Part I: Terrestrial Zoology: Encyclopedia Arctica 3: Zoology (Excluding Birds)

Author Stefansson, Vilhjalmur, 1879-1962

Part 1: Terrestrial Zoology

Vertebrates

Terrestrial Mammals

(EA-Zoo. A. L. Rand)

TERRESTRIAL MAMMALS

CONTENTS

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Page
Cenozoic Migrations 1
Effects of Glaciation on Present Range 3
Habitats 6
Abundance of Individuals 8
Fluctuations in Numbers 9
Bodily Adaptations 9
Food 16
Storage of Food 18
Migration 21
Hibernation 24
Shelter 27
Mammal Interrelationships 30
Mammals in Relation to Man 31
Synopsis of Northern Mammals 35
Insect Eaters: Insectivora 36
Bats: Chiroptera 37
Flesh Eaters or Carnivores: Carnivora 37
Hares, Rabbits, and Pikas: Lagomorpha 39
Rodents: Rodentia 39
Hoofed Mammals: Artiodactyla 41
Bibliography 42

EA-Zoo. (A. L. Rand)

TERRESTRIAL MAMMALS
Terrestrial mammals have been able to occupy the North as far as there is land on which grows the vegetation they, or the animals on which they prey, feed. The geological history of the land had its part in determining the course of evolution of the mammals there, and the climate and its effects have been a filter, determining what animals may invade the area. The rigorous northern conditions, with widely contrasting seasons, have demanded modifications in bodily structure and in behavior.
These same great seasonal changes, sometimes with yearly deviations, have given a biological situation not yet in balance, so that great changes still may occur from year to year. The relatively small variety of life in the North has resulted in the development of some close interrelationships, including the intimate dependency of man on the other mammals.
Cenozoic Migrations
One of the striking things we see in going from the northern part of the Old World to that of the New is that the mammals are much the same; familiar types of shrews, bears, weasel, mink, marten, otter, squirrels, voles, beaver, caribou, and sheep may be considered different species by the zoologist, but their close similarity indicates they are closely related and have had a common history.
This similarity between the mammals of northern North America and of northern Eurasia certainly indicates a closer land connection in the not-far-distant past,

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a land connection large enough and of long enough duration for the mammalian forms to develop together. As the differences increase as one goes south, the connection must have been in the north. Land connections across the Atlantic have been postulated, but Simpson’s study of the fossil evidence indicates that a land bridge across the Bering Sea from Alaska to Siberia accords much better with the evidence and is sufficient to explain both present-day distributions and those of the past as indicated by fossils. The effects of this land bridge on various faunas have been traced through geological time from the early Eocene to the Pleistocene epoch, and is still to be seen today, in the Recent. But it was not continually in existence. It evidently was interrupted a number of times, some of the interruptions lasting perhaps for several millions of years. The latest land bridge was in the Pleistocene. There may even have been some land connection and some small local migrations between Alaska and Siberia across the Bering Sea area during the Recent glaciation, whose distance in the past is probably measurable in tens of thousands of years rather than in millions, as with some of the former connections.
Where mammals first originated is still shrouded in mystery. In the early Eocene the fossil evidence indicates that representatives of all the large groups were involved in the movements across the Bering Sea between North America and Eurasia. But later exchanges were of smaller scope and concerned only northern groups already common to the two continents.
In the early Eocene the direction of the migration is unknown. In the late Eocene there are indications that the migration was more from North America to Eurasia rather than the reverse. After this, though the faunal exchanges involved migrations in both directions, the evidence indicates that more groups probably migrated from Eurasia to North America. This later preponderance of an eastward

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migration is correlated with the larger area of the temperate zone Eurasian land mass compared with that in North America, and the impact on its fauna of aggressive immigrants from both Africa and Asia, as well as from North America. Thus the Eurasian fauna was probably better adapted to migrate and survive changing condi– tions, while the North America fauna, developing in a smaller area and with aggressive new immigrants affecting it only from Eurasia, had a more tranquil history.
The Bering Sea land bridge apparently always had a cold, rigorous climate, operating with active selectivity and allowing only cold-tolerating animals to use it.
In the latest epoch, the Pleistocene, the mammal migrants were mostly types so similar to present-day living forms that they are classified in the same genera. Some of them that perhaps came from Eurasia to North America are: the hare, pika, tree squirrel, bear, wolverine, moose, caribou, bison, and sheep. Some that perhaps migrated from North America to Europe are: the marmot, banded lemming, brown lemming, red-backed mouse, vole, jumping mouse, and fox.
Effects of Glaciation on Present Range
The ranges now occupied by northern mammals have been occupied only recently, just since the last ice epoch, their age counted in tens of thousands of years. Since the last land bridge across the Bering disappeared, glaciers have covered practically all the arctic and subarctic areas with ice. Mammal life was impossible on it then as it is today on the ice fields of Greenland and Novaya Zemlya. The mammals moved southward ahead of the glaciers on both continents, or perhaps in a few cases remained ^ in isolated ^ ice-free refugia ^ refugia ^ , made into islands by the surrounding ice. The one in the Yukon-Bering Sea area was probably the largest and best authenticated.
With the melting and retreat of the ice, the cold-tolerating mammals, which had survived along the southern fringe, spread northward following the retreating ice.

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At the northern edge of the mainland however, the extent of water in summer and of ice in winter acted as a barrier in preventing some mammals from reaching the arctic islands. In North America where there is a whole arctic archipelago this is particularly apparent. Notable among the mammals that have not spread north of the mainland are the grizzly bear, the ground squirrel, and the cinerous shrew. The brown lemming, on the other hand, is famous for its mass migrations which may move over sea ice. One would expect this lemming to be widespread in the arctic islands, but it seems not yet to have reached Greenland or Ellesmere Island.
The caribou, musk ox, banded lemming, and arctic hare as well as the wolf, arctic fox, weasel, and polar bear have all spread across the ice from island to island to reach northern Ellesmere Island and Greenland.
But on Greenland the icecap covering the interior cane to the coast in the northwest and in the east, and some of the mammals that reached North Greenland have not been able to get across the icecap and colonize the coastal, ice-free areas of southern Greenland, notably the banded lemming, the weasel, and the musk ox.
The polar bear wanders far over the ice; it is carried by moving ice and swims long distances, and has been seen swimming at sea 30 kilometers from the nearest resting place. For such an animal there are few barriers in the Arctic, so long as food is present, and the polar bear gets much of its food from the sea. The arctic fox has similar habits; in winter it wanders widely on the sea ice; it shares in the polar bear’s harvest of food from the sea, as well as snatching some for itself (small fish or invertebrates); and it also is carried by the moving ice. Both polar bears and arctic foxes are brought as far south as the Gulf of St. Lawrence some years by this means, and one arctic fox even reached Nova Scotia.

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The arctic here ranges freely over the sea ice, reaching islets 40 kilometers or more offshore, so it is not surprising to find it, too, in South Greenland. One would think that the musk ox would also have been able to make the journey, but for some reason it did not do so. The banded lemming has been found some kilometers out on the ice, sometimes frozen to death, but it has never reached South Greenland. Perhaps the journey across the glaciers is too much for it, although it has been recorded as traveling 57 kilometers over the ice in North Greenland. The absence of the weasel in South Greenland can be correlated with the absence of its main prey, this same lemming.
The colonizing of the north from the south is still going on. In the last 60 years or so ^ , ^ the coyote has spread from south of our area (which is still not ^^ clearly defined) and has now reached the arctic coast, both in the forested country of the Mackenzie and in the tundra of Alaska. The red for as now ranged out onto the barrens since our first knowledge of the area, but in recent years has increased and spread there, to the consternation of some of the trappers who fear that in some areas it may usurp the place of the more valuable arctic fox. Recent records indicate that the red fox may become established in Baffin Island; the wolverine is there, and there are even surprising records of the ordinarily forest-dwelling lynx reaching southern Baffin Island, evidently traveling on drifting ice from northern Quebec.
In the Old World there is no similar archipelago and the islands lying in the polar sea there are few and scattered. Spitsbergen has been reached by the polar bear, the arctic fox, and the reindeer, which must have traveled over the sea ice. Nov ^ a ^ ya Zemlya, much closer to the mainland, has the brown and the banded ^^ lemming, the arctic fox, the polar bear, and the reindeer, according to Gorbundoff.

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Habitats
In the North there are distinctive types of landscape, each occupied by mammals peculiar to them. The treeless country north of timber line (the tundra of E a urasia, the so-called barren grounds of America) that forms a circumpolar ^^ ring around the tops of both continental masses and sends arms w ^ s ^ outhward down ^^ the crests of the mountain masses has its characteristic mammals. The northern edge of the coniferous belt, the taiga of Eurasia, and the Hudsonian zone forest of America, composed chiefly of open stands of spruce with thickets of willow and dwarf birch that forms the second circumpolar belt, also has its character– istic mammals. The taiga sends northward extensions along the river valleys.
The treeless country has: ( 1 ) polar deserts, which are the permanent ice and snow fields (as in Greenland, northern Baffin Island s , and the islands to ^^ the north of it, Nov ^ a ^ ya Zemlya and Spitsbergen), destitute of mammal life except ^^ for wanderers; ( 2 ) desert tundra, where the rocks are nearly devoid of vegetation, as in large areas of Baffin Island, and mammal life is scanty; and ( 3 ) the grass and lichen tundra, the main home of the polar mammals such as caribou, musk ox (North America only), arctic hares, banded and brown lemmings, wolf, arctic fox, and weasel (called ermine in the Old World). On the tundra near timber line, locally, are other species that also occur in timbered areas: shrews (genus Sorex ), brown and grizzly bears, wolverine, red fox, ground squirrels, red-backed mice, and voles of the genus Microtus ; above timber line in the mountains are sheep, pika, and marmots.
The coniferous forest belt does not begin suddenly but changes gradually through shrubs, such as willows and scattered trees, to forest. Taiga and tundra also interdigitate, especially where the Mackenzie River valley carries forest far north in America, and along the north-flowing rivers in Siberia. No more

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do the mammals suddenly change from those of barren ground to those of the forest. The caribou and the arctic fox, e x ^ s ^ pecially in winter, enter the forest and even ^^ the musk ox, in summer, favors the willow thickets where they are present.
The T ^ t ^ aiga is richer than the tundra in species of mammals, corresponding ^^ to its richer and more varied plant life. Feeding among the trees, in the air, are a few bats, especially in the Old World; in the trees live tree squirrels and flying squirrels and on the ground are moose, varying hare, some voles, chipmunks, lynx, red fox, mink, marten, sable, and black bears. Several insectivorous shrews occur; and in the freshwater streams are muskrats (America only), beaver, otter, and mink, all of which find their headquarters in this forest belt.
Only a few species range widely in both forest and tundra, among them notably the wolf and the weasel.
The increase in the number of species of mammals as one goes from ^ the ^ polar regions to the tropics is a general phenomenon, and its actuality is well illustrated by giving in Table I the numbers of land mammals reported from a series of stations in our area, arranged in a north to south series as far as possible. Note par– ticularly the great difference between the number of species inhabiting the taiga and the tundra.

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Table I.
Area Type of country Land mammals recorded, number of species
Northeast Greenland Tundra 8
Southern Baffin Island Tundra 11
Perry River, Queen Maud Gulf Tundra 14
Mackenzie Delta Taiga, tundra, and mountain 36
Wood Buffalo Park, Canada Taiga 44

EA-Zoo. Rand: Terrestrial Mammals

Abundance of Individuals
Although the number of species in the North is limited, the number of indi– viduals in a certain species may be great. When voles or lemmings are plentiful, the tundra may be so riddle s ^ d ^ with their burrows and marked by their digging that ^^ it has a tattered appearance. For long stretches the ground may be undermined with their tunnels and sown with their holes. In walking over the tundra the tiny rodents may scurry over the ground ahead of one. The vast numbers to which the brown lemmings attain are best seen when they make one of their irr ^ e ^ gular mass movements. In the Queen Maud Gulf area, Gavin saw “The sea ice was covered with a moving mass of lemmings.” He estimated an average density of one lemming to ^ the ^ ^^ square yard, and indicated the movement extended over a 60-mile front and lasted for 10 days.
The caribou numbers, when these animals move en masse, are even more striking. From a rise in the country west of Hudson Bay, sometimes many herds of from 100 to 2,000 animals may be seen at once, and an estimated 20,000 caribou have been reported in view at one time. Such herds, when forced into a small compass, as at a river crossing or a pass, may leave the ground as bare mud and dust with hardly a spot untrampled by their hoofs or with a bit of vegetation showing. Such aggregations of animals have been the basis of estimates in the past of herds of “millions” of caribou. David Thompson, Joseph B. Tyrrell, and Vilhjalmur Stefansson are ^ a ^ mong those who have published large estimates — Thompson up to three million, Tyrrell and Stefansson “several hundred thousand.”
In the forest too, sometimes, such species as the varying hare become so common that many of them may be in sight at one time in every willow and bush thicket they frequent. Their ravages on the vegetation, the bare stems from which the bark has been gnawed, and the stems ^ t ^ rimmed in their feeding may be conspicuous features of the landscape.

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But not all northern species are numerous. The least weasel, Mustela rixosa , though a widespread circumpolar species, seems always rare. Certain voles, such as those of the genus Phenacomys , and shrews of the genus Microsorex are usually scarce or rare.
Fluctuations in Numbers
Numbers are an unstable thing in the North. Some years a mammal may be in swarming abundance; in other years the species may be so scarce that it is difficult to find a single individual. This is well illustrated by the varying hare and the lemmings, animals that have become almost classical examples, with the corresponding fluctuations in numbers of the animals that depend on them largely for food, the lynx and the arctic fox especially. The ramifications of the effects of these fluctuations are much more widespread than the prey– predator relationships, extending as far as the differential growth of certain vegetations and the prosperity of trappers. There is much evidence to show that these periods of abundance and scarcity rec o ^ u ^ r with a regular rhythm, and with a ^^ periodicity of different length for some species; notably about four years for some of the smaller rodents and ten years for some of the larger animals. There is also evidence for long-term changes in abundance in some of the largest species, such as the moose and the wolf. Whether or not these have a regular periodicity is unknown. It used to be though these fluctuations were world wide, but much recent data indicate they do not synchronize over wide areas, and the animals may be common in one area while scarce in another but a short distance away (see “Population Cycles”).
Bodily Adaptations
The severity of northern c onditions has had its effect in directing the ^^ evolution of the animals living under them. No one factor will account for

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everything, but cold, deep snow, and type of cover and food available, as well as other less tangible factors have molded the bodily form; sometimes one factor is dominant, sometimes another.
It has been laid down as a zoological rule (Bergman’s rule) that when closely related animals occur in warm and cold climates, those from no ^ r ^ thern climates tend ^^ to be larger. This has been correlated with the larger animal having a propor– tionately smaller body surface to unit of body weight, and so reducing radiation of heat, an important factor in arctic climates.
When we examine this for arctic mammals, there are so many exceptions and contradictions that it seems other factors, probably ecological, have played more important parts in determining the direction of evolution of arctic mammals.
First it must be pointed out that both very large and very small mammals occur in the subarctic forests. In North America the pigmy shrew ( Microsorex ^ Microsorex ^ ) , ^^ less than four inches long, weighing only a few grams and one of the smallest mammals, and the big brown bear ( Ursus ) and the moose ( Alces ), weighing over 1,000 pounds and among the largest of North American mammals, all occur in the Subarctic. In the Eurasian Subarctic there is also a tiny shrew, an enormous bear, and a moose.
In some species the size is greater in the north than in the south: the northern short-tailed weasels tend to be larger than more southern ones; the arctic hare is larger than the more southern varying hare; the northern coyotes tend to be larger than southern coyotes. But there are many obvious exceptions: the largest musk ox are not to be found in the north; the southern caribou are larger than the most northern caribou; the northern wolves are not as large as some of the more southern ones; the northern chipmunk is a small species. The tundra shrew is smaller than its closest relative to the south. The common meadow

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mouse or vole of eastern North America in the east shows a decrease in size in going northward; in the west, going from southern Canada to the Mackenzie Delta it shows a decrease through northern Alberta and southern Mackenzie, and then an increase in size farther north. Therefore, it would appear that in gross size no general rule applies.
Correlated with a cold climate, appendages tend to be short (Allen’s rule), presumably an adaptation for the conservation of heat. We see it in the small ears of the arctic fox as compared with the ears of more southern foxes in both Eurasia and North America; the shorter ears of the arctic and varying hares as compared with the long ears of more southern forms such as the jack rabbits.
The tail, too, has undergone a reduction in the north; it is very short in the moose, caribou, and musk ox; the lemmings have the tail very much reduced; the arctic fox has a relatively shorter tail than the red fox; the red squirrel has a relatively shorter tail than does the more southern gray squirrel. However, when it comes to a question of feet, which according to Allen’s rule should be relatively shorter in cold climates, we find other environmental factors have been more effective. The moose has developed very long legs, presumably for wading through bogs and deep snow, though the musk ox, living farther north, does have feet relatively shorter than its more southern relative, the bison. The adaptations in feet are more pronounced in ^ ^ characters adapted to the physical ^^ aspects of the environment. The long legs of the moose have been mentioned; the caribou solves the problem of getting over deep snow and soft bog in another way, by developing “snowshoes.” Its hoofs spread widely when the weight is placed on them, and serve to keep the animal from sinking deeply into the soft medium. The same adaptation is also very apparent in the snowshoe rabbit, with its long broad hind feet which have given it its name, and in the lynx. Although the lynx is

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almost the same size as the bobcat, its nearest relative further south, its hind feet are almost 10 inches long and disproportionately broad, while the hind feet of the bobcat are only about 8 inches long and much less bulky. Another striking peculiarity in foot development peculiar to the Arctic is in the seasonal development of the foreclaws of the banded lemming. In the summer they are of normal size, but in winter the nails become greatly enlarged, the enlargement being shed in the spring. This is usually assumed to be a n unique adaptation for burrowing in snow, although the brown lemming that does not have this seems to get along just as well.
Other ecological adaptations for special environments as in other parts of the world are also found; the long feet or legs for running (deer); short broad feet with sharp claws for climbing (squirrels, martens, sable); fringes on sides of the feet for swimming (muskrat, some shrews); webbed feet for swimming (otter); and broadened forefeet with stout claws for digging (moles, marmot, ground squirrel). The spring coat of the Canada porcupine, with spines which stick into the mouth of a would-be predator and discourage it, is a striking case of defensive armor, but this type of adaptation is more common in temperate and tropical climates.
A prime necessity for an animal living in a cold climate is a heavy coat of fur for warmth. This we find reaching its highest development in the Sub– arctic and the Arctic. The hairy covering of many animals tends to consist of long, coarse, stiff guard hairs, serving to protect the underfur, and the shorter, denser, finer underfur, furnishing the warmth. In the musk ox these are strikingly contrasted, where the long guard hairs may reach a length of 24 inches or more, while the underfur is a woolly coat 2 or 3 inches thick, close to the skin. In the caribou there is not this sharp distinction, the

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insulating effect being achieved by a very dense coat of hair of about uniform length. The hairs are thicker at the tip than at the base, are hollow and filled with air, and are so closely spaced they will not lie down. The caribou skin is universally considered one of the warmest of skins for covering. The roughness of the pelt and the brittleness of the hair, however, make it un– suitable for civilized use.
Our finest furs for the fur trade come from the Arctic and Subarctic. Many of them are of the weasel and wolf tribe; their sleek, glossy, dense guard hairs and warm underfur make them especially prized. But some rodents also provide pelts of note ^ , ^ of which the beaver is the finest. From the north of North America important animals for the fur trade include the fisher, marten, lynx, beaver, and arctic fox, which supply the most costly furs, and also the red fox (whose color phases, the cross and silver, no longer command a fabulous price since they are being extensively raised on fur farms), wolf, weasel, squirrel (of secondary importance), otter, mink, and muskrat (a staple fur, now important for the large number taken). From the northern part of the Old World come such fur ^ s ^ as sable (related to marten but with finer fur), otter, lynx, kolinsky or yellow weasel, ermine (the weasel of North America), arctic fox, red fox, here (a much better fur than that of American hares and used to simulate arctic fox), burunduk (a chipmunk), baum marten (related to sable), otter, squirrel (related to the American red squirrel but of finer pelt), and wolf.
Not only is the fur coat of northern mammals warmer than that of those living in warmer climates, but the animals tend to be more completely furred. The deer of southern climates have bare areas on the nose; these areas are more fur-covered in the moose and caribou; the same is true of the musk ox.

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The feet of many northern mammals are also much more heavily furred than their southern relatives; the bottom ^ s ^ of the feet of the polar bear and ^^ arctic fox in winter are completely covered with fur, and the lemmings and some of the voles have furry feet.
A storing-up of fat under the skin as reserve food to carry the animal over the lean winter period and to serve as insulation against the cold has often been postulated. There is no doubt that many animals do get fat in the fruitful days of summer, but the value of this as a factor in overwinter sur– vival (except in hibernators) is probably overestimated. This is particularly true of the caribou, in which an old male may have 50 pounds of fat laid down as a blanket over its hips, just under the skin. Far from being a reserve for winter, this fat is used as a reserve food for the mating period in the fall, when for some weeks the male does not eat, and consequently enters the winter lean and in poor condition.
That the color assumed by arctic mammals is an adaptation to their environment has been questioned. Many arctic and subarctic mammals are some shade of brown, as shrews, voles, some bears, most of the weasel family, and beaver; the moose and the musk ox are blackish; some voles, the pika, and some caribou are grayish; [: the ] brilliant colors are approached in the red-brown of the red fox, red squirrels, red-backed mice, and banded lemmings. Contrasting patterns are found in only a few: the stripes of the chipmunk; the spots of some ground squirrels; the rump patches of sheep; the rump, neck, and flank markings of some caribou; and the lateral stripes of the wolverine.
But the striking thing about coloration of arctic mammals is the tendency of some of the tundra animals to be white. This is apparent in the polar caribou of both hemispheres. The most northern wolves also tend to be white

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(although the black phase of the red fox, a forest animal, tends to be most common in the northern edge of its range), and this whiteness reaches its greatest development in the polar bear, which is white the year round.
This whiteness is also acquired by another method; a seasonal change so that the animal is white in winter when the ground is snow-covered, and brown or gray when the land is not snow-covered. This seasonal color change is shown by some species of both predatory animals and those that are preyed upon. The hares of the forest have this double molt; the arctic fox is bluish- brown in summer and white in winter; the weasels and the banded lemmings (but not the brown lemming) show this change. With the arctic hare in North America, while the southern ones become slate-colored in summer, the northern ones stay white or nearly white the year round.
The functional value of a white coat to these animals is still being debated. The first and obvious correlation is that, with the snow changing the dark background to white, the color-changing food animals are less con– spicuous to predators; and color-changing predators are less conspicuous to their prey. This may be true, but white coloration reappears also in antarctic birds where there are no effective enemies.
The mammals that change color with the seasons have two molts a year. But with most arctic mammals there is but a single annual renewing of the fur. This molt takes place in a short space of time. The fur is often all shed about the same time, the incoming coat pushing off the old fur over the animal’s whole body. Before the molt the animal may be covered with a long dense coat; during the molt the caribou or the arctic fox may present a very ragged appearance, with great flakes of old fur peeling off. When the molt is first complete and the incoming hairs still short, the long-haired animals such as the arctic fox may appear to be much smaller than when they were wearing their winter coat.

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Food
Mammals all depend on vegetation for their food, but some get this at first hand while others get it by eating other animals which eat plants.
On the tundra, the caribou, the musk ox, the hare, and the banded and brown lemmings are the herbivores, feeding on grasses, leaves, and twigs. Preying on them are the wolf, the arctic fox, and the weasel. The polar bear has closer touch with the sea, whence it draws most of its food.
In and near the taiga with its more varied vegetation, one finds various types of herbivores: the voles, hares, ground squirrels, marmots, sheep, and moose grazing and browsing; voles tunneling for roots; some mice, tree squirrels, and chipmunks favoring seeds and fruits; beaver and porcupine eating much bark of trees but getting it in quite different ways (the beaver felling the trees by gnawing through them at the base and then eating the bark, the porcupine climbing the trees and gnawing off the bark in situ ); and the muskrat and the moose feeding to some extent on aquatic vegetation. As a natural correlation the greater the number of herbivores ^ , ^ the more predators there are: the wolf ^^ and weasel; but there is also the red fox feeding on mice and hares; the lynx preying on the woodland hare; the marten chasing the squirrel in the trees; and the otter chasing fish in the streams (and it is said to attack beaver and take muskrats). Here also are shrews, hunting insects and other invertebrates in the ground cover, and a few bats catching them in the air overhead.
The bears are special cases in regard to food: they are terrestrial carni– vores in structure, but the polar bear has invaded the domain of the sea, feed– ing on fish and on seals; and the black grizzly bears much of the time fill their stomachs with vegetable material but prefer meat, never losing an opportunity to secure a meal of it, even if it means eating carrion.

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Some mammals, such as the musk ox, sheep, voles, and lemmings feed on much the same types of vegetation in summer and winter. But the caribou that feeds on grass during the summer turns to lichens for the winter; the snowshoe rabbit, eating grass and herbs in summer, turns to bark and twigs in winter; the shrews, insect - ^ ^ eaters in summer when insects are plentiful, ^^ may eat seeds as well in winter; the arctic fox that in summer feeds on lemmings and birds (and their eggs and young), finding the birds gone and the lemmings hard to reach under the snow in winter, may go along the beach picking up whatever appears from the sea or may follow the polar bear for scraps.
The black and grizzly bears that feed on herbaceous matter (grass, roots, and horsetail) during the summer change to berries when they are plentiful in the fall; grizzlies change to fish diets when salmon run in the Pacific streams.
Adaptations in feeding habits have to be made. The caribou that grazes may find its food covered by snow and have to dig for it. On some wintering ranges, caribou have been reported digging pits four feet deep through the snow to the browse underneath. Legend has it that the caribou digs with the flattened brow antler, but recent observations indicate it uses its forefeet only.
Arctic hares, feeding on wind ^ - ^ swept areas, are usually able to push aside ^^ the light layer of snow covering their food by using their noses and scraping with their forepaws. Sometimes, however, there is a frozen crust over the snow, and then the behavior of the hare follows a different pattern. The sense of smell apparently enables it to locate its hidden food, and then over the chosen spot it hammers the crust with blows of its forepaws. With the crust broken, the hare may remove the larger pieces with its mouth, and then, with nose and forepaws, gets at its food in the usual way.

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Storage of Food
In a country where food is abundant for a short season and scarce or difficult to get for the rest of the ^ ^ year, a logical development is that of ^^ harvesting the food in the season of plenty and storing it until the season of scarcity. This we find in many of our mammals, both flesh - ^ ^ eaters and ^^ vegetarians. In some, this storage is a highly developed behavior, and probably essential for the animal’s winter survival, as with the beaver, pika, and voles; in others it is more haphazard, and its survival value is probably not so generally important.
The meat - ^ ^ eaters are able to store food in the cold climate where it ^^ decays slowly. Bears sometimes bury their surplus from a meal, and even the polar bear may cut out large blocks of snow with his forepaws and cover the remains of a seal. Wolves often chew off parts of a large mammal they have killed and carry the pieces away and bury them. These, however, seem to be temporary rather than long-range storage.
The arctic fox, locally at least, makes caches for winter use. In northern Greenland these foxes lay up stores of little auks or dovekies. They are laid together in an orderly manner, always with the heads eaten off, all the tails pointed the same way, and laid in rows or arranged in heaps. Large numbers of eggs are also collected. In one cache, 27 little auks and 40 eggs were re p ^ c ^ or t ^ d ^ ed. ^^
The mink stores such food as muskrat, fish, Squirrels, and birds. One mink cache in a hollow long was found to contain 13 freshly killed muskrats. Weasels often kill more than they need and this is sometimes considered a bloodthirsty habit. But it is no more a lust for killing than is the woodsman’s foresight in providing his larder with meat for the winter. Weasels when undisturbed [: ] o not leave their prey scattered about, but carefully store it

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away and, in many cases, bury it. Not only will the weasel carry [: ] off prey that it has killed, but will carry off a supply of animals already dead, as was shown when one rifled a collector’s tent and carried off not only mice and birds but also chipmunks that had been made into specimens. This storing instinct of weasels does not seem so definite as that of some rodents, and it is said much of the stored meat probably decays.
The vegetarians store a number of types of food; seeds, roots, herbage, and bark. The store may vary from a mushroom casually stuck in a fork of a tree by a squirrel to the elaborate cache of cut sticks made by the beaver.
In the coniferous forests, the tree squirrels become busy in the autumn cutting down the fresh crop of cones for the seeds which they contain and caching the cones in little holes dug in the forest floor under a favorite tree, and here they also deposit quantities of berries. The squirrels, or a succession of squirrels, use the same feeding and storing place year after year and a whole heap or mid ^ d ^ en of the cone scales accumulates. These middens ^^ may be conspicuous things, up to 9 feet high and 12 feet across. It is in this pile of debris that the squirrels bury their winter’s food supply.
The little chipmunk is also a diligent storer of food. Soon after he emerges from his winter home in the spring and until he retires in early winter, he devotes much of his time to gathering and storing away seeds for food. These are stored in various places - - some in crevi s ^ c ^ es in a rock, ^^ some in his winter home in an underground burrow.
Several species of voles store roots for winter. On the American tundra, the root of the plant Hedysarum boreale or “licorice root” is gathered by voles and stored in underground chambers. These roots are tasty to humans, and are stored in such quantities that some Eskimos search them out with the

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aid of dogs, and appropriate the store for their own use. Above timber line in the mountains of the New World, another species of vole excavates chambers just below the surface of the ground and stores quantities of carefully cleaned starchy rhizomes of a sedge, individual stores amounting to a double handful or more apiece. In Asia the same habit prevails with certain voles. In the lower Kolyma region of Siberia, the ^ a ^ farinaceous root is laid up in large ^^ quantities by voles in their burrows, and here too the local people seek out these stores as additions to their own food supply. Although these roots would be present in the ground all winter, the ground would be frozen, and this source of food would be unavailable to the voles if not gathered before the freeze-up.
The pikas that live in rock heaps and talus slopes above timber line make “haystack” caches. In late summer and early fall, the animals indus– triously gather the stems, leaves, and flowers of a wide variety of plants growing in or near their rocky homes. They pile their cuttings into miniature “haystacks” on shelving rocks, in spaces between rocks, or more rarely under logs or stumps, where they are exposed to the air but are well protected from rain by overhanging rocks or logs. These stacks average about the size of a bushel measure, and, safe under the snow, are the pika’s winter food supply.
The beaver makes an underwater store of its favorite food, usually of branches such as aspen or willow. The branches and sticks are cut on the shore, then pushed, rolled, or dragged to the water. Once afloat, the beaver grasps the stick in its teeth and lets it float alongside as it swims to the food pile. This food pile is usually located near the beaver’s lodge, handy for winter use. The first pieces brought to the store may be forced into the bottom of the pond to anchor them; on later trips, the beaver dives with its

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load and forces the material into that already there. The final store is an entangled mass of waterlogged brush and sticks that may reach considerable size, up to 124 feet in circumference. When the pond freezes over, the beaver has its food at hand. When it wants to feed, it leaves its house by its ^ the ^ underwater entrance, swims under the ice to the food pile, detaches a length of stick, swims back to its ^ the ^ house, and devours the bark from the stick at its leisure, without having to expose itself to the elements or to it ^ s ^ enemies.
Migration
A yearly seasonal change in environment, a change from a mild growing season of plenty to one that is severe, with snow and without plant growth, demands seasonal adaptations in the animals’ behavior. One of the most obvious is that of migration, when the animal moves from an area with unfavorable con– ditions to one of more favorable conditions. The question of migration in land mammals in the Arctic usually directs attention to the caribou and the brown lemmings with their well-known movements of sorts; perhaps incipient and rudi– mentary migration are indicated in a number of others. Voles of the genus Microtus tend to summer in low, wet , places where the growing vegetation on which they feed is lush; but in winter they may move, perhaps only a matter of a hundred yards or so, to a drier slope. One effect of this is seen in the spring when the snow melts, and vole signs completely cover a restricted area on a slope where they have wintered and eaten all available vegetation. Musk ox, one of the most stationary of arctic mammals, makes a similar biannual change of territory, from lowlands in summer to higher land in winter. Perhaps this is correlated with depth of snow in the lower places being greater than on wind-swept exposed places. Depth of snow is certainly correlated with movements of sheep in the Yukon and Alaska mountains, where sheep winter on areas of

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little snowfall or where the wind sweeps away the snow, and spend the summer spread out over a much wider area. The polar bear, which follows the sea ice, is scarce or absent in summer where there is no ice, and the female polar bear that has to leave the sea ice and move to an area of land to hibernate and to bring forth her young also shows incipient migration.
The brown lemmings, though their movements in northern Europe, and in North America to a lesser extent, have been widely publicized, represent a special case of migration. Periodically, but apparently without regularity, a population of lemmings may become very dense and then vast numbers of them move away from the densely populated area. In Norway they may swarm down the mountains and into the sea; on the Murmansk coast they sometimes swim out to sea; in Arctic America the most notable accounts are of cases where they cross the sea ice. The lemmings appear to move continuously in a certain direction, perhaps determined by slope, the animals going downhill which explains their going to the ocean. Steadily the movement presses on, as illustrated by Gavin’s observations on the frozen Queen Maud Gulf: “they were all headed in an easterly direction. They stopped at nothing. Untold thousands plunged over the ice into the water of a lead… They perished in large numbers in these leads, but here and there they found passages up the ice and blindly continued their journey without deviation and without hesitation.” But these movements differ from real migration in there being no return. The animals migrate en masse and perish. Only those few left behind survive to carry on the species, and start the population building up again to where another mass emigration occurs.
The caribou is another wanderer whose movements are often called migration but which really fall in another category. Uncertainty and irregularity are

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are the peculiarities of caribou movements. The general statements that caribou move toward the arctic coast of Siberia in summer and southward away from the coast in winter, and that similar movements occur in northern Canada tell only a part of the story.
The caribou in summer favors rich grass, but in the long winter it lives on the abundant lichens (“reindeer moss,” Cladonia ). The moss is slow-growing, taking perhaps 15 years to recover from severe grazing. (It has been demon– strated that arctic tundra will support at most 10 caribou per square mile.) The caribou is extremely gregarious and the herd units sometimes contain 100 to 2,000 head where the animals are plentiful. Thus they graze close-herded. If they stayed long on an area they would exhaust the grazing for many years. The only adaptation possible within this animal’s habits is a roving life, during which it stays but a short time in any place. This is just what has happened. Most of the caribou are continually moving, summer and winter; the rut takes place during migration, and even during calving there is only a temporary halt.
In the special case of caribou, the migration seems to be an adaptation for conserving the range; the animals return to certain favorite summering areas where they feed on grasses, but for the rest of the year they wander, getting a bite here and another there. To do otherwise would unduly deplete the range. The direction taken in migration seems to be determined in part by topographical factors; in part by a line of good pastures. This last, of course, may be determined by former use, as well as by chance, and we find that caribou may use an area for a number of years and then go elsewhere for a number of years. The direction may be north and south, as in Siberia and in the main range in Arctic Canada. The former migration to and from Victoria l Island to the mainland is a good example. In southern Canada, on the barrens and

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the forest edge, the migration, while north and south, tends to be circular, perhaps an adaptation to cover more range during the year; with the Yukon- Tanana herd, it retraces its route hammed in by mountains. In Baffin Island much of the migration is from the summer range of the grass plains and marshes of the west to the higher eastern country where lichens are abundant in winter. In northern Yukon the movement may, in the fall, first be north to the coast, then east to the Mackenzie River, then south, and west again through a mountain pass to the range from which they started (some years they don’t use this route at all). But all guesses may fail, and, as Murie writes, after pronounced movement caribou are still everywhere and it is difficult to say where they’ve gone; some caribou seem to winter some years in almost every part of the range, and some populations, as the present ones on Victoria Island, on Spitsbergen, and probably many others, do not seem to migrate at all.
The only clear-cut migration in Arctic-inhabiting mammals is that of the bats that have ventured into the northern forest. One, at least, the hoary bat, that has been recorded north to Southampton Island, makes definite migrations by flying to warmer climates in winter and back again in the summer.
Hibernation
Another way in which northern mammals are adapted to the winter is through hibernation, spending the winter in a dormant or semidormant state. The hiber– nating animal typically enters this winter sleep in a fat condition with the stomach empty. Some bears are said to purge out their intestines with special foods and then, both in North America and in Eurasia, to eat a quantity of fibrous material which plugs the intestines and remains in place during the winter. In its dormant condition, body temperature may drop to a few degrees above freezing and bodily processes slow down greatly, so that little of the

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accumulated body fat is used as a reserve of food during the winter. Most of this fat apparently serves as insulation and as a store of food for the animal when it wakes up in the wpring, when little fresh food is as yet avail– able. This winter sleep appears not to be continuous even with the most pro– found hibernators, the formant animal rousing at intervals throughout the winter. Possibly this is an adaptation to keep it from freezing; for with waking, bodily processes speed up and body temperature increases. Violent shivering often accompanies this waking, apparently another heat-inducing phenomenon.
While the most profound hibernators such as the marmots and the ground squirrel go into hibernation in the fall and do not emerge until the spring, there are other species such as the red squirrel which disappear into their shelters and remain quiet for longer or shorter periods, a few days or a week or two, in severe weather. Probably all should be called hibernators, of varying profundity. Since hibernation is an adaptation for passing over a period of unfavorable conditions, one might expect it to be commonest in the Far North. But this is not the case. Hibernation is particularly a phenomenon of temperate and arid climates; a larger percentage of mammals living in these climates hibernate than in either arctic or in very warm climates. The following illustrates the decrease in the number of hibernating mammals as one goes from the Temperate Zone to the Arctic: In an area in southern Alberta (about latitude 49°), 16 of th d ^ e ^ 55 mammals hibernate; in southern Mackenz ^ i ^ e (about latitude 60°), 9 of the 43 species hibernate; at the Mackenzie Delta on the Arctic Sea, only 5 out of about 36 mammals hibernate; while on the Canadian Arctic Islands, only 1 out of 9 species hibernates (the polar bear), and that only partially.

