Fauna (фауна). The present fauna of Ukraine began to develop in the late Eocene epoch, if not earlier. Only a few fossil remains of land animals of the Eocene and Paleocene epochs have been discovered in Ukraine. Among the mammals at the end of the Paleocene were the piglike Anthracotheriidae and the hornless rhinoceros Chilotherium; among the birds were the cormorant, sea gull, stork, wild duck, and owl. The rivers were inhabited by crocodiles and the seas by whales (zeuglodons) and many forms of mollusk and fish related to modern representatives of these groups.

In the Neogene period, when the climate was subtropical, the Hipparion fauna flourished on the extensive steppes. It was represented mostly by herbivores such as the Hipparion, the ancestor of our horse, rhinoceros (Dicerorhinus orientalis and D. etruscus), mastodon (Anancus, Mastodon), elephant (Archidiskodon meridionalis), giant deer (Eucladocerus pliotarandoides), archaic camel (Paracamelus), saber-toothed tiger (Machairodus), and beaver-like trogontherium (Trogontherium cuvieri). Beside them lived families and species that still exist (or existed until recent times) both inside Ukraine—for example, the desman (Desmana moschata), pika (Ochotona), hedgehog, fox, brown bear, rabbit, great bustard, chicken, partridge—or outside Ukraine—for example, the monkey, giraffe, porcupine, ostrich, and marabou. In the late Pliocene most of the present invertebrate and vertebrate species were to be found in Ukraine.

During the several ice ages in the Pleistocene epoch much of Ukraine's territory was covered with glaciers. In the tundra bordering them lived species adapted to the cold climate. In the interglacial periods the forest fauna returned to the lands from which the ice sheets retreated. After each glaciation the Tertiary species diminished greatly: some died out and others (eg, the monkey, ostrich, and giraffe) migrated south, but many withstood the existing conditions. Thus, in the period of maximum glaciation, there was a characteristic mixed mammoth fauna in Ukraine. Beside its typical representatives—the mammoth, woolly rhinoceros, cave-dwelling bear and lion, spotted hyena—lived such arctic animals as the musk-ox (Ovibos moschatus), arctic fox (Alopex lagopus), and true lemming (Lemmus lemmus); such steppe animals as the saiga (Saiga tatarica), bobak marmot (Marmota bobak), ground squirrel or suslik (Citellus); and such forest animals as the beaver. Besides these, wild horses, bisons, deer, foxes, wolves, brown bears, hares, wild ducks, geese, woodcocks, and many other extant species inhabited Ukraine.

In the Middle Holocene the climate became similar to what it is today. Many species of the earlier period gradually perished (mammoth, rhinoceros); some species moved farther north (musk-ox). In the relatively short dry period that succeeded the retreat of the glaciers, the steppe species advanced far north.

Human activities—hunting, herding, agriculture—contributed to the impoverishment of the mammoth fauna. The cultivation of the steppes, destruction of the forests, and draining of the marshes contracted the habitat of many species and contributed to their migration from Ukraine or to their extinction. Within historical times reindeer ceased visiting Ukraine during their winter migration. In the 16th century the Asiatic wild ass (Equus hemionus) and in the 17th century the aurochs and bison became extinct. In the 19th century the wolverine and flying squirrel disappeared from the forest zone; the tarpan, saiga, and ruddy ground squirrel disappeared from the steppe and forest-steppe zone; the alpine marmot (Marmota marmota), willow grouse (Lagopus lagopus), and white hare (Lepus timidus) vanished from the Carpathian Mountains; and the wild boar disappeared from the Crimean Mountains. Some species have become rare: the little bustard (Otis tetrax), great bustard (Otis tarda), ruddy sheldrake (Tadorna ferruginea), and swan. Some valuable fish species—sturgeon and eel—have disappeared from the rivers. Some species, for example, the suslik, have spread with the cultivation of the steppes; these include pests (mouse, hamster). As well, humans have added to the variety of fauna by acclimatizing such new species as the nutria, raccoon, silkworm, and others.

