, calculated only for the ungulates considered in this study, increases with the number of ungulate species in the community. There is also a significant rela­tionship between ungulate community breadth and food niche breadth of wolves. Food niche breadth of wolves, however, does not achieve very high values even in the richest ungulate communities. Wolves easily adapt to locally abundant food of anthropogenic origin (livestock, garbage). The level of predation on livestock may be a result of different husbandry practices (eg use of livestock guarding dogs) rather than of differences in availability of wild and domestic prey. Available data from Europe suggest that wolves likely limit density of red deer and moose in some areas. Roe deer density can be decreased locally by wolves but is limited mainly by lynx Lynx lynx. Wild boar density is more influenced by mast crops of Quercus spp. and Fagus siluatica (and to a lesser extent by snow depth) than by wolf predation.

Słowa kluczowe

EN

ungulate community; forest ecosystem; trophic ecology; wolf; red deer; wild boar; Alces alces; Fagus sylvatica; Sus scrofa; Capreolus capreolus; Quercus; prey preference; predation; Cervus elaphus; Canis lupus; moose; roe deer; Europe; Bialowieza National Park

• Wydawca: -
• Czasopismo: Acta Theriologica
• Rocznik: 1995
• Tom: 40
• Numer: 4
• Opis fizyczny: p.335-386,fig.

Twórcy

autor

Okarma H.

• Polish Academy of Sciences, 17-230 Bialowieza, Poland

Bibliografia

• Abdunazarov B. B. and Taryannikov V, I. 1991. [Volk in the western Tien-shan!. [In: Ekologiya, okhrana i akklimatizatsiya mlekopitayushchikh v Uzbekistane. 0. P. Bogdanov, ed], Izdatelstvo AN UzSSR, Tashkent: 42-48. |In Russian]
• Almasan H., Scarlatescu G., Nesterov V. and Manolache L, 1970. Contribution a la connaissance du regime de nourriture du loup (Cants lupus L.} dans les Carpathes roumaines. Transactions 9th IUGB Congress, Moscow 1969: 523-529.
• Apollonio M., Randi E. and Toso S. 1988. The systematics of Italian wild boar. Bollettino di zoologia 3: 213-221.
• Aragon S., Braza F. and San Jose C. 1995. Socioeconomic, physiognomic, and climatic factors determining the distribution pattern of roe deer Capreolus capreolus in Spain. Acta Theriologica 40: 37-43.
• Atlas Świata [Atlas of the World], 1962. PWN-Polish Scientific Publisher, Warszawa: 1-523.
• Ballard W. B., Whitman J. S. and Gardner C. L. 1987. Ecology of an exploited wolf population in south-central Alaska. Wildlife Monographs 98: 1-54,
• Ballard W. B., Whitman J. S. and Reed D. J. 1990. Population dynamics of moose in in south-central Alaska. Wildlife Monographs 114: 1-49.
• Bannikov A. G., Zhirnov L. V., Lebedeva L. S. and Fandeev A. A. 1961, [Biology of saiga antelope], Izdatelstvo Selskokhozyaistvennoi Literatury, Zhurnalov i Plakatov. Moskva: 1-336. ¡[In Russian]Baskin L. M. 1979. Ecology and behaviour of European bison. |In: European bison, morphology, systematics, evolution, ecology. V. E. Sokolov, ed], Izdatelstvo Nauka, Moskva: 442-470. [In Russian]
• Beiyanin V. N. 1979, [Wolves in Zhigule Mountains], [In: Ekologicheskie osnovy okhrany i ratsio- nalnogo ispolzovaniya khishchnykh mlekopitayushchikhl. Izdatelstvo Nauka, Moskva: 86-87. [In Russian]
• Bergerud A. T. and Elliot J. P. 1986. Dynamics of caribou and wolves in northern British Columbia. Canadian Journal of Zoology 64: 1515-1529.
• Bergerud A. T. and Snider J. B. 1988. Predation in the dynamics of moose populations: a reply. Journal of Wildlife Management 52: 559-564.
• Bergerud A. T., Wyett W. and Snider J. B. 1983. The role of wolf predation in limiting a moose population. Journal of Wildlife Management 47: 977-988.
• Bibikov D. I. (ed). 1985. The wolf. History, systematics, morphology, ecology. Izdatelstvo Nauka, Moskva: 1-606. Iln Russian]
• Bibikov D. I., Kudaktin A. N. and Ryabov L. S. 1985. Synanthropic wolves: distribution, ecology. Zoologicheskii Zhurnal 64: 429-441. [In Russian with English summary]
• Bjárvall A. and Isakson E. 1982. Winter ecology of a pack of three wolves in northern Sweden. [In: Wolves of the world, F. Harrington and P. C. Paquet, eds]. Noyes Publications, New Jersey: 146-157.
• Blanco J. C., Cuesta L. and Reig S. (eds) 1990. [The wolf (Canis lupus) in Spain!. ICONA, Madrid: 1-118. [In Spanish with English summariesi
• Blanco J. C., Reig S. and Cuesta L, 1992. Distribution, status and conservation problems of the wolf Canis lupus in Spain. Biological Conservation 60: 73-80.
• Bobek B., Borowski S. and Dzięciołowski R. 1975. Browse supply in various forest ecosystems. Polish Ecological Studies 1: 17-32.
• Bobek B., Perzanowski K. and Śmietana W, 1992. The influence of snow cover on the patterns of selection within red deer population by wolves in Bieszczady Mountains, Poland. |In: Global trends in wildlife management. B. Bobek, K. Perzanowski and W. Regelin, edsl. Transactions of 18th IUGB Congress, Kraków 1987. Świat Press, Kraków-Warszawa, Vol. 2: 341-348.
• Boitani L. 1982. Wolf management in intensively used areas of Italy, [In: Wolves of the world. F. Harrington and P. C. Paquet, eds]. Noyes Publications, New Jersey: 158-172.