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While there is great change in the number of hibernating species, going from north to south, it is not a habitat difference; polar bears, grizzly bears, ground squirrels, and marmots hibernate north of the timber line, and one must go some distance south in the forest before the number of hibernators increases greatly.
It is sometimes said that hibernation is impossible in the Arctic because with permanently frozen ground the animals cannot find a frost-free place to hibernate. However, this is obviously not the case, as both ground squirrels and marmots hibernate near the edge of the Arctic Sea; the polar bear is said to hibernate in snowdrifts; and farther south, the black bears and certain bats hibernate in resting places where the temperature of their immediate surroundings goes below freezing.
A hibernating species usually hibernates for a longer period in the northern than in the southern part of its range; for example, the black bear in Florida does not hibernate, in Ontario it goes into hibernation in December, while at the Mackenzie Delta it goes into hibernation in October. But this does not hold with the polar bear, which is a partial hibernator in the Hudson Bay area, but hibernates little, if at all in the northern Greenland.
The age of the animal, its sex, and its physical condition seem to affest its hibernation. Where the polar bear hibernates, it is said that more females than males hibernate, or perhaps only gravid females. Of animals that ordinarily hibernate, such as the grizzly bears, individuals abroad during the winter are lean, perhaps not having been able to accumulate sufficient fat to enable them to go to sleep for the winter.
The average dates of mammals going into hibernation and emerging from it in the spring are not sufficiently well known in the Arctic for us to tabulate.

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Data from elsewhere are not applicable, as this varies locally with climate. In general, the most profound hibernators retire in the fall and are not seen again until the spring. Those with the longest period of hibernation are probably the ground squirrels. In the Arctic they have been recorded abroad from April to December but these are extreme dates and the average period of activity is probably May to September at most, for farther south one species is recorded with a hibernation period of 9 months.
The place of hibernation is usually an underground chamber dug by the animal and line s ^ d ^ with vegetation, but bears may scratch out only shallow dens, ^^ and polar bears are said to hibernate in snow burrows. The young of the black, f ^ g ^ rizzly, and brown bears are born while the females are in hibernation, and ^^ suckle and female while she is in a dormant state. Even in the farthest north where the polar bear hibernates little, the female retires to a snow cave at the time of the birth of the young and remains there for some time, allowing the small, weak young to develop somewhat before venturing abroad.
When hibernators emerge in the spring they are still fat, only a small part of the fat having been used during the winter sleep, and this store of fat which served so well as insulation during the winter becomes an important food reserve for the animal until food becomes plentiful later in the spring.
The mammals in our area with long hibernating periods include the bats, bears (especially the black, grizzly, and brown), badgers, skunks, ground squirrels, chipmunks, marmots, and jumping mice.
Shelter
Despite the severity of the arctic winter, many of the arctic mammals are active throughout it, with no more than the sheltered side of a hill, some bushes, or some rocks to break the wind. Among these are the deer, the musk ox, the

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rabbits, and many of the carnivores. Indeed, for some of these grazing animals the wind is a friend, for it sweeps the snow from their grazing grounds.
Many of the mammals that do make tunnels and use shelters apparently do it as much for escape from their enemies , or for feeding purposes as for ^^ protection from the elements.
Lemmings live much of their lives in burrows, underground or under the snow, partly correlated with their feeding, partly with their need of pro– tection from predators, and partly for shelter from the elements. On low fertile country in northern Greenland, long stretches are quite undermined by banded lemming tunnels, and sown with their holes. In the depth of winter the banded lemmings are rarely found above the surface of the snow, rather carrying on their winter life in the system of passages between the layer of snow and the surface of the ground. Here they find their food and make their nests. Only when spring approaches do they begin to wander over the surface of the snow and then, sometimes, a frozen crust traps the animals, preventing them from digging down into the snow again, and they perish.
Both food and shelter lead the weasel under the snow in the depth of winter in the Far North. It digs down through the snow to the system of lemming tunnels, between the snow and the earth, and ^ , ^ following these, preys on the lemmings. It ^^ has been suggested , that the shelter of the snow blanket protecting these animals from the cold is essential, and a winter with little snow may prove fatal to many lemmings and weasels.
The pika of the rocky areas spends most of its life in and about ^ the ^ crevices of the rocks, only venturing a little distance into the meadows in search of herbs and grasses and then darting back to shelter. During the winter, with

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its store of food laid by, it does not have to leave the shelter of its rocky fastness at all. The ground squirrels and the marmots make deep burrow ^ s ^ , about ^^ which their lives center. These may be largely for protection from enemies in summer, but are also their retreats for the long sleep of winter.
In the taiga, voles and shrews make tunnels and runways through the soil, the vegetation, and the snow, and generally live in cracks and crannies; but tiny shrews may be abroad on the snow even when it is far below zero, as their lacy tracks show. Tree squirrels make warm nests of twigs and fiber in trees in the more southern parts of the taiga, but in the north depend more on under– ground burrows for sleeping places. Muskrats and beaver spend much time in the water and make burrows with underwater entrances; both make dome-shaped “houses” in the water, but the beaver with its larger size and requirements make ^ s ^ the more elaborate system. To ensure that there will be enough water for its needs, the colony of three to ten animals (typically a beaver family consists of the two adults, the two or three yearlings, and the several young of the year) make a dam of sticks, other vegetation, and mud, above which a pond full of water accumulates. Out in this pond, resting on the bottom, the beaver builds its house of sticks, which may be six feet or more across, and as many high above the water. Hollowed out inside, above the water level, is the ne x ^ s ^ t chamber, ^^ with an underwater entrance and exit. In winter, the beaver is completely shut off from the outside world by his frozen roof and the frozen surface of the pond, and, if it were not for human trappers, it would be most secure from every enemy and the elements.
Mammals which ordinarily do not use shelters in their everyday lives may make them for the breeding season, as places in which the young are raised. This is well illustrated by the deep dens of the wolves and foxes.

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Mammal Interrelationships
In an area where species are few, some close interrelationships between animals develop, recalling the well-known associations between animals and plants such as the dependence of the caribou and reindeer on reindeer moss (lichens) and squirrels on conifer seeds. Wolves follow both caribou and reindeer herds; the presence of these animals probably makes possible the presence of wolves on the tundra. The abundance of the arctic fox is so closely correlated with the abundance of lemmings that the arctic fox increases and decreases with the increase and decrease in the numbers of lemmings. A natural check has demonstrated the validity of this: in southern Greenland where there are no lemmings, the foxes may fluctuate but there is no regular variation in numbers corresponding to those in the arctic fox in Baffin Island opposite where lemmings do occur. The patterns in a relationship may vary from place to place, giving interesting correlations.
The red fox, in Ungava, seems to depend for food on the lemmings and wolves ^ voles ^ , ^^ with a four-year cycle of abundance; the red fox also has a four-year cycle. In Ontario, where the fox depends more on the snowshoe rabbit, with a ten-year cycle of abundance, the fox, too, tends to follow the ten-year cycle of its favorite prey. The dependence of the lynx abundance on that of the varying hare has been mentioned under “Fluctuations in Numbers,” p.000.
The mountain sheep is an animal of the mountains, where it can easily out– distance the wolf. On level country, the sheep is at a disadvantage and it is probabl y ^ e ^ that the pressure of wolf predation on mountain sheep has been an ^^ important factor in restricting it to its present habitat in the mountains, and has perhaps been a factor in its evolution as a mountain animal.

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Herds of caribou and reindeer attract arctic foxes, both summer and winter, and, during the fawning season of reindeer at least, the foxes occasionally attack weak fawns.
In summer, with lemmings common and easy to secure, the arctic fox lives bountifully; in winter, with food scarce, the arctic fox may attach themselves to polar bears and follow them out onto the sea ice, trying to get part of the meals of the polar bear or taking his leavings, and even eating their badly digested excrement with voracity. When a polar bear is lying asleep, the attendant arctic fox may curl up on an ice block nearby.
On the tundra where some hummock makes a resting place and lookout for birds of prey, their droppings enrich the soil there and the vegetation s grows more luxuri eua ^ a ^ ntly, making a good feeding place for lemmings. The same has been ^^ recorded for the mounds at the entrance to arctic fox burrows on the Siberia tundra. An interesting result is that the carnivores unwittingly provide a good feeding area for lemmings and attract them to places where they are more quickly found.
When lemmings are plentiful, the wolves may prey entirely on them, leaving the caribou in peace; thus the lemmings may affect caribou numbers.
Mammals in Relation to Man
Advantages . The mammal life of the sea and the land has made human life possible in the North. While sea mammals and fish have been an important food source, some of the natives have depended entirely on land mammals, and all have drawn heavily on them. The mammals, especially the caribou and the moose, supplied staple meat, although some Indians used hares extensively and one group became known as : ^^ Hare Indians.” The meat, killed in abundance when the ^^

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animals were plentiful, was eaten fresh, raw or frozen, or cooked; it was preserved for future use in caches where the cool climate retarded its dis– integration; and some was wind-dried (by Eskimos) or smoke-dried (by ^ ^ Indians) for better preservation. To make a still more nourishing and lasting fare, dried meat was sometimes pounded and mixed with fat to form pemmican that would keep for years.
No part of the animal was wasted; what is often regarded as waste (the blood, marrow, and intestines) was eaten, and with the caribou even the con– tents of the paunch. Not only are “game” animals so used, but even foxes, wolves, ground squirrels, and lemmings may be eaten. Besides eating the animals themselves, the Eskimos would sometimes seek, with the aid of dogs, the roots stored by the voles on the American and Eurasian tundra to add these tasty items to their own fare.
Not only did the land mammals provide food; they supplied clothing too: the caribou hide supplied material for warm, light garments, ideal for arctic winter wear and for which no satisfactory substitute has yet been found. Other skins were also used, especially polar-bear skins for pants by the Eskimos; Indians used mooseskin for tanned leather; on occasions an inland mountain– dwelling Eskimo group might dress completely in sheepskin; skins of marmots and ground squirrels were also used for jackets and robes, and skins of hares where woven into robes. The mammals also supplied the material for sewing, sinews still being the most satisfactory material for sewing skins into clothing.
The sinews and strips of hide provided material for snares and fish nets; musk-ox horns were spliced together to form bows, and horn and bone were used for tipping spears.

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In Eurasia, the wild reindeer provided the ancestor of the domestic reindeer, on which an important culture of the Eurasian tundra exists.
Not only was the land mammal fauna important to the original human in– habitant, but early traveling white men, explorers, and scientists would have been unable to make the travels and the studies they did without the caribou and the musk ox to supply them with meat, and some prospectors in the North still depend on the country for their meat.
With the advent of traders and the market for furs, the residents of the Arctic and Subarctic grew to depend on civilization for many things, such as flour, tea, tobacco, and weapons of the chase. To pay for them, the fur trade became important and is still the main support of many northern people and the main economic productiveness of large areas.
Disadvantages . While the land mammals are an important scource of support for the arctic dweller, they affect man adversely at times.
Unprovoked attacks on man, of course, stand out although they are rela– tively few in number. Wolves in the New World are not known to attack man unless the wolf is obviously deranged, though they have been known to kill sledge dogs wandering too far from camp. In the Old World, however, wolves have long been a real danger to human life. In Russia, the wolf was man’s chief enemy in the animal world, and the peasants pronounced a spell on St. George’s Day as a protection against the wolf. The present-day prejudice and fear of the wolf in the New World is probably based, in part, upon the very real fear in which it was held in the Old World.
Bears are always potentially dangerous and especially so when they have cubs. Probably a bold front will usually avert an attack, but not always. Richardson, while saying that grizzly bears usually ran away from man, gave

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one account of a grizzly which molested a party of voyageurs . They “had been employed all day in tracking a canoe up the Saskatchewan, had seated themselves in the twilight by a fire, and were busy in preparing their supper when a large grizzly bear sprang over their canoe…and seizing one of the party by the shoulders carried him off.” One of the men followed and rescued the bear’s victim by shooting the bear. The polar bear has been known to come to the edge of the ice and watch an approaching boat full of hunters with the apparent intention of throwing itself in among the hunters; the polar bear has been recorded as stalking humans and killing them apparently for food. However, it has been suggested that many cases of polar bears approaching humans are the result of curiosity.
Any large strong mammal may be dangerous at close quarters, but only two others need mention here. the musk ox has been recorded as making unprovoked attacks on men, and the bull moose, in the rutting season, occasionally attacks man.
Where supplies of food are put down in caches for future use, the destruct– tion of these caches can be extremely serious. On the tundra, polar bears may do this. They may enter huts, break up boxes, smash cans of meat and eat the contents, and even drag away the stovepipe. The arctic fox is also a per– sistent robber of meat caches, squeezing through almost incredibly small crevices between the stones covering the cache, and the unfortunate travelers have returned to find their meat supply gone.
On the barrens and in the forest, the wolverine is universally known as the worst and most persistent robber of caches, cabins, and trap lines. If a wolverine has time to work undisturbed, there are few caches he cannot enter; he rolls away heavy stones and logs; he gnaws through fixed timbers; he climbs

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to elevated caches and excavates buried once. He eats what he can and carries away not only food but articles he cannot use, such as guns and spyglasses. Although he is very destructive and very difficult to catch, some northern dwellers have come to consider a certain amount of wolverine predation as inevitable, and, like the annoyance of mosquitoes, take this as a matter of course.
Black bears in the forest country have a deservedly bed name for destroying unattended camps, breaking open and ravaging cabins, and smashing canoes, doing much more damage than the mere rifling of the stores would necessitate. Trappers, returning to their trap lines in the fall, have had their plans seriously upset by finding a black bear has paid their outfit a visit.
The smaller beasts, shrews, mice, and small carnivores, may all levy toll on man’s stores. Generally, because of their smaller size, these activities tend to be more annoying than serious, although red-backed voles have removed 40 pounds of rolled oats in a 3-week period, and squirrels, in addition to eating delicacies, may take buttons off underwear or chew up clothing for nests.
The only poisonous item in the land mammals of the North is the liver of the polar bear. It has a probable excessive amount of vitamin A, which may cause violent sickness when eaten by humans.
SYNOPSIS OF NORTHERN MAMMALS
As stated previously, from the northern Arctic southward, the number of species tends to increase. Each species has its own tolerance and its own requirements; the environmental conditions change gradually from tundra to forest, from northern forest to southern forest. Some mammals have their headquarters in one or other of the habitats. Others, more common elsewhere,

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ranging into northern Europe, they barely get north into our area.)
Bats: Chiroptera
These flying mammals are chiefly found in the tropics, where species and individuals swarm. Insect-eating species would find an abundance of food in the mosquitoes of the arctic summer, and some more northern species hibernate or m o ^ i ^ grate. Bats are nocturnal and the lack of darkness in the North may re– strict their northward range. In America they are rare (genus Myotis ) or accidental in our area. However, in Eurasia at least one species (genus Eptesicus ) ranges to the arctic coast in the west and onto the tundra in Siberia.
Flesh- ^ E ^ aters or Carnivores: Carnivora
Bears (Family Ursidae) . The white polar bear (genus Thalarctos ), drawing much of its sustenance from the Arctic Sea, is circumpolar in distribution. The brown bears of Europe and Alaska and the grizzlies of western North America (genus Ursus ) range commonly on the barrens and the forest. The black bear of North America (genus Ursus or Euarctos ) is a forest bear and ranges to the timber line.
Wolves and Relatives (Family Canidae) . The wolf (genus Canis ), of both tundra and taiga, is circumpolar. The arctic fox (genus Alopex ), a true polar animal among those ranging farthest north, is also circumpolar in distribution. And the red fox (genus ( Vulpes ), very similar in Eurasia and America though ^^ sometimes considered as different species, is a widespread forest animal extend– ing its range onto the edge of the tundra. In the New World, the coyote ( Canis latrans ), a more southern animal of open forests and plains, has in recent years extended its range greatly into our area and now reaches the arctic coast in the west. In Eurasia, the red dogs (genus Cuon ) of Asia are more southern, though ranging north to the Amur district of Siberia.

EA-Zoo. Rand: Terrestrial Mammals

Cats and Relatives (Family Felidae) . Only the lynx (genus Lynx ), one species in America and one in Asia, are truly boreal. They are animals of the taiga ranging into the neighboring tundra in years when they are plentiful. In America, the puma ( Felis ^ Felis ^ concolor ) strays north almost to our area; in Asia, the tiger ( Felis tigris ) ranges north only to about the latitude of Lake Baikal.
Weasels and Relatives (Family Mustelidae) . Two species of weasels (genus Mustela ) are circumpolar: the least weasel and the short-tailed weasel or ermine. In North America, there is the mink (genus Mustela ) which extends north to timber line; in Eurasia the kolinsky or yellow weasel, a large brownish weasel (as fur it ranks just below the mink) extends from the Sea of Okhotsk to European U.S.S.R. in the forest area; in Europe, the European mink ranges north to the Arctic Sea.
In the New World there are one or two species of ^ marten ^ (genus Martes ); in the ^^ Old World there are two; the sable, ranging from Kamchatka to European U.S.S.R., and the pine marten of northern Europe. All are closely related and are largely geographical representatives. In North America another close relative, the fisher, barely extends north into our area.
The European badger (genus Meles ) ranges north to reach the Arctic Sea at the White Sea; the American badger (genus Taxidea ) is more southern.
In North America, the otter (genus Lutra ) of the waterways is a rare animal of the streams of the forest taiga, extending north to the Mackenzie Delta; in Eurasia, related species occur.
The wolverine (genus Gulo ) is ^ a ^ circumpolar species group (or two closely related species), more typical of the taiga but wandering commonly into the barrens.
The skunk (genus Mephitis ) is a New World group with one species, the striped skun i ^ k ^ , barely intruding into the southern part of our area. ^^

EA-Zoo. Rand: Terrestrial Mammals

Hares, Rabbits, and Pikas: Lagomorpha
Hares and Rabbits (family Leporidae) ^ (family Leporidae) ^ . The arctic hares of North Americ [: ] ^ a ^ ^ ✓ ✓ ^ and the snow hares of Eurasia are all closely related animals (genus Lepus ) of the tundra. In North America, there is also the varying hare or snowshoe rabbit (genus Lepus ) of widespread distribution in the taiga.
Pikas or Rock Rabbits (Family Ochotonidae) . These small relatives of the hares (genus Ochotona ) occur in both Eurasia and America. In the New World they are confined to the mountains of the west; they also occur in the mountains of Europe and Asia, and north to northern Siberia where they extend nearly to sea level and the Arctic Sea.
Rodents: Rodentia
Squirrels and Relatives (Family Sciuridae) . Tree squirrels (genus Sciurus ) are common important taiga animals, ranging north to the limit of trees. They are represented in America by the red squirrel and in Eurasia by the common squirrel. A flying squirrel (genus Glaucomys ) inhabits the American taiga, and a distantly related species (genus Pteromys ) occurs in that of Eurasia, going north of the Arctic Circle.
Chipmunks (genus Eutamias ) are small, active, terrestrial squirrels with stripes. In America, one species ranges northward into the western part of our area; in Eurasia, a related species ranges north of the Arctic Circle in Siberia.
In America one species of ground squirrels ^ (genus Citellus ) ^ occurs in the Western Arctic; ^^ in Eurasia related species (called suslik ) range well north in Siberia.
Marmots (genus Marmota ), in America called woodchucks or ground hogs inhabit the meadows and open forests and extend into our area in the west. The hoary marmot ranges, in Alaska, north to the mountains facing the Arctic Sea, and in Siberia and Kamchatka. Other species occur eastward into the mountains of Europe,

EA-Zoo. Rand: Terrestrial Mammals

Beaver (Family Castoridae) . Related species (genus Castor ) occur in the streams of the taiga of Eurasia and America.
Voles and Relatives (Family Cricetidae) . This family of rodents contains two subfamilies: the Microtinae (voles and lemmings) and Cricetinae (wood m mice). The voles and lemmings, small vegetarians, have two circumpolar speci al ^ e ^ s groups ^^ on the tundra: the banded lemming (genus Dicrostonyx ) that changes to a white winter coat, and the brown lemming (genus Lemmus ) that does not. Other voles, of the genera Microtus and Clethrionomys (meadow mice or voles, bank voles, water voles, red-backed voles, etc.) and related forms are plentiful as to species and individuals in the taiga and spread out onto the tundra of both the Old and the New Worlds. The most striking of these is the huge (for the group) muskrat (genus Ondatra ) of the American taiga, which has been introduced into parts of Eurasia.
The white-footed mice (genus Peromyscus ) of America range through much of the taiga, and a pack rat (genus Neotoma ) ranges in the mountains of the west. In Eurasia their place is taken by the distantly related wood mice (genus Apodemus ), which tend to be more southern.
Jumping Mice (Family Zapodiae) . In America one species intrudes into the Mackenzie (genus Zapus ); related species in Eurasia are more southern, restricted to the Chinese area.
American Porcupines (Family Erethizontidae) . The Canada porcupine (genus Erethizon ), a forest animal, ranges north to the limit of trees. It is a member of a New World group with headquarters in the American tropics. In the Old World, the porcupines, belonging to quite another group (family Hystricidae) do not range as far N north as our area.

EA-Zoo: Rand: Terrestrial Mammals

Hoofed Mammals: Artiodactyla
Deer (Family Cervidae) . Caribou and reindeer of several species (genus Rangifer ) are the chief deer of the circumpolar tundra and range into the taiga; the circumpolar species group Alces , called moose in America and elk in Eurasia, is restricted to the taiga.
In the New World the mule deer (genus Odocoileus ) intrudes into the southern edge of the taiga; in the Old World the musk deer (genus Moschus ) extends north almost to the Arctic Circle.
Sheep and Relatives (Family Bovidae) . Mountain sheep (genus Ovis ), in America, occur only in the west above the timber line; in Kamchatka and the mountains of Siberia are related species.
Bison of America (genus Bison ), which still exist under protection, used to extend into the southern part of our area; in Europe the related European bison or aurochs (wisent) is about extinct.
The musk ox (genus Ovibos ) is an animal of the American tundra, with no living Eurasian representative. The mountain goat of western American mountains (genus Oreamnos ) occurs only in the southern part of our area; its relatives are in the mountains of central Asia and Europe.

EA-Zoo. Rand: Terrestrial Mammals

BIBLIOGRAPHY

1. Anderson, R.M. “Mammals and Birds,” Bethane, W.C. ed., Canada’s Western Northland , Ottawa, Patenaud ^ e ^ , 1937, pp.97-122. ^^

2. ----. “Mammif e ^ è ^ res de la Province de Qu e ^ é ^ bec,” Soc. Provancher, Quebec. ^^ Rapport Annuel, 1939, pp.37-111.

3. Anthony, H.E. Field Book of North American Mammals . N.Y., Putnam, 1928.

4. Brehm, A.E. Brehms Thierleben, Allgemeine Kunde des Thierreichs. Vol. 1. Die S a ^ ä ^ ugethiere. Pts.1-2 . Grosse Ausg. 2. ^^ umgearb. und verm. Aufl. Leipzig, Verlag des Bibliographischen Institut e ^ s ^ , 1876-77. ^^

5. Hesse, Richard, Alee, W.C. and Schmidt, K.P. Ecological Animal Geography . N. Y., Wiley; Lond., Chapman & Hall, 1937.

6. Kobelt, Wilhelm. Die Verbreitung der Tierwelt . Leipzig, Tauchnitz, 1902.

7. Miller, G.S. Catalogue of the Mammals of Western Europe (Europe Exclusive of Russia) in the Collection of the British Museum . Lond., The Museum, 1912.

8. Ognev, S.I. Zveri ^ S ^ SSR i Prilezhashchikh Stran; Zveri Vostochnoi Evropy i ^^ Severnoi Azii . (The Mammals of Russia (USSR) and Adjacent Countries; the Mammals of Eastern Europe and Northern Asia.) Moscow, Akademii Nauk, ^ S ^ SSR, 1928-40. Vols.1-4. ^^

9. Rand, A.L. Mammals of Yukon, Canada . Ottawa, Cloutier, 1945. Nat.Mus.Can. Bull . 100.

10. Sclater, W.L. and Sclater, P.L. The Geography of Mammals . Lond., Paul, Trench, Trübner, 1899.

11. Seton, E.T. Lives of Game Animals . N.Y., Doubleday, 1925-28. Vols.1-4.

12. Simpson, G.G. “Holarctic mammalian faunas and continental relationships during the Cenozoic,” Geol.Soc.Amer. Bull . vol.58, pp.613-88, 1947.

13. Tate, G.H.H. Mammals of Eastern Asia , N.Y., Macmillan, 1947.

14. Winge, Herlaf. “Gr o ^ ø ^ nlands Pattedyr,” Medd.Grønland , vol.21, pp.319-521, ^^ 1902.

A. L. Rand

Amphibians

EA-Zoology (Karl P. Schmidt)

AMPHIBIANS

PHOTOGRAPHIC ILLUSTRATIONS
With the manuscript of this article, the author submitted one photograph for possible use as illustration. Because of the high cost of reproducing as halftones in the printed volume, only a small proportion of the photo– graphs submitted by contributors to Encyclopedia Arctica can be used, at most one or two with each paper; in some cases none. The number and selection must be determined later by the publisher and editors of Encyclopedia Arctica . Meantime all photographs are being held at The Stefansson Library.

EA-Zoology (Karl P. Schmidt)

AMPHIBIANS
Amphibians and reptiles whose body temperature fluctuates with that of the environment must be able to hibernate safe from frost. Complete freezing, i.e., freezing of the heart, kills them. It appears that the extremities of cold-hardy frogs and salamanders may freeze without permanent injury; and it is likely that the freezing point of the body fluids, concentrated by the freez– ing of the extremities, is well below 0°C. Thus, north of the line of perma– nently frozen subsoil, amphibians find no safe refuge, except in hot spring ^ s ^ . ^^ Cold hardiness in amphibians requires that the development of the eggs, de– posited in water, may take place at or near the freezing point.
Among amphibians, only two species of frogs range into the Subarctic in the Western Hemisphere. Both of these reach Great Bear Lake, which is crossed by the Arctic Circle. These are: Pseudacris migrita septentrionalis , the northern swamp tree frog, and Rana sylvatica latiremis , the northern wood frog. These forms are characterized by the shortness of their legs, and they merge with a uniform gradient of increasing leg length (a geographic cline) into the related forms at the south. Thus the establishment of a southern limit for the northern subspecies is entirely arbitrary. In a broad sense the swamp tree frog ranges south to the coast of the Gulf of Mexico, and the wood frog to Arkansas.
The swamp tree frog does not appear to be known from Alaska or from Labrador.

EA-Zoo. Schmidt: Amphibians

The wood frog ranges to the Yukon and Mackenzie deltas and has been recorded also from the Labrador coast at Jack Lane Bay, at about latitude 56° ^ ^ N.; it is ^^ still abundant at Fort Chimo, Ungava Bay, north of 58°.
The Rocky Mountain toad, Bufo boreas boreas , reaches southern Alaska and the Peace River district of Alberta. A northern race of the American toad, Bufo terrestris copei , reaches Hudson Bay, as do the mink frog, Rana septentrionalis , and the leopard frog, Rand pipiens .
The Old World amphibians that range to the 60th parallel or farther north include several species of salamanders and frogs and toads.
Salamanders
Hynobius keyserlingii , a salamander with no vernacular name, is found from the northern Urals to Kamchatka, ranging north of the Arctic Circle at Verkhoiansk.
Triturus cristatus cristatus , the crested newt, inhabits central Europe and goes as far no ^ r ^ th as 60° ^ ^ N. in Scandinavia and Soviet Russia. ^^ ^^
Triturus vulgaris vulgaris , the common European newt, ranges through central and northern Europe, north to 63° N. in Scandinavia.
Frogs and Toads
Bufo bufo bufo , the common European toad, ranges throughout northern Europe and Asia, north to the 60th parallel; the eastern limits are unknown.
Rana arvalis arvalis , the moor frog, inhabits northern Europe and Asia, north to the Arctic Circle. Another subspecies, Rarvalis issaltschikovi , has been described from Archangel.
Rana ridibunda ridibunda , the common European pond frog, is found in central Europe and north to th t e 60th parallel in Soviet Russia. ^^

EA-Zoo. Schmidt: Amphibians

Rana temporaria temporaria , the European grass frog, ranges through central and northern Europe and across Asia to Kamchatka, north to North Cape in Scandinavia, and to Verkho l ^ i ^ ansk in Siberia. ^^
Of the Old World frogs, the grass frog is by far the most abundant northern species, ranging well beyond the Arctic Circle. This species is directly related to the American wood frog.

EA-Zoo. Schmidt: Amphibians

BIBLIOGRAPHY

1. Boulenger, G.A. The Tailless Batrachia of Europe . London, Ray Society, 1898.

2. Hildebrand, Henry. “Notes on Rana sylvatica in the Labrador Peninsula,” Copeia , 1949.

3. Mertens, Robert, and Müller, Lorenz. “Die Amphibien und Reptilien Europas,” [: ] Senckenbergische Naturf.Ges. Abbandl . no.451, pp.1-56, 1940.

4. Patch, C.L. “Notes on northern woodfrogs,” Copeia , 1949.

5. Schmidt, K.P. “A geographic variation gradient in frogs,” Field Mus. Nat.Hist. Zool.Ser . vol.20, pp.377-82, 1938.

6. Werner, Franz. “Die nordlichsten Reptilien und Batrachier,” Fauna Arct ., Jena, vol.4, pp.527-44, 1906.

Karl P. Schmidt

Reptiles

EA-Zoology (Karl P. Schmidt)

REPTILES

Reptiles in northern climates like amphibians must be able to hibernate safe from frost or at least from severe frost. It is likely that their north– ward range can extend only slightly beyond the line of permanently frozen sub– soil. No reptile appears to range much beyond the 50th parallel in North America. In the Old World, however, no less than six species range north of latitude 60° N. and some of these pass the Arctic Circle.
Two of the northern species, the lizard known as the blindworm and the European viper, hibernate in aggregations, and some of these overwintering groups are mixed lots of the two species, no doubt with occasional representa– tives of the smooth snake and perhaps of other species present.
There is an evident advantage of viviparity for breeding in the North, the gravid mother being enabled to follow the sun and thus maintain herself and her developing young at a higher temperature than is available for eggs deposited in the ground. This appears to be correlated with the fact that the two species that range well beyond the Arctic Circle, the mountain lizard and the viper, produce living young, as does the blindworm, which ranges nearly as far north.
The scientific and common names of the northernmost reptiles, and their approximate distributions are given below.
Lizards
Lacerta agilis agilis , the sand or fence lizard, inhabits central Europe northward beyond 60° N. in Sweden, Finland, and western Soviet Russia.

EA-Zoo. Schmidt: Reptiles of the Arctic Region

Lacerta vivipara , the mountain or viviparous lizard, is found throughout central Europe across northern Asia to Sakhalin Island, north to North Cape in the Scandinavian Peninsula and to the Arctic Sea in Soviet Russia.
Anguis fragilis , the blindworm or Blindschleiche , is found in Europe, north to the Arctic Circle in Sweden and Finland.
Snakes
Natrix natrix natrix , the common European water snake, is found in central and northern Europe to central Asia, northward in Sweden and Finland to 64° N.
Coronella austriaca austriaca , the smooth snake, exists in Europe, north to 63° N. in Norway, and north to 64° N. in Sweden.
Vipera berus berus , the common European viper, is located in central and northern Europe and across northern Asia to Sakhalin Island, and north beyond the Arctic Circle in the Scandinavian Peninsula.

EA-Zoo. Schmidt: Reptiles

BIBLIOGRAPHY

1. Hecht, Günther. “Zur Kenntnis der Nordgrenzen der mitteleuropäischen Reptilian,” Berlin. Univ. Zool.Mus. Mitt . Vol.14, pp. 501-95, 1929. Text fig. 1-15, pl. 3.

Karl P. Schmidt

Caribou and Reindeer

Caribou

(EA-Zoo. A. W. F. Banfield)

CARIBOU
CONTENTS

Scroll Table to show more columns

Page
Status of Geographical Herds 2
Barren Ground Caribou 2
Alaska Peninsula Caribou 3
Osborn Caribou 3
Stone Caribou 4
Ungava Caribou 4
Polar Caribou 5
Western Woodland Caribou 6
Eastern Woodland Caribou 7
Newfoundland Caribou 7
General Aspects 7
Range 7
Physical Characteristics 8
Habits 9
Disease ^ s ^ 12 ^^
Enemies 13
Bibliography 16

EA-Zoology (A. W. F. Banfield)

CARIBOU
When the prairies of western North America were first explored by white men, they supported vast herds of bison, which formed the basis of the economy of the Indian tribes. By 1890, with the advance of agriculture, ranching, and the transcontinental railway, the bison had been reduced from its millions to the verge of extinction.
The first explorers to the arctic grasslands of North America also found innumerable herds of another big-game species — the caribou (Rangifer arc–tious). They, too, provided she ^ l ^ ter, clothing, and food for the native popu– lations of Eskimos and Indians. The bison has been extirpated from its mid– continental plains habitat with the exception of a few herds in national parks, but the caribou still roams the arctic prairies in numbers roughly comparable to primitive conditions, largely because its domain has not yet been fully exploited by civilization.
The caribou is still the cornerstone of the economy in large areas of northern North America. It serves primarily as a staple source of food. The hides supply arctic clothing, beddings, and shelter; other parts, such as the antlers and sinews, are utilized. Residents of isolated inland settle– ments in northern Canada and Alaska, both native and white, are largely de– pendent on the caribou for their continued existence in remote areas.