Zoogeographically, the fauna of Ukraine belongs to the Euro-Siberian zone of the Palaearctic subregion of the Holarctic region. Only the Crimean Mountains and the southwest part of Caucasia belong to the Mediterranean subzone. Because of Ukraine's border locations and the absence of natural barriers, its fauna is intermediate between the fauna of Europe and Central Asia, and between the fauna of the forest belt and the steppe belt and that of the subtropics. The western boundaries of the habitats of many eastern species run through Ukraine; for example, of the yellow suslik (Citellus fulvus) and pygmy suslik (Citellus pygmaeus). The eastern and northern limits of the habitats of many western European and Mediterranean species are found in Ukraine: Bechstein's bat (Myotis bechsteini), the wildcat (Felis silvestris), the eel (Anguilla anguilla), the Baltic sturgeon (Acipenser sturio), and others. The Dnipro River is an important natural barrier to the east-west distribution of animals. The habitat of many northern species—elk, lynx, brown bear, white hare, capercaillie, black grouse, hazel hen, and others—extends as far south as Ukraine. The northern boundary of many southern species, including the lesser mouse-eared bat (Myotis oxygnatus), long-winged bat (Miniopterus schreibersi), and many insects, lies in Ukraine. There are few endemic species in Ukraine.

In general, there are about 28,000 species on the territory of Ukraine, among them more than 690 species of vertebrates (101 mammal, 350 bird, 21 reptile, 19 amphibian, and over 200 fish, 110 of which are freshwater species), over 1,500 protozoan species, over 700 species of worm, 400 crustacean species, 330 mollusk species, about 3,300 arachnid species, 20,000 other species of insects, and about 1,080 others.

The zoogeographical regions of Ukraine coincide with the natural biogeographical zones: Polisia, forest-steppe, steppe, semidesert, littoral, mountain (Carpathian Mountains, Crimean Mountains, and western Caucasia), and marine (the Black Sea and the Sea of Azov).

The fauna of Polisia is known for its variety of forest and swamp species. In the past Polisia had large numbers of elk, lynx, bear, capercaillie, black grouse, and hazel hen, which today can be found only in the remotest areas. Wild boars, wolves, foxes, roe deer, forest martens, and other species are quite common. Valuable fur-bearing animals such as the beaver (on reservations along the Teteriv River, the Horyn River, the Prypiat River, the Dnipro River, and the Desna River), mink, otter, and ermine have survived here. The red-backed mouse (Clethrionomys glareolus), field vole (Microtus agrestis), striped field mouse (Apodemus agrarius), and red-toothed shrew (Sorex) live in the forests, swamps, and meadows. Untilled, open fields are inhabited by the common mole, gray common vole (Microtus arvalis), wood mouse, and other species. Among the birds are found the capercaillie, black grouse, hazel hen, tit, rock dove, spotted eagle (Aquila clanga), short-toed eagle (Circaëtus ferox), and tree pipit (Anthus trivialis). Along rivers and swamps wild ducks, snipes, bald coots, black storks (Ciconia nigra), and common cranes (Grus grus) thrive. At crystalline cliffs and outcrops along the rivers the bee-eater (Merops apiaster) and rock thrush (Monticola saxatilis) can be found. Among the reptiles, the forest grass snake, mud tortoise, asp, and such lizards as blindworm (Anguis fragilis), fast lizard (Lacerta agilis), and viviparous lizard (Lacerta vivipara) are quite common. Amphibians such as the newt, common fire-bellied toad (Bombina bombina), bullfrog, and frog are widely distributed. Rivers and lakes sustain such fish as the carp, tench, and pike. Various species of beetles, bugs, mosquitoes, cicadas, and other insects live here.