• Boitani L. 1992. Wolf research and conservation in Italy. Biological Conservation 61: 125-132.
• Boutin S. 1992. Predation and moose population dynamics: a critique. Journal of Wildlife Manage­ment 56: 116-127.
• Boyd D. K., Ream R. R., Pletscher D. H. and Fairchild M. W. 1994. Prey taken by colonizing wolves and hunters in the Glacier National Park area. Journal of Wildlife Management 58: 289-295.
• Brana F., Campo del J. C. and Palomero G. 1982. Le loup au versant nord de la Cordillere Can- tabrique. Acta Biologica Montana 1: 33-52.
• Brangi A., Rosa P. and Meriggi A. 1991. Predation by wolves (Canis lupus L.) on wild and domestic ungulates in northern Italy. Proceedings of a conference "Ongules/Ungulates 91", Tolouse, France: 541-543.
• Brtek L. and Voskár J. 1987. IFeeding ecology of the wolf in Slovakian Carpathians]. Biologia (Bratislava) 42: 985-990. [In Slovak]
• Bubenik A. B. 1966. [Impact of lynx Lynx lynx L. and wolf Canis lupus L. on population structure of roe deer and red deer]. Lynx 6: 7-10. [In Czech with German summary]
• Buchalczyk T. 1980. The brown bear in Poland. [In: Bears - their biology and management. C. J. Martinka and K, L. Mc Arthur, eds[. Bear Biologists Association Conference Ser 3: 229-232.
• Bunevich A, N. 1988. [Dynamics of numbers and diet of wolf in Belovezha Primeval Forestj. Zapo- vedniki Belorussii 12: 108-113. [In Russian]
• Bützler W. 1986. Cervus ela.ph.us Linnaeus, 1758 - Rothirsch. [In: Handbuch der Saugetiere Europas. Band 2/11. J. Niethammer and F. Krapp, eds|. Aula-Verlag, Wiesbaden: 107-139.
• Caboń K. 1958. Das Massensterben von Wildschweinen im Naturstaatspark von Białowieża im Winter 1955/56. Acta Theriologica 2: 71-82.
• Carbyn L. N. 1983. Wolf predation on elk in Riding Mountain National Park, Manitoba. Journal of Wildlife Management 47: 963-976.
• Carbyn L. N., Oosenbrug S. M. and Anions D. W. 1993. Wolves, bison and the dynamics related to the Peace-Athabasca delta in Canada's Wood Buffalo National Park. Circumpolar Research Series No 4, Canadian Circumpolar Institute, University of Alberta, Edmonton: 1-270,
• Centofanti E. 1994. [Presence of the wolf Canis lupus L. and other wild canids in relation to ugulate populations in the Casentinesi Forests], Unpublished report, Gruppo Lupo Italia: 1—11. [In Italian!
• Chapman D. and Chapman N. 1975. Fallow deer. Terence Dalton Limited. Lavenham-Suffolk: 1-271.
• Cowan I. McT. 1947. The timber wolf in the Rocky Mountain National Parks of Canada. Canadian Journal of Research 25: 139-174.
• Cuesta L,, Barcena F,, Palacios F. and Reig S. 1991. The trophic ecology of the Iberian wolf (Cants lupus signatus Cabrera, 1907). A new analysis of stomach's data. Mammalia 55: 240-254.
• Danilkin A. A. 1989. [Structure of geographical range of roe deer in USSR and factors responsible for its limits], Ekologiya 4: 46-51. [In Russian)
• Darman Yu. A. 1986. [Wolf and ungulates in Khingansk reserve). Trudy 4 Sezda Vsesoyuznogo teriologicheskogo obshchestva No 3, AN SSSR, Moskva: 45. [In Russian]
• Dubrovski'i V. Yu. 1980. [Remarks on summer ecology of the wolf in taiga of the Kostromska oblast], [In: Behaviour of the wolf. D. I. Bibikov, ed]. AN USSR, Moskva: 123-129. |In Russian]
• Dzięciolowski R. 1969. The quantity, quality and seasonal variation of food resources available to red deer in various environmental conditions of forest management. Forest Research Institute, Warszawa: 1-295.
• Dzięciolowski R. and Pielowski Z. 1993. [Moose], Anton-5, Warszawa: 1-208. [In Polish!
• European bison pedegree book 1993. 1995. National Council for Nature - Białowieża National Park, Białowieża: 1-61.
• Fadeev E. 1984. [Sika deer in the USSRI. Okhota i okhotniche khozyaystvo 5: 18-21. [In Russian!
• Faliński J. B. 1986. Vegetation dynamics in temperate lowland primeval forest. Dr. W. Junk Pub­lishers, Dordrecht. Geobotany 8: 1-537.
• Fedosenko A. K., Zhiryakov V. A. and Grachev Yu. A. 1978. Some data on the ecology and behaviour of wolves in the northern Tien-shan and Dzhungara Alatau, Byulleten Moskovskogo Obshchestva Ispytatelei Prirody, Otdel Biologicheski 83(3): 5-18. |In Russian with English summary!
• Fico R., Morosetti G. and Giovannini A. 1993. The impact of predators on livestock in the Abruzzo region of Italy. Revue scientifique et technique. Office international des epizooties 12(1): 39-50.
• Filonov K. P. 1980. Predator-prey problems in nature reserves of the European part of the RSFSR. Journal of Wildlife Management 44: 389-396.