EA-Zoo. Banfield: Caribou

The recent increased interest in northern development in Alaska and Canada has drawn attention to the precarious state of many bands of caribou. In the past decade (1940-49), improved northern transportation facilities and renewed interest in the Arctic have made possible the collection and publication of data on mammal distribution. It is now possible to take stock of this important natural resource more accurately than at any previous time. The present status of North American caribou has been reported by Banfield (3).
The present classification of the genus Rangifer in North America is generally considered unsatisfactory because of the scarcity of representative specimens. The most generally accepted classifications are those of Anderson (1) and Murie (15). They in turn have accepted the work of Jacobi (9). Within the area covered by this Encyclopedia, nine ^ seven ^ races of four ^^ species are recog- ^ ][: Ok FB] Au. O.K? ^ nized. The present status of these races is discussed below.
Status of Geographical Herds ^ ][: Ok FB]? ^
Barren Ground Caribou ( Rangifer arcticus arcticus ). This race, which is by far the most numerous, includes the several large herds of continental arctic prairie caribou of the Mackenzie and Keewatin Districts of the Northwest Territories. Basing his calculation on the carrying capacity of the land, Anderson (2), in 1938, estimated a population of approximately 3,000,000 ani– mals. A recent preliminary aerial survey (1949) has indicated that, though present numbers are probably less than this estimate, primitive abundance in the central portion of their range is still undiminished.
The nomadic migrations of these caribou during the past ten years have brought them in winter well into the northern portions of the p ^ P ^ rairie p ^ P ^ rovinces and west to the Mackenzie River valley. In the northeast section of their

EA-Zoo. Banfield: Caribou

range, caribou have been greatly reduced in numbers by heavy hunting pressure of the Eskimos. Over large areas, caribou are reduced to small nomadic bands which no longer carry out the characteristic summer migrations to and from the lower arctic islands and peninsulas. According to Manning (12), scattered bands are still to be found on Melville Peninsula. Caribou are still fairly plentiful along the arctic coast, from the mouth of the Back River west to the Horton River. Gavin (7) relates that in the Perry River area large num– bers appear in the early summer and bear their fawns on small coastal islands. In recent summers, caribou have occurred in increasing numbers on the Kent Peninsula and a few have crossed to Victoria Island. On that island, however, only a few individuals are regularly observed.
According to Manning (12), the population of the species on Southampton Island has been similarly reduced to isolated bands. Nearly Coates Island, which has no resident native population, supports a fairly large herd of caribou from which numbers are taken from time to time by journeying Eskimos.
The caribou population on Baffin Island is found chiefly along the western coast, bordering Foxe Basin. The Eskimo population is largely distributed on the eastern and southern shores of the island. Movement of Eskimos to the western coast to utilize these remaining herds has been intentionally dis– couraged.
Alaska Peninsula Caribou ( Rangifer arcticus granti ). This race of caribou occurs, according to Murie (15), from Unimak Island east to Port Heiden and Becharof Lake.
Osborn Caribou ( Rangifer arcticus osborni ). The Alaska Highway has not only increased our knowledge of the distribution of this race of caribou in northern British Columbia and southern Yukon Territory, but has greatly increased

EA-Zoo. Banfield: Caribou

its vulnerability to hunters. It seems that these caribou are restricted to isolated bands inhabiting alpine ranges. The seasonal movements of this race are altitudinal or local in nature.
In western Yukon Territory this race of caribou occurs, according to C. H. D. Clarke, in the Kluane Reserve area, from Donjek River north to the White River, where their range is occasionally overrun by winter incursions of Stone caribou. To the east, bands occur in central Yukon, along the Canol Road to the Macmillan Pass (17). S am ^ ma ^ ll bands occur on the Mackenzie District ^^ side of the pass, along the Keele River. An aerial reconnaissance of the upper Nahanni River, in early 1948, indicated a moderately numerous caribou population.
Stone Caribou ( Rangifer arcticus stonei ). This is the migratory caribou of the Alaska mainland and arctic coast, east to the Mackenzie River delta. According to Murie (15), there are several separate herds in the following areas: Alaska range, arctic coast ^ , ^ and Yukon-Tanana region. Dufresne (5), in 1942, estimated a population of about 4 ^ 5 ^ 00,000 caribou in Alaska. More recent reports suggest a decrease in this number.
There are at present two centers of abundance in the Yukon Territory, Canada. The northern herds are found on the plains between the mountains and the arctic coast during the summer. During autumn they migrate south through the Mackenzie Mountains, west of Aklavik and Fort McPherson, to the upper Porcupine and Peel rivers. In recent years these herds have shown only a gradual decline in numbers. In the second center of abundance, west of Dawson, numbers have been greatly reduced. The Yukon River crossings at Forty - mile, Dawson, and Stewart are no longer in frequent use.
Ungava Caribou ( Rangifer caboti ). Migratory caribou were formerly

EA-Zoo. Banfield: Caribou

abundant on the unforested lands of the Ungava Peninsula and were an important natural resource for the Eskimos. If the peninsula is considered as a whole, a gradual decrease in numbers has occurred (13). In several areas the decrease was comparatively rapid, due in part to emigration. Recent fluctuations in local populations are believed to be caused by local movements of small herds.
Large forest fires during the latter part of the nineteenth century have been generally blamed for the decrease in population. Using information ob– tained by aerial reconnaissance, Manning (11) estimated, however, that only 8 per cent of the caribou lichens were destroyed by fires in the wooded area. In excellent winter caribou range bordering the treeless country, he saw prac– tically no sign of fires.
The present range of the Ungava caribou seems to have two fbci of abundance. On the east side of Hudson Bay, caribou are to be found in small numbers in an area that extends from the headwaters of the Povungnituk River south to the Seal lakes, and includes the headwaters of the Larch and Leaf rivers. Bands also occur in an area that includes the vicinity of the Quebec-Labrador boun– dary at latitude 56° to 57° N., part of the George River basin, and the upper Whale River basin.
Polar Caribou ( Rangifer pearyi ). The range of this small white caribou is restricted to the northern arctic islands. It has suffered less human interference than any other species of caribou. It was previously thought it was ^ to be ^ present in considerable numbers over most of its range. Recent explora– tions, however, have indicated that much of this territory is unsuited for caribou range because of the lack of vegetation on mountains and the presence of extensive icecaps.
Polar caribou are reported from Banks Island and no ^ r ^ thern Victoria Island,

EA-Zoo. Banfield: Caribou

but the center of abundance seems to be on the islands north of Lancaster Sound. They occur in some numbers on Prince Patrick and Melville Islands. On Ellesmere Island, caribou occur in small numbers on the small erosion plains at the heads of the numerous fjords.
Caribou also are reported from the northwestern coast of Greenland as far north as the Thule area, but they are practically extinct there now. Distribution along the west coast ranges from Upernivik District southward to the Frederikshaab District.
Along Greenland’s northeast coast, caribou have entirely disappeared, ^ — Au: oK? [: Ok FB] ^ although they were frequent there in former days, notably around Scoresby Sound, where the Ryder Expedition (1891-92) encountered numerous herds. The Danmark Expedition (1906-08) found traces of caribou (old, castoff antlers and ex ^ c ^ rements) as far north as Holm Land (80° 24 w ^^ N.), and the First Thule ^^ Expedition (1912) found similar remains as far north as Adam Biering Land and Vildt Land, at the head of Independence Fjord, southern edge of Peary Land. The finds of the Thule Expedition have been confirmed by the Danish Peary Land Expedition (1948-50), as indicated by a preliminary account of its geographic work given by Fristrup (6). The account states specifically that caribou formerly inhabited Peary Land. So far, no traces of caribou have been found in upper Peary Land, that is along its northern shores (10).
Peary, who hunted caribou both on Ellesmere Island and in the Thule District of northwest Greenland, found that the Greenland specimens differed somewhat from those on the Canadian side of the Smith Sound route. “I have seen many winter coats of the Greenland Caribou and they are pronouncedly darker than the Ellesmere specimens” (16).
Western Woodland Caribou (Rangifer caribou sylvestris). The main centers

EA-Zoo. Banfield: Caribou

of abundance of the woodland caribou are south of the region treated in this Encyclopedia. In the Mackenzie River valley of the Northwest Territories, woodland caribou occur in small isolated bands. They have suffered from range destruction by forest fires. Heavy hunting pressure has further restricted their range to areas remote from settlements and river routes. They are to be found in the western half of Wood Buffalo Park. A recent aerial reconnais– sance (1949) has indicated a population of caribou south from the Liard River to the British Columbia boundary. They are also known from the Trout Lake area, west of Fort Providence. East of the Mackenzie River they are reported in the Horn Mountains, the Fish Lake area, and the Willow Lake area.
Eastern Woodland Caribou ( Rangifer caribou caribou ). The eastern woodland caribou has suffered habitat destruction by agriculture, lumbering, and forest fires. Its range in North America is no longer continuous, the species having been reduced to isolated herds. In Labrador, the woodland caribou are reported from the Cartwright area, the upper Hamilton River valley, and in the Mealy ^ Au: oK? [: Ok FB] ^ Mountains to the south. In Quebec, there are scattered herds north of the Gulf of St. Lawrence and in the Rupert River drainage. The discovery of large deposits of iron ore on the Quebec-Labrador boundary constitutes a new threat to the survival of caribou in the Ungava Peninsula. A herd occurs in the Gasp e ^ é ^ Peninsula and in 1949 the population of this herd was estimated at 1,500.
Newfoundland Caribou ( Rangifer caribou terra n enovae ). In the 25,000- square-mile caribou range in Newfoundland, and estimated number of 15,000 caribou was reported in 1949. There has been no decrease noted in recent years. The annual kill by residents is less than 300 animals.
General Aspects
Range . The range of caribou, in general, is farther north than that of

EA-Zoo. Banfield: Caribou

Any other member of the deer family (Cervidae) and extends to the northern limits of land, except that they now appear to be extinct in Peary Land. The range of the genus Rangifer ^ Rangifer ^ , like that of the wolf, is circumpolar. The members of the genus in Europe and Asia have been called reindeer; but recently the tendency, in North America at least, is to restrict the designa– tion reindeer to the various domestic subspecies, using caribou for the wild animals.
Physical Characteristics . The caribou is specially adapted to its northern habitat. The pelage is thick and long and the muzzle is furred. Another northern deer, the moose ( Alces Americana ), overcomes the difficulty of deep snow with its long legs. The caribou, moderately long legged, has little deep snow to contend with north of the tree line. When the caribou run, the hooves click. If a large herd of caribou is passing by, this “click-click” noise is loud and distinct.
In fresh autumn coat the caribou are cinnamon-brown with a white neck, “beard,” and a lateral stripe across the shoulder and along the flank. The abdomen, rump, tail, nose, and a ring above the ho f ^ o ^ fs are also white. The legs, chest, and lower flanks are darker brown. By late spring the old coat is much worn and has become pale-buff in color. At a distance the caribou then appear almost white. A new dark pelage grows during the summer months.
There is considerable range in the size of caribou in different parts of northern North America. Differences of size are among the characteristics that form the basis for separation of the caribou into several races. The Alaskan races are the largest, while the polar caribou ( Rangifer pearyi ) is the smallest. Murie (15) states that the average live weight of six Alaskan males in 366 pounds. The average dressed weight of eighteen males was 247

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pounds. The females are much smaller. The average total length of male cari– bou from the interior of Alaska is about 78 inches and the average height of shoulder is about 46 inches.
Both sexes carry antlers; those of the bucks are much larger than those of the does. Each antler consists of a main beam which sweeps back and out from the burr. Close to the burr are two tines which point forward. The first, which grows medially forward over the face, is called the brow tine. The two opposite brow tines grow close together and usually one is dominant and palmate in form. The second tine on the beam is called the bez tine. It, also, is generally palmate and grows anteriorly. Beyond these two tines the beam turns sharply up and gives rise posteriorly to the ^ a ^ number of terminal tines. There is great variation in antler formation; some are palmate, others more digitate. It is a general characteristic that the beams of cari– bou ( Rangifer arcticus ) are circular in cross section while those of the wood– land caribou ( R. caribou ) are generally flattened. The bucks grow and shed their antlers much earlier than the does.
Habits . Caribou possess a keen sense of smell and rely largely on this sense for warning of danger. Their eyesight is fair. They notice movements quickly but can easily be stalked upwind. The normal reaction when they are suspicious of danger is to circle to a position downwind from the object of curiosity. They do not seem to associate noise with danger. Caribou are well known for their insatiable curiosity. When alarmed they flee, but usually soon pause to look over their shoulders at the intruder. In large numbers they seem even more curious and bolder and sometimes may be approached closely.
Caribou are strong swimmers and readily cross large bodies of water in migration. In the water they float with the back and rump above the surface.

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It is suggested by Clarke (4) that the sound of running water attracts caribou to shallow crossings.
The caribou uses a variety of gaits. It seems to be a restless animal by nature. While slowly feeding along a ridge, it will suddenly break into an easy trot, only to resume a walk again after a few seconds. When alarmed, the caribou trots with head held high and legs thrown loosely forward and out. This gait can be changed to a fast pace with which the caribou covers great distances with seemingly little effort. When fully alarmed, the caribou will break into a gallop, but this gait is never maintained for a long period.
During the summer the bucks put on considerable back fat before the rutting season, which takes place during the autumn. Murie (14) observed small bands of does, each with a buck in charge, during the period September 14 to October 7. When the rut is on, the big bucks hard the does, spar with opponents, and often neglect to eat. This means that they enter the winter in poor condition. In Alaska the fawns are born between May 15 and June 15. They are reddish in color, with a black muzzle and dorsal line, and lack the spots of other fawns of the deer family. This coat fades to a buff color in July. A little later this coat is lost and fawns take on a darker coat similar to that of the adults.
The fawns are precocious and soon follow their mothers. Within a few days they can run as fast as the does, and by midsummer they are grazing for them– selves. The caribou does are solicitous mothers and seldom desert their young when in trouble.
During the short arctic summer, when the herds are on the open lands beyond the tree line, their food consists chiefly of green forage. During this season, the caribou seem to feed chiefly by browsing — stripping the leaves from the shrub willows ( Salix sp.), birth ( Betula glandulosa ), and

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bilberry ( Vaccinium uliginosum). They also graze on the grasses and sedges in the arctic swales. Typical plants utilized are: bent reed grass ( Cala–magrostis sp.), bluegrass ( Pea sp.), sedges (Carex sp.), and cotton grass (Erøophorum sp.). In late summer, mushrooms are eaten when found.
During the winter, lichens assume the major role in the caribou diet. The caribou easily paw through the snow to secure the ground vegetation. Some of the more important lichens are: Cladonia , Cetraria , Usnea , and Alectoria . Dried grasses, the leaves and twigs of many heaths, such as bearberry (Arcto–staphylos sp.) and crowberry (Empetrum nigrum), and the twigs of shrubs, such as willow ( Salix sp.) and aspen ( Populus tremuloides ), are also eaten.
Local movements seem to be governed by the food supply and the avail– ability of routes of travel. The lichens which from the major portion of the winter diet of caribou may take as long as fifteen to twenty years to recover from heavy utilization. Long migrations are a common characteristic of the caribou in North America. Along the routes the terrain is covered with parallel trails which the caribous follow in single file.
The populations of aborigines have learned to depend on these seasonal movements for their supplies of meat and clothing. Many theories have been advanced to explain to movements of the caribou. Marie (14) states that the migrations are probably due to several factors. There is the midsummer search for fresh feeding grounds, then the change from the grassy summer ranges to the winter lichen range. Coupled with these factors, the rutting activity stimulates movements in the autumn and the animals retrace ancestral routes through habit. In the spring there is a tendency to leave the forest because of the desire for a change to a grass diet and the urgency of the ap– proaching fawning season.

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Disease ^ s ^ . Caribou are subject to a variety of parasitic diseases. The most extensive parasitic condition of the caribou herds is that caused by the warble fly (Oedemagena tarandi). The warble fly is a large orange-and-black beelike fly which deposits its eggs on the hair of caribou during the summer. The larvae hatch, bore through the skin, and, by autumn, have moved to the back region. Here they become encapsulated under the skin, with small breathing holes through the skin. By May, the larvae are about three-quarters of an inch long. When mature, they enlarge the breathing holes and squirm through, dropping to the ground, where they pupate. From the pupae the adult flies emerge early in the summer. The effect of a heavy infection of warble larvae is to render the hide useless for clothing during a large part of the year.
A second parasitic fly which attacks caribou is the nostril fly (Cophe-^[: u: for trompe?][] Hadwen(1922) gives both for reindeer I dont know! FR^ nemyia nasalis). This is a large, dark fly. The larvae are deposited in the nostrils during the summer by the adult fly, which is viviparous. These migrate backward and lodge in the nasopharyngeal region. By early spring they have become much enlarged and form tight clusters in the throats of the animals. Caribou are often observed coughing^ ^ and sneezing at this period, probably because of the discomfort due to the presence of these larvae. The larvae detach themselves and drop out early in the summer, to pupate on the ground, where they develop into adult flies.
In the Arctic, there is a close association between many of the large mammals, including man, through a predator-prey relationship. Several para– sitic flatworms seem to have taken advantage of this close relationship to complete their complicated life histories. Thus the tapeworms ( Taenia hy–datigena, T. krabbei, and Echinococcus granulosus), which occur as adults in the intestines of wolves, coyotes, and domestic dogs, appear as larval bladder- worms in the livers, lungs, and muscle of caribou. The predators become

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parasitized by feeding on the infected caribou which had previously become parasitized by eating eggs of the tapeworms washed out of wolf or other f a eces onto the vegetation.
The close relationship of natives to both caribou and their domestic dogs has introduced the possibility of a secondary cycle, including man, which is distinct from the natural caribou-wolf-caribou cycle.
Several other diseases have been observed among caribou. Hadwen (8) ^ (7a) ^ recorded an infestation of a protozoan of the Sarcosporidia order. Murie (14) reported findings skulls showing necrosis of the jawbones. It seems very likely that caribou suffer from a disease similar to necrotic stomatitis.
There is some loss of life due to accidents. Waterfalls and rapids take a natural toll during migrations. Clarke (4) reported the findings of more than 500 carcasses at the foot of a falls on the Hanbury River in the Northwest Territories.
Enemies . With the exception of man, the wolf is the most important preda– tor of caribous throughout the greater portion of their range. In recent years North American caribou have undergone serious reductions in range and popula– tions. In several cases, the ranges are no longer continuous and the isolated herds are vulnerable to extirpation.
It is popularly believed that wolf predation has been the major factor in the caribou decrease. This view, when subjected to critical analysis, can– not be maintained. In many areas the decrease in caribou started before any local influx of or increase in wolves. It seems significant that the Barren- Ground caribou, which has had less contact with civilized man than the majority of other races, still occurs in numbers comparable to those found under prim– itive conditions, in spite of the presence of a relatively uncontrolled wolf

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population. It should also be remembered that caribou and wolves shared the same arctic ranges for countless years before the arrival of European man, yet the latter found caribou there in abundance.
Wolves take their greatest toll of caribou during the winter months, when they follow the migrating herds. During the summer months there is less wolf pressure on caribou because of the presence of other forms of prey, such as waterfowl and other nesting birds on the grasslands, ground squirrels ( Citel lus parryii ), and the young of other mammals, such as arctic hares ( Lepus arcticus ), arctic foxes ( Alopex sp.), and lemmings ( Lemmus sp.). These animals act as buffer species between the wolf and the caribou. ^ Au: OK? Ok FB ^
The second period when the caribous is particularly vulnerable to predation is during the fawning season. During the first week of their lives, caribou fawns are most vulnerable to the attacks of wolves ( Canis lupus ), as well as coyotes ( Canis latrans ), lynx ( Lynx canadensis ), and golden eagles ( Aquila chrysaëtos ). There are doubtful claims that the wolverine ( Gulo luscus ) preys on fawns and extremely doubtful ones that the grizzly bear ( Ursus sp.) does too. Although all these predators, but not including the wolf, manage to capture a few fawns at this season, these animals are so few in numbers and local in distribution as to have little effect on the survival of the caribou herds as a whole. Observations of grizzly bears and golden eagles hunting newborn fawns are given by A. Murie (14) and observations concerning the relation of the lynx to the caribou are given by O.J. Murie (15). Clarke (4) has estimated that a single wolf kills, on the average, twelve caribou per year, but that would be for sections where the wolf has many sorts of provender. In the Far North, where the caribou is practically the only food available through two-thirds of the year, the rate would be higher.

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Since the writings of Darwin, the theory of the survival of the fittest has achieved wide acceptance. In removing the less-fleet caribou, which are usually the aged, sick, or injured animals, the predator benefits the species by improving the stock. When wounded or aged animals are commonly observed hobbling in the rear of a migrating herd of caribou, one can justifiably con– clued that the wolf pressure is not excessive.
Harper (8) has pointed out the interesting fact that the Queen Charlotte Island and Spitsbergen were the only areas where the caribou did not share their range with the wolf. The races of caribou inhabiting these areas were the runts of the tribe. The Queen Charlotte Islands caribou ( Rangifer arcticus dawsoni ) is believed to be extinct. In achieved this status without the as– sistance of the wolf.

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BIBLIOGRAPHY

1. Anderson, R.M. Catalogue of Canadian Recent Mammals . Ottawa, 1946. ^ Nat. Mus. Can. Bull . 102. Biological Ser . no.31. ^

2. ----. “The present status and distribution of the big game mammals of Canada,” North American Wildlife Conference, 3d. Baltimore, 1938. Trans . pp.390-406.

3. Banfield, A.W.F. “The present status of North American caribou,” North American Wildlife Conference, 14th, Washington, 1949. Trans . pp.447-91.

4. Clarke, C.H.D. A Biological Investigation of the Thelon Game Sanctuary . (Ottawa, Patenaude, 1940.) Nat.Mus.Can. Bull . No.96. Biological Ser . No.25.

5. Dufresne, Frank. Mammals and Birds of Alaska . Wash.,G.P.O., 1942. U.S. Fish and Wildlife Serv. Circ . No.3.

6. Fristrup, Børge. “A preliminary account of geographical work of the Danish Peary Land Expedition,” Geogr.Tidsskr . vol.49, pp.41-66, 1948/49.

7. Gavin, Angus. “Notes on mammals observed in the Perry River district, Queen Maud Sea,” J.Mammal . vol.26, no.4, pp.226-30, 1945.

7a. Hadwen, Seymour, and Palmer, L.J. Reindeer in Alaska . Wash.,G.P.O., 1922, p.68. U.S. Dept.Agric. Bull . no.1089

8. Harper, Francis. Land of the Caribou . Natural History Magazine, 1949. Manuscript. 18(5): 224-231, 239 ^ vol. 18,no.5, pp. 224-231, 239, 1949. ^

9. Jacobi, Arnold. “Das Rentier, eine Zoologische monographie der Gattung Rangifer,” Zoologischer Anz . B.96. Erganzungaband . 1931.

10. Jensen, Ad.S. “The fauna of Greenland,” Greenland. Commission for the Direction of the Geological and Geographical Investiga– tions in Greenland. Greenland, Vol.1. The Discovery of Greenland, Exploration and Nature of the Country . Editors: M. Vahl (and others). Copenhagen, Reitzel; Lond., Milford, 1928, pp.320-23.

11. Manning, T.H. “Bird and mammal notes from the east side of Hudson Bay,” Canad.Field Nat . vol.60, no.4, pp.71-85, 1946.

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12. ----. “Notes on the coastal districts of the eastern barren-grounds and Melville peninsula from Igloolik to Cape Fullerton,” Canad.Geogr.J . vol.26, pp.84-105, 1943.

13. ----. “Preliminary report on a background study of the caribou,” Quebec. Assoc. for the Protection of Game and Fish. Annual Report vol.80, pp.20-21, 1948.

14. Murie, Adolph. The Wolves of Mount McKinley . Wash.,G.P.O., 1944. U.S. National Park Service. Fauna of the National Parks of the United States. Fauna Series no.5.

15. Murie, O.J. Alaska-Yukon Caribou . Wash.,G.P.O., 1935. U.S. Biological Survey. North American Fauna no.54.

16. Peary, R.E. Nearest the Pole . Lond., Hutchinson, 1907.

17. Rand, A.L. Mammal Investigations on the Canol Road, Yukon and Northwest Territories, 1944 ^ Territories, 1944 ^ . Ottawa, 1945. Nat.Mus.Can. Bull . 99. Biological Ser . No.28.

A. F ^ W ^ . W ^ F ^ . Banfield
Economic Zoology of Caribou

EA-Zoology (Olaus J. Murie)

ECONOMIC ZOOLOGY OF CARIBOU
From early times, throughout the world, venison has been a favored food for primitive people. Not only have the members of the deer family served as food for man, ancient and modern, but they have contributed importantly to the diet of large carnivores. Probably in adjustment to this ecological state of affairs, the deer of the world have developed characteristics such as speed, alertness, large [: ] size, or fecundity, or combinations of these, to meet the hazards of their environment.
In their special adaptations, the caribou of the Arctic and Subarctic are unique in several respects. In response to the rigors of their climate they have developed an unusually warm cost of hair, with hairiness extending even over the nose. Instead of the trim feet of southern deer, they grew rounded hoofs that spread when necessary, the better to walk on snow or boggy ground. They respond to cold with heavy layers of fat. And they exist in large herds. Incidentally, for some reason, the females bear antlers, and even the fawns grow spike antlers in the first summer.
Some fo ^ of ^ these are the very zoological attributes that have been such a boon to man in the Arctic. The caribou skin makes warm clothing; the excessive fat is a godsend to fat-hungry man in the winter cold; and the fact that caribou run in herds, and normally are numerous, makes it possible

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to depend on them as a source of food. I say this as a general rule, having in mind that not always have the herds been dependable in specific instances!
To begin with, the north country of America was people s ^ d ^ by two principal groups, The Eskimos were primarily a coastal people, finding much of their subsistence in the sea, though taking advantage of the caribou and other land animals at every opportunity. In exceptional instances Eskimo groups live ^ d ^ inland, away from the sea. The North American Indian was a man of the forested country (and the southern plains and deserts). There was accordingly a different human eco p ^ l ^ ogy for the coastal Eskimo and the Indian of the northern forest. The two peoples were not entirely exclusive of each other, and their interests clashed, with some hostility, at the borders of their natural territories.
For both of these ethnic groups the caribou had an economic significance in many respects similar to that of the bison for the Plains Indian farther south. Both animals traveled in herds, wore numerous, and furnished an important source of food, clothing, and shelter.
With the crude weapons at first available to them both Eskimos and Indiana found it necessary to summon great ingenuity to capture the animals. The bow and arrow were, of course, standard equipment for stalking game. Among the Eskimos, at least, this had its drawbacks in open country, but by various ruses, taking advantage of the animal’s temporary curiosity, or its grazing habits, or some local topographic features, the hun g ^ t ^ er was able to get within bow-shot of caribou.
But driving was a favorite method. The process has been described in several variations, but essentially the plan was as follows. The hunters would choose an area frequented by caribou, such as a migration route, and

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there two lines of cairns were placed, consisting of rocks, piles of turf, perhaps topped with sticks and even with a piece of garment that would flutter. These lines would converge on a narrow defile, or the shore of a lake. As the animals wandered between the two “wings” of the trap, hesi– tating to approach those prominent objects on either side, women and children might appear from behind the cairns the animals had already passed. As the caribou moved on, urged from behind, more people appeared, until the animals came into the narrows where concealed ^ hunters ^ had an opportunity to shoot, or into the water where hunters in kayaks would dart out after them and spear the swimming animals. Large numbers were sometimes killed by this method.
In various localities snare fences were built. In northwestern Alaska it is reported that snares were sometimes simply placed in gaps between willows, where caribou were accustomed to pass. But in interior Alaska, in the Tenana and upper Yukon region, long pole fences were erected across well-known migration routes, sometimes extending for several miles. At intervals there was a gap in the fence, in which was place a twisted raw– hide snare. As late as 1921 I saw some of these snare fences still in good repair, the poles and posts being held firmly with willow withes. One of these led directly to the old corral, where the animals were at one time thus guided into a definite trap. When thus enclosed they were shot by arrows, or sometimes speared through the bars with a knife fastened at the end of a pole.
In this general area there were also lookouts for spying migratory g herds. One of these was a platform built high in a spruce, with the limbs cleared away for an unobstructed view. Another, more elaborate structure,

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was built of poles tied with willows, in the form of a huge stepladder with a platform on top.
Snare fences were reported also by ^ ^ Lowe (3) near Boulder Creek, in Copper River Valley, and a reindeer herder reported snare fences formerly built by Eskimos in the lower Kuskokwim River region. In 1920 at least five caribou were caught in snare fences near Tanana Crossing; in this case old aban– doned telegraph line was used for snares. As late as 1926 rawhide snares were still being used on the Old Crew River, in norther ^ n ^ Yukon Territory, mostly by the old men or hunters lacking ammunition. MacFarlan d ^ e ^ (4) speaking of woodland caribou of the Northwest Territories, remarks that at one time Indians used twisted rawhide snares to capture them, and that the Indians near Fort Anderson, on Anderson River, obtained a few of the barren ground caribou in the same manner. Similar snare fences were used in parts of Alaska to capture moose.
These we d ^ r ^ e the primitive methods by which the Eskimos and Indians supplied their needs. The meat was a staple, obtainable at certain seasons in quantity. Seals, walrus, fish, and rabbits were the other more or less dependable food resources of these northern people.
The caribou meat was eaten fresh or dried. The contents of the rumen, rich with lichen remains, were eaten fresh or, when possible, frozen and stored for future use. The colon was relished, and it may be significant that this is also sought out by certain carnivores. The marrow of the bones was greatly enjoyed, and of course the quantity of fat which the caribou accumulates in the fall is a prize in the North. I have never found caribou meat tough, and a large fat bull is excellent eating. The meat has a high water content and some of us who drove dogs found that twice as much caribou meat was required by a sled dog as compared with mou ^ n ^ tain sheep meat. (The latter was fed to dogs only in real emergency.)

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Caribou skins were universally used for clothing — warm park [: ] s, leggings, moccasins, mittens, and other miscellaneous a d ^ r ^ ticles. In the Hudson Bay country I found that moose-hide moccasins were somewhat warmer in the cold dry snow but that caribou-hide moccasins were better moisture repellents when the weather was somewhat warmer and the snow was damp. In Hudson Bay, also, the Indians used the unborn caribou fawns for food, and made light caps the bags from their skins. Caribou rawhide, or babiche, was used for the wea b ^ v ^ ing of snowshoes, and hides were used for tents. Caribou antlers were used for many handy tools. Possibly no other northern animals has produced a greater variety of economic uses for the original inhabitants of the North. Add to this the fact that this hardy deer inhabited some of the most “barren” portions of the Arctic, and it will be realized T ^ w ^ hat an important animal this proved to be in arctic economy.
Stefansson (11) has given us a dramatic example of some of the bleakness of the surroundings in which you might find caribou in the northern fringes of its range. He had reached Isachsen Land with two of his companions seriously ill. “I walked that day twenty miles across one of the very few stretches of entirely barren land that I have seen in the Arctic. Underfoot was gravel without a blade of grass.” Yet next morning (in a more favorable grazing area) he came on a band of caribou, which saved the day for his party. Here was a group of men, exploring some of the northern fringes of our con– tinent, reaching into the polar sea, where one might expect little life to be found. Yet there were caribou.
Caribou are not always dependable, or predictable, and Indians have on occasion suffered hardship through the un d ^ c ^ ertainties and vagaries of caribou movements. They are erratic travelers and while generally they follow

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established routes, they may be slow in arriving, they may veer away from a former course, or they may not go as far as in former years.
In the early spring of 1921 I reached Tanana Crossing, in Alaska, and found that the Indians were having a difficult time. They were awaiting the caribou herds which come back over the Alaska Range to drop their fawns in the Tanana River country. Fish were scarce and I was having great diffi– culty in obtaining food for my dogs. Finally, on April 28, the caribou same — long lines of them leading down the snowy slopes into the valley of the Tanana. The Indian village came to life and soon caribou meat was hung to dry on numerous racks, something like fourteen or fifteen per family. It is often famine or feast in caribou country.
With the coming of the whi l te man in the North, and his rifle and trade goods, the caribou-man ecology changed. For the explorer, the whaler, the trader, and particularly the pioneer trapper and prospector, caribou meat became a staple food. Hunting became easier. The native people relinquished their primitive methods and accepted firearms, and they were able to kill more game. They killed more than they needed, and found it profitable to supply the white men. The annals of the North are filled with accounts of barter in caribou meat and skins. In the enthusiasm over the new hunting facilities, many animals were wasted.
E. W. Nelson (6) writing of the Bering Sea coast of Alaska, said: “When the Americans first obtained control of the Territory firearms were unknown among the natives, and when the natives first obtained guns they kept the traders supplied with meat at the rate of two charges of powder and ball for a dear. One winter, just preceding the transfer of the Territory, an enormous herd of Reindeer [caribou[ passed so near St. Michaels that a

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6-pounder loaded with buckshot was fired at them, killing and wounding a number of them.”
Speaking of Alaska Peninsula and adjacent coasts, he says further” “In the summer of 1880 one men from Point Barrow took about five hundred skins, and many others took nearly as large a number. Only a few stragglers now remain on the Kaviak Peninsula and in the country between the Yukon and Kuskokwim Rivers.”
Again: “When Mr. Dall came down the Yukon in the spring of 1867, he saw over four thousands skins of reindeer fawns hanging up in a village near Anvik, and at present scarcely half a dozen deer, old and young, are [: ] r ^ t ^ aken yearly in that district.
“The skins taken in summer are valued at about one dollar each among the fur traders, who buy them in one part of the country and trade them for furs in other parts.”
James Wickersham (12) writes of interior Alaska in 1900: “Captain Farnsworth, then in command at Fort Egbert, had established a hunting camp on the divide between Mission creek on Forty Mile river, about fifty miles from Eagle, and had sent hunters and a squad of soldiers there to kill caribou for the post.” Mr. Wickersham mentions seeing fifty caribou hung on racks.
Osgood (7) reports that in the fall of 1901, in east central Alaska, a white man killed 62 caribou in one day, and immediately afterward a party of Indians killed 42. Next day a hunter killed 7 cripples in that vicinity and followed the bloody trail of the herd for some miles.
Edward A. Preble (8) referring to the report of A. J. Stone, says: “He considers that the animals are fast being exterminated in that quarter

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[east of the Mackenzie River delta], principally on account of the demand for meat at the trading posts, and at the wintering places of the whalers along the Arctic coast.”
MacFarlane (4) commented: “In the course of the company’s five year occupation of Fort Anderson, we received considerable quantities of venison and many skins of the Barren Ground reindeer from the Eskimos and Indians who reported thereto for purposes of trade.”
In 1917, at Fort Chimo, Ungava Bay, I learned that the Naskapi Indians would occasionally bring some caribou meat to the trading post, th r ough these animals had already become extremely scarce on the Labrador Peninsula. Our party had crossed through the interior without seeing one. Three years ^ ^ later, in interior Alaska, I found that mountain sheep, moose, and caribou were being sold to the construction camps along the route of the government Alaska Railroad. At the restaurants in the towns one could order moose or caribou from the menu. A number of years previously professional hunters had been supplying caribou to the local markets, and one of these men gave me many details of their hunting technique.
Originally the numbers of caribou were more or less in adjustment with their environment, which included the scattered human population using primitive weapons. When the white pioneers came into the North — explorers, whalers, traders, prospectors, trappers, adventurous souls with that innate urge to experience what lies beyond — bringing with them more lethal weapons and a large market for game, the drain on the caribou herds became greater than their population dynamics could withstand. Still later came the more permanent settlements, with increase of white population. Today even a grater influx of people is taking place, with roads, railways, and aircraft.

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Thus the original prospect in northern lands has virtually disappeared. Industrial civilization is reaching northward, with modern convenience and huge demand on natural resources.
It must be kept in mind that during the early ^ ^ pioneer period the caribou served on important purpose. W ti ^ it ^ hout the caribou — a herd animal existing in large numbers — with its potential for abundant meat and materials for clothing and similar necessities, life in the North would have been much more difficult. Even today, in much of the territory, caribou contribute importantly to the personal economy of many people — Eskimos, Indians, and whites.
However, no longer does the supply appear inexhaustible. The caribou have completely disappeared from many areas, such as the extensive portions of Alaska bordering on Bering Sea, and the area east of the Mackenzie River delta. The woodland caribou has disappeared from former ranges in northern states and parts of lower Canada, and they have become extremely rare on the Labrador Peninsula. In many other areas they are reduced in numbers.
In Siberia and northern Europe the reindeer was domesticated long ag l o, so extensively that in parts of their former range wild reindeer are scarce or no longer exists. The North American caribou were never domesticated. But with the disappearance of the wild caribou in the areas bordering on Bering Sea, domestic reindeer from the Old World were substituted, together with Lapp herders to instruct the Eskimos. In many ways this was a help to the Eskimo, but there is much confusion about the project. Proper reindeer ^ hording ^ requires a nomadic life.
Consider, for example, the natural feeding habits of caribou. In my study of the Alaska-Yukon caribou (5) some years ago, I came to the following

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conclusions: “…the fact that the caribou do not linger in any one locality, but are nearly always on the move, keeps them distributed over a wide territory and tends to preserve their winter food.”
Studies conducted by the U.S. Biological Survey in the nineteen twenty’s indicated that the lichen growth, so beneficial for caribou and domestic reindeer, once it is destroyed requires about as long to recover as does a destroyed forest. These considerations led me to suggest further in my published report (5):
“Apparently each species adopts a particular migratory habit to suit its food requirements. In the case of the caribou the need appears to be an avoidance of concentration, and the need for greater dispersal of the caribou has been met by greater restlessness, resulting in a greater and more varied migration. Throughout its circumpolar range, the caribou normally seeks lichens for winter forage, and throughout its entire range it is a wanderer. These two facts appear to be related, and in the absence of any other solution it seems reasonable to conclude that the caribou’s peculiar taste has made the species the restless wanderer that it is.”
It may prove to be a fact that the Alaskan reindeer ranges are overgrazed. Certain white owners undertook to raise domestic reindeer for market, and since commercial enterprise naturally requires large quantity for profit, such operations no doubt provided an additional impact on the native caribou herds and the original range.
It is axiomatic that a reindeer industry and wild caribou herds cannot occupy the same area. The migratory caribou draw with them on their journey sizable numbers of reindeer that are thus lost to the owner. For this reason, in reindeer territory it has been the custom to attempt to eliminate any

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stray caribou that come into the range of the domestic animals.
At one of the sessions of the International Technical Conference on the Protection of Nature, meeting at Lake Success under the auspices of UNESCO in the summer of 1949, a Canadian delegate introduced the subject of preserv– ing some of the Canadian species of caribou, particularly those of the Labrador Peninsula. There has been consideration of substituting the domestic reindeer. To meet this difficulty some considerations of caribou biology may be worth while.
There has been the belief that the North American caribou, being migratory, cannot be domesticated. Yet I have seen a caribou calf, pretty well grown when it was captured in the fall, so thoroughly tamed in less than two weeks that it would follow its owner about like a dog. An elk calf, caught when a few days old in the highly migratory Jackson Hole elk herd in Wyoming, grew up perfectly tame and, although up to more than four years of age it had perfect freedom and in winter mingled with the wintering wild elk, it never showed any desire to migrate.
In areas where native caribou have been reduced to dangerous levels, experiments would be worth while to raise a herd of the animals native to that area, rather than replacing the remnant with the domestic reindeer. It is entirely possible that the native caribou could be given sufficient domestication for controlled herding as in the case of domestic reindeer. Such a procedure would tend to insure the perpetuation of the native stock, a serious consideration among the many people, scientists as well as other groups of conservationists, whose aims are typified by those of the International Union for the Protection of Nature. Under such a plan escapes from the domestic herd would find their own kind, and there would be the

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hope of restoring the original wild population.
Wolves, disease, and perhaps other factors have always been a drain on the caribou herds. These cannot be ignored. But the most critical of all factors, as we have found in modern game management, is the destruction of habitat. In the future, if it is our decision to retain the caribou, there must be such land-use planning that proper range, extensive range, may be designated for caribou. The more intensive types of land use could well be segregated into the more favorable areas. The caribou ranges thus preserved could be dedicated to multiple use, of the kinds not incompatible with the needs of caribou. Such uses would be the wilderness type of recreation (which is being given so much attention today among conservationists), certain field sports, scientific studies, particularly in ecology, preservation of other wildlife that shares similar habitat, and like categories.
By proper planning and suitable regulations, caribou may still be an economic asset to many people, and of great recreation value as well.

EA-Zoo. Murie: Economic Zoology of Caribou

BIBLIOGRAPHY

1. Boas, Franz. “The Central Eskimo,” U.S. Bureau of Ethnology. Annual Report , 6th, 1884-’85. Wash., G.P.O., 1888, pp.409-669.

2. Jenness, Diamond. “The life of the Copper Eskimos,” Canadian Arctic Expedition, 1913-1918. Report, vol.12, pt.A, pp.1-277, 1922.

3. Lowe, P.G. “From Valdez Inlet to Belle Isle, on the Yukon,” Glen, E.F., and Abercrombie, W.R. Reports of Explorations in the Territory of Alaska (Cooks Inlet, Sushitna, Copper, and Tanana Rivers) 1898 . Wash., G.P.O., 1899, pp.353-87. U.S. War Dept. Adjutant General’s Office. Doc . no.102. Military Intelligence Div. Publ . no.25.

4. MacFarlan d ^ e ^ Roderick. Notes on Mammals Collected and Observed in the Northern Mackenzie River District, Northwest Territories of Canada, with Remarks on Explorers and Explorations of the Far North . Wash., G.P.O., 1905, U.S.Nat.Mus. no.1405.

5. Murie, O.J. Alaska-Yukon Caribou. Wash., U.S. Dept.Agriculture, Bur. of Biological Survey, 1935. North American Fauna no.54.

6. Nelson, E.W. Report upon Natural History Collections Made in Alaska between the Years 1877 and 1881. Wash., G.P.O., 1887. U.S. Army. Signal Service. Arctic Series of Publications no.3.

7. Osgood, W.H. Biological Investigations in Alaska and Yukon Territory . Wash., G.P.O., 1909. North American Fauna .no.30.

8. Preble, E.A. A Biological Investigation of the Athabaska-Mackenzie Region . Wash., G.P.O., 1909. North American Fauna no.27.