The forest-steppe is a transitional zone in which steppe species live side by side with forest species. Certain forest species that inhabit wooded river banks penetrate far into the steppe, for example, the squirrel and the pine marten (Martes martes). Among the forest species that inhabit the forest-steppe are the roe deer, hazel mouse (Muscardinus avellanarius), forest dormouse (Dyromys nitedula), gray dormouse (Glis glis), red-backed mouse, and field vole. The steppe species that can be found deep in the forest-steppe are the steppe polecat (Mustela eversmanni), birch mouse (Sicista subtilis), mole rat (Spalax), and gray hamster (Cricetulus migratorius). The Left-Bank forest-steppe provides a habitat for the steppe lemming (Lagurus lagurus) and the large jerboa (Alactaga jaculus). The spotted suslik (Citellus suslica) is common in the entire forest-steppe, while the European suslik (Citellus citellus) lives in the southwestern section. The black-bellied hamster (Cricetus cricetus) and gray vole are fairly widespread. The red kite (Milvus milvus), stock dove (Columba oenas), ringdove (Columba palumbus), common turtledove (Streptopelia turtur), green woodpecker (Picus viridis), thrush nightingale (Luscinia luscinia), and other species inhabit oak forests. The common quail and partridge are widely distributed. Since the Dnipro River is a migration route and the forests, ponds, and fields of the forest-steppe are stations for migratory birds, wild geese, cranes, and various species of wild duck can be found here. Among reptiles, the Aesculapian snake, viper, common and tree snakes, mud tortoise, and lizard are the most common. The amphibians are represented by the pond frog (Rana esculenta), common newt, crested newt (Triturus cristatus), and others. Among the insects of the forest-steppe are such pests as the owlet moth (Agrotis segetum), sugar-beet weevil (Bothynoderes punctiventris), and lamellicorn (Lethrus apterus).

The Tysa Lowland of Transcarpathia is a unique region of the forest-steppe. Here are found species that are rare or non-existent in other regions of the forest-steppe: the greater horshoe bat (Rhinolophus ferrum-equinum), Ikonnikov bat (Myotis ikonnicovi), lesser mouse-eared bat, and long-winged bat. There are many Mediterranean species among the insects.

In the steppe the most typical species are rodents that are adapted to open plains and an arid climate. As in the forest-steppe, so in the steppe the western, central, and eastern sections differ from one another. The mammals that are typical of the steppe are the large jerboa, pygmy suslik, steppe vole, mole-vole (Ellobius talpinus), and three-toed sand jerboa (Scirtopoda tellum), all of which live east of the Dnipro River. The western steppe is inhabited by the spotted suslik, mole rat, and steppe polecat. The lesser mole rat (Spalax leucodon) is found in the southwest, while the marbled polecat (Vormela peregusna) is limited to the southern part of the steppe and is retreating gradually eastward into the Asiatic steppes. The bobak marmot, which formerly was common in the west, still survives on reservations along the Donets River. In the eastern steppe one can still encounter the corsac fox (Vulpes corsak) and long-eared hedgehog (Hemiechinus auritus). The birds that are typical of the steppe are rare today: the great bustard, calandra lark (Melanocorypha calandra), demoiselle crane (Anthropoides virgo), black-winged pratincole (Glareola nordmanni), tawny eagle (Aquila rapax), and little bustard. The most common reptiles are the yellow-bellied coluber (Coluber jugularis), steppe viper (Vipera renardi), four-striped snake (Elaphe quatuorlineata), and green lizard (Lacerta viridis). The steppe lizard (Eremias arguta) is less common. A great variety of insects, including Italian and Asiatic locusts, inhabit the steppe.

In the east the steppe changes into semidesert, where desert species from the Aral-Caspian deserts exist side by side with steppe species.

The steppe north of the Black Sea and the Sea of Azov belongs to the littoral region, in which water fowl and aquatic animals are prominent. Many bird species are found here: the herring gull (Larus argentatus), tern (Sterna), Kentish plover (Charadrius alexandrinus), avocet (Recurvirostra avosetta), and spoonbill (Platalea leucorodia). The Old World white pelican (Pelecanus onocrotalus) and the eastern glossy ibis (Plegadis falcinellus) nest at the mouth of the Danube River. Migratory birds pass through or winter in this region: in spring and fall innumerable geese and wild ducks come here. The muskrat, European otter, mink, ondatra (acclimatized), goose, wild duck, occasional swan, gadflies, gnats, and mosquitoes inhabit the river floodplains. In the limans and deltas is found a mixture of sea and freshwater fish species. Mullet, medusa, flounder, and other sea species live alongside migratory species.