• Filonov K. P. 1983. [Variability of mortality factors in populations of wild ungulates], Ekologiya 2: 57—64. [In Russianl
• Filonov K. P. 1989. Ungulates and large predators in wildlife reserves. Izdatelstvo Nauka, Moskva: 1-251. [In Russian]
• Filonov K. P. and Kaletskaya M. L, 1985, [Influence of wolf predation on wild ungulates], (In: The wolf. D. I. Bibikov, ed], Izdatelstvo Nauka, Moskva: 336-354. [In Russian]
• Flerov K. K. 1979. Systematics and evolution. [In: European bison, morphology, systematica, evo­lution, ecology. V. E. Sokolov, ed], Izdatelstvo Nauka, Moskva: 8-127. [In Russian]
• Fonseca F. P. 1992. Population trends of wolves in Portugal. [In: Wolves in Europe. Ch. Promberger and W. Schroder, eds]. Proceedings of the workshop "Wolves in Europe — current status and prospects", Oberammergau 1992. Miinich Wildlife Society, Ettal: 111-112.
• Forbes G. and Theberge J. B. 1992. Importance of scavenging on moose by wolves in Algonquin Park, Ontario. Alces 28: 235-241.Formozov A. N. 1946. [Snow cover and its importance in the life of mammals and birds of SSSR]. Izdatelstvo Moskovskogo Universiteta, Moskva: 1-287. [In Russian]
• Fritts S. H. and Mech L. D. 1981. Dynamics, movements, and feeding ecology of a newly protected wolf population in nortwestern Minnesota. Wildlife Monographs 80: 1-79.
• Fruziński B. 1992. [Wild boarl. Cedrus, Warszawa: 1-248. [In Polish]
• Fuller T, K. 1989. Population dynamics of wolves in north-central Minnesota. Wildlife Monographs 105: 1-41.
• Fuller T. K. 1991. Effect of snow depth on wolf activity and prey selection in north central Minnesota. Canadian Journal of Zoology 69: 283-287.
• Fuller T. K. and Keith L. B. 1980. Wolf population dynamics and prey relationships in northeastern Alberta. Journal of Wildlife Management 44: 583-602.
• Gaross V. Ya. 1979. [Impact of lynxes and wolves on roe deer population in Lithuania], [In: Eko- logicheskie osnovy okhrany i ratsionalnogo ispolzovaniya khishchnykh mlekopitayushchikh], Izdatelstvo Nauka, Moskva: 257-261. |In Russian]
• Gasaway W. C., Boertje R. D., Grangaard D. V., Kelleyhouse D. G., Stephenson R. O. and Larsen D. G. 1992, The role of prédation in limiting moose at low densities in Alaska and Yukon and implications for conservation. Wildlife Monographs 120: 1-59.
• Gasaway W. C., Stephenson R. O., Davis J. L., Shepherd E. K. and Burris O. E. 1983. Inter­relationships of wolves, prey, and man in interior Alaska. Wildlife Monographs 84: 1-50.
• Gatikh V. C. 1979. [The role of wolves in biocenosis of the Belarussian Polese], [In: Ekologicheskie osnovy okhrany i ratsionalnogo ispolzovaniya khishchnykh mlekopitayushchikh], Izdatelstvo Nauka, Moskva: 94-95. [In Russian] Gavrin V. F. and Donaurov S. S. 1954, [Wolf in Belovezha Primeval Forest], Zoologicheskiï Zhurnal 33: 904-924. [In Russian]
• Genov P. 1981. Food composition of wild boar northeastern and western Poland. Acta Theriologica 26: 185-205.
• Genov P. 1992. The wolf Cam's lupus L. in southwestern Bulgaria. [In: Global trends in wildlife management. B. Bobek, K. Perzanowski and W. Regelin, eds]. Transactions of 18th IUGB Congress, Kraków 1987. Świat Press, Kraków-Warszawa: 359-362.
• Geograficzny Atlas Świata [Geographical Atlas of the World], 1991. Państwowe Przedsiębiorstwo Wydawnictw Kartograficznych. Warszawa-Wrocław: 1-274.
• Golgofskaya K. Yu. 1970. The growth of wild ungulate populations and the state of grasslands in the Caucasian preserve. Byulleten Moskovskogo Obshchestva Ispytatelei Prirody, Otdel Biologicheski 75(4): 17-21. |In Russian with English summary]
• Gordiyuk N. M. 1986. [Selectivity of wolf hunting in the southern Ural Mountains depending on depth and condition of snow layer], |In: Rol krupnykh khishchnikov i kopytnykh v biotsenozakh zapovednikov], CNIL Glavokhoty RSFSR, Moskva: 54-59. [In Russian]
• Gordiyuk N. M. 1991. Interrelations between wolves and ungulates as registered by observation of the marked wolf pack in the southern Urals, Byulleten Moskovskogo Obshchestva Ispytatelei Prirody, Otdel Biologicheski 96(1): 3-10. [In Russian with English summary]
• Gromov E. I. 1979. [Diet of wolves in Sikhote-AlinskiV Reserve]. [In: Ekologicheskie osnovy okhrany i ratsionalnogo ispolzovaniya khishchnykh mlekopitayushchikh], Izdatelstvo Nauka, Moskva: 101-102. [In Russian]
• Gunchak N. S. 1980. Diet of wild boar in the Ukrainian Carpathians, Vestnik Zoologii 5: 73-78. [In Russian with English summary] Gyimesi J. 1971. [Migrations of moose to Czechoslovakia], Polovnicky Zbornik 1: 217-224, [In Slovak with German summary]
• Haglund B. 1968. Winter habits of the bear (Ursus arctos L.) and the wolf (Canis lupus L.) as revealed by tracking in the snow. Viltrevy 5: 217-361. |In Swedish with English summary]
• Heidemann G. 1986, Cervus dama (Linnaeus, 1758) - Damhirsch. [In: Handbuch der Saugetiere Europas, Band 2/11. J. Niethammer and F. Krapp, eds], Aula-Verlag, Wiesbaden: 140-158.