9. Stefansson, Vilhjalmur. The Friendly Arctic . N.Y., Macmillan, 1921.

10. ----. Hunters of the Great North. N.Y., Harcourt, Brace, 1922.

11. ^ ----. ^ My Life with the Eskimo . N.Y., Macmillan, 1913.

12. Wickersham, James. Old Yukon. Wash., Washington Law Book Co., 1938.

Olaus J. Murie
Reindeer Herding in Canada

EA-Zoology (Richard Finnie)

REINDEER HERDING IN CANADA
The first successful experiment in reindeer husbandry in Canada began in the spring of 1935, when 2,370 animals from Alaska were delivered to a reserve at the east side of the Mackenzie Delta. Despite many difficulties and setbacks, the reindeer steadily increased and, by 1947, numbered 6,400, even though between 3,000 and 4,000 had be slaughtered for food and clothing during the intervening years, and still others had strayed away.
In 1908, Dr. Wilfred Grenfell established 250 Lapland reindeer at St. Anthony in northern Newfoundland; within ten years they had increased to 1,200 but had begun scattering for lack of attention. In 1918, all that were still available, 126, were given to the Canadian Government and moved to the north shore of the Gulf of St. Lawrence, near St. Augustin, Quebec. At first they multiplied, but here again inadequate care took its toll and, in 1923, the 145 remaining were shifted to Anticosti Island and liberated. There they were stricken with disease and, by 1941, only 9 were left.
This was not the first attempt to introduce reindeer into Canada. Back in 1911, the Department of the Interior had bought 40 breeding does, 6 stags, and4 geldings (to be trained as draft animals) from the Grenfell herd at St. Anthony, and had shipped them to Fort Smith, Northwest Territories. The journey was a hard one for them: by boat to Quebac k , by train to Edmonton,

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by wagon to Athabaska Landing, and by scow to the Fort Smith vicinity. Fifteen died on the way from injuries or lack of suitable food. By the fall of 1913, only three were left, one having been killed by dogs, the rest having taken to the woods when tormented by flies or having succumbed to some undiagnosed malady.
In 1918, the same year that the ill-starved experiment was being under– taken in Quebec, Vilhjalmur Stefansson returned from five years in the Arctic with the conviction that there lay an opportunity to develop a vast new source of wealth with domesticated reindeer and musk oxen. He set about trying to induce the Federal Government to go in for reindeer breeding as a public enterprise. The Right Honorable Arthur Meighen, who was then Minister of the Interior, soon became sufficiently interested to arrange the appoint– ment, in May 1919, of a Royal Commission to investigate the possibilities as outlined by Stefansson in an address before both Houses of Parliament. The commission consisted of J. G. Rutherford, Railway Commissioner (chairman); James S. McLean, manager of the Harris Abattoir Co., Toronto; James B. Harkin, Commissioner of Dominion Parks; and Dr. Stefansson. During the next year the commission questioned 35 expert witnesses and finally brought in a favorable report. In March 1920, Stefansson withdrew from the commission, partly because he felt that he was prejudiced and partly because of a new idea he had: that a reindeer industry might be expedited with private backing.
In October 1919, Stefansson opened negotiations with the Hudson’s Bay Company. He next made formal application to the Department of the Interior to lease grazing rights over a large area of southern Baffin Island for a fifty-year period, with the privilege of transferring it if granted. Meanwhile the Hudson’s Bay Company agreed to support his reindeer venture, and later

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a musk ox venture, too, should he be granted his lease.
The lease was formally signed on June 11, 1920, giving Stefansson exclusive rights for reindeer grazing over 113,900 square miles of southern Baffin Island for 30 years. The Hudson’s Bay Reindeer Company was incorporated and to it Stefansson transferred his lease, in consideration for which he was retained solely in an advisory capacity.
Stefansson’s plan was to buy a selected herd of a thousand reindeer in northern Norway and ship them to Baffin Island, and he recommended Storker T. Storkerson, a member of his Canadian Arctic Expedition, as permanent manager. Storkerson, a Norwegian, visited the prospective grazing grounds during the summer of 1920, was well pleased, and the following winter journeyed to London to report to the head of the Hudson’s Bay Company with the expectation that he would now have the responsibility of selecting and buying the breeding stock in Norway. This was denied him, however, and he at once resigned in protest.
This was the beginning of a chain of misfortune ^ s ^ that led to disaster. The Hudson’s Bay Company’s supply vessel ( Nascopie , which was sent to Norway to fetch the reindeer, had space for only 687 animals, 60 of which died or were lost before the herd was loaded. Rough weather en route took another 77, so that only 550 remained to be put ashore at Amadjuak Bay, Baffin Island. No accommodation for the Lapp herders who were employed had be d ^ e ^ n provided, nor had provision been made to receive the deer; so while the herders were trying to erect dwellings, the herd dispersed. A year later, in June 1922, there were 210 reindeer left, including new calves. The Lapps departed for home and were replaced by local Eskimos who had not yet acquired enough experience as herders. By 1925, the entire herd had vanished.

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However, the sanguine report of the Royal Commission upon the possibilities of the reindeer industry in the arctic and subarctic regions, published in 1922, was not forgotten.
The report recommended that a number of experimental herds be estab– lished in the most suitable locations. It was believed that the development of such herds would provide reliable and economical food and clothing supplies for the natives, both Indians and Eskimos, and guard against actual starvation due to failure to find wild game. The commissioners added that the herds might become a food source also for white men engaged in developing the natural resources of the country, and lay the foundation for a future commer– cial meat industry.
The Northwest Territories and Yukon Branch, Department of the Interior, which had been created in 1921, made further inquiries about reindeer herding, especially in Alaska. This was intensified by the rapidly changing economy of Canadian Eskimos. Nearly all had acquired rifles with which they were decimating the d ^ c ^ aribou and altering their migration routes; and al were turning more and more to the trapping of white foxes. It was feared that fluctuations in the numbers of fur bearers and the uncertainties of the fur market might seriously affect the natives.
The Government observed that similar conditions had prevailed in Alaska forty years earlier, and that the natives’ lot had been improved by the introduction of domestic reindeer from Siberia. The original herd of 1,280 animals introduced between 1891 and 1902 had increased to more than 750,000. In addition more than 200,000 had been slaughtered to supply meat and clothing. Two-thirds of the Alaskan reindeer were nor owned by natives.
It was the desire of the Canadian Government to promote the introduction

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of an industry which the natives could develop rather than to establish a government industry, and it was decided, before incurring the expenses involved in introducing reindeer, to have a thorough investigation made. This was to cover the country between the Alaska-Yukon boundary on the west, the Coppermine River on the east, Great Bear Lake on the south, and the Arctic Sea on the north.
It was to be a general botanical reconnaissance with special reference to reindeer pasture, carrying capacity, and other general conditions of importance to a future reindeer industry. Since in Alaska the reindeer industry had become such a success, it was considered that previous to the field work in Canada the investigators should, with the permission of the United States Government, spend a season in Alaska to make an exhaustive study of the reindeer industry and its effect on the country and people.
Chosen for this assignment were two brothers, A. Erling ^ P ^ orsild, a trained botanist, and R. T. Porsild, a zoologist, both of whom had spent many years in North Greenland, were experienced arctic travelers, and could speak the Eskimo language. The investigation lasted from May 1926 to November 1928, during which the Porsilds traveled 15,000 miles by dog team, canoe, motorboat, and on foot. They returned with 15,000 herbarium specimens of vascular plants and nearly 5,000 specimens of cryptogams, plus some zoological specimens and many photographs.
In his report, Erling Porsild indicated a half-dozen sections he considered suitable for reindeer pasture: ( 1 ) the arctic coast from the Alaska-Yukon boundary eastward, including Herschel Island, to the Mackenzie Delta; ( 2 ) the Mackenzie Delta and its islands; ( 3 ) the arctic coast and hinterland from the Mackenzie River to Cape Bathurst; ( 4 ) the plains north

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of Great Bear Lake; ( 5 ) the Dease Valley; and ( 6 ) the Dismal Lakes and Kendall River valleys.
In summing up, Porsild estimated that there was a total of 15,000 square miles of grazing land in the coastal area from the Alaska-Yukon boundary to Cape Bathurst and from the mouth of the Anderson River to Point Separation at the south end of the Mackenzie Delta. Allowing 40 acres per head for this type of country, he considered that it would have a year-round carrying capacity of 250,000 reindeer. The pasturable country north and east of Great Bear Lake comprised 38,000 square miles which he believed would support 300,000 reindeer, giving each one 60 acres. (In 1930, he made a grazing survey of the central Keewatin District, west of Hudson Bay between Churchill and Chesterfield Inlet, ascertaining that, while the greater part must be classified as summer pasture only, it still contained suitable reindeer ranching areas exceeding those of the Mackenzie District, along the coast from Nelson River north to Dawson Inlet.)
On the basis of the Porsild report, the Canadian Government proceeded to acquire the breeding stock for establishment in the Mackenzie Delta. By an Order in Council of May 1, 1929, the Minister of the Interior was authorized to purchase from Lomen Brothers, Nome, Alaska, 3,000 head of adult reindeer. Under the terms of a contract subsequently made, Lomen Brothers agreed to furnish this number of the largest and sturdiest animals in their possession, with a proper proportion of bucks and does, and to deliver them at the Mackenzie Delta in the spring of 1931.
In the fall of 1929, Erling Porsild went to Nome to represent the Government in the selection of the herd and the beginning of the drive, while his brother was given charge of arrangements for the reception of the

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of the reindeer in the Mackenzie Delta. In December, 2,890 does and 307 bucks were selected in the Buckland Valley, and to these were added about 250 steers for food and draft purposes. Lomen Brothers placed Andrew Bahr, a veteran Lapp herder, in charge of the drive, assisted by other Lapps and several Eskimos.
The drive got under way in December 1929. After the initial effort of detaching the selected animals from the main herd, many unforeseen difficulties were encountered. At first many of the deer broke away and were not recovered. Blizzards, intense cold, and depredations of wolves, the distraction of wild caribou, and the stampeding of the reindeer, all impeded progress along the 1,500-mile route. The trek continued, however, with losses being recouped in some measure each fawning season. The deer reached Canadian territory in 193 8 ^ 38 ^ , but an unsuccessful attempt to get them across the Mackenzie Delta in the winter of 1933-34 delayed delivery until March 1935.
The number of deer delivered was 2,370, consisting of 1,498 does of all ages, 3 ^ 2 ^ 89 bulls, 322 male fawns, and 261 steers. Only a fifth of these were original stock from the Buckland Valley herd in Alaska.
Long before the trek ended, a corral had been constructed at Kittigazuit on the mainland just east of the Mackenzie Delta, while headquarters for the reindeer staff was established, with dwellings and warehouse, about 40 miles inland at the foot of the Caribou Hills along the East Channel of the delta, 70 miles by water from Aklavik.
In 1931, three Lapp herders and their families were brought from Norway to assist in controlling the herd upon its arrival and in training the local Eskimos. These men joined the herd during the winter of 1932-33, for the last stage of the drive.

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In December 1933, the Government officially created a reindeer reserve comprising about 6,600 square miles, including both summer and winter ranges, immediately east of the Mackenzie Delta. Two years later this area was photographed from the air, and the resulting maps proved invaluable to the herders in their travels and in the selection of fresh grazing grounds.
During the first summer on the reserve, the herd grazed in the coastal area in the vicinity of the corral at Kittigazuit, where a roundup was held in midsummer. A count then made showed that there was an increase of 800 healthy fawns. In early winter the herd was moved inlan t ^ d ^ to the environs of the main station, where the plateau extending back from the river carries a good cover of reindeer moss for winter feeding. The following spring the reindeer were taken to Richards Island, adjoining the Kittigazuit range. The island proved wholly satisfactory for summer grazing and has been used thus ever since. Here a corral with holding pens and a lead fence was constructed for the annual roundup.
When alarmed, the reindeer from into a compact body and begin to circle. Herds consistently move clockwise or counterclockwise, according to the tendency of the parent herds. The Canadian deer mill counterclockwise, as do their Alaskan forebears, and this peculiarity has to be taken into considera– tion in the construction of corrals and holding pens.
The roundup procedure is to count all the deer, mark all that have not previously been marked (with a V-shaped notch cut in the right ear), and check the sexes, enumerating does, fawns, and yearlings. Bulls over four years old are castrated. All diseased or maimed animals are killed at once, while old does and surplus bucks (a proportion of 10% of the latter is maintained) are selected for slaughter later in the year.

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The handling of the reindeer at roundup time and on other occasions tends to prevent them from becoming wild and intractable. Some of the mature steers in the herd are broken to harness for transporting supplies and firewood, moving camp, and the like.
Of fine grain and classed between beef and mutton, reindeer meat is palatable, nourishing, easily digested, and free from gamy flavor. The skin, with hair intact, is unsurpassed for cold-weather bedding and clothing. Contrary to the practice in Norway, the milking of Canadian reindeer has not yet been attempted to any extent, although the milk is among the richest kinds known, with a butterfat content of 2 3 ^ 2 ^ %.
In the summer of 1939, an inspection of the Canadian reindeer was made by Dr. Seymour Hadwen, Director of Pathology and Bacteriology, Ontario Research Foundation, Toronto, and formerly Chief Veterinarian and Parasi– tologist of the United States Bureau of Biological Survey. He reported the herd to be in excellent condition and practically free from disease and parasitism, and was struck by the fertility of the deer as shown by the bearing of fawns by some of the does only one year old. He noted that there had been a gradual increase in the dressed weight of slaughtered steers. Some carcasses now weighed as much as 200 pounds, and the average was 167 as against 150 in 1935. The dressed weight of the aged does was about 135 pounds. He concluded that the excellent range and favorable climate, as well as good management, were responsible for the development of large and vigorous stock.
The Canadian deer are allowed to graze freely over an area of several square miles, the herders watching for straying individuals — which, after a winter storm, sometimes have to be tracked some distance and returned to

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safety. On the semiannual drive between summer and winter grazing grounds, the main herd covers about 75 miles in easy stages. Reindeer moss ( Cladonia rangiferina ) is the principal food in winter, and although it is eaten to some extent in summer too, it is not nearly so nourishing as the grasses, willows, and other shrubs that are then available. If overgrazed, the moss may take many years to recover, but the other food renews itself annually. A system of rotating the winter grazing areas guards against moss depletion.
Like all other northern land mammals the reindeer are harassed by flies in summer, but by moving them 15 to 20 miles away from an area at the time warble-fly grubs emerge from the hides and drop on the the ground, the herders are able to minimize reinfestation.
There was steady though unspectacular growth in the size of the herd, the official totals at roundup time being 2,980, including 815 fawns, in 1935; 3,750, including 936 fawns, in 1936; 4,092, including 1,181 fawns, in 1937; 4,631, including 1,281 fawns, in 1938.
In December 1938, about 950 deer were separated from the main herd and transferred to a location in the vicinity of the Anderson River. This became the nucleus of a new herd under the management of Eskimos under government supervision, and the following yea d ^ r ^ it had increased to 1,196. A roundup, in August 1940, showed that the new herd numbered 1,559 animals, including 448 fawns. Meanwhile, the main herd on Richards Island and grown by 1940 to 5,076, including 1,486 fawns, despite the reduction in 1938.
The training of young natives as herders proceeded, and a second subsidiary herd was started in 1940. To qualify as independent herders the Eskimos had to serve as apprentices for three years, then they received the deer under a lending arrangement subject to the return of a similar number

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of animals as the herds increased.
Both the training program and the industry itself suffered a serious setback in 1944, when the custodians of the native herds, together with their families and a white supervisor, lost their lives in the wreck of a schooner in a storm off the arctic coast. The native herds became scattered, and all the animals that could be recovered were maintained as a government unit in the Anderson River area.
The reindeer industry in northwestern Canada is still in its infancy, and the main concern is to train young natives as herders and at the same time build up new breeding stock. Still, there has been available for dis– posal each year a number of surplus animals, chiefly steers and old does. About 300 deer in the main herd have been slaughtered annually for food and clothing. The slaughtering operations have been conducted for the most part in the late autumn and early winter, the meat being distributed mainly to the mission hospitals and residential schools in the Mackenzie Delta. ^ The remainder of the meat goes far relief and camp purposes, with a limited quality for local sale. ^
Fawn skins which become available at the annual roundup, usually from injured or defective animals, are turned into parkas and other clothing for the herders. Experiments in tanning have been carried on with hides from mature animals. Some of the skins of adult reindeer killed for food have been shipped as far as the Canadian Eastern Arctic, when caribou were scarce, to be used for bedding and the making of winter clothing.
In the spring the main herd is moved from the winter range to the coastal area, where fawning occurs between the first of April and early June. This interlude is followed by the driving of the deer over the ice to the northern part of Richards Island, where the winds help to lessen the scourge of flies. Later the deer are shifted gradually toward the corralling

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grounds. The roundup is usually held during the last week of July, and, if the winter is favorable, finishes within four or five days. Afterward the herd grazes over the summer range until it is once more assembled for the annual slaughter of surplus stock.
All herders share in an annual bonus of fifty cents per head. Two men are usually on duty concurrently, each having a dog to help him, for a period of 24 hours, at the end of which time they are relieved. Under ordinary conditions the herder’s life is fairly placid, but it [: ] becomes strenuous during emergencies such as storms, or the threat of wolves; and when the herd must be transferred from one range to another, or rounded up.
Two of the Lapp herders returned to Norway in 1936 and 1938, but the third has remained. In the first years of the building up of the herd all of the Eskimos employed or apprenticed belonged to the Mackenzie Delta, but, with the placing of native herds farther east, young Coronation Gulf Eskimos were encouraged to take training.
The Canadian reindeer industry was beset with problems from the first, of which the most formidable were not with the deer themselves but rather with people. While capable white men can be found to act as supervisors, it is always hard to maintain a full complement of native herders and apprentices.
This difficulty was anticipated as long ago as 1920 by Dr. Diamond Jenness, Chief of the Division of Anthropology, National Museum of Canada, when he appeared before the Royal Commission and stated that he was not sure that it would be an easy task to convert the Eskimos into efficient herders. He pointed out that while they were intelligent and trustworthy, they had always been hunters and fishermen, and as long as caribou and fish

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and seal were plentiful they might not readily turn to herding. If game became scarce, however, he thought that they would develop into herders as Eskimos had done in Alaska.
He did not mention the further complication of fur trapping which in those days was still a minor occupation of the Eskimos. In succeeding years the Mackenzie Delta Eskimos, particularly, trapped white foxes almost to the exclusion of hunting, and a number of them became comparatively wealthy. Such people have been reluctant to forego opportunities to make quick profits with foxes in favor of long-term investment in reindeer herds.
Familiar complaints of native herds are that they are onerously tied down to their deer, with no time for trapping or even putting up enough dog feed, and that they are obliged to hire other Eskimos to help them but lack the means to pay them. To cap the climax, from their point of view, the number of deer they may slaughter is rigidly restricted. The principles of conservation are hard for them to grasp.
^ { ^The successful continuation and expansion of the reindeer industry will therefore depend on the employment of an ample number of keen and competent white supervisors, careful over-all management, and a suitable program of education for the natives, plus a profitable market for meat and hides to keep pace with the expanding herds. ^ } ^
Richard Finnie
Reindeer Breeding in U.S.S.R.

EA-Zoology (V. J. Tereshtenko)

REINDEER BREEDING IN THE U.S.S.R.
CONTENTS

Scroll Table to show more columns

Page
Definition and Classification 1
Historical Background 5
Reindeer Breeding Before the 1917 Revolution 14
Reindeer Breeding After the Revolution 17
Reindeer Herding 24
Utilization of the Reindeer 28
Draft Animals 30
Meat 37
By-Products 40
Bibliography 44

EA-Zoology (V. J. Tereshtenko)

REINDEER BREEDING IN THE U.S.S.R.
Definition and Classification
The English term “reindeer breeding” is liable to misinterpretation when applied to conditions in the Soviet Union, and may easily cause con– fusion among foreign students who do not have sufficient mastery of the Russian language or do not inform themselves as to views on the zoological classification of reindeer.
There are two sources of possible confusion, the first of which comes from Russian terminology and nomenclature applied to reindeer breeding. Berthold Laufer, in The Reindeer and Its Domestication (27), falls victim to this difficulty, for he complains that the Russians do not discriminate between stag and reindeer, calling both indifferently olen . But any Russian-English dictionary indicates that olen means “deer” in English. The English word “reindeer” and its equivalents in other languages are translated into Russian by adding a qualifying word to olen . Thus “reindeer” in Russian is severnyiolen , which literally means “northern deer.” The only Russian equivalent of the term “reindeer breeding” is olenevodstvo , the ending vodstvo implying the breeding or cultivation of something, thus the word may signify the breeding of any deer. As a matter of fact, in

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the territory of the U.S.S.R. the breeding of other species of deer as well as reindeer has been developed. Accordingly, although the word olenevodstvo often conveys the meaning of “reindeer breeding,” this is not always the case, and therefore if taken out of context the word may lead to misunderstanding.
A second source of possible confusion to the foreign student of Soviet reindeer breeding is the lack of precision in the existing classification of reindeer in Russian literature and even more so in the works about reindeer written outside of Russia.
For instance, the first 16 reindeer purchased in 1891 for the introduction of reindeer breeding in Alaska, and the second group of 171 purchased in 1892. were obtained from Cape Sertse Kamen and South Head, Siberia. The initiator of the plan was Dr. Sheldon Jackson, United States General Agent of Education in Alaska. Lieutenant B. P. Bertholf was later sent by Jackson to purchase reindeer, with instructions to get a better breed than those “usually met with on the northeast coast of Siberia.” In the report on his mission, Bertholf stated that the Tungus deer “are much larger and sturdier and have longer legs than the deer of the northeast coast of Siberia”, that “the interior Chukchi and those some distance to the west of the east cape have deer which are larger than those on the coast”; and that Koriak deer are smaller. In the light of the modern zoological classification, Bertholf’s mention of the “Tungus deer” may perhaps be interpreted as a reference to reindeer which are described by Soviet zoologists as a subspecies rather than a “better breed”.
Rangifer , the genus which in English commonly includes all species of of domestic and wild reindeer and caribou, but which is treated by some Soviet writers as a species, is represented in the Soviet Far North by Rangifer tarandus , the Russian severnyi olen . Within this group the following subgroups

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are distinguished, the Russian equivalents of the Latin names being formed by adding one or two qualifying adjectives to severnyi olen: ( 1 ) Rangifer tarandus tarandus , an inhabitant of Lapland and eastern European taiga ; ^ , ^ Laplandskii severn ^ y ^ i olen in Russian; ( 2 ) Rangifer tarandus pearsoni , which is found on Novaya Zemlya and called Novozemelskii severnyi olen in Russian; ( 3 ) Rangifer tarandus sibiricus , or Sibirskii tundrovyi severnyi olen , in Russian (Siberian tundra reindeer); it lives in the eastern European and Siberian tundra, on the Novosibirskie Islands, and sometimes is found also in the nor h thern taiga; ( 4 ) Rangifer tarandus valentinae , an inhabitant of the woodland area of Siberia and Ural up to Tataria in the west, northern Mongolia in the south, and the Stanovoi Range in the east; it is called Sibirskii lesnoy severnyi olen (Siberian woodland reindeer) in Russian; ( 5 ) Rangifer tarandus phylarehus , found on the Kamchatka Peninsula, on the coast of Okhotsk Sea, in the Amur region, and called Okhotskii severnyi olen in Russian; ( 6 ) Rangifer tarandus setoni , Sakhalinskii olen in Russian; it is met on the island of Sakhalin; ( 7 ) Rangifer tarandus angustrirostris , found in the mountains of the Transbaikal region, and in the Barguzin Range in particular, its Russian name is Barguzinskii severnyi olen .
The wild reindeer does not occur between the middle course of the Yenisei and that of the Ob. It is found on Sakhalin, but not in the center of Yakutia. It is found also on Novaya Zemlya, Vaigach, and the islands of Novosibirskie, Belyi, and Medvezhii. In the past, the bor ^ d ^ er line went farther south than that of the present time and reached such cities in C ^ E ^ uropean Russia as Novgorod and Kazan, and the middle course of the Kama.
As in the whole deer family, reindeer horns differ from those of other

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ruminants in being solid, generally branched, shed, and renewed annually. In contrast to other deer, the reindeer has antlers in both sexes, those of the males larger and more branched. In the early stages of growth they are soft and covered with a downy, vascular covering — the “velvet” — which later is shed, or rubbed off. The antlers are cast every year; the males usually cast them between November and December, the gelded reindeer and the yearlings from March to May, and the females in May and June. At first the growth of antlers proceeds slowly; toward the end of May, and especially in June, the antlers of the mature stags grow rapidly. It is said that in the whole kingdom of vertebrates there is no other tissue known which grows as rapidly as reindeer antlers (4). According to some observations made in the Pechora region, antlers grow at a rate of 0.39 inch and even as much as 0.79 inch a day at the beginning of June. Speaking of the antlers of Cervus elaphus , another species of the deer family, Maiseven (4), asserts that “their growth cannot be compared even with the growth of malignant tumors. It far exceeds the latter.”
Depending upon climatic conditions and food supply, reindeer differ considerably in weight and size. North of the tree line the reindeer is dark brown in color, short and stocky (the height being about 39 to 43 inches), but lighter in weight than the woodland reindeer which is long, big-framed, with long legs and narrow chest (height about 43 to 46 inches) and of steel-gray color. As a rule reindeer increase in weight from northwest to southeast. The Nentsi reindeer are smaller than those met in the Norilsk district, while the reindeer of the Tungus are much [: ] larger, weighing up to 330 pounds. On the northern slopes of the Saian Mountains ( Karagasskii olen ) [: ] it reaches a height of 55 inches. It is very strong, capable of carrying

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more than 200 pounds on its back over the most difficult roads. (According to data of the U.S. Dept. of Agriculture, the average reindeer in Alaska stands 42 to 44 inches high, measures 5 1/2 to 6 feet from nose to tip of tail, and has an average dressed weight around 150 pounds.^)^ The reindeer of the Tafalarsk district (Irkutsk region) is considered one of the strongest and largest. The weight of the male reaches 440 to 507 pounds, and it can easily carry a load of 175 to 220 pounds on its back. At the All-Union Exhibition in 1939-40, some 4-month fawns from the Tafalarsk district were shown weighing 185 pounds. The reindeer in Buriat-Mongolia, in the Amur region, and on the island of Sakhalin are also strong and larger than those of Yakutia. In the Chukhotsk district they are rather small, the carcass weighing 130 to 150 pounds. The reindeer of European Russian (such as those found in Malozemelskai Tundra, Bolshezemelskaia Tundra, and the Iamal s — Iamal reindeer) are a little larger than those of the Chukhotsk district but smaller than the Tafalarsk, Amur or Yakut reindeer. The average weight of the Kolguev reindeer is about 300 pounds for the buck and 250 pounds for the doe.
According to the views of some, the domestic reindeer of a given region are in size and other characteristics similar to the wild caribou (wild reindeer) of the same or neighboring districts.
Historical Background
The origin of reindeer breeding is still doubtful. According to some writers, it falls within historic times; by other investigators, domestication is believed to date from prehistoric time. D. M. Wilcken in his work on the history of domestic animals Grundzüge der Naturgeschichte der Haustiere

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(Leipzig, 1905), asserts that the domestication of reindeer occurred in prehistoric times. Recent Soviet investigators accept this, as in the article on reindeer breeding in the Great Soviet Encyclopedia.
There are many authors, however, who do not share the prehistoric theory. For instance, L. Reinhardt in his Kulturgeschichte der Nutztiere (Munich, 1912) maintained that the domestication of reindeer “could not have taken place much earlier than 500 years ago.” ^ Berthold Laufer for (27) suggests that ^ Reinhardt’s figure should be multiplied “at least by three.” Laufer concedes that the wild reindeer (caribou) was among the game hunted by Paleolithic man, but does not see any evidence that man of that age attempted to domesticate the animal. Laufer suggests that since the domesticated reindeer is lacking in aboriginal America, its domes– tication cannot be of very ancient date. Further proof of this, according to Laufer, is the fact that the classical authors, at least in western Europe, have left us no account whatever of the domesticated reindeer. The first good description of wild reindeer ( bos cervi figura ) under the name of tarandus , was by Julius Caesar. King Alfred of England (849-899) knew that in his time the Norwegians had domestic reindeer in the north of their country (26A).
Archaeological evidence does not shed much light on the question. In Scythian and Siberian antiquities of the Bronze Age, as well as on burial stones in Mongolia, figures of elk have been found, but they are shown as wild animals, and no representation of domestic reindeer accompanied by men has yet been discovered.
The question of where the first domestication began leaves less room for speculation than the question of time. It is true that the wild reindeer or caribou had formerly a much wider geographical range than at present. Its remains are found associated with hippopotami in Pleistocene formations much

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farther south in Europe than the location of the Hercynian forests, where they we e ^ r ^ e found by Caesar. All investigators agree that the domesticated reind d ^ e ^ er was the cultural property of the Old World and the domestication of the animal occurred somewhere in the territory which may be described at present as the Soviet North, together with the adjoining region of Lake Baikal. A former popular notion that the animals were tamed at first only by the inhabitants of the northern plains was banished long ago. Laufer (27) considered that “the records referring to the woodland reindeer are much older than those pertaining to the tundra reindeer of the maritime coasts,” and that “the woodland reindeer is the first in point of time to be domesticated and spread from the south into other regions, gradually developing into the tundra reindeer through infusion with the blood of wild forms of the tundra.” There is no historic evidence that reindeer breeding was restricted to a certain ethnic group. Apparently it was bound originally to certain localities with a suitable floristic environment.
The earliest records in existence containing references to the domestic– cated reindeer are Chinese. The Chinese were well acquainted with tribes in the north and northwest of their country, and although no Chinese author of the pre-Christian era made any allusions to reindeer, a story dating back to 499 A.D. contains such a reference. It was found in the Annals of the Liang dynasty ( Liang shu , ch. 54, p. 12) in a story told by the Buddhist monk, Huei Shen. In 499 A.D., J ^ H ^ uei Shen returned after a long journey to the Liang and gave a fabulous account of the land of “Fu-sang,” allegedly located far from the “northeastern ocean.” He reported that in Fu-sang people had vehicles drawn by horses, oxen, and stags, that they raised deer as oxen were reared in China, and that cream was made from deer milk. The

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whole description of the land of Fu-sang is considered a concoction placed together from heterogenous ^ ^ elements without any coherent geographical value, nevertheless, the allusion to reindeer is accepted as authentic. Since the breeding of horses, cattle, and reindeer ^ ^ is found concurrently only in the Baik e ^ a ^ l region, there is every reason to identify the oldest Chinese account of the tame reindeer with the culture of that region.
The Annals of the T’ang dynasty (618-906) contain another reference to reindeer breeding. Speaking of the tribe Wu-huan, which lived either east or southeast of Lake Baikal, the Annals state that those people “have neither sheep nor horses, but keep reindeer [stags] in the manner of cattle or horses. These animals subsist only on moss. They are trained to draw sledges [carts]. Moreover, reindeer skins are utilized as material for clothing.” ( T’and shu , ch. 217 B, pp. 7a-b).
Marco Polo’s travels also contain s a reference to reindeer. Speaking about a tribe called Mescript, he states: “They are a very wild race and live by ^ ^ their cattle, the most of which are stags and these stags, I assure you, are used to ride upon.” (Yule and Cardier. The Book of S e ^ i ^ r Marco Polo , Vol. I., p. 269). Marco Polo’s “Mescript” is identified with the Merkit in the country of Barga, near Lake Baikal.
In the history of the Mongols, written by the Persian historian Rash i ^ ī ^ d al-D i ^ ī ^ n in 1302, an allusion to reindeer breeding is understood by some specialists in the text where the author speaks of a tribe, “Woodland Uryangkit,” living in the o forests northeast of Lake Baikal. While the oldest Chinese records refer to reindeer breeding in the Baikal region, the Annals of the Mongol dynasty of the thirteenth century mention the existence of the domesticated reindeer at the source of the Yenisei and east of the

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river “Wu-se” (Is), an affluent of the Yenisei. ( Yuan shi , ch. 42, 63, p. 32 B; K’ien-lung ed.). These Annals speak of the Kirgiz living on the upper Yenisei, and mention a small territory “Han-ho-ha,” apparently inhabited likewise by the Kirgiz: “This region is accessible only over two mountain passes and abounds in wild game, while domestic animals are scarce….They transport their chattels on white deer and consume the milk of this deer.”
The Chinese noticed that reindeer were used also by the Tungusian tribe Oroci. The memoirs of the Manchu Tulishen’s embassy to the Kalmyk (1712-1715) describe briefly the reindeer among the Tungus living in the region of Irkutsk (G.T. Staunton, Narrative of the Chinese Embassy to the Khan of the Tourgouth Tartars ^ ) ^ . For the Yakuts the reindeer was a secondary acquisition. According to some historical records, the Yakuts were driven from the grass– lands near the Caspian and Aral seas by the Mongols of Genghis Khan and brought to the land of present Soviet Yakutia the steppe dwellers’ made of life. Their domestic animals were the horse and cow, while reindeer breeding was adopted by them from the Tungus. The Yakuts used to call e reindeer “Tungusian foreign cattle” and asserted that the Tungus were acquainted with no other domestic animals. (V. L. Seroshevski, Yakuty , vol. 1, pp. 146,307.) Historically, reindeer breeding has never occupied an important place in the culture of the Tungus and has spread mainly in the northern districts of Yakutia. The Yakuts did not adopt milking the reindeer and killed the animals for meat only on rare occasions.
Regarding the western centers of reindeer breeding in the Soviet North, the earliest reference is contained in the narrative of the Norseman Otter (Ohthere), who “said to his lord, King Alfred, that he dwelt farthes e ^ t ^ north

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of all Norsemen.” Around 870 A.D. ^ , ^ Ottar (26A) undertook several voyages in the course of which he navigated the White Sea, reached the south coast of the Kola Peninsula and became acquainted with the old Permians of the present northeast of European Russia. The account of Ottar’s travels is included in an Anglo-Saxon translation of the Hormista of Paulus Orosius and contains the following statement: “He [Ottar] was a very rich man in those possessions in which their wealth consists, that is, in wild animals. He still had when he came to the king, 600 tame deer, unsold. These deer they call ‘reindeer’; six of them were decoy-dear; these are much prized among the Finns [Lapps] because they capture wild deer with them.” (Alfred the Great. The Anglo-Saxon Version from the Historian Orosius. London, 1773).
No references to domesticated reindeer are found in the Finnish national epic poem, Kale y ^ v ^ ala , although the elk and caribou are mentioned in songs dating from a time prior to the Finns’ Christianization in 1151 A.D. Among the Russian sources, probably the earliest reference to reindeer is made in a document relating to 1499 (I.V. Shcheglov. Chronologicheskii perechen vazhneishikh dannykh iz istorii Sibiri 1032-1882 ; Published by the East Siberian Section of the Imperial Russian Geographical Society, Irkutsk, 1883, p. 12). In that year, Ivan the Great decided to complete the subjugation of “Yugra” - the territory of the Ural Mountains, inhabited by Vogul and other Ugrian tribes - and sent a military expedition there. In the description of this expedition, the following statements are found [translation]: “The military chiefs [voivody] slew 50 men of the Samoyed [Nentsi] on the Rock and captured 200 reindeer. …Proceeding from Lyapino they met the Yugor princes who came on reindeer from Obdor; from Lyapino the military chiefs traveled on reindeer; the army, however, on dogs.” (Lyapino, later called)

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Vorulsk, was a small town on the banks of the Sygma; Obdor was a settlement not far from the mouth of the Ob; “Rock” was a designation of the Ural; the distance covered from the Rock to Lyapino was about 290 miles.) This document makes it clear that the Uralic Ugrians wee acquainted with the domesticated reindeer at the end of the fifteenth century. Mention of domesticated reindeer is made in the epic traditions of Irtysh-Ostiak, traced to a period from the fourteenth to the fifteenth centuries (S. Patkov. Die Irtysch-Ostjaken ).
The credit for bringing the reindeer from Asia to C ^ E ^ urope is usually attributed to the Nentsi [Samoyeds] whom Laufer calls “the most skillful and successful reindeer breeders.” Their language has a very specialized nomenclature of reindeer and the equipment pertaining to reindeer breeding, and has special terms denoting the gelded and ungelded male, wild, domesticated, and hornless animals, calves (fawns) in various stages of growth, etc. Among the Nentsi, as among the Ostiaks, on sacrificial holidays the reindeer was slaughtered in honor of the gods. They also adopted, like all other reindeer- breeding tribes, the use of property marks for the purpose of recognizing their animals. While Chukchi used to bite a piece out of the fawns’ ears for this purpose, the Nentsi, Tungus and others cut marks on the ears of their reindeer. These tribes display [: ] great skill in lassoing the reindeer.
As to reindeer driving, it apparently developed as an imitation of driving dog sledges, which for centuries had been a distinctive method of transport– tation among such tribes of Eastern Siberia as the Koriaks, Kamchadals, Giliaks, and Yenisei-Ostiaks. An old statement regarding the use of reindeer with sledges is found in the information on Siberia gathered from oral accounts by the Jesuit Philippe Avril who visited Moscow in 1664, having

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been commissioned by the King of France to discover a new land route into China: “To make the reine-deer go more swiftly they tie a great dog behind, that scaring the poor beast with his barking, sets her a running d with his barking, sets her a running with that speed, as to draw her burthen no less than forty leagues a day. …But that which is more wonderful as to these sort of sledges, they are driven along by the wind sometimes over the land cover’d with snow, sometimes over the ice of frozen rivers, as our vessels that sail upon the sea. For in regard the country beyond Siberia is open and extremely level as far as Mount Caucassus, the people who inhabit it making use of this advantage to spare their beasts, have so order’d their sleds, as either to b y ^ e ^ drawn along by the reine-deer, or else to carry sails, when the wind favours ‘em.” (Jesuit Philippe Avril. Voyage en divers etats d’Europe et d’Asie . Utrecht, 1673, Paris, 1692. Also published in English under the title “ Travels into Divers Parts of Europe and Asia ”, London, 1693.)
The custom of decorating riding deer with ribbons to which glass beads or buttons are sewn has been widespread among the natives of Siberia, par– ticularly the Tungus. It has continued until modern times and as recently as September 1950, the Soviet Weekly reported on the traditional “Reindeer Day” festival held in the Nenets National Territory, at which the competition for the best decorated animals and sledges was a part of the program.
According to Laufer, reindeer breeding spread westward from the Nentsi to the Ugrian tribes of the Ural and the Lapps, and eastward to the Tungusians, from the latter to the Yakut, Chukchi, and Koriak. When the Nentsi moved northward beyond the forest, they took along their woodland reindeer. Their old stocks were gradually replenished by capturing the northern caribou, until a point was reached when the better bread predominated.

