For centuries, man has engaged in incessant and persistent struggle against the wolf, a highly organized, large predator preying on wild and domestic ungulates. This strife has resulted in wolf extermination in many Western European countries and in most of the United States. The existence of the species is endangered even in Europe's most protected mountain areas -- the Pyrenees, the Apennines, the Balkans, and the Carpathians.
Diminished wolf range and numbers appear to be inevitable with increasing human transformation of the landscape, population growth, and urbanization. This process is natural and in some ways justified, since in developed areas the wolf's regular functions are taken over by man. Game management evidence from some European countries shows that high numbers of wild ungulates and their good physical condition have been achieved in the absence of the wolf. There is also no doubt that farmers, cattle breeders, and hunters would be opposed to paying a tribute to the wolf by reintroducing it into its original range.
Recently, ecologists have given strong evidence of the wolf's positive role in maintaining equilibrium in ecosystems little disturbed by man. This role consists of control of ungulate numbers and maintenance of population fitness. But the cedrease of wild nature offers little chance of wolf preservation in most of its range.
In the USSR, the wolf is a common and widely distributed beast badly affecting cattle breeding and game management....In the Soviet Union, seven wolf subspecies have been distinguished, distinct as to size and coloration. Their ranges and morphological features call for further investigation.
The wolf is not an endangered species in the USSR. That is why control is still an urgent problem where the wolf [competes with humans], for example, in areas of extensive cattle breeding and management of ecologically important wild ungulates.
In the USSR, the current century has seen two peaks of wolf numbers, both of which followed years of war and economic difficulties when wolves were not controlled. At the end of the 1940s, the number of wolves in the country was roughly estimated at 200,000....Active control, with annual kills of 40,000 to 50,000 head, reduced the wolf population considerably by the early 1960s. The annual kill number was then fixed at 15,000.
However, the early 1970s saw a new rise in wolf numbers. In the Ukraine, in Byelorussia, and in central Russia, wolf populations increased twofold over a five-year span. The reasons were slackening control efforts, reduced wolf density, and popularization of natural conservation. In recent years, hunting agencies and societies have contributed considerably to increased efforts at controlling wolves. Conducive to control were such measures as an increase in the premium for killed wolves: for an adult it is now up to 100 rubles and for a young [wolf], up to 50 rubles. Wolves are killed throughout the year by all means, including shooting from helicopters, vehicles, and aerosledges.
Undoubtedly, wolf numbers will again drop below the early 1970s level; that is, there will remain fewer than 50,000 wolves in the USSR. Their range in the European USSR is likely to be reduced to half of what it once was. But wolf populations in the mountains of the Caucasus, Central Asia, and Siberia are not likely to decrease in the near future. These are sparsely populated areas, difficult of access, and no effective hunting techniques are available.
Wolf populations are most vulnerable in open landscapes -- tundra, steppe, and desert -- where hunters use aircraft. Wolf-control data for open landscapes indicate that the tundra canis lupus albus and desert C.l. desertorum subspecies are endangered. Preservation of genetic variety within any species being important, this can hardly be justified.
With a huge range and environmental diversity, wolf ecology in the USSR varies to a great extent geographically. First and foremost, this involves population densities and feeding. The wolf being a territorial animal...population density may exceed one individual per ten square miles only in such exceptional cases as in ungulate winter yards and in some reserves....About 100 wolves dwell in the Caucasian reserve, which has an area of 1,000 square miles. Territoriality limits wolf numbers in...reserves....
Wolves used to be abundant in the forest-steppe zone of central Russia, the Ukraine, Pvolzhie, and western Siberia, but systematic human pressure has considerable reduced these populations....In the tundra and desert, the wolf has always been scarce, the mean density not exceeding one to two individuals per 400 square miles. We have recorded wolf population densities of up to twenty individuals per 400 square miles in the range of the saiga antelope -- an animal that has recovered greatly from near extinction -- in the Aktyubinsk region of Kazakhstan.
The diet of the tundra wolf is wild and domestic caribou, and in the eastern part of the region, bighorn sheep as well. Taiga wolves feed mainly on elk. In Siberia and the Far East, the wolves' diet is supplemented buy deer, roe deer, and wild boar. When ungulates are scarce, blue hare is an important component of the wolf diet. In the western forest-steppe region, the wolf takes roe deer (the Baltic republics, the Ukraine), wild boar (Byelorussia), and deer. In the desert zone, the bulk of the wolf diet includes saiga and goitred gazelle, and in the river valleys, deer and wild boar. In mountain-forest ecosystems, the wolf consumes deer, wild boar, mountain goat (in the Caucasus); mountain sheep, roe deer, deer, wild boar, and mountain goat (in the mountains of Kazakhstan and Central Asia). The role of domestic ungulates, specifically sheep and goats, increases in the wolf diet where pasture livestock farming is well developed and wild ungulate resources are small.
....Studies of wolf relations with ungulate prey have been the focus of attention in the USSR. Naturally, a diversity of prey species and environmental conditions over cast areas has led to highly contradictory inferences. The wolf's role as a population quality improver is definitely not appreciated in areas where the predator under study lives on domestic animals. Many game management specialists regard the wolf as harmful....Ungulates acclimated in game preserves invariably lose their capacity to resist wolves. The selective role of the wolf...has been lost under such conditions.
Along with their American and Canadian colleagues, Soviet scientists have given conclusive evidence of the wolf's useful role for its prey populations in ecosystems slightly affected by man. In fact, studies in Crimean reserves have demonstrated a degeneration of red deer in an overdense population in the absence of wolves. Similarly, a sharp reduction of wolf numbers brought about a substantial increase in the number of sick caribous on the Taimyr. Similar evidence is provided by the Darwin reserve, where the elk population was healthy, its numbers increasing despite annual kills of about fifty animals by numerous wolves. Following the introduction of wolf control, the Darwin elk herd was reduced by bad skin diseases.