• Heptner V. G., Nasimovich A. A. and Bannikov A. G. 1961. [Artiodactyla and Perissodactyla]. [In: Mlekopitayushchie Sovetskogo Soyuza. V. G. Geptner and N. P. Naumov, eds]- Izdatelstvo Vysshaya shkola, Moskva: 1-776. [in Russian]
• Herre W, 1986a. Sus scrofa Linnaeus, 1758 - Wildschwein. [In: Handbuch der Saugetiere Europas. Band 2/II. J. Niethammer and F. Krapp, eds], Aula-Verlag, Wiesbaden: 36-66.
• Herre W, 1986b. Rangifer tarandus (Linnaeus, 1758) - Ren, Rentier. [In: Handbuch der Saugetiere Europas. Band 2/II. J, Niethammer and F. Krapp, eds], Auta-Vertag, Wiesbaden: 198-216.
• Holleman D. F. and Stephenson R. O. 1981. Prey selection and consumption by Alaskan wolves in winter. Journal of Wildlife Management 45: 620-628.
• Houston D, B. 1982. The northern Yellowstone elk, ecology and management. Macmillan Publishing Co., Inc., New York: 1-474.
• Huggard D. J. 1993a. Prey selectivity of wolves in Banff National Park. I. Prey species. Canadian Journal of Zoology 71: 130-139.
• Huggard D. J. 1993b. Prey selectivity of wolves in Banff National Park. II. Age, sex, and condition of elk. Canadian Journal of Zoology 71: 140-147.
• Huggard D. J. 1993c. Effect of snow depth on predation and scavenging by gray wolves. Journal of Wildlife Management 57: 382-388.
• Ionescu O. 1992. Current status and prospects for the wolf in Romania. [In: Wolves in Europe. Ch, Promberger and W. Schroder, eds]. Proceedings of the workshop "Wolves in Europe — current status and prospects", Oberammergau 1992. Münich Wildlife Society, Ettal: 9-14.
• Ivanov V. K. 1988. Observations on the behaviour and biology of wolves (Canis lupus L., 1758) in the Ihtiman Sredna Gora Mountains. Bulgarian Academy of Sciences, Ecology 21: 25-33.
• Jacobs J, 1974. Quantitative measurements of food selection; a modification of the forage ratio and Ivlev's Eiectivity index, Oecologia 14: 413-417.
• Jędrzejewska B., Jędrzejewski W., Bunevich A. N., Milkowski L. and Okarma H. 1996a. Population dynamics (1847-1993) and demography of wolves Canis lupus in Białowieża Primeval Forest (Poland and Belarus). Mammal Review, (in press)
• Jędrzejewska B., Jędrzejewski W., Okarma H„ Schmidt K., Bunevich A. N. and Milkowski L. 1996b. Population dynamics (1869-1994), demography and home ranges of the lynx in Białowieża Primeval Forest (Poland and Belarus). Ecography, (in press)
• Jędrzejewska B., Okarma H., Jędrzejewski W. and Milkowski L. 1994. Effect of exploitation and protection on forest structure, ungulate density and wolf predation in Białowieża Primeval Forest, Poland. Journal of Applied Ecology 31: 664-676.
• Jędrzejewski W., Jędrzejewska B., Okarma H. and Ruprecht A, L, 1992. Wolf predation and snow cover as mortality factors in the ungulate community of the Białowieża National Park, Poland. Oecologia 90: 27-36,
• Jędrzejewski W., Jędrzejewska B. and Szymura A. 1989. Food niche overlaps in a winter community of predators in the Białowieża Primeval Forest, Poland. Acta Theriologica 34: 337-348.
• Jędrzejewski W., Schmidt K., Milkowski L., Jędrzejewska B. and Okarma H. 1993. Foraging by lynx and its role in ungulate mortality: the local (Białowieża Forest) and the Palaearctic viewpoints. Acta Theriologica 38: 385-403.
• Kaletskaya M. L. 1973. [Wolf and its role as a predator in the Darwinskii" Reserve], Trudy Darvins- kogo gosudarstvennogo zapovednika 9: 41-58. [In Russian]
• Kaluziński J. 1974. The occurrence and distribution of field ecotype of roe-deer in Poland, Acta Theriologica 19: 291-300.
• Kartsov G. 1903. [Belovezha Primeval Forest. Its historical description, contemporary game manage­ment, and monarchial hunts in the Forest, A. Marks, St. Petersburg: 1-414. [In Russian]
• Kazenevskii P. F. 1961. On more case of death in the wild ungulates (winters 1955/56). Byulleten Moskovskogo Obshchestva Ispytatelei Prirody, Otdel Biologicheski 66(6): 147-148. [In Russian with English summary]Kazenevskii P. F. 1979. [Volk in the Khopersk Reserve], [In: Ekologicheskie osnovy okhrany i ratsio- nalnogo ispolzovaniya khishchnykh mlekopitayushchikh], Izdatelstvo Nauka, Moskva: 110—112. [In Russian]
• Keith L. B. 1974. Some features of population dynamics in mammals. Proceedings of 11th IUGB Congress, Stockholm 1973. National Swedish Environmental Protection Board, Stockholm: 17-58.
• Kerechun S. F. 1979. [Impact of carnivores on numbers of ungulates in a zone of the Carpathian Mountains]. [In: Ekologicheskie osnovy okhrany i ratsionalnogo ispolzovaniya khishchnykh mleko­pitayushchikh], Izdatelstvo Nauka, Moskva; 43-44. [In Russian]
• Kochetkov V. V. 1988. [Biology of the wolf in Verkhnevolzhe, a case of the Central-Forest State Reserve], Ph D thesis, Gosudarstvennyj Agropromyshlennyj Komitet SSSR, Moskva 1988: 1-20. [In Russian]
• Kochetkov V. V. and Sokolov A. A. 1979. [Wolf feeding in the Central-Forest State Reserve). [In: Ekologicheskie osnovy okhrany i ratsionalnogo ispolzovaniya khishchnykh mlekopitayushchikh], Izdatelstvo Nauka, Moskva: 112-114. [In Russian]
• Kotov V. A. 1969. A hard winter for wild ungulate animals in the Caucasian preserve. Byulleten Moskovskogo Obshchestva Ispytatelei Prirody, Otdel Biologicheski 74(1): 57-61. [In Russian with English summary]
• Kozlo P. G. 1975. I Wild boar], Izdatelstvo Uradzhai, Minsk: 1-223. [In Russian]
• Krasiński Z, A. 1967. Free living European bisons. Acta Theriologica 12: 391-405.