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Soviet scientists, in their attempts to determine the original area of the domestication of the reindeer, are less definite in their conclusions than were some of their western colleagues, particularly [: ] Laufer. For instance, volume 43 of the Great Soviet Encyclopedia (1939) states that the question of indicating the exact region where the reindeer were originally domesticated is highly controversial; the “two oldest centers of reindeer breeding” established with certainty are the Nymylanskoe in eastern Asia, and the Nenets in the west. The Soviet scientists also consider it proved that the larger ethnic groups of the north adopted reindeer breeding from the smaller tribes; the Yakuts took it from the Evenki, the Komi from the Nenets, etc.
Perhaps one reason for the doubt displayed by Soviet explorers regard– ing the origin of reindeer breeding is the confusion of names applied by historians to various reindeer breeding tribes of the Far North. Early Russian traders often gave these tribes names which were really terms of derision. Only in recent years have the correct names been restored or established with the help of modern ethnography and better knowledge of local languages and dialects. For instance the word “Samoyeds,” mentioned as early as 1096 in the Old Russian chronicle of Nestor, actually means “people who devour one another.” The tribe called Nentsi, living at present on the shores of the Arctic Sea, from the Mezen River to the Taimyr, constitute apparently the bulk of former “Semoyeds.” These Nentsi, however, should be distinguished from Nentsi (Gods) living in the Far East. Possibly also some of the Ostiaks should be included in the group of former “Samoyeds.” Some [: ] of the supposed old tribal names proved to be simply the native words designating “men” or “people.” Chukchi living in the Soviet Chukotsk-National

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District were recently renamed the Luoravetlans, since, “Chukchi” simply means “rich in reindeer.”
It has been said that reindeer are less dependent on man than other domestic animals such as horses and cattle. This is largely a matter of adaptation to environment. The reindeer respon ^ d ^ s to care and feeding as other animals do, but it is at home in arctic and subarctic regions; it can find its own food if necessary, and it requires no shelter from the cold, while sheep and cattle imported from warmer climates require stabling and other care. Large herds of cattle and sheep do live through severe winters on the plains of the western United States without shelter, but every now and then, in bad winters, the losses are severe.
Reindeer Breeding Before the 1917 Revolution
As an organized industry, reindeer breeding did not exist in Imperial Russia before the Revolution. It was considered an occupation of the primitive tribes who inhabited the Far North and the Russians did not display active interest in its study or development. When Lieutenant Berthof visited St. Petersburg on his assignment to purchase the reindeer for Alaska, he “was astonished at the apparent lack of information at the government’s disposal regarding reindeer matters in Siberia…. It was impossible to find out h where the deer were or the number of deer, and I got the impression that the officials didn’t much care.”
The Great Encyclopedia of F. A. Brockhaus and I. E. Efron, published in St. Petersburg in 1897, indicates that in 1888-89 the reindeer herds in Russia amounted to 533,000 head; of these 263,000 were in European Russia, 198,000 in Siberia, and 72,000 in the principality of Finland. In 1893, the

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number of reindeer in the Archangel Region was 270,000 and it increased to 367,128 in 1895.
A 1914 publication of the Resettlement Administration of the Imperial Government (22) stated that “There is no exact figure available as to how many natives in Siberia are engaged in reindeer breeding and what number of the reindeer they possess.” According to this source there were about 1,100,000 reindeer in Russian Asia in 1906; of these 515,000 were in two districts of the Tobolsk region, 95,360 in the Yeniseisk region (mostly in the Turukhansk district), 287,000 in four districts of the Yakutsk region, 176,000 in the Primorsk region (including Kamchatka), 2,000 in the Narymsk district of the Tomsk region, 2,400 in the Irkutsk region, about 1,500 in the Transbaikal region, and also a small number in the Amur region. “Neverthe– less,” states the above source, “these figures are not reliable and actually there should be many more reindeer, since the count was not made everywhere.”
The number of reindeer in Yakutia dropped to 120,900 by 1911, such decreases having occurred often owing to a complete lack of zootechnical and veterinary facilities, depletion of pastures, and frequent outbreaks of various epizootics. The “Siberian plague” (anthrax) was a real scourge to reindeer breeders, in the Bolshezemelskaia Tundra alone, 76,000 animals died of disease in 1898, 200,000 in 1907, and 100,000 in 1911.
Some influential Russians were concerned for the welfare of the reindeer industry. There were, for instance, the nineteenth-century projects of Senator Unterberg and N. L. Gondatti (later Governor-General of the Priamur region) aiming at the encouragement of reindeer breeding.
In the early 1900’s, the Russian government forbade the exportation of reindeer to Alaska after the first few purchases (1,280 head) made by the

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United States government. The reason for this unexpected veto has never been explained; It may have reflected the apprehension of the Russian government that further exports on an increasing scale could have depleted the East Siberian herd with damaging effects on the natives’ economy, or it may have been part of a general plan to discourage non-Russians from attempts to build up a commerce with the natives of northeastern Siberia.
Shortly before the end of the Imperial regime, there occurred a dramatic episode connected with the utilization of reindeer (25). In 1915 the Russian government was hastily building the new port of Murmansk and the railroad across the Kola Peninsula in order to arrange a channel of transportation for munitions and supplies coming from the western Allies, of which Russia was critically in need. As early as November 1915, a question was raised in St. Petersburg as to how the cargoes from Murmansk could be delivered to Rovaniemi, the nearest railway station in Finland, before the railroad con– struction was finished. Rovaniemi was roughly 150 miles from Kandalaksha, a Russian town approximately 135 miles south of Murmansk. About 12 miles of the railway line was already built from Kandalaksha toward Murmansk by January 1916. Under the auspices of the Department of the Navy, a meeting was called in Archangel to discuss with local businessmen the possibility of using reindeer. The consensus was that reindeer transportation would be too expensive and impractical. A navy captain named Roshakovski did not agree with the majority. With the help of a local man he made an investigation and reported his conclusions to the Tsar. All precedents were broken and he obtained official permission to use reindeer and an appropriation for the project.
In the meantime a few British ships loaded with munitions anchored in

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Murmansk, being unable to proceed to Archangel. On January 1 6 ^ 5 ^ , 1916, Roshakovski was back in Murmansk and on the same day the loading of the cargo on sledges began. About 15,000 reindeer with an adequate number of drivers was drafted. Day and night the caravans of sledges were dispatched from Murmansk under the supervision of noncommissioned officers, with one, two, or four reindeer carrying each sledge loaded with boxes of munitions. The reindeer were driven about 45 miles without rest and then released to pasture, while fresh reindeer assembled at established points were harnessed. At times 20 reindeer trains were in circulation. At a point reached by the railway under construction, the cargo was loaded on cars, delivered to Kandalaksha 12 miles away, again reloaded on sledges and carried to Rovaniemi. Some cargo was shipped directly to Petrozavodsk, over 500 miles south of Murmansk, but that distance proved to be too long. By the middle of April the thaw– ing of the snow interrupted this unusual transportation. By that time, however, the reindeer had delivered to the Russian army 250,000 rifles with adequate quantities of cartridges and a great quantity of hand grenades. It was revealed later that about 1,000 reindeer died as a result of exhaustion in the course of the spectacular performance.
Reindeer Breeding after the Revolution
The first years after the Revolution of 1917 did not create conditions especially favorable for the reindeer husbandry. Although there are con– siderable discrepancies among the available statistics, the depletion of stocks by 1933 is evident. Between 1926 and 1933, the reindeer of the U.S.S.R. decreased from 2,193,000 to 1,931,000 (P. V. Orlovski, “Agriculture in the Far North”, in Sovetski Sever , 1933, no.2, p.15). This reflects the general decline of Soviet animal husbandry in 19 3 ^ 2 ^ 9-33 as a result of

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collectivization and violent resentment to it on the side of the wealthier groups of the rural population (kulaks). Also, continuous losses from various diseases had their effect. For instance, in 1926-27 in the Kamchatsk region and on the Okhotsk coast 5% and 4.6%, respectively, of the reindeer were lost because of various diseases, while 1.8% and 3.0% disappeared from “unknown reasons.”
After 1939 the Soviet government ceased the regular release of statistics in absolute figures. The available statistics are usually given in percentages showing changes in comparison with a certain year for which no absolute figures are known. This makes it very difficult to estimate the present reindeer population of the U.S.S.R. or to determine the number of animals from year to ^ ^ year. Nevertheless, an attempt is made here to arrive at the figures sought. Table I may give an approximate picture of the reindeer stocks of the U.S.S.R. in 1933-36.
^ The ^ Great Soviet Encyclopedia states that in 1937 the U.S.S.R. possessed about 70% of the world’s reindeer population which were bred on an area of about 11,000,000 square kilometers; that year the share of Alaska was 16% and that of Scandinavia 14%. In 1948 the same Encyclopedia indicated that the total number of reindeer in the U.S.S.R. at the beginning of 1940 equaled “approximately 2,000,000” (against 2,138,000 arrived at in Table I.) The increase of reindeer after 1943 was interrupted by World War II. The war increased the demand for reindeer meat and consequently caused greater slaughter of the animals; also it led to the drafting of many experienced herdsmen.

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Scroll Table to show more columns

Table I. Reindeer Stocks of the U.S.S.R. a
Year In % of 1933 herd In millions
1933 100.0 1,615
1934 104.2 1,683
1935 110.5 1,785
1936 117.0 1,890
1937 122.9 1,985
1938 128.1 2,069
1939 130.8 2,112
1940 132.4 2,138
1941 133.3 2,153
1942 123.3 1,991
1943 118.2 1,909
1944 118.8 1,919
1945 119.8 1,935
1946 129.8 2,096
1
Table II gives a picture of the prewar distribution of reindeer herds in the Asiatic portion of the Far North:

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Table II. Reindeer in the Asiatic Far North (13).
Geographical regions 1926 1933 1937 1938 (according to the plan)
^Ob ^ Sever: 438,000 303,800 418,300 454,700
Ostiako-Vogul district 128,000 92,000 74,200 80,700
Yamalo-Nenets district 310,000 211,000 344,100 374,000
Narym district 5,000 --- 3,000 3,300
Yenisei Sever: 179,000 124,800 152,800 169,000
Taimyr district 119,000 61,300 80,300 90,000
Evenk district 49,000 52,200 57,100 63,100
Igarka and Turukhansk counties 11,000 11,300 12,700 ---
Yakutian A.S.S.R. 118,000 142,100 167,300 194,100
Khabarovsk Sever 857,900 578,000 610,900 669,000
Total 1,597,900 1,148,700 1,352,300 1,490,100
In 1940 the centers of reindeer breeding in the Asiatic Far North were the northern districts of the Khabarovsk and Omsk regions. The industry was less developed in Yeniseisk Sever and in Yakutia, where about 80% of the reindeer were found in the 16 northern districts (about 16% of the Yakutian reindeer are in the Aldan district). In the Narymsk district it was of no economic importance. Of the 610,900 reindeer in the Khabarovsk Sever in 1937, 498,000 or 80.2% were in the Kamchatka region (not so much, however, on the Kamchatka Peninsula itself) while only a little over 2% were on the island of Sakhalin. In the Chukhotsk district reindeer numbered 537,000 in 1934, while by 1940 about 62% of all the reindeer of the Asiatic Far North were concentrated here (13). No recent data are available on the value of reindeer production in rubles. In 1926-27 the total gross value of the products was given at 1,097,000 rubles.

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

In the Murmansk district, within its post-1920 frontiers, there were 74,000 reindeer in 1914, the figure dropped to 28,300 in 1921 and then increased to 54,000 in 1934. It was established that this stock would reach 82,000 in 1938, of which about 10,000 would be in the industrial district of Murmansk itself.
Attempts with domestic reindeer in Novaya Zemlya before the Revolution were unsuccessful, although wild reindeer are found here in great numbers. The first attempts to develop the industry here were made in 1928 by Sevkraigostorg (the Trading Office for the Northern Regions). This organi– zation sent a large herd of reindeer with an adequate number of herdsmen to Novaya Zemlya but lack of knowledge of the local conditions prevented success. When the Soviet Academy of Science came to the assistance of the local state farm in 1930, progress was made, especially after the 1932 survey of pastures. No data are available regarding the number of reindeer in Novaya Zemlya at present.
After the Revolution, one of the first undertakings of the Soviet authorities was an attempt to improve the bread, and to organize a scientific crusade against those diseases which were periodically depleting the herds and badly damaging the products of the reindeer industry. (For instance, one parasite, the warble fly ( Oedemagena tarandi ), sometimes depreciated the value of reindeer hides 75 to 80%.) A number of special studies were undertaken for the prevention of epizootics, veterinary facilities were provided on an ever-growing scale, mass vaccination of the reindeer was introduced, and experimental breeding farms were organized. In 1932 the Reindeer T o ^ r ^ ust opened courses to provide specialists in reindeer breeding; subsequently a number of technical schools were opened for the same purpose.

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

In the course of the First Five-Year Plan over 120 zootechnical and veterinarian aid posts were opened. As early as 1924 two special labora– tories were established in Salekhard and Izhma and assigned the task of studying methods of fighting anthrax. Later studies on the prevention of reindeer diseases were carried out by the Institute of Urogravidantherapy. It is reported that such diseases as anthrax no longer threaten the reindeer of the Soviet North.
In 1931 a special Institute of Reindeer Breeding was founded in Leningrad. It sponsors various projects to foster reindeer breeding, conducts research work, organizes expeditions to the Far North in order to study local condi– tions of husbandry, publishes literature on the subject, etc. Since 1936 the institute has functioned as a branch of the All-Union Institute of Polar Agriculture.
In 1936 Glavsevmorput (Northern Sea Route Administration) submitted to the government a project for the establishment of three scientific research institutes at Obdorsk, Igarka, and Yakutsk for the study of agriculture and animal husbandry in the Arctic. Within the framework of these institutions 14 district stations were to be opened, to embrace all Yakutia and the northern sections of the Ob-Irtysk and the Yenisei regions. This network of scientific institutions was to study the native animals, work on the problems of reindeer transportation, and train specialists from among the natives.
On the eve of World War II, 4 regional stations and 5 zootechnical bases in the Far North were devoting their work to the d problems of rein– deer husbandry exclusively. An inventory of better-bred stocks was taken in some districts, and the method of artificial insemination has been applied.

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

Concentrated fodders and portable corrals are used effectively in many places. Methods of more efficient protection of reindeer against wild animals, the wolf in particular, were studied. (In 1926-27 approximately 75,000 reindeer were victims of wild animals.) A special set of rules for reindeer breeding was worked out and decreed by the government, these decrees regulate the use of pastures, prescribe the obligatory zootechnical and veterinary measures, etc. A periodical called The Soviet Reindeer Industry is published several times a year, and is devoted to various problems of reindeer husbandry and technology
State farms and collective farms are considered by the Soviet govern– ment as the main channels for application of the achievements of science and better methods in reindeer breeding, and mass collectivization in the Far North began in 1929. However, the private segment in animal husbandry has been for years more important in the North than in any other part of the U.S.S.R.; this is especially g true regarding reindeer breeding in the most northern districts of the Asiatic Far North. In 1937, 85.5% of the reindeer on the Chukhotsk Peninsula and 59% of the ones in the Khabarovsk Sever were still the individual property of the natives. By January 1, 1937, 19 collective farms had been established by Glavsevmorput. Of “about 2,000,000 reindeer” indicated by the Great Soviet Encyclopedia for 1940, 436,000 are listed as the property of collective farms, while those of the state farms is estimated at 243,000. A 1939 publication of the Gosplan listed 36 “rein– deer state farms.” One of the largest of them is in Novaya Zemlya. Some of the reindeer state farms received from the government an allotment of 1,000,000 to 2,000,000 hectares of land.
It was reported that the best collective farms achieved considerable

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

success in the annual increases of their herds. These farms contributed also to better organization of reindeer transportation. At present the reindeer farms interested in incomes from transportation services submit their claims to the local district administration, which works out a plan of transportation operations and allocates the demand among the collective farms; then, according to the plan, orders for the required number of reindeer, sledges, drivers, etc., are issued.
Reindeer Herding
Efficient methods of herding and proper utilization of pastures constitute the basis for successful reindeer breeding. Almost 68% of the northern prairie is considered suitable for pasture in the summer; of this 39% is also suitable for winter grazing. At the end of summer the rein– deer are driven southward toward the forest border. In spring a northward shift takes place. Such seasonal drives in the Bolshezemelskaia Tundra sometimes take s the reindeer more than 300 miles to the south. At the Lena the distance between the extreme points of movement is usually from 110 to 125 miles.
Approximately 400 far northern plants, that is, about one-third of all plants known there, may be used by the reindeer as fodder; lichens are especially important, for they preserve their nourishing value all year round, which makes them the important winter food. The animals, perhaps by their sense of smell, find the lichens underneath the snow and uncover them by pawing. Some lichens, however, grow only 0.3 to 0.4 centi– meter a year. It may take ten years before they re-establish themselves on overgrazed land; hence overgrazing presents one of the chief difficulties

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

to reindeer breeders.
In the U.S.S.R. a systematic investigation of lichen growth was first undertaken by two stations of the Institute of Reindeer Breeding, Salaskin (at Oksino on the Kola Peninsula) and Igoshina (at Labytnang on the lower course of the Ob). This study made it possible to work out a theoretical basis for choice of pasturage. Grazing should not exceed a certain limit; a sufficient quantity of viable lichen bases must be left for recovery of the pasture. Also the propagation of such valuable and comparatively quick– growing fodder lichens as Cladonia rengifera and [: ] C. uncialis is practiced.
In 1932 the institute made surveys of pastures, sometimes from the air, and grazing grounds were divided by the government so as to avoid overgrazing. From 40 to 70 hectares of pasture per year is required for each reindeer, according to Soviet sources. (The United States Dept. of Agriculture studies of carrying capacity of pastures made by pasturing reindeer within fences in Alaska indicate 33 acres as the minimum year-long grazing area requirement for one reindeer.) Such norms made it possible to establish the capacity of each territory and apportion the number of rein– deer accordingly. The haphazard movement of peoples and herds were brought under control. Long journeys from winter to summer feeding grounds were reduced to a minim.
Reindeer have natural periods of loss and gain, with some variations dependent on age, sex, and castration. Winter is generally their lean season, while during summer they fatten, gaining as much as 3/4 of a pound a day in the Malozemelskaia Tundra. The increase in weight at peak-condition periods reaches as much as 60 pounds in steers and 40 to 50 pounds in other animals.

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

In winter the reindeer spend about 11 hours a day feeding. They may be maintained in good condition on cultivated fodder, such as hay, grain, fish, and bone flour. Over 13 pounds of fodder is required daily by a nonworking animal. About 4 pounds of salt a year is consumed by reindeer in pasture and 5 pounds in a feed lot. During World War II, when reindeer were used on the northernmost front, artificial fodders proved to be very useful.
Reindeer are herded much as sheep are on the large western ranges in the United States, by herders following the animals as they move about. In selecting pastures, not only the question of fodder is considered, but also the problem of the insects which attack the animals during the warm season. The reindeer requires protection from insects either by locating summer pastures on higher ground exposed to the wind, or by the use of fly repell a ^ e ^ nts and similar devices. A reindeer may lose as much as 125 grams of blood a day if exposed to mass attacks of mosquitoes, and cases have been recorded of animals dying from insect bites.
A reindeer herd should be constantly watched. They are marked by cutting or notching the ears; or a metal tag or button may be used for identification. The natives are skillful in identifying each individual animal; it is said that in one of the nomad languages there are 16 different words to designate various shades of the gray-brown color of the local reindeer.
Trained dogs are useful to herders. When the United States made a first appropriation of $6,000 for the introduction of reindeer in Alaska Dr. Shel [: ] ^ d ^ on Jackson sent a notice to the Scandinavian newspapers in the United States, seeking the services of experienced Scandinavians acquainted

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

with methods of reindeer breeding in Lapland. In the great majority of the 250 answers received, it was stated that the trained dogs of Lapland were necessary for herding. When the first superintendent of the Alaskan Teller Reindeer Station, William A. Kjellman, was sent to Lapland, his assignment was to bring not only Lapps but dogs.
However, Soviet herders rely on dogs less than do the Scandinavian Lapps. The Nenets and Komi herders use dogs more than anyone else in the Soviet Far North, the Evenki and Yakutians seldom use them. Generally, the farther from the Kola Peninsula toward the Lena, the less use is made of dogs, and east of the Lena up to the Chukhotsk Peninsula, the reindeer breeders have never used dogs in herding until recently. Shortly before World War II, the utilization of dogs in herding somehow became known in the Bulun district of Yakutia, in Chukhotsk and Koriak districts, and in the Khabarovsk region. With a few exceptions, dogs are not used even now in herding in the T ^ t ^ aiga, although experiments have proved that they can be useful there.
Reindeer are castrated to produce steers for meat and to reduce the number of bucks to a proper proportion of the herd. Fawn crops average between 50% and 60% of the adult doe population. Male and female fawns occur in about equal numbers. Twin fawns are rare. The average doe breeds to an age of 10 to 12 years. The newborn fawns weighs from 7 to 16 pounds.
The size of a herd may vary from 100 or 150 head to 3,000. The desirable size for the Komi and Nenet herds, as advocated by Soviet specialists, ranges from 1,200 to 1,500 animals. While larger herds in open country are not objectionable ; ^ , ^ on land even partially forested, herds of from 250 to 600 animals are more desirable. From 5 to 6 herdsmen are required to take care

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

of 2,000 Lead according to experience in the Murmansk region.
Utilization of the Reindeer
For centuries reindeer breeding has been one of the main occupations (sometimes the only one) of a great many tribes living on the northern prairie and at the northern border of the forest. The nomads had mastered the art of making full use of the “all provider.” However, the emphasis placed on various forms of reindeer utilization differs from region to region, for instance, riding or milking is not practiced by all the natives of the Far North. On the Kola Peninsula, in the Archangel region, and in the region of Omsk, the utmost importance is attached to the reindeer’s meat and skin; but in the Krasnoiarsk region and forested districts of Yakutia the reindeer has been appreciated mainly as a draft animal. In some forest districts of the Krasnoiarsk region, and in the Tobolsk district of the Omsk region, reindeer milk is considered important.
The folk tales of the Nentsi express colorfully the natives’ apprecia– tion of the reindeer harnessed to a sledge and say that it is “second to none in the world.” What the reindeer meant for the natives was emphasized in Russian Asia (22), published by the Resettlement Administration of the Imperial Government in 1914 [translation]: “[the reindeer] is so useful that in the whole animal kingdom you may find hardly another creature which might equal it. In the full meaning of the world there is nothing in this remarkable animal which is not used in the everyday practice of the natives of the North who are blessed by its possession. Its fatty milk, similar to cream, serves as an excellent beverage and is used in the preparation of a very tasty cheese. Meat, internal organs, marrow, brains, and the blood are used

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

for food; also the antlers, before ossifying, may be used for food; jelly can be made of them, which is considered not only tasty but also as a medicinal food. The skin is used for making the natives’ clothes as well as for shelters. All kinds of household items are made of the bones and the ossified antlers: knives, forks, spoons, sewing needles, etc. Hollowed– out hoofs are used instead of cups and glasses. The sinews are used for thread and the hair for spinning and weaving. The intestines are used in the preparation of sausages stuffed with a mixture of blood and various roots. Candles are made of the fat. The stomach is dried and serves as a substitute for kitchen utensils. Even the content of the stomach, if used immediately after the reindeer is slaughtered, is not wasted; the natives consider it a delicacy and are only too glad to use it as food. Considering the complete absence of roads in the tundra, the reindeer serves as the only pack and transportation animal which can move with ease over the most boggy ground and through snows of any depth.”
Three main types of reindeer breeding are distinguished by the Soviet specialists at present: ( 1 ) Breeding which emphasizes meat as the main product and the skin as a product of secondary importance. This type of breeding requires the maintenance of a proportionally large number of gelded reindeer rather than fawns or does (about 37%); it prevails in the eastern tundra, on the Chukhotsk Peninsula and in the Koriak district. ( 2 ) Breeding with emphasis on skin production; it is developed mainly in the Murmansk region, in Nenets and Iamalo-Nen ^ e ^ ts districts, and in Komi A.S.S.R. The best results here are achieved by slaughtering fawns in the fall and keeping herds with a possible maximum number of does (up to 45%). ( 3 ) Breeding with emphasis on the reindeer as a draft animal, which calls for a smaller number of does

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

in the herds (about 30%); it prevails in the forest zone where reindeer milk is also widely utilized.
Since the importance of reindeer transportation on the northern prairies is great, Soviet authorities consider it their ta ks ^ sk ^ to obtain a bread of animals which will be equally good as suppliers of meat and as draft animals. This is especially important for reindeer breeders in Yakutia, Chukhotsk Peninsula, Ismalo-Nen ^ e ^ ts and Taimyr districts.
Draft Animals . Without the reindeer, remarkable for endurance, strength, and speed in drawing sledges, much of the Soviet Far North could hardly be inhabited permanently. The chief advantage of reindeer over dog transporta– tion is that a grazing animal finds its own food, while provisions must be carried for dogs and their maintenance often proves difficult. Also, in an emergency the reindeer is a source of food to its master. Windy weather is difficult for the dog travel, especially when heading into the wind, while reindeer naturally travel to windward, even in a blizzard, this being a trait they have from their wild forebears, the caribou. The reindeer does not mind a temperature of −80°F.; its broad, deeply cleft hoofs fit it admirably for traveling over the broken snow, and it makes it way through areas of deep snow more easily than even a horse. Gelded reindeer between the ages of 4 and 12 are preferred for drawing sledges. The training of animals for this service starts with the third winter.
The Reindeer Breeding Branch of the All-Union Institute of Polar Agriculture undertook a number of projects for improvement of the technique of reindeer transportation. Serious defects were found in the traditional equipment used by the natives, which f ^ v ^ aried from place to place. For instance, on the Kola Peninsula and in Karelia the natives use harness with a collar,

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

but breast straps or neck bands, with all kinds of local modifications, are used in the rest of the Far North. When the Samoyeds, ^ harness ^ a few animals to draw a sledge, they place the foremost one on the left side; placing the leader on the right side is called the dolganski , method of harnessing. To urge the reindeer to go faster, the natives prod them rather cruelly in the back with a long birch pole which has a bone button affixed to its end. The Nentsi almost never use reindeer for riding. The sledge used for transport– ing heavy loads is called narta . It may be 3 meters long; the distance between runners is up to 80 centimeters. In building their narta the natives formerly used ropes and wooden rails only, since iron nails were unobtainable, and, besides, iron breaks easily under the low temperatures of the Arctic. Freight up to 770 pounds may be loaded on such a narta , drawn usually by four reindeer.
Newly devised equipment enables the adoption of reindeer for services to the Red Cross, such as carrying stretchers and transporting the sick, while a new harness recently devised makes it possible to use reindeer for work which until recently has been considered too heavy for them, such as plowing and harrowing.
The Encyclopedia Americana indicates that full-grown reindeer can draw a sledge with a load of 300 pounds and travel at the rate of 100 miles a day. w ^ v ^ arious sources of information, however, give different figures in this respect, since much depends upon the size of the reindeer in a given region, the habits of the natives, and the care they display regarding their animals. According to the Soviet Resettlement Administration, the reindeer, without being fed, easily makes 65 miles in 10 to 12 [: ] hours, provided it is permitted to stop for 1 or 2 minutes every 6 or 7 miles; however, cases

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

of reindeer covering a distance of more than 130 miles in 24 hours are not at all rare. N.N. Urvantsev reported in 1928 that in the course of his expedition he traveled on one occasion from Norilsk to Dudinka in 5 hours and 37 minutes, having covered a distance of about 62 miles.
Interesting records were established at special reindeer races organized periodically by the Soviet government. In their primitive form, such races have been popular among the tundra natives from time immemorial and are still on the programs of the yearly reindeer breeders’ festivals. The names of the victors are retained in the memory of reindeer breeders for years. Reindeer races were introduced in their modern form in the 1920’s. At the races held in the Leningrad hippodrome between December 6, 1931, and February 7, 1932. the record of the winning troika (three reindeer harnessed abreast) was 1,600 meters in 2 minutes and 39 seconds. These reindeer were from the Kola Peninsula. In 1933, at the races in Narian-Mar, Nenet National District, the best results were achieved by four reindeer of local breed, harnessed abreast, which covered 1,380 meters in 2 minutes and 10 seconds. At the race in Lovozero, Murmansk region, on December 7, 1938, the distance of 1,600 meters was covered by four reindeer harnessed abreast in 2 minutes and 40 seconds.
In tests of the reindeer’s endurance and weight-drawing capacity the average records achieved at the races in Yakutia in 1936 and 1937 were as follows: for endurance, a weights of 900 to 1,000 pounds were pulled 12.4 miles by two reindeer harnessed abreast in 1 hour and 20 minutes; for maximum weight, two reindeer harnessed abreast pulled a load of 2,200 to 5,500 pounds a distance of 400 meters.
The following speed and average load were recommended in 1948 by the Administration of the Far North of the Ministry of Agriculture of the

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

R.S.F.S.R. for reindeer continually employed for transportation services.

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Table III. Norms Recommended for Reindeer Transportation (26).
Type of road Load on sledges with 2 reindeer, in kg. Speed, km. per hr. Distance per day, in km.
Good smooth road 300 4-5 25-30
150 7-8 45-50
120 9-10 55-60
Average tundra road 200 4-5 25-30
100 7-8 45-50
80 9-10 55-60
Poor tundra road 100 4-5 25-30
50 7-8 45-50
40 9-10 55-60
The Administration also worked out a set of recommendations which should be followed in the use of reindeer for commercial transportation. A new harness, devised recently by a member of the staff of the All-Union Institute of Polar Agriculture, S. P. Popov, makes it possible to increase the load up to 700 or 800 and even 1,000 kilograms in nartas used on a good smooth road. The winter reindeer transportation train ( argish ) usually consists of 24 to 30 nartas , 70 to 85 reindeer (of which 8 or 10 are kept in reserve), and 4 or 5 men. Strings of sledges in greater numbers are considered less convenient because of the possible difficulties of finding food for a larger number of reindeer; if a string consists of only a few sledges, the utilization of the drivers becomes less economical.
The utilization of reindeer for riding and carrying pack loads is important in forests and among mountains, as well as on the [: ] prairie during the summer. Under normal conditions the reindeer may be used for

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

these services for a period of 5 to 6 years, up to an age limit of 10 years.
A 1934 United States Dept. of Agriculture publication on reindeer breed– ing in Alaska indicates from 60 to 70 pounds as a normal pack load for an animal which covers from 20 to 25 miles a day. The natives in Siberia usually load as much as 60 kilograms (more than 130 pounds) on a reindeer’s back. The Soviet Administration of the Far North, however, suggests using the following norms for continued employment of animals for this type of service.

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Table IV. Norms for Pack-Load Carrying (26).
Type of road Load on one rein– deer, in kg. Speed, km. per hr. Distance per day, in km.
Good smooth trail 50 4-5 25-30
35 6-7 40-45
Average broke country 40 4-5 25-30
28 6-7 40-45
Very broken country 30 4-5 25-30
20 6-7 40-45
Experiments with pack equipment have shown that the center of gravity of the reindeer’s body lies considerably farther forward than in other transport animals. Hence the usual type of pack equipment intended for horses proved unsuitable, as it overloads the hind legs. A new type of saddle and pack were cons ^ t ^ ructed at the Bulun Reindeer Station in Yakutia in 1937, which allows the reindeer to carry loads up to 50 or 60% of its own live weight, with a maximum not exceeding 80%.
The reindeer as a draft animal has proved important for Soviet hunters and ^ trappers ^ in North. Trap lines are long and the Siberian hunter sometimes has

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

to cover from 30 to 45 miles a day in order to inspect them, besides, these trap lines may be several hundred miles from the hunter’s home. Under these conditions reindeer transport becomes as indispensable basis for the fur trade, and possession of reindeer strongly affects the incomes of members of the collective farms engaged in hunting and trapping. For instance in 1945, the Evenki National Region, the yearly income from hunting averaged only 165 rubles in the case of a farmer without reindeer; for those who had less than 40 reindeer the income was 2,499 rubles; and for those with more than 40 reindeer, 3,469 rubles.
Observations of reindeer under saddle and pack established the fact that animal the [: ] ^ animal ^ has a constant length of stride on a level trail and changes it with strict regularity, depending upon the angle of slopes and ascents. This discovery made it possible to use the reindeer stride as a measure of length of the roadway traveled by geological, topographical, and other research expeditions.
The modernization of transportation methods does not decrease the reliance on reindeer transport in the Soviet North; on the contrary, its importance has been constantly increasing. A publication of Glavsevmorput, 1939, says: “Numerous large scale projects of the Far North were material– ized almost exclusively with the aid of the reindeer transport. In many regions of the Far North, reindeer transport is the basic means of conveying people and freight. Also in strengthening the defense of our country, the reindeer may occupy not the last place.” A 1948 publication by the Far North Administration calls the reindeer transportation “irreplaceable” and states that “in the majority of the regions of the Far North the demand for reindeer transportation has been increasing yearly.”

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Carrying good ^ s ^ from polar sea ports southward across the prairie and even into the forest, delivering meat, fish, skins, furs, and a number of other raw materials, is done by reindeer transport. Thousands of reindeer and men are engaged in this work. In 1936, in the northern regions of Yak ^ u ^ tia, 2,680,000 rubles, or 21.8% of the gross incomes of collective f o ^ a ^ rms, came from reindeer transport services. The corresponding figures for the Iamalo-Nenet ^ s ^ district was 403, 0 ^ 1 ^ 00 rubles or 15%; for the Ostiako-Vogul district, 638,700 rubles or 16%. In 1946 the income from reindeer transporta– tion constituted 17% of the total income of the collective farms in the Far North, while for some of them it [: ] was the basic income.
As a matter of curiosity it may be added here that the elk ( Alces alces ) may become a serious competitor to reindeer in some parts of the northern forest. In accordance with an order of the Yakutian government, whose country is very rich in elk, experiments were made in 1936 at the Yakutsk Agricultural Experimental station in training and driving the elk. The tests were continued in 1937 and proved that the elk, if caught at the age of one month, is easily handled and domesticated, readily fed, and as a draft animal shows high ability.
In the northern districts of the Tiumen region, E ^ K ^ rasnoiarsk Krai, Yakutia, and other districts of the Far North, there are established reindeer fr ^ e ^ ighting lines. Every 40 or 50 kilometers special shelters have been built — chumy , made of skins for winter and tarpaulins for summer. They are equipped with stoves and provide comfortable resting places for the drivers; also feed for reindeer is kept in some of them. These freight– ing lines are so planned that the overnight stopping places are at good pastures. The length of some of these lines reaches 1,000 [: ] kilometers.

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Meat . Meat is another entry on the list of articles and services offered by reindeer breeding. The full-grown reindeer yields 90 to 110 pounds of meat on the average, the autumn-killed fawn 45 to 55 pounds. The animal slaughtering is usually at the end of summer. In their estimates of meat sources of the Arctic, Soviet statisticians do not overlook the meat of the wild reindeer (caribou). It is regarded as a potentially important item of food on the Taimyr Peninsula as well as in the territory between the rivers of Iana and Indigirka. Workers and employees of the governmental industrial project in Nordvik were b ^ g ^ etting about 6,000 caribou a year from the natives in 1940; it is said that an additional 3,000 or 4,000 could be easily used without damage to the local herds. Between the Iana and Indigirka, approximately 30,000 caribou were used in a year; in 1940 the officials estimated that delivery of about 1,000 tons of meat per year could be easily arranged in this district. Better methods of hunting and utilization of airplanes for scouting the caribou herds were suggested in order to increase the yield.
In composition reindeer meat does not differ much from beef or veal of the same grade, but it differs slightly in flavor. It is fine in texture and most of the meat is tender. The Russians consider that the most desirable cut is the round, which they use for steaks or roasting; the bottom round, which is less tender, can be used best for pot roast or in soup. Reindeer meat finds a good market far to the south; somewhat as Scandinavian reindeer meat does in Oslo, Stockholm, and Copenhagen. Its importance in the Asiatic Far North may be seen from the following table:

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

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Table V. Consumption of Reindeer Meat in Relation to Total Consumption of Meat in the Asiatic Far North (13).
Geographical Regions All meat, in tons Percentage of reindeer meat
Omsky Sever: 3,650 73.7
Ostiako-Vogul district 1,651 44.4
Iamalo-Nenet district 1,998 97.9
Yeniseisk Sever: 1,299 75.8
Turukhansky district 271 33.6
Igarka district 101 4.0
Evenki district 392 94.4
Taimyr district 535 97.2
Yakutsk North (16 districts) 1,528 52.2
Chukhotsk district 2,591 98.9
Total 9,068 77.6
As of 1940 the yearly slaughter of reindeer amounted to about 300,000 animals. In the Murmansk district the reindeer meat production as planned for 1937 amounted to 607 tons.
Stressing nutrition and calorie content, Soviet authorities consider reindeer meat one of the best meats. Soviet tests claim that it is several times richer in vitamins than beef or veal; it is rich in vitamin B 2 . The [: ] October 1950 issue of the Soviet magazine Priroda asserts that recent investi– gations showed 4.2 mg. of ascorbic acid as a minimum found in reindeer meat.