Observations in the Caucasian reserve revealed population growth in deer, wild boar, and mountain goat during a period of intensive wolf control, the number of wolves being reduced to between twenty-five and thirty by 1960. The number of ungulates exceeded 30,000, and winter feeding yards were destroyed under the impact of too many animals. From 1965 to 1971, few wolves were killed, and their population increased. Ungulate populations subsequently began to decrease. From 1971 to 1975, the predator-prey ratio was stabilized at 1 to 300. This allowed an average take of 4,800 pounds of prey per wolf per year. Both pastures and the physical condition of the deer have improved. In the area under study...deer six years of age and older (59.2 percent) and fawns (25.1 percent) were preferentially killed by the wolves. Comparison of evidence indicates that wolves have a less deleterious effect on ungulate reproduction than human hunting, which eliminates the most productive animals.
The wolf's epizootic role is not yet clearly understood. Many authors emphasize that wolves eliminate sick members, which makes the prey population healthier. A considerable body of evidence is available, Soviet sources included, on the wolf's pathogenic role, particularly in regulating helminthoses. Also imperfectly understood is the role of the wolf in retaining the rabies virus. The wolf carries and transmits this infection in nature; however, the virus can exist in the absence of the wolf. Foxes and raccoon dogs are prime carriers in some Western European countries. It should be taken into account that in the presence of the wolf, fox numbers are not high.
Wolves hunting saigas exhibit a pronounced prey selectivity. During the rut, for instance, wolves are known to take weakened and exhausted males most frequently. Similarly, during fall saiga migrations we observed wolves keeping near small antelope groups that follow the multithousand major herds. In such small groups, the proportion of inferior and wounded individuals is high (up to 5 percent). Creating test situations through pursuit of small groups, the wolves overtake the inferior individuals lagging behind....
Wolves are not always so selective when preying on saigas. For instance, when wolves take pregnant females and newborn young it is a matter of chance as to which will fall prey. The death rates of pregnant females and young are negligible, however, because of the large concentrations of saigas. It appears that the concentration of many thousands of calving females in a territory of a single wolf family is an adaptation of the saiga to wolf predation....
No selectivity in wolves taking saigas appears to be operating at river crossings. The antelopes are aware of the water obstacle hazard and approach it with great caution, avoiding reeds and other cover that wolves use. If the predators manage to attack saigas during a crossing, losses are bound to be great: dozens of saigas of both sexes are commonly killed.
There occur situations when wolves kill quite healthy and efficient individuals. This happens under deep snow-cover conditions when frozen crust handicaps ungulate escape. The possibilities of useful selection are also limited in wild boar and Caspian seal populations, since wolves largely kill young individuals. Thus the problems of wolf-ungulate management are very complicated, no unambiguous solutions existing.
....As early as the beginning of the current century, the wolf's optimum range was in the forest-steppe zone where it preyed mostly on domestic animals -- wild ungulates having been long exterminated. The wolf's range widened following human distributional patterns. The change in range can be best traced in the plain-taiga zone, a landscape unfavorable for the wolf because of its deep snow cover in winter. The expansion of the wolf range in this zone was fostered by the following factors: an increase in the road network due to logging (which facilitates the wolf's movement under deep snow conditions), new settlements, and better feeding conditions at the expense of wild and domestic animals. The wolf penetrated to the arid deserts of Turkmenia and Uzbekistan thanks to irrigation and formation of water reservoirs near artesian wells....
Under conditions of human harassment, familial wolf groups tend to break up and total population numbers are reduced. The wolves acquire greater mobility; they migrate frequently and at longer distances. Our observations of wolf packs in the Smolensk region revealed that in winter wolves would often migrate over distances of 18 to 24 miles, leaving their territory completely during a single night. They would return after a few weeks, but commonly only for a short time....
In addition to large-scale migrations and a partial loss of territory conservatism in some wolves, a substantial change in the pattern of using killed prey has emerged in recent years. Where previously, a pack kept near the dead prey for a few days, wolves now only confine themselves to a single feeding, after which they move long distances. This new behavior precludes the use of bait to keep the wolves at the site for better hunting.
Generally, wolf behavior is most conservative. This is particularly true of their habit of howling. It would seem that an animal that changes its behavior flexibly to protect itself would find it natural to get rid of this habit, which attracts hunters and hence is fatal to survival. However, this does not happen. The communicative significance of howling must be so high that this behavior cannot be rearranged even at the hazard of species survival.
The situation in regions where the structure of the wolf population has been depleted through extensive control is quite peculiar. Unable to find a mate, lone wolves (both males and females) mate with dogs, which yields cross populations. This is the most unusual adaptation to extreme conditions created by man. Currently such crosses have been recorded in more than ten regions in the USSR, and their numbers and ranges have been increasing....Crosses are closer to true wolves ecologically, but they are less robust, preferring to attack small livestock, fowl, and dogs. By the early 1970s nine populations of wolf-dog crosses were recorded in the Voronezh region. By 1976 only three of them remained following a recovery in wolf numbers from 22 to 26 to 230 to 250. The earlier reduction in wolf numbers followed the occupation of the woodlands by wild dogs, which badly affected all living things, deer included. With wolf population recovery, wild dog numbers were reduced....
....In recent years wolf ecology studies in the USSR have intensified....The regional projects of wolf control will reduce the wolf's range in the USSR. This reduction is necessary in many areas, but preservation of the diversity of ecological and geographical wolf forms should also be stressed.