• Krasiński Z. A. 1978, Dynamics and structure of the European bison population in the Białowieża primeval forest. Acta Theriologica 23: 3-48,
• Krasiński Z. A. 1983. The first 10 years of free-living bison in the Białowieża Primeval Forest 1952-1961. Parki narodowe i Rezerwaty przyrody 4(2): 39-50. [In Polish with English summary]
• Krasiński Z. A., Bunevich A. N. and Krasińska M. 1994, Characteristics of lowland European bison populations in Polish and Belarussian parts of Białowieża Forest. Parki narodowe i Rezerwaty przyrody 4: 25-68. [In Polish with English summary]
• Kuchmel S. V. 1994. [An assesment of wolf impact on ungulate populations]. Materiały 7 Zooto- gicheskoi Konferentsii "Problemy izucheniya, sokhraneniya i ispolzovaniya biologicheskogo raznoobraziya zhivotnogo mira", Minsk 1994. Izdatelstvo Navuka i Technika, Minsk: 256-257. [In Russian]
• Kudaktin A. N. 1978. Selective wolf predation on ungulates in the Caucasian nature reserve. Byulleten Moskovskogo Obshchestva Ispytatelei Prirody, Otdel Biologicheski 83(3): 19-28. [In Russian with English summary]
• Kudaktin A. N. 1982, The hunting habits of wolves in western Caucasia. Byulleten Moskovskogo Obshchestva Ispytatelei Prirody, Otdel Biologicheski 87(1): 24-30. [In Russian with English summary]
• Kudaktin A. N. 1986. [Influence of the wolf on ungulates in the Caucasian reserve!. [In: Rol krupnykh khishchnikov v biocenozakh zapovednikov. Sbornik nauchnykh trudov CNIL Glavokhoty RSFSR. A. K. Fedosenko, ed|. Moskva: 21-35, [In Russian]
• Kuzmin I. F, and Khakhin G. V. 1980. [Impact of wolves on ungulates in the middle of the European part of USSR]. [In: Kopytnye fauny SSSRI. Izdatelstvo Nauka, Moskva: 171-172. [In Russian]
• Larsen D. G., Gauthier D. A. and Markel R. L. 1989. Causes and rate of moose mortality in southwest Yukon. Journal of Wildlife Management 53: 548-557,
• Lavov M, A. 1972. [Impact of wolves and lynx on wild ungulates in western Siberia], Materiały k nauchnoi konferentsii, posvyashchennoi 50-letiyu Instituta. VNIIOZ Kirov: 112-115. [In Russian]
• Lebedeva L. S. 1956. [Ekological characteristics of the wild boar in Białowieża Primeval Forest], Uchenye Zapiski Moskovskogo Gorodskogo Pedagogicheskogo Instituta im. V. P. Potemkina 61: 105-271. [In Russianl
• Lehmann E. von and Sagesser H. 1986. Capreolus capreoius Linnaeus, 1758 - Reh. [In: Handbuch der Saugetiere Europas. Band 2/II. J. Niethammer and F. Krapp, eds]. Aula-Verlag, Wiesbaden: 233-268.
• Lequette B., Houard T., Del Corso C., Mari F. and Boscagli G, 1994. [Wolf return to the Marittime Alps: Preliminary data]. Abstracts of 1st Italian Theriological Congress, Pisa, October 1994: 87. [In Italian]
• Levins R. 1968. Evolution in changing environment. Princeton University Press, Princeton: 1-120.
• Litvinov V. P. 1981, The wolf (Canis lupus) and wild boar (Sus scrofa) in the Kyzyl-Agach State Reservation, Zoologicheski'i Zhurnal 60: 1588-1591. [In Russian with English summary]
• Litvinov V. P., Litvinov V. F. and Tikhanskii A. D. 1981. [Feeding of wolves in Berezinskii and Pripyatskii* Reserves], Zapovedniki Belorussii 5: 98-102. [In Russian]
• Magalhaes C. Paixao de and Fonseca F. P. 1982. The wolf in Braganca country. Impact on cattle and game. Transactions of 14th IUGB Congress, Dublin 1979: 281-286.
• Makridin V. P. 1962. The wolf in the Yamal north. Zoologicheski'i Zhurnal 41: 1413-1417. [In Russian with English summary]
• Makridin V. P. 1979. [Polar wolves and their impact of reindeer breeding and wild ungulates). [In: Ekologicheskie osnovy okhrany i ratsionalnogo ispolzovaniya khishchnykh mlekopitayushchikhl. Izdatelstvo Nauka, Moskva: 122-125. [In Russian]
• Matteucci C. 1992. Preliminary data on the ecology of a wolf Canis lupus L. population in northern Italy. [In: Global trends in wildlife management. B. Bobek, K. Perzanowski and W. Regelin, eds|. Transactions of 18th IUGB Congress, Krakow 1987. Swiat Press, Krakow-Warszawa, Vol. 2: 367-370.
• Mattioli L., Apollonio M., Mazzarone V. and Centofanti E. 1995. Wolf food habits and wild ungulate availability in the Foreste Casentinesi National Park (Italy). Acta Theriologica 40: 389-404.
• Mech L. D, 1966. The wolves of Isle Royale. Fauna of the National Parks of the United States, Fauna Series 7: 1-210.