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Table VI. Chemical Content of Reindeer Meat (26).
Water Albumen Fat Carbohydrates Minerals Calories per kg.
Meat of a gelded rein– deer (average fatness 63.74 18.61 16.05 0.60 1.0 2,269
Meat of a fawn (average fatness) 70.76 21.31 5.95 074 1.24 1,452

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

The Tafal [: ] rsk reindeer found in the Irkutsk region is one of the best producers of meat; the 4 or 5-month-old fawns are 80 to 90 pounds dressed weight. The gelded reindeer of the Chukhotsk Peninsula gives about 130 pounds, sometimes up to 180 pounds of meat. At an outdoor temperature of 2 or 3°C. fresh reindeer meat can be kept safely in the open for 4 or 5 days. If frozen, it can be preserved for a year. Salted meat may be shipped and retained for one year at a temperature of ^ + ^ 3 to −1°C. Steaks are used either fresh, dried, salted, or smoked; salted meat is used most often. Natives usually dry the ribs and legs in the open air and then slightly smoke them; in this form the meat is easily carried on long journeys. The tongues are considered a great delicacy; they contain from 8 to 25 mg. ^ % ^ of ascorbic acid, according to the article in Priroda .
Until relatively recent times reindeer liver was not fully appreciated. For instance, a 1929 booklet of reindeer recipes, published by the Bureau of Home Economics, United States Dept. of Agriculture (30), refers to it rather modestly as a product which “when properly prepared compares favorably in tenderness and flavor with other kinds.” The conclusion of Soviet specialists, however, bases upon the research of the Murmansk technological expedition of the Institute of Reindeer Breeding in 1932, are ^ is ^ that a pat e ^ é ^ of such high quality may be made of the reindeer liver that it “undoubtedly could serve as an item for export.” According to Priroda , reindeer liver contains from 60 to 137 ^ % ^ mg. of ascorbic acid, while that of cattle contains 6 to 20 ^ % ^ mg. only. Proper methods for utilizing the kidneys, brain, lungs, heart, and head as food have been devised recently. The reindeer’s melted fat is widely used; in taste, it is similar to lamb fat.
Hides . Reindeer hide, after proper dressing, is very soft and pliable.

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

It provides the principal clothing material for the natives. From it they make garments, including boots, mittens, socks, leggings, trousers, sleeping bags, etc. It is used also for bedding, being warm and easily transported; an average skin weighs only from 4 to 6 pounds. Fawn hides are used as winter clothing with the fur on the outer side; such furs are very warm. Reindeer hides provide an excellent leather for the industrial manufacture of kid gloves, leather jackets, etc. the “chamois” prepared from the reindeer skin, especially that of the fawn, has a high value on the world’s market; it is used in the manufacture of precision instruments, optical instruments, and those used in aviation.
By-Products . As a result of years of technological research of the Institute of Reindeer Breeding, reindeer by-products are widely utilized in industry at present. The blood is salted and as a product rich in [: ] albumin is used for food and as an admixture in concentrated forages. Together with bones and viscera it is used in the preparation of dog food. It is also used in the manufacture of glue for plywood and veneer. As a matter of fact, the reindeer if properly slaughtered gives proportionally more blood than any other domestic animal (on the average, 6.76% in the case of females and 5.44% in the case of males). No industrial utilization was made of reindeer intestines until in 1932 when the Institute of Reindeer Breeding made the first tests in Leningrad. Then the expedition of 1935, sent by the Institute to organize the reindeer slaughterhouses in the Nenet region, proceeded with the experiments which were repeated in 1936. The studies proved that certain parts of reindeer intestines do not differ in their wall strength from those of cattle, and the manufacture of sausage casing from them can be carried out with the usual methods.

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

The strength of the intestinal walls was found to increase considerably if immediately after removal from the carcass they are washed with water and kept for 25 or 30 days in salt. However, a disadvantage of the casings manufactured from reindeer intestines is their small diameter which does not conform with the standard requirements of the market.
Reindeer hair is also highly valued because each hair is hollow, which makes it useful for manufacturing life preservers and upholstery. The investigation made by the Textile Research Institute in 1929 and con– tinued by the Institute of Reindeer Breeding since 1932 proved that the reindeer down (the soft woolly part of the hair) is equal in quality to the highest grade of merino wool. However, the presence of a large quantity of dead, coarse, brittle fibers in the hair made it impossible for use in textile manufacturing. Finding rational methods of separating down from brittle fibers was a task set for Soviet scientists; no information is available as to what progress has been made toward solving the problem.
The sinews of the reindeer are also utilized. They are separated from the meat and dried. From them the natives skillfully make thread which they use for sewing both shoes and other clothing.
Reindeer hoofs are used for the production of keratin glue, plastics, hoof meal, hoof coal, and ferrocyanide of potash. They cannot be used, however, in the manufacture of buttons and other notions because of the insignificant thickness of their walls (0.1 to 0.2 cm.).
Antlers are ^ a ^ nother important by-product of the ^ ^ industry. They are not only employed for decorative purposes, but also for making knife handles and parts of harness; they are also used in the manufacture of glue. As early as 1866, in the Archangel Region, a merchant names Volodin made the first
p. 42 of Tereshtenko Reindeer Breeding
Chese and milk [: neg]- 5th line from bottom ch. 4 [: butter ] from same?

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

successful attempt to manufacture glue from reindeer antlers for the market. The glue obtained from the antlers is of very high quality. As a result of the technological research of the Institute of Reindeer Breeding the methods of glue production were improved to the point, where, by its quality, it could easily compete with the standards established for exported glue. The weight of the product reaches from 10 to 20% of the weight of the antlers. The weight of the antlers collected per year between 1933-36 was estimated by various authors at 1,412 to 4,740 tons. As another by-product, photogelatin was obtained from the antlers; no information is available about its 2 / quality.
In some regions of the Far North, reindeer milk is considered a valuable product. From 0.2 to 0.3 liters of milk a day can be obtained. Karagask reindeer in the Saian ranges give as much as 1 liter of milk a day. In A ^ a ^ ppearance and taste it compares with cream. Its chemical content, according to Soviet data of 1948, is as follows: water, 67.7%; fat, 17.1% casein, 10.9%; milk sugar, 2.8%; minerals, 1.5%. Thus, it contains about 3 times more minerals and 4 or 5 times more fat than cow’s milk. From reindeer milk both butter and cheese are prepared. In chemical content the butter contains 15% water, 83.5% fat, 1.4% albumin and other organic substances, 0.1% minerals; thus it approximates butter prepared from cow’s milk. It melts at a temperature of 36 to 40°C. On the average, 16% of butter can be obtained from reindeer milk and 25 to 28% of cheese. The cheese reminds one of the sheep’s-milk cheese called brynza in Russian.
Soviet authorities assert that the reindeer industry in the U.S.S.R. is in its initial stage only. On the basis of recent geographic surveys of pastures the government estimates that the Soviet Far North may provide

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

fodder for about 10,000,000 reindeer, and that the task is not only to improve the breed of reindeer but also to at least triple the available herds. It is said that whether this goal will be reached will depend upon the general economic development of the Soviet Far North; but this, in turn, may depend considerably upon the progress of the reindeer industry.

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R.

BIBLIOGRAPHY

In Russian:

1. Akademia Nauk S.S.S.R. Trudy Poliarnoi Komissii (Works of the Committee for the Polar Regions), vol.14, Leningrad, 1934, pp. 54-69.

2. Arkticheskogo Instituta, Trudy . (Transactions of the Arctic Institute), 1935, Vol.22.

3. Bobrinskoi, N.A., ed. Opredelitel Mlekopitaivshchikhy S.S.S.R. (Identi–fication of the Mammals in the U.S.S.R.). Moscow, 1944.

4. Bol, V.K. and Nikolaevski, L.D. “O Roste i Sbrasyvanii Rogov u Severnogo Olenia.” (Regarding the Growth and Casting of the Reindeer Antlers) Sovetskoe Olenevodstvo , vol.8, 1936, pp.45-59.

5. Bunakov, E.B. “Economicheskoe Obosnoyanie Razvitia Olenevodstva Murmanskogo Kraia” (The Economic Foundation of the Reindeer breeding in the Murmansk District). Sovetskoe Olenevodstvo , vol.4, 1935, pp.107-162.

6. Debel, D.B. “Ispolzovanie Rogovogo Bashmaka Severnogo Olenia” (Utilization of the Reindeer Hoof). Sovetskoe Olenevodstvo , vol.8, 1936, pp.177-180.

7. Dushechkin, V.I. “Olenii Pastbishcha v Kharaulakhskykh Gorakh (Yakutia)” (Reindeer Ranges in the Kharaulakh Mountains (Yakutya)). Trudy Arkticheskogo Instituta, vol.63, pp.209-243.

8. Eikhfeld, I.G. and Chmora, N.Y. “Selsko-Khoziaistvennoe Osvoenie Krainego Severa” (Agricultural Reclamation of the Far North). Materialy Soveshchania po nauchvo-izsledovatelskoi rabote Na Krainem Severe, 27/II-3/III, 1936. Moscow, Vsesoiuznaia Akademiya Selsko-Khoziaistvennykh Nauk, Moskva, 1937.

9. Geptner, V.G. and Tsalkin, V.I. Oleni S.S.R. Sistematika i zoogeografia (The Reindeer of the U.S.S.R. Classification and Zoogeography). Moscow, Moskovskoe Obshchestvo Ispytatelei Prirody, 1947.

10. Gorodkov, B.N. “Itogi Izuchenia Prirosta Lishainikov” (Results of the Study of the Growth of lichens). Sovetskoe Olenovodstvo , vol.8, 1936.

11. ----. ed. “Sistemy Vypasa Olenei i Pastbishchnoe Khoziaistvo Murmanskogo Okruga” (Reindeer Pastures and Vegetation Cover of the Murmansk District). Trudy Arkticheskogo Institut, vol.72, Leningrad, 1936.

12. Govorukhin, V.S. “Sezonnye Pastbishcha Severnogo Olenia” (Seasonal Pastures in the Far North). Kalendar Prirody S.S.S.R., vol.2, Moscow, 1949, pp.313-338.

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R. - Bibliography

13. Khrapal, A.A. Selskoe Khoziastvo Aziatskogo Severa . Moscow, Glavsevmorput, 1940.

14. Liverovski, Y.A. and Kolesnikov, B.P. Priroda Yuzhnoi Poloviny Sovetskogo Dalnego Vostoka. (The Nature of the Southern Part of the Soviet Far East). Moscow, Akademia Nauk S.S.S.R. Institute Geografii, 1949.

15. Lukashevski, V.A. and Kharin, P.S. “Svoistva Oleniei Shersti-Lenki” (The Properties of the Reindeer Hair). Sovetskoe Olenevodstvo , vol.4, 1935, pp.163-176.

16. K Mirovich, A.F. “K Voprosu ob Ispolzovanii Kishek Severnogo Olenia dlia Kolbasnogo Proizvodstva” (On the Utilization of the Intestines of Reindeer in the Manufacture of Sausage Casing). Sovetskoi Olenevodstvo, vol.8, 1936.

17. Mutovin, M.E. and Purin, A.A. “Bogatstva Kamchatskoi Oblasti i ikh Expluatatsia” (Natural Resources of the Kamchatka Region and Their Exploitation). Kamchatka Petropavlovska na Kamchatke . Shankhai, 1940, pp.155-195. “Slovo” publisher.

18. “Novoye v Ispolzovanii Severnogo Olenia i Losya dlia Transporta” (News Regarding the Utilization of the Reindeer and the Elk for Transportation). Trudy Nauchno-Izsledovatelskogo Instituta Poliarnogo Zemledelia, Zhivtnovodstva i Promyslovogo Khoziaistva. Trudy , Seriia Olenevodstva , vol.6, 1939.

19. Pinegin, N.V. Novaya Zemlya , Sevkraiz, 1935.

20. Rodionov, K.V. “Materialy po Izuchaniu Technologicheskikh Svoistv Severnogo Olenia.” (Materials Regarding the Study of the Technological Properties of the Reindeer Antlers). Sovetskoe Olenevodstvo , vol.6, 1936, pp.51-62.

21 Rodionov, K.V. “Voprosy Tekhnologii Produktov Olenevodstva” (Questions of the Technology of the Reindeer Products). Sovetskoe Olenevodstvo, vol.4, 1935.

22. Russian Asia , vol.1, 2. St. Petersburg, Resettlement Administration, 1914.

23. Sdobnikov, V. and Romanov, A. “O Myasnykh Respursakh v Arktike” (Concerning the Meat Resources in the Arctic). Sovetskaia Arctica . no.9, 1940, pp.23-36.

24. Sochava, V.G., ed. “Olenevodstvo. Olenii Pastbishcha i Priemy Vypasa Olenei v Anadyrskom Krae” (The Reindeer Ranges and Methods of Pasturing of the Reindeer in the Anadyr Region). Transactions of the Arctic Institute of the U.S.S.R., vol.62, Leningrad, 1936.

EA-Zoo. Tereshtenko: Reindeer Breeding in the U.S.S.R. - Bibliography

25. Varnek, P.A. “Russkii Sever v Pervuiu Mirovuiu Voinu” (The Russian North in the First World War). Morskie Zapiski , Pub. by Obshchestvo Byvshikh Ofitserov v Amerike. Vol.7, no.1, March 1949, pp.23-25.

26. Zhigunov, P.S. and Terentiev, F.A. Severnoe Olenevodstvo (Reindeer Breeding in the North). Ministerstvo Selskogo Khozuaistva R.S.F.S.R. Upravlenie Raionov Krainego Severa. Moscow, 1948.

In English:

26a. Bosworth, Joseph. A Description of Europe and the Voyage of Ohthere and Wulfstan , written in Anglo-Saxon by King Alfred the Great. London, 1855.

27. Laufer, Berthold. “The Reindeer and its Domestication.” Memoirs of the American Anthropological Association, vol.4, 1917, pp.91-147.

28. Palmer, L.J. “Raising Reindeer in Alaska.” U.S. Dept. of Agric., Misc . pub . no.207, 1934.

29. Report of the Royal Commission Appointed by Order in Council of Date May 20 , 1919, to Investigate the Possibilities of the Reindeer and Musk-Ox Industries in the Arctic and Sub-Arctic Regions of Canada . Ottawa, 1922.

30. Stanley, Louise. “Reindeer Recipes.” Bureau of Home Economics, U.S. Dept. of Agric., Leaflet no.48, 1929.

V. J. Tereshtenko

Biology of Ovibos

EA-Zoology (A. L. Rand and Vilhjalmur Stefansson)

BIOLOGY OF OVIBOS

CONTENTS

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Page
Description 2
Range 4
Habits 5
Utilization 8
Bibliography 10

EA-Zoology (A. L. Rand and Vilhjalmur Stefansson)

BIOLOGY OF OVIBOS
The bison, through v ^ f ^ alse analogy miscalled the buffalo, and the ovibos, through misunderstanding or perhaps through cupidity miscalled the musk ox, are NOrth America’s great representatives of the family Bovidae, the one occupying the prairies of the midcontinent, the other those of the Far North.
The ovibos ranks with caribou, seal, and whale as one of the four animals most important to man in relation to the Far North. Its one land rival, the caribou, may seem to lead in the competition, because some caribou have been domesticated as reindeer; but friends of the ovibos reply that its Latin-derived name, from ovis for sheep and bos for cattle, truly implies that this animal possesses, as a candidate for domestication, the combined merits of the sheep and the cow, and is thus potentially the greatest of assets for northward colonization, being already resident in even the most northerly arctic lands.
Ovibos , appearing in northern literature under many names, among them northern ox or northern cattle, arctic ox or arctic ca f ^ t ^ tle, musk ox or musk cattle, polar ox or polar cattle, is, as said, a member of the family Bovidae, to which belongs our domestic cattle, as well as sheep and goats. Much has been written of these immediate relatives, and at one time the ovibos was

EA-Zoo. Rand and Stefansson: Biology of Ovibos

considered a bison specialized for arctic conditions. More recent studies show that the ovibos of northern America and the takin of the mountains of southern Asia are the sole living representatives of a once more widespread and diversified group, only some of which were adapted for arctic conditions. This group is most closely related, on the one hand, to the goats and sheep, and, on the other, to the goat antelopes, such as the Rocky Mountain goat and the chamois.
The original home of the ovibos group seems to have been in Eurasia, where five fossil genera are known from the Pliocene. By the next epoch, the Pleistocene, the five genera had disappeared, another fossil genus had evolved in Eurasia, and the group had invaded North America, where three fossil genera appeared. Also in the Pleistocene, the two modern genera arose, Ovibos (musk ox, polar ox) in Eurasia and America, and Budorcas (takin) in Asia. With the close of the Pleistocene only these two modern genera were left. Budorcas is still restricted to Asia. Ovibos lingered on in Eurasia until the Recent period; in Europe it existed in prehistoric times along ^ with ^ the last Neanderthal and Cro-Magnon men, the mammoth, and the woolly rhinoceros; in Siberia it may have survived longer. By historic times Ovibos existed only on the northern plains and arctic islands of America.
Description
Ovibos moschatus is a rather stout, compact ungulate with short legs and a very short tail. The pelage is long and shaggy, contributing materially to a bulky appearance. The underfur is a fine, soft, woolly layer, 3 to 6 inches thick; the abundant guard hairs are very long and coarse and may reach a length of 24 inches or more. In adult males the horns sweep downward,

EA-Zoo. Rand and Stefansson: Biology of Ovibos

outward, and upward in sharp hooks, the base of the horns enlarging with age until they cover the forehead; the horns of the female are less expanded at the base. The color of the pelage is generally deep brown to black, with light-colored saddle and legs; varying geographically, there is some white in the forehead and about the head, more pronounced in the females and young; the color of the horns is dark brown to creamy white.
The adult male totals 96 inches in length; tail 4 inches; hind foot, 19 inches; height at shoulder, 59 inches; and horns on outside curve up to 29 inches in length. The males weigh up to 900 pounds; the females are somewhat smaller. The northern animals are somewhat smaller than the more southern.
The calf is covered with short, dark-brown, curly hair. A new-born calf is recorded as weighing 16 pounds. The first trace of a horn core appears at 6 months. Adult size and well-grown horns are reached at about 6 years of age.
Three subspecies are recognized, owing to differences in size and color.
The Ovibos moschatus moschatus , the “barren-ground musk ox,” is a large brownish subspecies with dark-colored horns and no white in the forehead, occurring on the Canadian arctic mainland.
The Ovibos moschatus niphoecus , the “Hudson Bay musk ox,” is a black subspecies, smaller than the subspecies O. moschatus moschatus . Their horns are light-colored; the females are white about the head. They occur to the northwest of Hudson Bay.
The Ovibos moschatus wardi , the “white-faced musk ox,” is a grayer race. Their horns are of a creamy-white color. Both o the male and the female have whiter faces, and they are longer and more slender than the

EA-Zoo. Rand and Stefansson: Biology of Ovibos

subspecies O. moschatus niphoecus . They occur in Greenland and the Canadian Arctic Islands.
Range
The present range is mainland Arctic Canada between Hudson Bay and Horton River, the Canadian Arctic Islands to the north (but not Banks, Baffin, or Southampton islands), to northern Ellesmere Island; also in northern and northeastern Greenland.
As said, ovibos disappeared from Eurasia before historic times , ^ ; ^ in America a much wider range than at present occupied is indicated by fossil remains of Ovibos from the central United States (perhaps correlated with the ice age) and from the Yukon Valley. Within historic times the ovibos was known from the arctic coast of Alaska, where it probably became extinct about 1860 or 1870. The southern limits of the historic range in Canada originally coincided with the northern edge of the forest west of Hudson Bay. Minor withdrawals have occurred and the animals no longer range west of the longitude of western Great Bear Lake. The records from northern Manitoba are all old. Parts of this area from Hudson Bay to Horton River are still occupied by ovibos. In Banks Island, the last were killed by Victoria Island Eskimos around 1912.
In the early part of the twentieth century it seemed as if the ovibos might be exterminated; but extinction no longer seems to be threatened. A population ^ estimate ^ made in 1930 gave the following figures: Canadian mainland, 500 animals; Canadian Arctic Islands, 12,420; North and East Greenland, 1,500; total, 14,420 animals. However, the Greenland estimate has been questioned as too small, and estimates as high as 10,000 animals have been made for the northern east coast alone for about the same period. Though ovibos ha s ^ d ^ been reduced perhaps 1% of their primitive number by 1930, the animals on the

EA-Zoo. Rand and Stefansson: Biology of Ovibos

Canadian mainland had doubled in number by 1940, and the reports from farther north indicate s that protective laws have been successful in conserving the species.
Habits
Though ranging the northern forest in early historic time, as the wood bison still does, ovibos is like bison in being essentially a prairie animal, frequenting the marshes, plains, and lower, wetter country in the summer. In the southern part of its present range where there are willow thickets, as in the Thelon Game Sanctuary, these are also favored. In winter they tend to frequent higher, drier and more exposed ground, presumably where the wind will sweep away the snow. The favored foods are sedges and grasses; willow twigs and herbaceous plants are also eaten. There is not the pronounced change from summer food of grass to winter food of lichens as with the caribou , ^ ; ^ some lichens (reindeer moss) are eaten, probably mostly in winter. When the food is covered with snow, the ovibos digs through with its forepaws, and pushes aside the snow with its no c ^ s ^ e and head. With hoofs and nose the animal may work through four feet of snow, but it does not seem to resemble the caribou in actually preferring to dig for food under snowbanks.
Although there may be slight seasonal changes in the habitat occupied, the ovibos is, in general, an inactive animal. If a herd is seen in one area one year, the chances are that it will be there the next. Individuals appear to move about but little. Old bulls have been recorded as not moving a half mile all summer; a band stayed in sight of one camp for five days.
Ovibos are placid animals, usually found in small herds or medium-sized bands, apparently spending their time grazing and resting during the day. During

EA-Zoo. Rand and Stefansson: Biology of Ovibos

the hot, bright arctic summer they may prefer to feed during the night. The belief, expressed by some writers, that ovibos take shelter in valleys, or behind boulders, from the blizzards of winter, is doubted by some observers.
Ovibos bands seem typically to consist of about 10 to 15 animals, although smaller groups of 5 or 6, as well as lone bulls, are often reported. Where the animals are common, herds of as many as 200 have been reported. The social organization of these herds is imperfectly known. There seems to be some seasonal change, the herds tending to be larger in winter. In spring herds of bulls only may be found, but x during much of the year more than one bull may be found in one herd. The proportion of calves in the herds is always low, compared with many other hoofed animals. This is correlated by some with a deduced slow rate of reproduction.
The late summer or early fall is the time of breeding. The ovibos is polygamous and old males fight fiercely, bellowing challenge d ^ s ^ , pawing the ground, and rushing at each other and meeting head on with a tremendous crash. They also rip each other with their sharp, hooked horns, sometimes breaking their horns in th o ^ e ^ se fights. Apparently bulls sometimes kill each other, for skulls spilt open, presumably by the force of fighting heads meeting, have been found. Not all fighting is connected with the rut, as bulls sometimes fight and then later consort amicably in the herd.
According to present information, reproduction is slow, presumably adapted to the relative immunity of the species to wolf predation. It is believed that the females do not breed until 3 or 4 years old, and that ordinarily they bear young only every other year. The usual number of young at a birth is one, born in April, May, or perhaps June. Victoria Island Eskimos told Stefansson that calves born early in the season, in April when temperatures

EA-Zoo. Rand and Stefansson: Biology of Ovibos

may drop to −50°F., often f e ^ r ^ eeze to death while still wet, immediately after birth. The variation in the size of calves [: ] ^ seen ^ with the herds indicates a long breeding and calving season. The cows may retire from the herd at the calving season; some herds for a time appear to consist solely of cows and calves. The calves are active from birth, or shortly thereafter, and follow the mothers. They suckle to some extent until August at least.
The defense behavior of ovibos is very effective against wolves. The band prefers to take a stand and fight on high ground. The animals range themselves in close formation in more or less of a circle, heads out; they may rub their noses and horns on their forelags, and paw the ground, perhaps to intimidate attackers. If calves are in the herd, they may be in the d ^ c ^ enter of the formation or they too may face out. The bulls do most of the fighting, but all adults and even yearlings may take part. As the wolf or dog approaches, a bull rushes out 10 to 15 yards, trying to catch the enemy on its horns, then wheels about and returns, backing into place in line. The number of animals that dash out depends on the number of attackers; sometimes all the bulls may be out of line at one time. Dogs, when caught by the horns in this way, have been severely injured, partly through being tossed high in the air. A lone ovibos at the approach of danger may run to a cliff or a rock against which it puts its back, ready to fight.
This defense of the ovibos, while adequate against wolves and dogs, is the weakness of the animal when it encounters man. At the approach of hunters, especially if they have dogs, the ovibos do not flee but adopt their d ^ c ^ ustomary formation, in which the dogs can hold them indefinitely. Hunters can then kill them at leisure with rifles, or tease them with arrows until they charge, and then kill them with spears.

EA-Zoo. Rand and Stefansson: Biology of Ovibos

Ovibos have been introduced into Norway and Sp t ^ i ^ tsbergen, apparently with some success. In Alaska, the United States introduced 34 animals in 1930, with the object of restocking suitable areas; this work is still in the experimental stage.
Utilization
The hide of ovibos is too thick and hard for general use as clothing, although it has sometimes been used for boot soles. The woolly character of the fur, which makes it hard to clean, is another drawback to its domestic use. Although lacking the qualities that appeal to ordinary fur markets, there did develop for a while in the nineteenth century such a demand for ovibos skins that Indians of the northern Canadian forest made numerous excursions to “no man’s land” beyond the tree line to secure skins for sale to the Hudson’s Bay Company; some Eskimos also came south into this region (usually avoided because of mutual Eskimo-Indian dread of each other and therefore previously a sanctuary to ovibos), and a few of the resulting skins got into the hands of traders.
Most of the skins sold in London were sent to the United States and Canada where they were prized for sleigh robes. This trade flourished from about 1870 to 1900, and was important to certain groups of Indians who depended on these hides for credit with the posts. The Company’s average annual sale for the 20-year period preceding 1902 was 200 to 250 pelts, although, in 1891, 1,358 pelts were sold in London.
The leisurely habits of the animals and the ease with which they are killed have made them dependable sources of food, but these very features make it impossible for them to survive, while left to their own resources.

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where many men live. So long as they are in a wild state, it would seem we must be content to have them survive in the remote, unsettled arctic islands, and in sanctuaries on the mainland where they seem to be prospering at the present time after years in which their existence was in jeopardy.

EA-Zoo. Rand and Stefansson: Biology of Ovibos

BIBLIOGRAPHY

1. Allen, J. A. “Ontogenic and other variations in muskoxen, with a systematic review of the muskox group, recent and extinct,” Amer.Mus.Nat.Hist., Mem . n.s. vol.1, pt.4, 1913.

2. Clark, C.H.D. “A Biological Investigation of the Thelon Game Sanctuary,” Nat.Mis.Can., Bull . no.96, 1940.

3. Hone, E. “The Present Status of the Muskox in Arctic North America and Greenland with Notes on Distribution, Extirpation, Transplanta– tion, Protection, Habits and Life History,” Amer.Comm.Internat. Wild Life Prot., Special Pub l. no.5, 1934.

4. Stefansson, V. “The Domestication of Ovibos,” in The Northward Course of Empire , Harcourt, Brace and Co., New York, 1922.

5. Sverdrup, Otto. New Land. Four Years in the Arctic Regions , Longmans, Green. London, 1904.

A. L. Rand and Vilhjalmur Stefansson

Biology of the Polar Bear

EA-Zoology (A. L. Rand)

BIOLOGY OF THE POLAR BEAR

CONTENTS

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Page
Range 2
Habitat 3
Habits 4
As Object of the Hunt 10
Edibility of the Meat 12
Bibliography 15

EA-Zoology (A. L. Rand)

BIOLOGY OF THE POLAR BEAR
The polar bear, Thalarctos maritimus , of the family Ursidae, is a large bear which inhabits the arctic regions of both the Old and New Worlds, occurring in every part of the Arctic Sea and its margins, occasionally found on adjoin– ing land, but rarely far from sea ice. The scientific name doubly states this fact, being a combination of the Greek words, thalassa , sea, and arktos , bear, to which has been added the Latin word, maritimus , of the sea. It is perhaps the second largest of the bears, next after the Kadiak, the male averaging 8 feet in length, 900 pounds in weight; the female averaging a little more than 6 feet in length, 700 pounds in weight. However, males often attain more than 1,000 pounds, and a record of 1,600 pounds has been claimed. The fur is yellowish white, dense, and hair practically covers the soles of the feet, leaving naked only the black lips and nose tip. Apparently there is one molt a year, during the summer months. Old males develop a sort of ma in ^ ne ^ and noticeably longer fur on shoulders and forelegs (14). The polar bear is distinguished from other bears by its pelage, by its semiaquatic nature, and by structural differences, having a longer neck, narrower skull, and relatively small molar teeth. Subspecies have been described, but are generally thought not sufficiently well characterized to justify recognition ( 2 ^ 1 ^ ).

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Partly owing to the hairy soles of its feet, the polar bear walks sure– footedly over the sea ice (which incidentally is not as slippery as lake ice) and can, when hurried, attain a surprising speed, in spite of its lumbering gallop. It is a powerful swimmer, said nearly to equal the speed of an eight– oared European whaleboat, although readily overtaken by the exceedingly swift Eskimo kayak.
Polar bears are solitary, except at mating time. When two are roving the same locality, they tend to avoid each other. The young sometimes follow the parent female well into the second year. Exceptionally, as many as 50 bears have been reported on a few square miles of land, gathered to the neighborhood by a stranded whale carcass. Also where they are extremely abundant locally (as on St. Matthew Island, where several hundred were sighted within one month on the 22-mile-long island), the younger males may consort in small groups of three or four (6).
Range
The species has been recorded as ranging on land north to Spitsbergen and Novaya Zemlya (8), to beyond 82° N. latitude on the north coast of Ellesmere Island, and to 83° N. in the Franz Josef Archipelago (6). Records at sea show an even more northerly ordinary range, or they may indicate that occasionally individual bears may wander almost anywhere in the polar area. Among the high– latitude records are the tracks of a bear seen by Peary, March 24, 1909, near 86°30′ N. (11), and a bear with cubs seen by Papanin near 88° N. on August 1, 1937 (10). Both of these were in the Greenland-Ellesmere sector.
The southern limits of the Pacific range are, apparently, near the middle of Bering Sea; bears are common around the St. Lawrence and St. Matthew Islands.

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On the Asiatic coast of Bering Sea they have been observed on drifting ice floes in the vicinity of Plover Bay; on the American coast they have rarely been observed as far south as the mouth of the Yukon River (6). In Atlantic waters the species has been observed on floating ice to the north of Norway, where they reach land once or twice in a century, and on the ice off Iceland, where they come ashore once every ten or twenty years. Occasionally they are reported as far south as Newfoundland, the north shore of the Gulf of St. Lawrence, and in James Bay, the southerly arm of Hudson Bay, at about 55° N. latitude.
Habitat
Sea ice, especially when moving, provides the environment most favorable to the polar bear’s existence, and has earned for him, in some localities, the name “ice bear.” In the pack, he seeks narrow open leads, which provide him most readily with his favorite food, the seal. In winter, the bear wanders over the frozen sea, seeking leads or the edge of the ice and open water. When compelled by ice conditions to do so, he can swim long distances. A polar bear has been reported swimming 20 miles or more from land, out of sight of any sea ice. On the other hand, during the short summer, he occasionally wanders inland, where he may secure some food from vegetation of the grasslands fringing the Arctic Sea. Polar bears have been observed at a distance of 100 miles inland, but such an occurrence must be rare.
Because of the movements of the ice, the polar bear, unlike most mammals, lacks a definite home range. When the sea freezes in autumn the range grows more southerly, for it is coextensive with the ice; in spring it shrinks with the shrinking ice.
The presence or absence of polar bears during a season in any region may depend on whether or not the heavy pack comes in that year. Occasionally, bears

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have been found stranded on islands and mainland shores, having been caught there by mischance when the ice moved away. Then, too, the bear may find himself “stranded” in the ocean, when the ice melts unexpectedly beneath him, as on contact with the Gulf Stream. What may be farthest-form-land record is that of a bear seen by the Cherevichnyi expedition on April 16, 1941, at 78°27′ N., about 500 miles from Wrangel Island, the nearest known land (3).
The polar bear distributes itself throughout its range in what is generally an irregular fashion and dependent upon the displacement and accidents of the ice. Further study of the species, however, may disclose a tendency toward seasonal migrations. In the Bering Sea, Nelson observed that, as the ice closes in the north of Bering Strait in October and November, large numbers of bears are brought down on the drifting pack, pass through the strait, and reach St. Lawrence and St. Matthew Islands, where they are said to winter (6). When spring comes, according to this observation, the bears, following along the border of the pack ice, pass north through the strait and into the Arctic Sea. Such a movement, toward land in the autumn, toward the sea in spring, may be considered as at least incipient migration. Apparently some bears come to land to hibernate, but the polar bears’ travels seem controlled, as said, by the seasonal expansion and contraction of the floating ice, beneath which lives the game he subsists by, the seal that he can catch by plunging from above though he cannot overtake it by swimming in open water.
Habits
To what extent the polar bear hibernates is still a moot question. Their absence from any habitual location, as has been said, does not necessarily imply migration, nor does it imply hibernation. It may merely indicate that the moving ice floes have not brought the bear to that locality. Apparently it is usual for

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gravid females to hibernate; some claim that last year’s half-grown cubs, while still unable to fend for themselves, hibernate with the mother. Probably mature males remain active throughout the winter. In northern Alaska and north ea ^ we ^ stern Canada, according to Stefansson, the Eskimo belief was that only gravid females hibernate (15; 16).
It can safely be assumed that the habits of the bear are similar throughout its circumpolar range; but until closely coordinated studies have been made, it is impossible to make a flat statement to this effect. They have been reported as active throughout the winter in northern Greenland where, it is said, the pregnant females retire merely to give birth to their young and where, during spells of severe weather, females with young may take shelter in snow caves. For hibernation, they have been known to travel some miles inland and are said to excavate burrows in the snow, in which they sleep from December to March.
The spring is the mating season. In the extreme North, this period is during the months of April and May; farther south, it may occur later on. During that season the males are embattled. When couples have formed they are believed to remain in pairs for perhaps two weeks. After this, they drift apart, reverting to their normal way of life, which is solitary. The male reaches puberty at the age of three; the female at about the age of two. Some claim that the female mates but once every two or three years, but this makes it hard to explain why they are so often reported as accompanied by cubs of two litters which are usually taken to be this year’s and last year’s.
The young, one or two in number, are born in midwinter, the gestation period being 10 or 11 months (9). The newborn cubs are incredibly small in size, not larger than a well-grown rat, the weigh only about 650 grams. They develop quickly, however, and at about the age of three months are able to follow the

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mother bear. It is reported that about one month after the young have opened their eyes, the mother starts migratory life with them. If this be so, then it can be supposed that at this stage the mother bear would merely transport her young in her mouth to another locality. They stay with their mothers until they are able to fend for themselves, usually for about one year. Up until the age of five months the young are still so helpless that they remain with the mother even when she is slain by hunters. At a later age, faced with such danger, they will flee. The solicitous female, pursued by hunters, has been observed to urge forward her young, with pushes, nudges, and blows (14). It has been recorded that, in the water, the young sometimes seize the female’s short tail in their teeth and are thus borne along by the swimming parent. When the young are unable to clamber out on the ice after their mother, she may reach down and lift her cub out, seizing him by the nape of his neck in her teeth. Not only must the female look after her cub, teaching him to swim and to hunt, but she must also protect him from roving males, who, it is reported, sometimes devour the young (14).
The young are playful and mischievous, which sometimes impedes the mother when stalking a meal. Even adult bears have been observed to display mischievous tendencies along with other characteristics of the upper mammalia. Calmness, coolness, and sagacity are supposed to be the attributes of the polar bear. But when injured or thwarted, he displays what could be called fits of rage.
Stefansson was able to confirm the Eskimo report that cub bears frequently and grown bears occasionally will climb a snow slope repeatedly, to slide down it on their haunches. When objects are thrown at bears they may bat them with their paws and, if they roll well, may pursue them a while, batting them two or a few times along the ice or land, reminding of a cat playing with a ball. In

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one reported case, the object thrown, intended to scare the bear away so that shooting him would not be necessary, was an empty five-gallon kerosene tin. The bear acted as if he were interested in the noise this can made when he batted it back, seemingly enjoying sound as well as sight of ^ as ^ it bounced along in front of him.
As evidence of its sagacity, the polar bear has been credited with using a boulder as a weapon is attacking a walrus, with using a piece of ice or snow as a moving shield when stalking a seal on the ice, and with holding up its paw to hide its black nose from its intended prey (14). Such reports may be merely a part of the body of myths and legends that have grown up about the bear. But there are many reports that may be accepted of his persistence and ingenuity in breaking into caches that would be safe against any other animal.
In his quest for food, the bear depends least upon his sense of hearing, more upon the senses of smell and sight. He scents his accustomed food at long distances. Seals (chiefly the ringed seal, Phoca hispida ) provide his staple; but when there is need he feeds on a variety of animal life and eve berries and roots and other vegetation. There are reports that walrus are sometimes slain and eaten and that the bears may get fatally wounded in such encounters. Nordenskiold tells of having seen, in Spitsbergen on two occasions, the blood and hair of reindeer which he inferred had been killed by bears (7).
Opinions and reports on the normal feeding habits of the bear are many and often conflicting, as is the case with any wild animal not as yet thoroughly studied in its natural habitat. For example, the polar bear is reported to feed upon vegetation when it comes out from hibernation (6; 14). In what month the period of hibernation ends i d ^ s ^ doubtful; but since the young are born in midwinter and are able to follow the mother at about the age of three months, it is