In the Black Sea dolphins and the rare white-bellied seals (Monachus monachus) are found. The fish of the Sea of Azov and the coastal waters of the Black Sea are very similar. Yet there are some local species: the Azov herring, Azov anchovy, Azov percarina, sardelle, great plaice, sprats, and gobies are found in the Sea of Azov, and the Black Sea trout, Black Sea herring, seahorse, tunny, mackerel, sturgeon, and others are found in the Black Sea.

The Carpathian Mountains contain mostly forest fauna. The vertical distribution of animals is to a great extent related to the vegetation. High mountain species—the snow vole (Microtus nivalis), alpine shrew (Sorex alpinus), water pipit (Anthus spinoletta), alpine accentor (Prunella collaris), and certain insects—are confined to the subalpine and alpine zone of the Carpathians. Some taiga species are found in the mountain forest zone: lynx, capercaillie, hazel hen, black grouse, and others. But most of the fauna consists of Central European forest species, which appear in all regions of the Carpathians: the now rare wildcat and brown bear, and roe deer, forest marten, ermine, Carpathian squirrel, wild boar, wolf, fox, golden eagle, hawk, owl, rock pipit, woodcock, and others. There are many species of reptiles and amphibia (Carpathian newt, spotty salamander, smooth snake, etc) and many European mountain species of insects, mollusks, and other invertebrates.

The fauna of the Crimean Mountains, especially that of the southern coast of the Crimea, is Mediterranean. The fauna is insular in character: species that are typical of the forest or steppe belts are absent. Instead, there are many endemic subspecies. The forests are inhabited by the Crimean red deer and roe deer, Crimean mountain fox, badger, bats, and other species. The squirrel and mouflon are acclimatized. Of the bird species, we find the griffon vulture, tawny owl, Crimean jay, and tomtit. The Crimean scorpion (Euscorpius tauricus) and spider solifug (Galeodes araneides) represent the arachnids, and the scolopendra (Scolopendra cingulata) represents the centipedes. There are many Mediterranean species, which arrived in the Paleogene when the Crimea was connected to the Balkans and western Caucasia, for example, the Crimean gecko (Gymnodactylus kotschyi Danilevskii), Crimean cicada (Cicada taurica), Crimean beetle (Procerus tauricus), and Crimean mantis (Ameles taurica). Many Mediterranean species of mollusk are found here. Of the pests introduced from the outside, the phylloxera is harmful. Mediterranean fauna appears also on the Black Sea coast of western Caucasia.

The ichthyofauna of Ukrainian rivers consists mainly of members of the Cyprinidae family. The main European watershed between the Baltic Sea and the Black Sea limits the distribution of certain fish species. In the rivers of Ukraine that flow into the Baltic are found the eel and Baltic sturgeon. The rivers flowing into the Black Sea contain gobies (Gobiidae), vyrezub (Rutilus frisi), sterlet (Acipenser ruthenus), other sturgeons, and the Ukrainian lamprey (Lampetra mariae), along with other species. In the Tysa River, the Cheremosh River, and mountain tributaries of the Danube River are found the Danube salmon (Hucho hucho), striped ruff (Acerina schraetser), and little zingle (Aspro zingel). Mountain streams contain trout and grayling (Thymallus thymallus).

To protect what remains of the rich flora and fauna of Ukraine from extinction, a network of nature preserves (zapovidnyky) that provide full protection to wildlife have been set up, including four biosphere reserves (biosferni zapovidnyky) and seventeen national parks. There are also 2709 wildlife refuges (zakaznyky) that provide partial protection. Many species—the beaver, lynx, muskrat, bobak marmot, elk, great bustard, and others—are under special protection.

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Edvard Zharsky

[This article was updated in 2008.]

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