• Mech L. D. 1970. The wolf: the ecology and behavior of an endangered species. Doubleday, New York: 1-289.
• Mech L. D. and Frenzel L. D., Jr 1971. An analysis of age, sex, and condition of deer killed by wolves in northeastern Minnesota. USDA Forest Service Research Paper NC-52: 35-50.
• Mech L. D., Frenzel L. D., Jr and Karns P. D. 1971. The effect of snow conditions on the vulnerability of white-tailed deer to wolf predation. USDA Forest Service Research Paper NC-52: 51-59.
• Mech L. D. and Karns P. D. 1978, Role of the wolf in a deer decline in the Superior National Forest. USDA Forest Service Research Paper NC-148: 1-23.
• Mech L. D., McRoberts R. E., Peterson R. O. and Page R. E, 1987. Relationship of deer and moose populations to previous winters' snow. Journal of Animal Ecology 56: 615-627.
• Meriggi A., Rosa P., Brangi A. and Matteucci C. 1991. Habitat use and diet of the wolf in northern Italy. Acta Theriologica 36: 141-151.
• Messier F. 1991. The significance of limiting and regulating factors on the demography of moose and white-tailed deer. Journal of Animal Ecology 60: 377-393.
• Messier F. 1994. Ungulate population models with predation: a case study with the North American moose. Ecology 75: 478-488.
• Messier F. and Crete M. 1985. Moose-wolf dynamics and the natural regulation of moose populations. Oecologia 65: 503-512.
• Mityk J. 1978. [Physical geography of the parts of the world]. PWN-Polish Scientific Publisher, Warszawa: 1-503. ¡In Polish]
• Murie A. 1944, The wolves of Mt, McKinley. U. S. Dept. Interior, U.S. National Park Service, Fauna Ser. No 5: 1-238.
• Nasimovicb A. A. 1955. The role of the regime of snow cover in the life of ungulates in the USSR. Izdatelstvo Akademii Nauk SSSR, Moskva: 1-403. ]In Russian]
• Naumov N. P. 1967. [The wolf], [In: Mlekopitayushchie Sovetskogo Soyuza, vol. 2. V. G. Heptner and N. P. Naumov, eds], Izdatelstvo Vysshaya shkola, Moskva: 123-193. [In Russian!
• Nelson M. E. and Mech L, D. 1986. Relationship between snow depth and gray wolf predation on white-tailed deer. Journal of Wildlife Management 50: 471—474.Nesterenko V. V. 1988. [The role of wolves in ecosystems of nature reserves]. [In; Populyatsionnye issledovaniya zhivotnykh v zapovednikakh. V. E. Sokolov, ed], Izdatelstvo Nauka, Moskva: 139-144. [In Russian]
• Nieminen M. and Leppaluoto J. 1988. Prédation in the reindeer husbandry area in Finland during 1976-1986. Rangifer 8: 25-34.
• Novikov G. A. 1956. [Mammalian predators of USSR fauna], Izdatelstvo AN SSSR, Moskva­-Leningrad: 42-52. [In Russian]
• Novikov G. A. and Ivanov P. D. 1970. The appearance of elks CAlces alee s L.) in the precincts of Leningrad. Byulleten Moskovskogo Obshchestva Ispytatelei Prirody, Otdel Biologicheski 75(6): 27-35. [In Russian with English summary]
• Nowak E. 1971. The range expansion of animals and its causes. Instytut Ekologii PAN, Zeszyty Naukowe No 3: 1-255. [In Polish with English summary]
• Nygren K. 1986. Alces alces (Linnaeus, 1758) - Elch. [In: Handbuch der Sàugetiere Europas. Band 2/II. J. Niethammer and F. Krapp, eds]. Aula-Verlag, Wiesbaden: 173-197.
• Okarma H. 1984, The physical condition of red deer falling prey to the wolf and lynx and harvested in the Carpathian Mountains. Acta Theriologica 29: 283-290.
• Okarma H. 1991. Marrow fat content, sex and age of red deer killed by wolves in winter in the Carpathian Mountains. Holarctic ecology 14: 169-172.
• Okarma H. 1992a. [The wolf - monograph of a species]. Białowieża; 1-168. [In Polish with English chapter]
• Okarma H, 1992b. Status, distribution and numbers of lynx in Poland. Proceedings of a seminar on the situation, conservation needs and réintroduction of lynx in Europe. Neuchâtel, Switzerland, October 1990. Council of Europe Press, Environmental Encounters No 11: 23-25.
• Okarma H. 1993a. Status and management of the wolf in Poland. Biological Conservation 66: 153-158.
• Okarma H. 1993b. Protection and management of large carnivores (wolf, lynx) in the Białowieża Primeval Forest, Poland. Proceedings of the seminar on the management of small populations of threatened mammals, Sofia, Bulgaria, October 1993. Council of Europe Press, Environmental Encounters No 17: 56-59.
• Okarma H., Jędrzejewska B. and Jędrzejewski W. 1995b. Białowieża Primeval Forest: habitat and wildlife management. Proceedings of a conference "Biodiversity conservation in transboundary protected areas", Bieszczady-Tatry 1994. US National Academy of Sciences, Washington, (in press)
• Okarma H., Jędrzejewska B., Jędrzejewski W., Mitkowski L. and Krasiński Z. 1995a. Roles of hunting, prédation, and few other factors in the winter mortality of ungulates in Białowieża Primeval Forest, Poland. Acta Theriologica 40: 197-217.
• Okarma H., Milkowski L., Jędrzejewska B. and Jędrzejewski W. 1993. Importance of wolf prédation among several factors of ungulate mortality in the exploited part of Białowieża Primeval Forest, Poland. International Symposium on the Wolf of the Wolf Specialist Group SSC-IUCN. Leon 1993, Spain: 51-52.