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questionable if there would be much vegetation available at this time. Besides, some of the cubs appear to be born on the sea ice far from land, where there would be no obtainable vegetation.
According to Stefansson, more than 99% of the polar bear’s food is seal (16). Usually, the only thing he recognizes as food by the sense of smell is something that smells like seal. When the meat is old, it develops the odor of decay that is common to all meat. Apparently this odor overcomes the natural fresh odor in all meats. Thus, the polar bear has been reported to leave untouched a cache of fresh caribou, passing to leeward without even swerving from his course of sniff at the meat more closely. This would indicate that he does not recognize caribou, in its fresh state, as food; but he is known to recognize it if even slightly high.
However, there are in the books many cases where individual polar bears have learned to try many different kinds of food, especially when they have had a chance to forage in scrap heaps thrown out on the sea ice by ships wintering in the Arctic. Such food may have numerous and pungent odors; and bears which have had experience of these varied tastes and smells will go after whatever has any smell, devouring it as avidly as if it were seal meat. The sophisticated polar bear, who has once learned that seal is not the only food, will enter camps of travelers, even when out in the pack ice, break open caches, eating greasy cloth, rope, paper. Men have been killed and eaten by polar bears; and the carcasses of their own kind have been consumed.
Bears marooned on an island or on a mainland shore, by a gale that takes all the ice away, will get hungry. If the season is the least bit warm, a good deal of decay takes place along the sea beaches; and a bear ^ of ^ keen appetite, not able to reach his accustomed sealing grounds among the pack ice, will give the impression of preference for a varied diet; for he eats mussels, starfish, and

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various crustaceans, the eggs of sea fowl, or anything edible washed up by the sea - even seaweed.
The polar bear hunts the seal in various ways. Sometimes he swims stealthily along until a seal is encountered basking on the edge of the ice, when he will rise suddenly between it and the water, cutting it off from escape, and crush its skull with a blow. The bear also stalks basking seals on the ice, creeping up slowly, silently, unobserved until it is too late. He may wait patiently at the edge of a lead, to plunge from above upon a seal swimming by. He also secures the very young of the ringed seal in their natal snow caves, hidden from above by a roof of snow and visited by the female through the b f ^ r ^ eathing hole contrived in the ice below. These caves the polar bear searches out by scent. Apparently he prefers adult to young seals, for on occasion he may kill several baby seals and leave their carcasses uneaten. He also prefers the skin and fat of seals to their flesh, and may gorge on the blubber and skin alone. (It has been said that the bear stores food under blocks of ice, but the report should be taken with caution.)
From combining what Eskimos told him with evidence deciphered from blood, footprints, and sleeping lairs, Stefansson worked out what may be a typical feeding procedure. If the seal killed is small, the whole of it is consumed, with bones and entrails. If a little larger, the bear eats all the skin and blubber, with some of the lean. From a very large seal he will eat perhaps a third or half the blubber, twenty to thirty pounds, and most of the skin but none of the lean. Satiated, he then walks off fifty or a hundred yards and goes to sleep. When he awakens he goes off without investigating his kill, usually traveling upwind.
Seemingly the bear, on waking up, feels as if he would never be hungry again,

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at any rate scorning the cold remnants of his warm meal. But there may not be anything much left to scorn; for usually a bear is trailed by several foxes that depend on him for food. They hop around him in excited circles while he is eating, now and then coming so close they have to dodge when he makes passes at them. When he has had his fill and goes off to sleep, the foxes come in for their share. If it is summertime, the foxes have the same trouble keeping the gulls away that the bear had keeping the foxes away.
Foxes often have difficulty following a bear, for if he comes to an open lead he swims across and they may be unable to find a bridge anywhere near. It was under such conditions, and in winter when there were not gulls, that Stefansson occasionally found a dead seal from which nothing had been eaten except skin and blubber. On other occasions he found seals at which foxes had been gnawing, showing they were not present at the kill and arrived only when the remnants of the bear’s feast had been hardened by the frost. At other times he found remnants of frozen seals that had been devoured by wandering hungry bears that came along hours or days late.
As Object of the Hunt
Man is bear’s only enemy. But being of little economic value as a source of hides, furs, and food, and bear has never been hunted systematically on a large scale. They are slain by travelers chiefly as c ^ s ^ ource of food.
The skin is utilized for clothing to some extent, but though strong, it is heavy and not commensurately warm. Most of the skins exported are for rugs, or merely as trophies. The lighter hides are used for semiwaterproof trousers by some Eskimos; the heavier hides as bedding. Light skins may be used by Eskimos for mittens, and skins of any weight are cut up into strips for dog harness. The

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long hairs are esteemed for ornamental purposes by the women in some Eskimo districts.
Particularly in the eastern Canadian Arctic and around Greenland, Eskimos hunt the polar bear with dogs (11; 14). When the bear is sighted, the dogs are let loose. Then, when they have brought him to bay, he is shot. In former times, the Eskimos of Greenland and the Chukchis of Siberia killed bears with spears, sometimes made by lashing hunting knives to walking sticks. Occasionally a hibernating female is located in its snow cave with the aid of dogs, when she is speared or shot.
Much of the information we have on polar bears in the wild state derives from the reports of travelers and hunters. For instance, as to the age which these animals naturally reach, we have only the reports of men who have killed bears showing by their pelage and their worn teeth that they must be old. In captivity the polar bear has attained the age of 33 years.
From reports of travelers, also, we get the many conflicting statements regarding the ferocity of the bear. According to some who have encountered them, these animals without natural enemies are, nevertheless, somewhat shy and timid. They are also curious, and will approach a man fearlessly. They will come boldly into occupied camps, knocking the dogs aside. From the accounts of Barents and other polar explorers of the sixteenth the seventeenth centuries, the bear emerges as a most ferocious and dangerous beast. But M’Clintock’s Greenlandic guide, Petersen, is quoted by Nordenskiold as thinking the bear as little dangerous as a sheep.
Sometimes a polar bear will stalk a man who is crawling on the ice, hunting seals. Stefansson believes this is because the bear mistakes for a seal the man’s recumbent figure. But the polar bear has been known to stalk an upright man, and

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Stefansson has reported a narrow escape he had on one occasion (16). The books contain stories of men who saved themselves from belligerent individuals by lying down on the ice, when the bear would merely sniff curiously at them, doing no harm. And whereas, according to most reports, the polar bear flees when shot at, in some accounts, bears, when fired upon or disturbed at a meal, turned and attacked their hunters. Perhaps the bear who approaches a men in harmless curiosity is merely a bear without previous experience of his enemy. In the same way, the bear that flees, or the bear that assaults, may be one with experience which has taught him to recognize his enemy.
The curiosity and ingenuity of the bear are proverbial, as is his strength. Certainly, it is difficult to make a cache which is secure against him. Bears have been known to break into the scatter about caches of every kind, even when they contained no food, although it may be surmised that the object of the search was food. Stefansson has described the making of an ice cache which is more or less polar-bear-proof (15; 16). It could be constructed by digging a pit into an ice hummock, placing in it the things to be stored, filling the rest of the pit with boulders of ice, tamping in snow between the chunks and pouring water on the whole that has been secured from beneath the ice.
^ Edibility of the Meat ^
Various opinions are held as to palatability of the meat. Some white men have found it good eating, preferring it to caribou; others have found it rank, unpleasant, and even unwholesome. A few say they prefer [: ] to eat the flesh raw or frozen; most prefer it cooked. Freuchen describes the flesh as pink, sweet, and savory, but says it becomes unpalatable when too exclusively eaten . ^ (2). ^ But Nansen and Johansen, who probably had most experience of all explorers in living on polar bear meat, found they could eat it exclusively, day after day for several months, without tiring of it. “We consumed large quantities,”

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writes Nansen, “at every meal, and strange to say, we never grew tired of this food, but always ate it with a ravenous appetite” (5). They usually ate it “boiled in soups,” he tells us. The paws, heart, tongue and brains are con– sidered delicacies.
Many arctic travelers have reported that eating the liver of the polar bear may produce various unpleasant symptoms; drowsiness, sluggishness, irritability, severe headache, vomiting, and peeling of the skin. It has been claimed that older, presumably experienced, sledge dogs will refuse to eat fresh bear liver, although young dogs will do so, but that frozen liver is eaten by young and old dogs alike.
Rodahl has made a vitamin assay and reports that polar bear liver is very high in vitamin A (13). Of three specimens tested, chemically and biologically, he measured two as containing 18,000 International Units of vitamin A per gram, and the third 13,000. The high concentration of vitamin A might be the cause of the toxicity, leading to hypervitaminosis A. The bear feeds upon the seal, and seal liver was found to vary in its vitamin A content according to the time of the year; so the liver of the bear would perhaps vary similarly according to the season, thus accounting for some of the conflicting reports as to its toxicity. Or the eating of several livers at successive meals might produce a headache when a single small meal produced no symptoms.
Recently it has been discovered that trichinosis is rather prevalent among carnivorous arctic mammals, among them the polar bear. During World War II, several Germans on a secret mission to the Franz Josef Islands were taken ill, invalided to Norway (which was then in German hands), and found to be suffering from trichinosis, readily traceable to bear meat. Since then trichinae have been found in bear meat from various parts of the Arctic, and trichinal infection

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has been determined from bearskins preserved as specimens in museums. Tests have revealed that some men now living are afflicted with trichinosis derived from bear or other arctic meat eaten years ago, the organism being now in an encysted state. A number of epidemics in the Far North and many separate deaths, either previously written down as mysterious or else falsely diagnosed, are now traced to this cause. A common wrong diagnosis has been typhoid. It is now believed, too, that much of the baffling dog sickness of the Arctic has been trichinosis.
Trichinae are killed by freezing, though not as quickly as by boiling. In the Arctic, most bear meat that is eaten has previously been frozen, which may have lessened materially the incidence of human and dog trichinosis. (The U.S. Department of Agriculture publishes tables showing how long meat needs to be kept frozen at various temperatures to be free from trichinae, the more intense the chilling the shorter the required period.)
Much has been written about the polar bear; some of it has now been discredited, some needs to be checked, and what remains still shows a wide range of variation in the habits of this animal.

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BIBLIOGRAPHY

1. Bobrinksoi, N. Opredelitel Melkopitaiushchikh ^ S ^ SSR (Mammals of the USSR), Moskva, Sovetskaia Nauka, 1944.

2. Degerbøl, Magnus and Freuchen, Peter. Mammals . Copenhagen, Gyldendalske Boghandel, Nordisk Forlag, 1935. Thule Expedition, 5th, 1921-24. Report vol.2, no.4-5, pp.102-19.

3. Karelin, D.B. “Vozdushnaia Ekspeditsia v vysokie Shiroty Arktiki v 1941 g.” (Air Expedition to the High Latitudes of the Arctic in 1941), Vsesoiuznoe Geogr. Obshch. Izvestia , vol.77, no.3, pp.164-69, 1945.

4. Miller, G.S. Catalogue of the Mammals of Western Europe in the Collection of the British Museum . London, Trustees, 1912, pp.297-303.

5. Nansen, Fridtjof. Farthest North. Being the Record of a Voyage of Exploration of the Ship Fram 1893-96 . Westminster, Constable, 1897. 2 vols.

6. Nelson, E.W. Report upon Natural History Collections Made in Alaska Between the Years 1877 and 1881 . Ed. by Henry W. Henshaw. Wash., G.P.O., 1887, pp.254-55. Arctic Series of Publications issued in Connection with the Signal Service U.S. Army, no.3.

7. Nordenskiold, A.E. The Voyage of the Vega round Asia and Europe . London, Macmillan, 1881, vol.1., pp.137-47.

8. Ognev, S.I. Zveri SSSR i Prilezhashchikh Stran . T. 3, [Khishohnye i Lasto– nogie] (The Mammals of the U.S.S.R. and Adjacent Countries, v ^ V ^ ol.3, Beasts of Prey and Pinnipedia.) Leningrad, M.L. ^^ Glavpushnine N.K.V.D. Biomedgiz, 1935, pp.124-39.

9. Ouwelhand, C.W. “Über Geburt and jugendentwicklung c ^ e ^ ines Eisbaren,” ^^ Zoologische Garten , Frankf o ^ u ^ rt, n.s., vol.2, pp.102-7, 1929. ^^

10. Papanin, D. Life on an Ice Floe . N.Y., Messner, 1939.

11. Peary, R.E. The North Pole , N.Y., Stokes, 1910.

12. Pedersen, Alvin. Der Eisb a ^ ȁ ^ r (T ^ h ^ alar e ^ c ^ tos maritimus Phipps ). København, Bruun, 1945. ^^

13. Rodahl, K., and Moore, T. “The vitamin A content and toxicity of bear and seal liver,” Bio-chem.J. vol.37, pp.155-68, 1943.

14. Seton, E.T. Lives of Game Animals . Garden City, N.Y., Doubleday, 1926. vol.2, “The Polar Bear,” pp.195-228.

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15. Stefansson, Vilhjalmur. The Friendly Arctic . N.Y., Macmillan, 1921.

16. ----. Arctic Manual. N.Y., Macmillan, 194 4 ^ 5 ^ .

17. Zalkin, V. “On the biology of the white bear of the Franz-Josef Archipelago,” Moskovskoe Obshch.Ispytat.Prir.Otdel.Biol. Bull . n.s., vol.45, pp.355-63, 1936.

A. L. Rand

Fur Bearers of North America

EA-Zoology (Leonard Butler)

FUR BEARERS OF NORTH AMERICA

CONTENTS

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Page
Arctic Fox 2
Muskrat 8
Red Fox 11
Wolverine 13
Beaver 14
Marten 14
Weasel 15
Mink 15
Arctic Hare 15
Arctic Wolf 16
Polar Bear and Musk Ox 16
Lynx 16
Otter 17
Squirrel 17
Bibliography 18

EA-Zoology (Leonard Butler)

FUR BEARERS OF NORTH AMERICA
The life of the Eskimo and Indian in the Far North in dependent upon hunting, fishing, and trapping. The Arctic and subarctic was formerly a vast preserve which annually produced a large fur catch for the scattered native population. These pelts were turned in at the trading stores for more and more white man’s good ^ s ^ until native life became almost completely centered around trapping. As population spread northward, exploitation of the fur resources of this frontier increased. Overtrapping was practiced and little done to protect the fur bearers. Under such conditions the more easily trapped and the less prolific animal ^ s ^ , such as beaver and marten, have been seriously depleted.
Trapping begins in November as soon as there is enough snow on the ground for traveling. A group of families usually camps together, living in tents or, in some parts of the North, after the real cold weather begins, in snowhouses. The men visit the traplines periodically to bring in fur and reset the traps, while the women and children remain in the central camp.
The land animals of the Arctic which are used for their fur consist of the following species: arctic fox, polar bear, musk ox, arctic hare, wolverine, and arctic f wolf. On the southern limits of the arctic area and in favorable localities extending up into the Arctic Circle, such as the Mackenzie River

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valley, the fur crop includes most of the bush species such as beaver, weasel, colored fox, lynx, marten, mink, muskrat, otter, and squirrel.
Arctic Fox
The bulk of the arctic fur crop comes from the arctic fox ( Alopex lagop y ^ u ^ s) . The relative number of pelts taken yearly from the different areas being:-

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White Blue
Alaska 5,000 10,000
Canada 50,000 1,000
Greenland 500 4,000
U.S.S.R. 80,000 1,500
The number of pelts taken fluctuates greatly from year to year. The Canadian catch crops to 18,000 pelts in times of fox scarcity and rises to 75,000 pelts in times of abundance. These periods of abundance occur every four years (see Population Cycles).
The arctic fox is circumpolar in distribution and occupies most of the land north of the tree limit in both the Old and New Worlds. In early times the range extended much farther south and included all of Europe and most of the northern part of Asia. There is no indication how far south the range extended in Canada. As climatic conditions changed, its range moved northward, and at the present time it is a typical representative of the tundra fauna of Europe, Asia, North America, and the islands scattered over the northern seas.
The arctic fox exists in two coat-color phases; white and “blue.” The white phase dominates in northern Canada and part of Alaska, the proportionate number of the blue varying from one blue to 500 whites to

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to one blue to 3,000 whites. In Greenland, the Aleutians and Pribilof Islands the blue phase predominates and white pelts are very scarce. On the islands off the Soviet coast the blues predominate but on the mainland the whites are in the majority. The original coat color was blue, and white is due to a recessive mutation. Thus it is possible for the same litter to contain both blue and white young ones and for two blue e foxes to be the parents of white cubs.
The fur of the arctic fox is very dense with almost woolly underfur; the guard hair is short and soft and the tail is thickly furred. The white phase has a snow-white winter pelt consisting of long white guard hairs and short, smoky-gray undefur. The underfur on the tail is darker and longer. Poorer quality pelts have a yellow tinge especially on the belly. The dark eyes and black nose stand out clearly against this background.
The blue phase varies in color from a smoky greyish-brown through the various shades of brown and gray. Many of the more desirable types have a sprinkling of white hairs scattered over the body, giving them a silvery appearance. They have no white tip to the tail.
The summer coat of both color phases is a smoky-brown color with no long guard hairs. At this stage whites and blues look much alike but can be distinguished by the expert. The autumn change in coat beings in September and lasts until December. The first signs of the spring molt appear in the second half of March. The molt begins at the head and gradually passes to the back, shoulders, and finally the belly.
The arctic fox belongs to the dog family and the genus Alopex . Many of the characteristics of this genus are halfway between Canis the dog genus and Vulpes the red fox genus. The main species is lagopus, meaning hare foot,

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and many authorities maintain that the rest have only subspecific rank. It is about two-thirds the size of the common fox and has an average weight of five kilograms. It resembles the red fox but its muzzle is broader and not as long. The ears are short and rounded and covered with hair so that they are hardly noticeable in the long winter coat. The nose and claws are black. The legs are shorter than those of the red fox and the pads of the feet are covered with hair. The average length of life is four to five years but in captivity some have been known to live for eighteen years.
The arctic fox makes its burrows in sandy or peaty soil or in favorable stony places on the tundra, generally not far from the coast. In soft soils the foxes dig burrows with numerous entrances while [: ] ^ in ^ stony places they make use of natural passages and crevices formed by large stones lying on top of one another. In these burrows in April and May the young are born. The size of the litter varies from 3 to 15, with authentic reports of litters as high as 22 pups. The average litter size is 7 or 8. The gestation period is 53 days, the rut extending from the middle of February to the middle of April, being a month later than that of the red fox. Both parents are said to take part in rearing the cubs. The mortality rate is very high especially before the young leave the burrow for the first time. In August the young embark on an independent life.
The main food of the arctic fox is the lemming, a large arctic rodent which fluctuates greatly in numbers. Other main sources of good are birds and their eggs in the nesting and hatching period, and fish and white whales thrown up by the sea. In the summer they [: ] usually live well off the eggs and young of ducks, geese, gulls, and other water birds. They vary

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their diet by taking berries and seaweed, crustace ^ a ^ ns, mollusks, and other small sea animals. In winter they are restricted to lemmings with an occasional arctic hare and ptarmigan. They sometimes devour the remains of a polar bear’s prey and old foxes will often persistently follow a bear for this purpose.
When food is scarce the arctic fox may migrate and at such times they come as far south as the tip of James Bay and into northern Manitoba and Saskatchewan. These southern migrations used to occur at each population high but since about 1936 there has been little of this southern movement.
The arctic fox has few enemies. Wolverines, wolves and occasionally polar bears hunt it but none of these animals, as far as the fox is concerned, is seriously predatory. In the past fifteen years the red fox has extended its range northward and infiltrated into the white fox territory. Wherever the red fox appears it is a serious enemy of the white fox. Because of its greater size and cunning the red fox is able to catch and kill many whites. The red also competes with the white fox for food and, from experience of the red for [: ] invasion of the Belcher Islands, it appears that a given area can support only a third as many red foxes as it can white ones. Since the red fox is worth only a quarter as much as the white fox and since it destroys many white foxes after they are caught in traps, the red fox can become a serious economic menace.
The arctic fox suffers from several diseases, the chief of which are paratyphoid fever, distemper, encephalitis, and rabies. Epidemics have been reported among northern sledge dogs with cyclic regularity when the fox population is at its highest and it is suspected that the arctic fox is the reservoir and carrier from one locality to another. At such times the

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foxes become bold, come close to dwellings and attack men, dogs, and reindeer. In the Soviet North this disease was classified as rabies but in Canada it was considered a form of encephalitis and was spoken of as “crazy dog disease.” Investigations at Baker Lake in the winter of 1947-48 definitely prove that ra v ^ b ^ ies was present in the dog population at this point and it may therefore be inferred that it was also present in the foxes.
Arctic fox trapping begins early in November as soon as the pelts become prime. Baits are scattered before the commencement of trapping, so that the foxes may get used to visiting this spot and not be suspicious when trapping commences. For bait, carcasses and lumps of the fat of the seal, bearded seal and Greenland seal, and arctic cod are used. Sometimes the arctic fox takes to addled guillemot eggs and they are used as bait. The arctic fox is usually much less trap-wary than is the red fox.
Most trapping is done with the double-spring trap. The trap is set in the immediate proximity of the bait. The chain is fastened to one of the stones that cover the bait, or to a peg driven into the snow. In the place designed for the trap, a disk of snow, corresponding in size to that of the trap with open jaws, is cut out with a knife. The bottom of the hole thus formed is made level and the set trap is lowered into it. Ten or more yards away from the place where the trap has been set a second disk of snow is cut out, a little larger than the first in size. This disk, which is about one inch thick, is brought to the hole in which the trap lies and carefully laid over it in such a way that the edge of the disk lies on the edge of the hole without spilling snow into it. After the disk is placed in position, snow is scattered along the edge. An area in the middle of the disk over the plate of the trap is scraped very thin but not enough to let the plate show through,

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Coming to the bait to eat it, the arctic fox breaks through the thin disk of snow, falls with its paws on the plate of the trap, thus springing it and getting its foot caught in the jaws of the trap.
A more primitive method of trapping is the deadfall which the trappers make with axe and knife out of driftwood or flat stones. It is a contraption of balanced levers with a trigger stick. The arctic fox enters the trap, pulls on the bait, which is attached to the trigger stick. This jerks the trigger stick out of place upsetting the balanced wood or stones which fall and crush the fox.
Many attempts were made in the Arctic to farm arctic foxes by the pen– raising methods used in silver fox ranching. Such attempts were mostly unsuccessful as food was costly to obtain in such localities and the fox did not breed very readily. Later, island farms were started in which mostly blue foxes are raised in a semiwild state.
When island farms were first established in North America the arctic fox was left to find food for itself. Now in the best farms the animals are fed periodically during the winter and early spring, which are the most difficult times for them.
The food is put in special cage traps, which act as both feeding places and live traps. The trap is in the shape of a little house made of planks. The food is put in the loft. There is a trap door at the point where the animal enters the loft. In the trapping season the catch on this trap door is released and the weight of the animal trips the door, dropping the fox into the lower part of the cage. The trap door is reset automatically, ready for the next fox. Animals in the cage below are culled and either pelted or marked and released for breeding purposes.

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Recently many silver fox farms in Canada and the United States have begun to raise the blue phase of the arctic fox. Because of its larger litters and lower consumption of food, it is becoming a profitable fur animal.
Since arctic foxes have been raised in captivity there have been several reports of hybridization with the red fox. These hybrids are almost as large as the red fox, bark like arctic foxes, and are stronger and more vicious than either parent. The ears are short like the arctic fox and the tail may or may not be white tipped. The fur has the shorter guard hair and heavy underfur of the arctic fox, but some of the guard hair has white bars like the silver fox. Crosses of white fox with silver have reddish flanks and go through a partial molt and color change. So far the only hybrid reported from the wild was in Iceland where an escaped silver fox vixen in the absence of a male of her own species mated with a blue dog. Most of these hybrids are sterile.
Muskrat
Next to the arctic fox, the most important fur from an economic standpoint is the muskrat (genus Ondatea ). Alaska and arctic Canada each produce 300,000 to 500,000 pelts na annually worth about one million dollars. Most of the muskrat catch of arctic Canada comes from the Mackenzie River Delta, where nature has created an exceptionally fine habitat for these small fur bearers, for here the Mackenzie breaks up into a great number of rivers, streams, and cut-off channels leaving innumerable saucer-shaped islands. The soil of the islands slopes downward to the center away from the river edge because of the large amount of sediment dropped when the stream overflows its banks. The islands thus invariably contain one or more lakes or marshy

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places. These lakes are subject to periodic flooding and drying up which makes an ideal home country for muskrats. Under these conditions muskrat food is abundant and the Mackenzie Delta ranks with the Mississippi and the Saskatchewan deltas as the great muskrat-producing areas of the North American continent.
Muskrats are caught by being trapped, either in their houses in the ice or in burrows along the stream banks. When trapping in houses, a thin spear is inserted to find the chamber and then a hole is chopped through the house wall at that point and the trap put in. Then the hole is filled to keep the house from freezing up The end of the trap chain is fastened to a stick and the stick is left protruding from the house to show the location of the trap. Bank trapping is much harder work because the runways are not as easy to locate and a great deal more chopping has to be done before the trap can be set. For this reason most northern trapping is done either in houses or else in open water after the ice goes out. Open-water trapping is not desirable, as the breeding season has begun by this time and many of the pelts are torn because of the fighting of males. A popular method of “ratt l ing” in the Mackenzie Delta is that of shooting the muskrats in the open water after the ice breaks up. Equipped with a light canoe and a .22 rifle a native can move quickly from one lake or stream to another. With such equipment a trapper can take 100 to 200 muskrats a night instead of 20 to 50 obtained by the other methods.
The muskrat has two litters a year ^ of ^ from three to twelve young. They are usually born in the period from June to the k ^ m ^ iddle of August. At first they are naked and helpless and their greatest protection is their aquatic habitat. The young are born either in the house or bank den or else in grass nests among the dense marsh vegetation.

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The muskrat, or “musquash” as it is known in the northern fur country, is three or four times the size of the common house rat to which it bears a superficial resemblance. It has short legs and strong hind feet which are partly webbed and modified for swimming. The long, naked, scaly tail is strongly flattened vertically and in the water serves well as a rudder. The fur is very fine and dense to protect its owner from the cold water in which much of its life is spent. In marshes or shallow lakes muskrats build roughly conical houses three to five feet high. These houses are made of roots and stems of plants with a mixture of mud. One or more oval chambers are chewed out of the interior above the water level to which entrance is gained by passageways opening under water.
The muskrat’s food consists of the roots and stems of such aquatic plants as cattail, bulrush, sweet flag, sedges, and pondweeds In summer it cuts the plants loose and then carries them in its mouth to ^ ^ a projecting stone or piece of driftwood and sits on this to eat, washing the food before eating. If no solid areas are available it builds feeding platforms of floating vegetation. In winter the muskrat does not normally come out into the open air above the ice because it is an easy prey for predators and also because its tail and feet freeze and it cannot withstand severe col t ^ d ^ . Occasionally when conditions under the ice are unfavorable many muskrats eme e ^ r ^ ge and wander around until they are killed or die. Trappers regard this as a sign that there will be extremely poor trapping in the spring. Since their winter range is thus confined to their house and the water under the ice, they must obtain their food from roots and plant material in the unfrozen marsh bottom. The food is taken to a spot above the water line to be eaten. This spot is either the regular house or a “push-up” which is a tiny house

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made some distance from the main house for the purpose of extending the feeding range.
Red Fox
The red or colored fox belongs to the genus Vulpes . The American species V. fulv e ^ a ^ exists in three well-defined coat-color phases, silver or black, cross, and red. The black or silver phase is the most valuable. The black phase has an almost jet black pelt with a white tail tip and a white line down the center of the belly. There are a few hairs scattered over the body which have white bars on them. In the silver phase these white-barred hairs become more numerous and give the pelt a silvery appearance. As the bar becomes wider or more of the hair becomes white the silver becomes lighter in color. In Platinum foxes the whole hair is white. So far there has never been a case of the platinum mutation reported in wild foxes. The red fox has yellowish-red hairs in place of the black hairs of the black phase and has a white brush (tail tip) and white belly. The cross fox has a mixture of red and w ^ s ^ ilvery black hairs. Usually both flanks are reddish yellow and there is a black line down the center of the back and another line across the shoulders. These two lines from a cross from whence this color phase gets its name. All phases have an exceptionally bus ^ h ^ y tail more than half the length of the body. They curl this around them when they sleep to prevent the sensitive nose the mouth from freezing.
The red phase is the original wild type and the silver is a mutation. At least two different mutations have occurred, one in eastern Canada and the other in the vicinity of Alaska. These two mutations look alike but are genetically different. If only one of these genes is present, a mating between a silver and a red gives all crosses. The red and the silvers are

EA-Zoo. Butler: Fur Bearers in North America

true-breeding. When both mutant genes are present, the inheritance is more complicated and the reds and silvers are no longer true-breeding. Litters of mixed parentage may contain all three color phases.
The proportion of silvers to cross and reds is not the same in all parts of the range. In general the proportion of silver and cross increase as one moves north. With the influx of foxes from the south this gradient has been maintained but the proportions have changed. For instance, in the Mackenzie River area the percentage of silver has fallen from 12% to 5% and of cross from 43% to 38%.
The colored fox is a solitary, predacious, mostly nocturnal animal which feeds principally on small birds and rodents such as mice, lemming, snowshoe rabbits, and arctic hare. It also eats eggs, insects, and fruit. After a gestation period of 54 days, the young are born in a burrow during the month of March. There are usually from three to eight young per litter and both parents help to take care of them. There is only one litter per year.
Red fox have a very wide range. Unlike most wild animals, they have even thrived in the close vicinity of man. The clearing of bushland has created better habitat for foxes and they have increased greatly in numbers during the past twenty-five years. Prior to this population increase, the colored fox was only of sporadic occurrence in the Barren Grounds and rarely bec o ^ a ^ me abundant in the fingers of bushland which stretch up into the Arctic. Since they have increased in numbers in the south they have spread northward, especially during the population peak of the ten-year cycle, and now compete with, and prey upon, the arctic fox in parts of its tundra range.

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The T ^ t ^ rapping methods for red are similar to those for white except that greater care has to be taken to keep the fox from becoming suspicious of the trapping site.
Wolverine
The wolverin g ^ e ^ is a circumpolar species belonging to the northern forested regions of both America and Asia. It has a low, squat, heavily built body, with strong legs and feet armed with sharp claws. The wolverine ( Gulo luscus , Gulo meaning glutton) is the largest and most formidable member of the weasel family.
The skunk-bear, or carcajou as the Canadian voyageurs called this animal, possesses a diabolical cunning and persistence. It frequently trails trappers along their trap lines, eating the bait from the traps they set and destroying the fur of animals caught in these traps. The wolverin g ^ e ^ is a great traveler and covers miles of country in its ceaseless search for food and mischief. Whenever t ^ i ^ t starts preying on a trap line the stouthearted trapper puts all his energy, resources, and ingenuity to work immediately to trap this marauder. The weakhearted trappers pull up their traps and move to another territory.
Because of its strength and cunning h the wolverin g ^ e ^ has earned a special place for itself in Eskimo folklore. Many hunters wear belts made from the skin of the head and legs of this animal in the fond belief that they will acquire some of the hunting prowess of the wolverine.
Its main food consists of fish,mice, birds and their eggs, berries, and bait which it robs from traps. If occasion demands, it will kill a deer or caribou. In May and June, after a gestation period of about ninety days, two or four young are born in a cave or burrow.

EA-Zoo. Butler: Fur Bearers in ^ of ^ North America

The fur is coarse, dark brown with lighter-colored stripes stretching from the neck along the sides of the rump. It is most used for parka trimming because it breaks the wind but does not colle d ^ ct ^ and hold moisture as easily as other furs.
Beaver
Arctic Canada produces 17,000 beaver ( Castor canadensis ) pelts annually, most of which come either from the Mackenzie River or from the more southerly rivers of Hudson Bay and James Bay. This largest of the rodents (an adult weighs 50 lbs.) builds houses and dams of sticks and mud and digs canals to float sticks from the place where they grow to the dam. They live chiefly on the bark, twigs, and young leaves of the aspen, willow, and birch. They mate in February and their litter of three to five young is born in May or June. The beaver does not reach maturity until it is two and a half years old. Because of its small litter, slow maturity, and ease of trapping, the beaver is easily trapped out unless stringent game laws are enforced or preserves established.
Marten
The Mackenzie River area, the Yukon, and the country around James Bay were formerly the best marten ( Martes americana ) country in North America. Overtrapping has made this valuable fur very scarce, the pelt numbers having fallen from over 30,000 pelts in 1850 to less than 5,000 today. The marten is extremely alert and inquisitive and very nimble and fast of foot. It feeds chiefly on squirrels and small mammals, birds and their eggs, frogs, fish, insects, and berries. It mates in June and July. The fertilized ovum does not embed for six or seven months (delayed implantation) and the

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two to four young are born in March or April. Experience on fur farms indicates that the marten is not prolific.
Weasel
The two species most commonly found in the Arctic are Mustela cicognanii , the short-tailed weasel, and M. rixosa , the least weasel. The former has a tail about four inches long, the last two inches of which are black, and the latter has a one-inch tail with a black tip. Both species change the color of their fur from brown in the summer to white in the winter. Little effort is made to trap this animal, the ermine (weasel) pelts turned in being taken in traps set for other animals or else by children and women trapping near the camp.
Mink
The mink ( Mustela vison ), a large weasel with partially webbed feet to adapt it for swimming, is never found far from the water. It is a blood– thirsty, tireless, solitary hunter that kills wantonly at times. It eats mice and other rodents, birds and their eggs, fish and muskrat young. Its ge ^ s ^ tation p r ^ e ^ riod varies from 30 to 65 days, the three to ten young being born in April or May. The choicest silky pelts are found in Labrador and northern Quebec; the pelts from the Mackenzie and the Yukon are larger and coarser.
Arctic Hare
Arectic hares ( Lepus arcticus ) resemble large jack rabbits and often attain a weight of more than ten pounds. They are more heavily built than the snowshoe rabbit and have shorter legs and ears. Their feet are furred and they change their coat color from agouti in the summer to snowy white

EA-Zoo. Butler: Fur Bearers of North America

in the winter. The trip of the ear is black. They usually have one litter a year of from one to eight young. The sex ratio may at times be predomi– nately female, which gives rise to travelers’ stories that arctic hares are produced by virgin births.
The pelts are soft and sil ^ k ^ y but the leather is too thin and easily torn to withstand hard wear. They are used for lining sleeping bags. Because of the scarcity of these animals and the low value of their pelts they are not extensively used by the fur trade.
Arctic Wolf
The arctic wolf ( Cani a ^ s ^ tundrarum ) is the largest of the North American wolves. The pelts often measure seven feet in length and have long sil ^ k ^ y guard hairs. The color is generally pale, almost white in some cases. The breeding and hunting habits are similar to those of other wolves except that the length of the arctic winter and the scarcity of food make it harder for them to survive. They feed chiefly on birds and their young, lemmings, hares, foxes, caribou, and musk ox.
Polar Bear and Musk Ox
Polar bear pelts are not used extensively by the fur trade. They are sold in limited quantities for rug and robe making. The musk ox has become so scarce that it is protected by law and its pelt is no longer an article of commerce. (See separate articles on these two animals.)
Lynx
Lynx ( Lynx canadensis ) are long-legged cats with tufted ears and a short “bobbed” tail. They have a thick coat of fur which gives them a fluffy appearance. The general color is gray with silvery white guard hairs and

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and the belly is pure white. Lynx have from two to five kittens which are born in the late spring and resemble the young of a domestic cat. The young usually hunt with their mother as a family group during most of the first year of their life. This animal periodically becomes very abundant in the Mackenzie River valley (see Cycles).
Otter
The otter ( Lutra canadensis ) is a short-legged, long-bodied animal which is more at home in the water than on land. Its fur is very strong and has greater wearing qualities than any other American fur. Otter are not prolific breeders and the number of pelts taken annually in Canada has fallen from over 20,000 to 12,000 pelts.
Squirrel
The squirrel ( Sciurus hudsonicus ) occurs as far north as the limit of trees. Its pelts were formerly so cheap that it was not hunted in northern areas. In 1942, however, there was a tenfold increase in pelt prices and there is now considerable pressure on the squirrel population. The northern squirrels do not reach the high population densities which occur in the aspen– grove regions of the prairies and are therefore more easily decimated by trapping. Since the squirrel feeds on vegetation and is the lowest animal in many food chains, it is doubtful if the trapping of squirrels in northern sections is a good practice.

EA-Zoo. Butler: Fur Bearers of North America

BIBLIOGRAPHY

1. Bar ^ a ^ bash-Nikiforov. “Material on the food habits of the Mednuii (Copper) Island arctic fox Alopex beringensis semenovi Ogn.,” Moskovskoe Obshch. Ispytat. Prir. Otdel Biol. Bull . n.s. vol.48, no.1, pp.78-80, 1939.

2. Butler, L. “The genetics of the colour phases of the red fox in the Mackenzie River locality,” Canad.J.Res. Sect.D, vol.25, no.6, pp.190-215, Dec., 1947.

3. Dubrovskii, A.N. “Pesets ( Alopex lagopus (L.)) i pestsovyi Promysel na Novoi Zemle.” (The arctic fox (Alopex lagopus and arctic fox trapping in Novaya Zemlya), Leningrad. Arkticheskii Nauchn. –Issled.Inst. Trudy vol.77, pp.7-31, 1937.

4. Elton, Charles. “Epidemics among sledge dogs in the Canadian Arctic and their relation to disease in the arctic fox,” Canad.J.Res . vol.5, pp.673-92, 1931.

5. Kirpichnikov, A.A. “On the biology of the arctic fox of the southwest coast of Taimyr,” Moskovskoe Obshch.Ispytat.Prir.Otdel Biol. Bull . n.s., vol.46, pp.52-57.

6. Lavrov, N.P. Pesets . (The Arctic Fox.) Moscow, Vneshtorgizdat, 1932.

7. Nelson, E.W. Wild Animals of North America. Wash., National Geographic Soc., 1930.

8. Robinson, M.J., and Robinson, J.L. “Fur production in the northwest territories,” Canad.Geogr.J . Jan., 1946, pp.34-46.

9. Plummer, P.J.G. “Further note on arctic dog disease and its relation to rabies,” Canad.J.Comp.Med . vol.11, pp.330-34, 1947.