• Papageorgiou N., Vlachos Ch., Sfougaris A. and Tsachalidis S. 1994, Status and diet of wolves in Greece. Acta Theriologica 39: 411-416.
• Paquet P. C. 1989. Behavioral ecology of sympatric wolves (Canis lupus) and coyotes (C. latrans) in Riding Mountain National Park. Ph D thesis, University of Alberta: 1-198.
• Paquet P. C. 1992, Prey use strategies of sympatric wolves and coyotes in Riding Mountain National Park, Manitoba. Journal of Mammalogy 73: 337-343.
• Paquet P. C. 1993. Summary reference document - ecological studies of recolonizing wolves in the Central Canadian Rocky Mountains. Parks Canada, Banff National Park Warden Service, Banff, Alberta: 1-215.
• Patalano M. and Lovari S. 1993. Food habits and trophic niche overlap of the wolf Canis lupus, L. 1758 and the red fox Vulpes vulpes (L. 1758) in a mediterranean mountain area. Revue d'Ecologie (Terre et Vie) 48: 279-294.
• Pereira M., Fonseca F, and Magalhaes C. 1985. Wolf ecology in Portugal. Actas 1 Simposio de Predadores, Lisboa: 122-167. Peterson R. O. and Allen D. L. 1974. Snow conditions as a parameter in moose-wolf relationship.
• Naturaliste Canadien (Que.) 101: 481-492.
• Peterson R. O. and Page R. E. 1983. Wolf-moose fluctuations at Isle Royale National Park, Michigan, USA. Acta Zoologica Fennica 174: 251-253.
• Peterson R. O., Woolington J. D. and Bailey T. N. 1984. Wolves of the Kenai Peninsula, Alaska. Wildlife Monographs 88: 1-52.
• Pielowski Z. 1977. Das Rehwild - Wild der Zunkunft in der Agrarlandschaft. Beiträge Jagd- und Wildforschung 10: 193-200.
• Potvin F., Jolicoeur H. and Huot J. 1988. Wolf diet and prey selectivity during two periods for deer in Quebec: decline versus expansion. Canadian Journal of Zoology 66: 1274-1279.
• Promberger Ch. and Schröder W. (eds). 1993. Wolves in Europe - status and perspectives. Proceedings of the workshop "Wolves in Europe - current status and prospects", April 1992, Oberammergau. Munich Wildlife Society, Ettal: 1-136. Pucek Z. 1986. Bison bonasus (Linnaeus, 1758) - Wisent. [In: Handbuch der Säugetiere Europas. Band 2/11. J. Niethammer and F. Krapp, eds]. Aula-Verlag, Wiesbaden: 279-315.
• Pucek Z. 1991. History of the European bison and problems of its protection and management. [In: Global trends in wildlife management. B. Bobek, K, Perzanowski and W. Regelin, eds]. Trans­actions of 18th IUGB Congress, Cracow 1987. Świat Press, Kraków-Warszawa: 19-39.
• Pucek Z. 1994. [Progress in and threats to the European bison restitution]. Kosmos 43(1): 147-169, [in Polish with English summary] Pucek Z., Bobek B., Łabudzki L., Milkowski L., Morow K. and Tomek A. 1975. Estimates of density
• and number of ungulates. Polish Ecological Studies 1: 121-136.
• Pucek Z., Jędrzejewski W., Jędrzejewska B. and Pucek M. 1993. Rodent population dynamics in a primeval deciduous forest (Białowieża National Park) in relation to weather, seed crop, and predation. Acta Theriologica 38: 199-232. Pulliainen E. 1965. Studies on the wolf (Canis lupus L.) in Finland. Annales Zoologici Fennici 2: 215-259.
• Reig S., Cuesta L. and Palacios F. 1985. The impact of human activities on the food habits of red fox and wolf in Old Castille, Spain. Revue d'Ecologie (Terre et Vie) 40: 151-155.
• Rukovski'f N. N. and Kupriyanov A. G. 1972. Some peculiarities of distribution and ecology of Canis lupus in the Onega peninsula. Zoologicheskii Zhurnal 51: 1593-1596. [In Russian with English summary]
• Rusakov O. S. 1979a. [Carnivores as a factor in dynamics of ungulate numbers in the northwestern USSR], [ln: Ekologicheskie osnovy okhrany i ratsionalnogo ispolzovaniya khishchnykh mleko- pitayushchikh], Izdatelstvo Nauka, Moskva: 61-62. [In Russian]
• Rusakov O. S. 1979b. [Peculiarities of wolf feeding in the northern part of Pskovskaya oblast], [In: Ekologicheskie osnovy okhrany i ratsionalnogo ispolzovaniya khishchnykh mlekopitayushchikh], Izdatelstvo Nauka, Moskva: 132-134. [In Russian]
• Rusakov O. S. and Timofeeva E. T. 1984. IWild boar], Izdatelstvo Leningradskogo Universiteta, Leningrad: 1-204. |In Russian]
• Ryabov L. S. 1974. The attitude of wolves towards domestic animals and wild ungulates in the region of the Khopersk nature reserve. Byulleten Moskovskogo Obshchestva Ispytatelei Prirody, Otdel Biologicheski 79(3): 6-15, [In Russian with English summary]
• Ryabov L. S. 1979. Canis lupus in the east of the Voronezh district. Zoologicheskii" Zhurnal 58: 564-569. [In Russian with English summary]
• Ryabov L. S, 1987. On the wolf synanthropy in the central Black Earth Belt. Byulleten Moskovskogo Obshchestva Ispytatelei Prirody, Otdel Biologicheski 92(1): 3-12. [In Russian with English summary]
• Ryabov L. S. 1993. [Wolves of the Black Earth Belt], Izdatelstvo Voronezhskogo Universiteta, Voronezh: 1-167. [In Russian]
• Ryabov L. S., Likhackii Yu, P. and Nikitin N. M. 1992, [The wolf in the Voronezh Nature Reserve (feeding interactions)!. [In: Krupnye khishchniki. Sbornik nauchnykh trudov Glavokhoty RSFSR], Moskva: 42-47. [In Russian] Ryabov L. S., Likhackiï Yu. P. and Nikitin N. M. 1994. [Interactions of wolves (Canis lupus L.) of the Voronezh Reserve with wild and domestic ungulates], Ekologiya 94(1): 53-59. [In Russian]
• Rykovskiï A. S. 1980. [Observations on peculiarities of wolf behaviour in Belarus). |In; Behaviour of the wolf. D. I. Bibikov, ed], AN USSR, Moskva: 103-110. [In Russian]
• Sablina T. B. 1955. [Ungulates of the Białowieża Forest]. Trudy Instituta Morfologii Zhivotnykh AN SSSR No 15. Izdatelstvo Akademii Nauk SSSR, Moskva: 1-191. [In Russian]
• Sacchi O., Alieri F. and Cesaris C. 1994. [Wolf Cants lupus diet in an area with high availability of domestic ungulates]. Abstracts of 1st Italian Theriological Congress, Pisa, October 1994: 150. [In Italian]
• Salvador A. and Abad P. L. 1987. Food habits of a wolf population (Canis lupus) in León province, Spain. Mammalia 51: 45-52,
• Seip D. R. 1991. Prédation and caribou populations. Rangifer, Special Issue No 7: 46-52.
• Seip D. R. 1992. Factors limiting woodland caribou populations and their interrelationships with wolves and moose in southeastern British Columbia. Canadian Journal of Zoology 70: 1494-1503.
• Sinclair A. R. E, 1989. Population regulation in animals. [In: Ecological concepts: the contribution of ecology to an understanding of the natural world. J. M. Cherrett, ed]. Blackwell Scientific Publications, Oxford: 197-241.
• Skogland T. 1991. What are the effects of predators on large ungulate populations? Oikos 61: 401-411.
• Sludskiï A. A. 1963. [Mass death of ungulates in steppes and deserts of Eurasia], Trudy Instituta Zoologii AN Kazakhskoi SSR 20: 5-88. [ln Russian]
• Suvorov A. P. and Lisenko E. V. 1988. [Some peculiarities of wolf ecology in spring-summer period]. Sbornik Nauchnykh Trudov, VASKHNIL, Sibirskoe otdelenie, Novosibirsk: 45-50. [In Russian]
• Szymczyk W. 1973. Occurrence of moose in Europe in Holocene. Przegląd zoologiczny 17(1): 89-93. [in Polish with English summary]
• Śmietana W. and Klimek A. 1993. Diet of wolves in the Bieszczady Mountains, Poland. Acta Theriologica 38: 245-251.
• Telfer E. S. and Kelsall J. P. 1979. Studies of morphological parameters affecting ungulate locomotion in snow. Canadian Journal ofZoology 57: 2153-2159.
• Telfer E. S. and Kelsall J. P. 1984. Adaptation of some large North American mammals for survival in snow. Ecology 65: 1828-1834.
• Tellerta J. C. and Sâez-Royuela C. 1989. [Ecology one Iberian population of wolves (.Canis lupus)]. Doriana Acta Vertebrata 16: 105-122. [In Spanish]
• Van Ballenberghe V, 1987. Effects of prédation on moose numbers: a review of recent North American studies. Swedish Wildlife Research Suppl. 1: 431-460.
• Van Camp J. 1975. Snow conditions and the winter feeding behavior of Bison bison in Elk Island National Park. Canadian Wildlife Service Report; 1-91,
• Van Haaften J. L., Peireira M. R. and Fonseca F. P. 1983. A wolf study in Portugal. Proceedings of 16th IUGB Congress, Strbske Pteso 1983, Czechoslovakia: 550-554.
• Vyrypaev V. A. 1980. On hunting behaviour of the wolf in Tien-shan. Zoologicheskiï Zhurnal 59:1870-1874. [In Russian with English summary]
• Wabbakken P. 1992. Wolves in Sweden and Norway. |In: Wolves in Europe. Ch. Promberger and W. Schroder, eds]. Proceedings of the workshop "Wolves in Europe - current status and prospects", Oberammergau 1992. Munich Wildlife Society, Ettal: 9-14.
• Weaver J. L, 1994. Ecology of wolf prédation amidst high ungulate diversity in Jasper National Park, Alberta. Ph D thesis, University of Montana, Missoula: 1-166.Zalozny D. V. 1980. [Some peculiarities of wolf ecology in Gurievskoi oblasti], [In: Behaviour of the wolf. D. I. Bibikov, ed). AN USSR, Moskva: 77-89. [ln Russian]
• Zavatskií B. P. 1986. [The role of wolves in biocenoses of Sayano-Shushenskii zapovednik], Sbornik nauchnykh trudov CNIL Glavokhoty RSFSR, Moskva: 35-54. [In Russian]
• Zheleznov N. 1992. Ecology of the grey wolf Canis lupus L. on Chukotka. [In: Global trends in wildlife management. B. Bobek, K. Perzanowski and W. Regelin, eds]. Transactions of 18th IUGB Congress, Krakow 1987. Świat Press, Krakow-Warszawa, Vol. 2: 381-384.
• Zheleznov N. K. 1994. [Data on wolf Canis lupus L. ecology in northeastern USSRJ. Sbornik nauchnykh trudov "Teriologicheskie issledovaniya v Yakutii", Sibirskoe otdelenie AN SSSR, Irkutsk: 28-50. [In Russian]
• Zhirnov L. V. 1982. [Brought back to life: ecology, conservation, and utilization of saiga antelope], Izdatelstvo Lesnaya promyshlennost, Moskva: 1-223. [In Russian)