L r ^ e ^ onard Butler

Terrestrial Fur Bearers of Greenland

EA-Zoology (Magnus Degerbøl)

TERRESTRIAL FUR BEARERS OF GREENLAND

CONTENTS

Scroll Table to show more columns

Page
Collared Lemming ( Dicrostonyx groenlandicus groenlandicus ) 4
Greenland Polar Hare ( Lepus arcticus groenlandicus , and L. arcticus porsil i di ) 6
Greenland Arctic Wolf ( Canis lupus orien; C. lupus eogroenlandicus ; C. lupus arctos ) 7
Greenland Fox ( Alopex lagopus groenlandicus ) 9
Greenland Ermine ( Mustela erminea polaris ) 10
Polar Bear ( Thalarct ^ o ^ us maritimus ) 11
Bibliography 14

EA-Zoology (Magnus Degerbøl)

TERRESTRIAL FUR BEARERS OF GREENLAND
The land mammals of Greenland comprise only eight species in all, including two rodents, two ungulates, and four carnivores, as follows: ( 1 ) Collared lemming ( Dicrostonyx groenlandicus groenlandicus ); ( 2 ) Gro^e^en–land polar hare ( Lepus arcticus groenlandicus, and L. arcticus porsildi); (3) Caribou (Rangifer tarandus groenlandicus); ( 4) Musk ox (Ovibos moschatus wardi); (5) Arctic fox ( Alopex lagopus groenlandicus ); (6) Arctic wolf ( Canis lupus orion, C. lupus eogroenlandicus, and C. lupus arctos ); (7) Greenlandermine ( Mustela erminea polaris ); (8) Polar bear (Thalarctos maritimus).
Of these mammals, the caribou and musk ox are not considered as fur bearers and will not, therefore, be dealt with specifically in this paper.
There can hardly be any doubt that these land mammals, possibly with the exception of Rangifer , originated from Arctic North America, from which country Greenland is separated only by narrow and frequently ice-covered straits. Compared with that of North America the land-mammal fauna of Greenland is poor, although, particularly with regard to South Greenland, the conditions are such that means of subsistence are at hand for more species than really occur there. The explanation of this nonoccurrence is that the migration road for land mammals lay across the high-arctic North America,

EA-Zoo. Degerbol: Terrestrial Fur Bearers of Greenland

over the narrow sounds just mentioned. Only the species which have survived this process of selection have got through. Several American species of mammals which could undoubtedly live in Greenland, such as Gulo , Lemmus , and Citellus , have after the glacial epochs not been able to spread so far to the north and have therefore not yet reached Greenland. As none of the Greenland land mammals is of hibernating habit, they also have possibilities of distribution when rivers and waters, which are otherwise obstacles in the way of migration, are covered by ice.
Four of the mentioned eight species: collared lemming, musk ox, arctic wolf, and ermine occur only in North and Northeast Greenland, south to the Scoresby Sound region. Nor does the polar hare occur on the east coast south of this region; on this southeastern coast we find only two species of land mam ^ m ^ als: the arctic fox, which is found all over Greenland, and the polar bear. The caribou is now restricted in its distribution to the west coast, but until about 1900 it l o ^ i ^ ved also on the northeastern coast.
The reasons for the northeastern distribution of the four species (collared lemming, musk ox, wolf, and ermine) must undoubtedly be sought in the climate and the possibilities of distribution.
In Greenland we have a high-arctic continental climate in the north and a subarctic, oceanic [: ] climate to the south. The boundary between these regions is the important July isotherm of 4 to 5°C. On the west coast this isotherm comes in at about latitude 73° N. (Upernivik) and crosses the east coast south of Scoresby Sound at about 69° N. The country south of Scoresby Sound, the Blosseville Coast, is a boundary area. The precipitation in Angmagssalik is nearly three times greater than in Scoresby Sound. Another factor of decisive importance is that the Blosseville Coast essentially

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consists of very steep basalt rocks, divided by a large number of productive glaciers, and, as distinguished from Northeast Greenland, there are no extensive plains where edible material can form, and where great land mammals, e.g., musk ox, can find sufficient food. To the northwest the range of these four continental species is barred by the more than 100- m^k^ilo–meter-wide Humboldt glacier, which debouches into the sea at this point, or by the Melville coast with its numerous glaciers.
It has been supposed that these northern mammals arrived in Greenland in fairly recent time. In 1822, the Scoresbys, father and son, were in the fjord which now bears their name, but saw no musk oxen or any signs of them. Nor did Clavering and Sabine, who in the following year were on the coast farther north, find any trace of the presence of the musk ox. The first expedition to see live musk oxen on the east coast of Greenland was the Germania expedition under Koldewey in the year 1869. This led some investi– gators to conclude that in the years 1822-23, the musk ox had not yet come to East Greenland south of latitude 75° N. However, new light upon this question was given in 1934 when musk ox bones were excavated from Eskimo house ruins at Clavering Island dating from about 1600; also the fact that the East Greenland musk ox belongs to a special ra d ^ c ^ e ( Ovibos moschatus wardi ) points toward its occurrence in East Greenland as dating rather far back.
Curiously enough, these explorers did not see any caribou in 1822-23 either. Excavations have proved, however, that this species has lived in East Greenland at least from the beginning of the 16th century through the 19th century. For the later expeditions of Koldewey ^ Koldewey ^ ^ ( ^ 1869 ^ ) ^ , Ryder ^ Ryder ^ ^ ( ^ 1891 ^ ) ^ , and a Nathorst ^ Nathorst ^ ^ ( ^ 1899 ^ ) ^ , found herds of caribou in East Greenland, but whe d n the

EA-Zoo. Degerbol: Terrestrial Fur Bearers of Greenland

Denmark Expedition 1906-08 came to the east coast the species had entirely disappeared, and since then it has not been met with in East Greenland.
COLLARED LEMMING 9 ^ ( ^ Dicrostonyx groenlandicus groenlandicus )
Distribution : North and Northeast Greenland, south to d’Aunay Bay, Blosseville Coast. On Melville Peninsula the Greenland collared lemming intergrades with the American race D. groenlandicus richardsoni .
In summer coat the Greenland collared lemming is less brightly colored than the American subspecies, and in adult specimens the dark dorsal stripe is missing. The total impression of the animal in summer pelage is that the back is grayish to grayish black with a faint reddish-brown tinge. A characteristic feature in the Greenland collared lemming is that the hairs on the flanks and the upper part of the breast are rusty red. As is well known, the collared lemming acquires a white winter coat, and the claws of the two central digits are then v a ^ e ^ ry large, sometimes exceeding half an inch in length, and have the appearance of being double, one on top of the other.
Most of the authors who have visited Greenland are of the opinion that this animal breeds once or twice during the summer. One author states that in good lemming years it also brings forth a litt l ^ e ^ r of young in early spring, March-April, in winter nests which are situated on the ground under the snow; the second litter is then born in May-June, and the third in July-August. The young that are born in early spring propagate in their first year. The usual number in the litter is three to five; this number seems to be smaller than in the case of the American collared lemming. Observations made on animals held in captivity in Denmark have, however, shown that the Greenland collared lemming is able to bring forth at least five litters in the course

EA-Zoo. Degerbol: Terrestrial Fur Bearers of Greenland

of the summer, the interval between each litter being about a month, and the number of young per litter varying between two and seven.
In old embryos the vibrissae and claws are visible. After birth the following characteristics have been observed. One day old: head and body 35 mm.; the animal has a dark back and a light-colored belly with a sharp line of demarcation between these two parts; the back proper is covered with rather long, black hairs, up to 2 mm. in length, which are, however, not so closely set as to conceal the dark skin. Three to four days old: head and body 43 mm.; the animals have a distinct dark dorsal stripe; the hairs on the back are so dense as to cover the skin completely; the color of the belly is still determined by the skin. About 14 days old: head and body 53 mm., the eyes open. Adult: and head and body 100-145 mm. (20 specimens measured). The maximum lifetime of this animal is two years.
Young born in the spring have very pronounced pads on the toes below the central digits. This clearly shows that the young, which in Greenland are born under the snow in late winter or early spring, are from the start supplied with a double claw before they have had an opportunity to dig. Therefore, in this case there can be no question of acquired qualities; the phenomenon is undoubtedly hormonally determined.
The pelage of the collared lemming is extremely soft and thick, and the animal looks like a small furry ball. With its broad-bellied shape, it resembles in miniature (particularly when sitting on its hind legs) a small “Bobak,” the marmot of the Arctic. The collared lemming prefers fertile localities, in particular where willows, its favorite food, grow thickly and form a layer of humus. The collared lemming plays an important part

EA-Zoo. Degerbol: Terrestrial Fur Bearers of Greenland

in the animal world of the Arctic. Since this small rodent mainly accounts for the occurrence of several carnivorous animals, for instance such a valuable fur animal as the arctic fox, it is obvious that the considerable fluctuations in the number of individuals of the lemming will also affect the occurrence of several other animals. These fluctuations, as is well known, show a period of approximately four years.
GREENLAND POLAR HARE ( Lepus arcticus groenlandicus and L.a. porsildi )
Distribution : All Greenland coasts except the southeast regions south of Cape Dauss e y Blosseville Coast.
There are two subspecies of hares in Greenland, the North Greenland L.arcticu groenlandicus on the east coast, whence it spreads to the northern part of the west coast, the region round Disko Bay, and the South Greenland L. arcticus porsildi , L. arcticus groenlandicus is charac g ^ t ^ erized by its forward-projecting and protruding (proodont) upper incisors, while porsildi has more normal (orthodont) incisors. A method for measuring the angle at which the upper incisors emerge from the skull (by means of a goniometer) has made it possible to demonstrate that there is a gradual diminution of the index when going southward. It is interesting that this change is not confined to a limited zone, but is apparently observable between any two series from different latitudes, a beautiful example of a “cline.” The Greenland hares have probably emigrated as typical groenlandicus from the north, from Ellesmere Island. Projecting incisors are well adapted for rooting among snow and stones for dwarfed plants, but are inefficient for ordinary regions farther south. The orthodont feature must, the ^ n ^ , have been achieved as an adaptation to the more luxuriant plant life in southern Greenland, and the animals have resumed the normal curving of the incisors. The evolution has thus been: orthodont incisors in the American races of

EA-Zoo. Degerbol: Terrestrial Fur Bearers of Greenland

ar ^ c ^ ticus west of Davis Strait; projecting incisors in groenlandicus in Ellesmere Island and North Greenland; orthodont incisors again in South Greenland.
L. ar ^ c ^ ticus groenlandicus remains white throughout the year, though the white summer pelage is duller and scantier than the winter pelage; at all seasons the ear tips are black. In L. ar ^ c ^ ticus porsildi the summer coat is shaded with a pale buffy. The color of the leverets is always grayish.
Greenland hares generally have only one litter in a year. The young are born in June and the usual number in a litter is five or six. The hares are very common in Northeast Greenland, but in most other districts they are rather sparse. They live from the outer coast to the inland ice and on the nunataks, preferring the inner parts of the ice- pile oaves ^ free area ^ . Especially in winter they keep to fairly steep slopes where grass and other vegetation are kept free from snow. Their favorite food seems to be the roots of salix and saxifraga .
The animal is of no great economic importance to the Greenlanders, who generally do not like its meat. In the Thule district the skins are used for stockings.
GREENLAND ARCTIC WOLF ( Canis lupus orion; C. lupus eogroenlandicus ; C. lupus arctos )
Bistribution : North and northeast coast, south to Scoresby Sound.
The arctic wolf is a very rare animal in Greenland. It is extremely cauti on ^ ous ^ and shy and has only occasionally been observed and killed by expeditions and hunters. Its occurrence on the east coast was first ascertained by the Nathorst Expedition, 1899, which prompted this explorer

EA-Zoo. Degerbol: Terrestrial Fur Bearers of Greenland

to set forth the view that the wolf had only quite recently immigrated to the east coast from the north. In the excavated material from Clavering Island there were no wolf bones. In this connection it should, however, be observed that this animal, which roams about alone or in small groups, and which has seldom been trapped by the Eskimos, most probably was not seen by the first explorers of East Greenland; therefore it was not represented in the excavated material. As in the case of the East Greenland musk ox, it should be emphasized that the East Greenland wolf belongs to a special d race ( C. lungus ^ lupus ^ eogroenlandicus ). It is characterized by the very large premolars, indeed so large that the second and third premolars overlap. Pocock (1935) has based a special subspecies Canis lupus orion on a skin and a skull lacking in posterior, from Cape York, [: ] Northwest Greenland. There is not much to be said about such scanty material; it will only be mentioned that in this skull, which the author has examined in London, the premolars are widely separated. Probably this animal was an immigrant from Ellesmere Island ( C. lupus arctos ^ C. lupus arctos ^ ).
It may be mentioned as a unique event that as early as the winter of 1868-69 two white wolves were observed on the west coast, in the northwestern district of Umanak; one was shot and its skin and skeleton are now in the Zoological Museum of Copenhagen. This specimen, too, has widely separated premolars and must be regarded as an accidental visitor. The year 1868 was a well-marked invasion year for white foxes from Canada.
In summertime the wolf has no difficulty in finding food — lemmings, leverets, musk ox calves, broods of birds, etc., but in winter things are quite different; the struggle has been especially difficulty since about 1900, when the caribou disappeared from the country. At this season hungry wolves

EA-Zoo. Degerbol: Terrestrial Fur Bearers of Greenland

at times attack, kill, and devour dogs.
GREENLAND FOX ( Alopex lagopus groenlandicus )
Distribution : All districts of Greenland.
From an ecological point of view the Greenland foxes have been divided into two categories: lemming foxes, which live on the north and northeast coast to the Scoresby Sound region, where lemmings occur; and “coast foxes,” which subsist largely on products of the sea. In lemming foxes there are regular fluctuations with maximum numbers at intervals of about four years correlated with the four-year cycle of the lemmings. In coast foxes there is no such regularity, and the food supply at the disposal of these foxes does not fluctuate so regularly. In coast foxes the blue phase is most common, in some small islands even approaching one hundred per cent. As a further ecological differentiation within this last group may be mentioned foxes resorting to bird cliffs with their more constant food supply. Under such circumstances the blue variety is especially dominant and the number of young in a litter is fairly constant from year to year.
Apart from the autochthonous lemming foxes in Northeast Greenland, there are immigrant lemming foxes which come from Canada via Smith Sound or on the drifting ice. As almost all the Canadian foxes are white, the arrival of these animals is indicated by a rise in the percentage of white foxes in the catch. In the Thule District this immigration takes place every winter, but in especially great numbers every four years, that is, the year of lemming minimum following the lemming maximum. From an economic point of view, these white foxes are less valuable than the blue ones.
The rutting season of the arctic fox in Northeast Greenland is in March

EA-Zoo. Degerbol: Terrestrial Fur Bearers of Greenland

and the beginning of April, and the young are born in May. As already mentioned, the collared lemming may bring forth its first litter of young below the snow in the winter or early spring, especially in lemming years. It is evident that these early litters are of great importance to the breed– ing conditions of the arctic fox. “Fox years” are conditioned by a good state of nutrition before and during the period of pregnancy. Most authors, with the exception of a few earlier ones, as Otto Fabricius ^ Otto Fabricius ^ , deny the possi– bility of more than one annual litter of arctic foxes, even in good lemming years.
In Greenland blue foxes predominate, but the proportion between blue and white varies from place to place. In the year 1947-48, the following number of fox skins was traded: in South Greenland, to Holstein ^ s ^ borg, 966 blue foxes, 547 white foxes; in North Greenland, 1,075 blue foxes and 622 white foxes; in East Greenland, 41 blue foxes and 160 white foxes. In the Thule District the skins are used for trousers and over-jackets, and in this district the meat is eaten during the winter, when the animals are fat. In other districts the skins are sold to the Greenland Trading Company.
GREENLAND ERMINE ( Mustela ermine polaris )
Distribution : North and Northeast Greenland.
The collared lemming is the chief food of the ermine; it is quite natural, therefore, that the ermine has the same distribution as the lemming.
The ermine of Greenland shows the greatest affinity to the ermine of the eastern North American Arctic; in bo g ^ t ^ h the tail is relatively thin and short and the black tip of the tail is relatively large, but the Greenland ermine differs somewhat with regard to the color of the summer skin, having a lighter

EA-Zoo. Degerbol: Terrestrial Fur Bearers of Greenland

appearance. Practically speaking, the Greenland ermine is intermediate in color between the New and Old World erminea . The fur is soft and thick, as in specimens of the ermine from the high north elsewhere.
In Greenland the ermine is of fairly common occurrence, but it is nowhere to be found in any great numbers. In summertime the animal strolls about in the open air, but in winter it generally lives under the snow, being less adapted to resist the severe cold than other terrestrial mammals of the region. Pairing takes place by the end of the March-April. In the years when lemmings are rare, the ermin d ^ e ^ may not propagate at all or it may eat its young.
POLAR BEAR ( Thalarct u ^ o ^ s maritimus )
Distribution : All coasts of Greenland.
The polar bear may ^ ^ occur along all the coasts of Greenland, but on the west coast, between Julianehaab and Upernivik districts, it is a very rare and occasional guest. On the north coast proper, in the area between Polaris Bay and Independence Fjord the polar bear is also rare. Its real residence is the northern part of the west coast, and the east coast south to Scoresby Sound. On the east coast it comes from this region with the drift ice in winter and spring down the Blosseville Coast in fairly large numbers; some mo r ^ v ^ e farther south along the southeast coast, around Cape Farewell, and up along the west coast to the southern part of Julianehaab District. Not all animals, however, are transported so far south; many go ashore in order to go north again to the fjords of Northeast Greenland. The polar bear is e s ^ x ^ tremely skillful at finding its way and may of f ^ t ^ en be seen to take the shortest cuts between two fjord systems across peninsulas or behind promon– tories, where regular “bear paths” may be seen.

EA-Zoo. Degerbol: Terrestrial Fur Bearers of Greenland

The everlasting wanderings of the polar bear up and down the coasts is partly determined by the supply of food, partly by the drift ice. Since it lives principally on seals, it seeks out places where these animals are to be found. In East Greenland its chief residence in winter is the drift ice, especially where openings in the ice are to be found outside the coast, but it goes far out to sea too. At this season, before the seals come up on the ice, the bear can catch adult specimens at the breathing holes. In March-April the polar bears wander to the breeding places of the ringed seal, and may go far into the fjords where seals are living, especially on the east coast between Scoresby Sound and Dove Bay. Also on the northwest coast of Greenland in spring they seek the breeding places of the ringed seal, especially around Melville Bay and Peabody Bay, in Kane Basin. For a more complete discussion of habits and characteristics, see article on “Biology of the Polar Bear,” by A. L. Rand.
To the Polar Eskimos in the Thule District polar bears are of great importanc d ^ e ^ ; the skins are used for trousers and for rugs on sleeping platforms, and the meat is eaten. In this district only a few skins are traded. As far as the remaining parts of Greenland are concerned, polar bears are of some importance to only four of the districts, namely, Scoresby Sound and Angmagssalik on the east coast and Upernivik and Julianehaab on the west coast. When the Scoresby Sound settlement was founded in the year 1925, no less than 102 bear skins were traded, but the number soon diminished, averaging 44 annually in the years 1930 to 1940 and 36 in the years1940 to 1950. At Angmagssalik the average number of bear skins traded (number of animals killed is not known, but, practically speaking, all skins are traded) are, per annum: 1890-1900, 56.6 skins; 1900-10, 68.6; 1910-20, 86.9; 1920-30, 86.7; and 1930-40, 45.8.

EA-Zoo. Degerbol: Terrestrial Fur Bearers of Greenland

In the last decades the number of bears has been strongly declining. On the northwest coast the following number of bear skins has been k ^ g ^ raded: Upernivik District; 1850-60, 36.4; 1860-70, 39.8; 1870-80, 19.5; 1880-90, 15.7; 1890-1900, 25.2; 1900-10, 28.7; 1910-20, 14.4; 1920-30, 17.6; 1920-40, 5.7. Julianehaab District: 1850-60, 6.7; 1860-70, 8; 1870-80, 16.8; 1880-90, 29; 1890-1900, 31.7; 1900-10, 31.1; 1910-20, 29.1; 1920-30, 18.5; 1920-40, 6.6

EA-Zoo. B ^ D ^ eger d ^ b ^ ol: Terrestrial Fur Bearers of Greenland

BIBLIOGRAPHY

1. Braestrup, F. W. “A study on the arctic fox in Greenland. Immigrations, fluctuations in numbers based mainly on trading statistics.” Medd.Grønland . vol.131, no.4. 1941.

2. Degerbøl, Magnus. “Animal bones from the Eskimo settlement in Dødemandsbugten, Clavering Island. A contribution to the immigration history of the musk ox and reindeer in East Greenland,” Larsen, Helge. “Dodemandsbugten,” Medd. Grønland , vol.102, no.1, 1934. Zoological Appendix, pp.173-80.

3. ----. “A contribution to the investigation of the fauna of the Blosseville coast, East Greenland, with special reference to zoogeography,” Medd. Grønland , vol.104, no.19, 1937.

4. ----. “Mammals. Part 1, Systematic notes,” Thule Expedition, 5th, 1921-24. Report , vol.II, no.4-5, pp.1-67, 1935.

5. ----, and Braestrup, Wimpfen. “The Geographical variation of the Greenland floras,” Dansk Naturhist. Foren, Copenhagen, Vidensk Medd . vol.98, 1934.

6. ----, and Mohl-Hansen, U. “Remarks on the breeding conditions and moulting of the collared lemming ( Dicrostonyx ),” Medd. Grønland , vol.131, no.11, 1943.

7. Fabricius, Otto. Fauna Groenlandiae . Rothe, Hafniae et Lipsiae, 1780.

8. Freuchen, Peter. “Mammals. Part 2. Field notes and biological investi– gations,” Thule Expedition, 5th, 1921-24. Report . vol.II, no.4-5, pp.68-278, 1935.

9. Jensen, Ad.S. Grønlands Fanua . Københaven, Bianco Lunos Bogtrykkeri, 1928.

10. Manniche, A.L.V. “The terrestrial mammals and birds of North-East Greenland,” Medd. Grønland , vol.45, no.1, 1910.

11. Müller, B. Vildtet og Jagten i Sydgrønland . København, Hagerup, 1906.

12. Pedersen, Alvin. “Beiträge zur Kenntnis der Säugetier- und Vogelfauna der Ostküste Grönlands,” Medd. Grønland, vol.68, no.3, 1926.

13. ----. Der Eisbär (Thalarctos maritimus Phipp a ^ s ^ ). Verbreitung und Lebensweise . København, Bruun, 1945.

14. ----. “Fortgesdtzte Beiträge zur Kenntnis der Säugetier- und Vogelfauna der Ostküste Grönlands,” Medd. Grønland , vol.77, no.5, 1930.

EA-Zoo. B ^ D ^ eger d ^ b ^ ol: Terrestrial Fur Bearers of Greenland - Bibliography

15. Winge, Herluf. “Grønlands Pattedyr,” Medd. Grønland , vol.21, no.2, 1902.

16. Young, S.P., and Goldman, E.A. The Wolves of North America . Wash., American Wildlife Institute, 1944.

Magnus Degerbøl

Fur Bearers of Arctic and Subarctic Europe and Asia

EA-Zoology (Sven Ekman)

FUR BEARERS OF ARCTIC AND SUBARCTIC EUROPE AND ASIA

CONTENTS

Scroll Table to show more columns

Page
The Lynx 2
The Pine Marten 3
The Sable 6
The Ermine 8
The Glutton or Wolverine 10
The Otter 13
The Wolf 14
The Eurasian Red Fox or Forest Fox 16
The Arctic Fox 18
The Brown Bear 20
The Arctic Hare 22
The Squirrel 23
The Beaver 25
Other Eurasian Fur Bearers 26

[EA-Zoo. Sven Ekman]

FUR BEARERS OF ARCTIC AND SUBARCTIC EUROPE AND ASIA
Considered zoogeographically, the Arctic comprises the zone of the tundra and that of the related highland health or mountain tundra, so that its southern boundary coincides with the northern limits of the forest zone. The number of genuine (endemic) arctic animals is small; of the Eurasian fur bearers, and excepting the marine species (whales, seals, polar bears), only the arctic fox belongs among them, strictly speaking. Nevertheless, reindeer and glutton may also be grouped with them; both include the northernmost coniferous forests in their range, but their main habitat is the tundra or at least the lightly wooded tundra. Equally at home in the tundra, although of a still more southerly range than the foregoing two, are the wolf, the red fox, the ermine, the small weasel, and the hare. However, as the scope of this work calls for consideration not only of the tundra animals but of the fauna of the northernmost forest zone as well, several other species, some of which consti– tute excellent commercial values, must be included in the survey to follow.

EA-Zoo. Ekman: Eurasian Fur Bearers

The Lynx
The only member of the cat family that belongs in the category under consideration is the lynx ( Lynx lynx ). In former times, the animal inhabited the forests of Europe down to the Mediterranean and those of Asia as far south as the Caucasus, northern Persia, and southeastern Tibet. In central Europe, it has long since been exterminated, and it is rare even in Scandinavia, where a few specimens are still to be found in Sweden, north of about latitude 63° N. The species is somewhat more common in northeastern Finland, but its main habitat is now in northern European Russia and in Siberia.
The lynx is not an arctic animal and does not form a part of the fauna of the tundra zone. In the northern Scandinavian mountains even the birch forests that circle the southern edge of the highland health (mountain tundra) areas lie outside the usual habitat of the lynx, although it has been observed here prowl– ing for game. Its young, however, are never born above the limits of the coniferous forests.
According to observation made in zoological gardens and in the wilderness, the family life of the lynx in the northern region is as follows. April is mating time. This is one of the rare occasions when the voice of the animal is heard, starting with a “meow” resembling that of the domestic cat and ending in a hollow bark or howl, or else in a low snarl. The gestation period [: ] averages ten weeks, after which time two, rarely three, young are born, which are blind for about two weeks. The den is in a rocky crevice or the like, and so far has been found only by very few people. The young follow their mother until the end of the winter, but the latter send them about their business at the beginning of the next rut.

EA-Zoo. Ekman: Eurasian Fur Bearers

The lynx may attack prey as large as an adult reindeer, but [: ] its favorite quarry is the hare. Lemmings and voles are also to its taste and when abundant constitute its main food. Hares and large-sized game are over– taken with a few very long leaps, usually eight or ten, but the hunt is quickly abandoned if not successful at once. However, its temper is that of a wanton destroyer, for when breaking into a sheep or goat pen if will kill as many animals as possible in order to suck their blood. It is therefore considered a hateful marauder to be destroyed at all cost; it is also hunted because of its beautiful pelt, which has high commercial value. The lynx is easily caught, for it lacks endurance and when pursued by fast dogs quickly takes to a tree.
The Pine Marten
The pine or forest marten ( Martes martes ) is no tundra inhabitant either, but its range extends northward as far as the northernmost limits of the coni– ferrous forest. In Europe, it is found as far south as central Italy, Sardinia, and the Balearic Isles, although in some of the more densely populated countries it is now nearly or altogether extinct. In Asia, it inhabits the taiga zone as far east as Manchuria, and it is said to occur in the Himalayas, south of the steppes and the deserts.
No other beast of prey is as skilled a climber of trees as the marten, and the facility with which it hunts and overtakes squirrels in treetops is well known. Squirrels are its favorite prey and in regions where a marten has settled the former soon disappear. Its preferred way of traveling is from treetop to treetop but it also takes some of its prey on the ground, sometimes chasing a hare over several miles. It hunts during the nig t ht and rests during the day,

EA-Zoo. Ekman: Eurasian Fur Bearers

preferably in a squirrel’s nest. Hunters who have shot up to a hundred martens report that the majority of their quarry was killed or roused in such nests. The marten will kill any warm-blooded animal not very much larger than its own size, whether it lives in trees or on the ground. Many instances are known where a marten attacked a capereaillie or a black cock in a tree and then “flew” to the ground in the back of the bird; in one instance, a distance of 40 meters was thus traversed, in another all of 300 meters. While prowling at night, the marten has no difficulties in finding the trees on which the birds are roosting, as they betray their presence by the droppings accumulating on the ground below. Hens nesting on the ground are also attacked. More often than not only a small part of the quarry is devoured before the killer is off on another hunt again to leave only a partly eaten spoil behind. The marten also feeds on birds’ eggs, the honey of wild bees, on fruit, such as apples, pears, and cherries, and finally on berries, including bilberries, sorb apples, and the berry of the juniper bush.
Like most predators, the marten often buries the food of which it has no need at the moment, and it is able to find its various caches even after a long period of time. One man who followed a marten in wintertime , noticed that it had entered a stone pile twice and twice reappeared with the egg of a capercaillie which was then eaten on the spot. A forester reported five similar incidents. A hunter ^ , ^ who is also an experienced zoologist, relates that he had quite often seen evidence of a marten digging here and there in the snow to recover eggs buried during the [: ] previous summer; the eggs were those of capercaillies, black grouse, ptarmigan, and mallards. Dead capercaillies, found in the nests of squirrels or of black woodpeckers, had obviously been hidden there by a marten.

EA-Zoo. Ekman: Eurasian Fur Bearers

The fertilization story of the species, although not altogether exceptional among the marten-like varieties, is a strange one in at least one regard. Up to 20 or 30 years ago, it was assumed that implantation took place late in winter and that the period of gravidity was about 2 months, for the young are usually born in April. Observations in zoological gardens, on animal farms, and even outdoors, have led to different conclusions. There is no doubt now that in almost all cases the main, effective mating takes place in July, rarely toward the end of June or at the beginning of August, and that the period of gestation is 8 to 9 months instead of 8 to 9 weeks. At the fur-bearing animal station at Pushkino, U.S.S.R., where marten and sable are bred experimentally, the sexes are permitted to mate only during the summer and are kept separate for the rest of the year. Yet the young are regularly born in April. Only on very few occasions have such births been preceded by a winter mating. The few known exceptions were very young females, hardly a year old, although the female is generally not fully matured until her second year. The period of gestation may therefore vary to a considerable extent. An explanation for this may be that the development of the embryo is slowed up if not altogether at a standstill after mating has taken place, as is the case with the badger which belongs to the marten family. The badger mates in the late summer or early fall, but the embryo develops at such slow rate that it measures only 3 mm. after 4 or 5 months; however, within another 8 weeks it is fully developed and ready for birth.
The den of the marten is usually in a hollow tree and is easily detected by its odor. The litter consists of 3 to 5 young.

EA-Zoo. Ekman: Eurasian Fur Bearers

The Sable
The sable ( Martes zibellina ), one of the best-known and most valuable fur bearers, is a close relative of the pine marten, except that the color of its fur is darker, almost black, especially on the back, while the light patch under– neath the throat is of a more rust-colored, sometimes of a reddish-orange tint, shading to a paler hue after the animal’s death. Its build is somewhat coarser than that of the marten, especially as regards the legs and feet. The colora– tion differs among the various subspecies. The finest furs are said to be those of the Transbaikal variety, the pelts of which are said to be “splendid, soft and silky, of a brilliant dark black brownish hue, and the underfur dark bluish gray with a brownish tint, the throat patch much reduced and commonly not visible” (Ognev). Largest in size are the Kamchatka race and the sables which inhabit the forest zone adjoining the open tundra. The pelt of the forest sable is less valuable than that of the mountain sable; it is coarser and the underfur is more yellowish in color.
A detailed description of the characteristics of the sable has recently been given by I. Kozhantchikov. Its main habitat is the Siberian taiga (coniferous forest zone) but its western limit is somewhat beyond the Ural Mountains, along the Pechora and Kama rivers, between latitudes 60° and 65° N. The southern limit of the species extends from the Altai Mountains eastward along the southern forested slopes of the border mountains between Siberia and Mongolia as far as southern Manchuria. The sable is also found in Kamchatka, on the island of Sak– halin, and on Hakkaido Island.
The numerical distribution of the sable varies in the mountain regions, and

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in the area between the Altai Mountains and Lake Baikal three different dis– tribution zones may be distinguished, in general, the sable is relatively rare or at least of spotty occurrence at altitudes of from 400 to 600 or 800 meters, where the European pine ( Pinus silvestris ), the common aspen ( Populus tremula ), and the larch ( Larix sibirica ) are characteristic trees and where the growing season is about 5-1/2 to 6 months. In contrast, it is numerous in altitudes of from 700 or 800 meters to 1,400 or 1,500 meters where spring starts in May or June, and where the snow cover in winter has a thickness of 1-1/2 meters; it is absent here only in the deciduous forests or in treeless areas. Charac– teristic trees here are the stone pine ( Pinus cembra ), the fir ( Abies sibirica ), and two members of the birch family, Betula pubescens and humilis . At still higher altitudes, between 1,300 or 1,400 meters and 1,600 or 1,700 meters, where stone pine and Siberian fir constitute the northernmost trees next to the mountain tundra zone, the sable occurs in only a few specimens. Spring here starts at the end of June. It may be said in general that the sable is most numerous in dense forests of stone pine and Siberian fir. It is thus truly a taiga animal. However, in all areas it is far less numerous than in former times.
The sable is not entirely carnivorous and its food varies according to the seasons. In the fall, bilberries, lingonberries ^ , ^ and the seeds of the stone pine are said to furnish the bulk of its diet, although small rodents and oc– casionally birds may also be taken. At that time, its favorite haunts are the pine forests where bilberries are especially abundant. Early in December, when the ground is covered with snow, it invades the fir forests and occasionally visits the deciduous forests where it feeds on sorb apples and hunts various

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birds which are more numerous than elsewhere. At this season, however, half of its food consists of mammals, foremost among them the pika ( Ochotona ). From the beginning of March, it feeds primarily on mammals, including the squirrel, and in the spring also on birds’ eggs. The impression is gained from reports of hunters and trappers that the sable is far less arboreal in habits than the marten.
As to the time of mating and the gestation period, sable and marten are alike. The litter consists of 3 to 5 young which are usually born in May.
The sable has long played an important part in Siberia’s fur industry, especially in the areas east of the Lena River where up to a few decades ago sables were still relatively numerous and constituted the main source of income of both natives and new settlers. Sables are caught either in traps or by means of snares and nets, or else hunted with dogs and a gun. At the beginning of the century the trade in sable furs totaled about 70,000 pelts annually.
The Ermine
Although small, the ermine ( Mustela erminea ) is valued for its winter pelt which is pure white except for its black-tipped tail, changing to brown only in the spring. However, the change in color depends on the climate. For instance, in southern Europe, south of the Alps, and in countries with a maritime climate, such as Ireland or southern England, the ermine does not change to white at all. In contrast subspecies inhabiting the Fur North are slow to change from white to brown and will keep their winter pelage the longer the more northerly their habitat. The under parts of the rump are sometimes tinted a light yellow or a yellowish green due to secretions from the anal glands.

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The ermine is very widely distributed, its range in Europe extending from the Polar Sea to the Mediterranean and in Asia as far as Kamchatka, Japan, and Tibet. Whether the North American and the Eurasian species constitute two distinct types or whether all are subspecies of one and the same circumpolar species has not been decided. The ermine is a very hardy animal and can tolerate many types of climate, ranging from those of the Temperate Zones to that of the Arctic. The animal is as common in the birch zone of the Scandinavian highlands, that is, in the tree zone bordering on the mountain tundra, as in the dense coniferous forests farther below. Its tracks are also found in the snow fields above the tree line, as high up as 1,000 meters above the nearest tree. Not only does it prowl in these barren areas for game, but the female will whelp here, especially in years when lemmings and voles are abundant. The same applies to the species that is found in the tundra of the Siberian North.
The ermine hunts birds and various small mammals, notably small rodents, easily tracking the latter to their very burrows thanks to its short legs and its long slender rump which is no wider than its narrow head. It also attacks larger prey, such as hares and ducks. On one occasion a capercaillie was seen flying through the air with an ermine fastened around its neck much like a white boa. The ermine attacks its prey in the neck or at the base of the skull and when its victim is a small rodent, its grip is so strong that the eyes are squeezed out. This form of attack is characteristic also of other members of the marten family, such as the marten and glutton, while members of the dog tribe (wolf, fox) attack their victims in the throat, provided their prey is not too small.
Recent experiments have shown that the ermine, like the marten and the sable, has two rutting seasons, one in summer and one in winter. In the case of the

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ermine, however, the winter or early spring rut seems the more important of the two. If summer mating (June/July) has taken place, gestation is practically at a standstill for the next six or seven months and proceeds more rapidly only during the two months preceding the birth of the litter, that is, subsequent to the period of the late winter rut. In case of winter-mating (usually March), the female whelps about two months later. The size of the litter depends on the availability of food. Usually there are from k 5 to 7 young, but in years when small rodents are abundant, the litter may be larger. On rare occasions, there have been up to 13 young, and up to 16 embryos have been found inside females.
Other varieties of the weasel family include the least weasel or mouse weasel ( Mustela nivalis ) which inhabits Europe and western Siberia and has a range here similar to that of the ermine. The variety is probably similar to the typical Mustela rixosa whose main range is farther to the eastward. Because of its small size, it has little commercial value.
The Glutton or Wolverine
The wolverine ( Gulo gulo ), although not a tundra inhabitant exclusively, is a more strictly northern species than any of the other mentioned fur bearers. In northern Scandinavia, it inhabits the treeless highland tundra, the birch zone, and the coniferous forests closest to the tree limit and visits the forests farther south only in winter and during its various marauding expeditions. Its range, in general, is limited to the cold temperate zone. In Siberia, too, it is more frequently found in the mountain tundra than in the forest zones. Its range here extends from the Arctic southward to the Altai Mountains and in– cludes the tundra proper, where, according to Middendorff, it habitually robs

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fox traps. The North American wolverine is distributed in similar manner. The latter is sometimes considered a separate species, but might better be called a subspecies of the Old World variety ( G. gulo luscus ).
The glutton is the largest of the marten-type animals. The thick, long– haired pelt of the adult is a deep black-brown while that of the young is charac– terized by hands of light brown running along each side of the rump. During the summer it feeds mainly on small rodents, sometimes adding berries to its diet, although to a far lesser extent than the marten and sable. In winter, when the pursuit of rodents is rendered difficult by the snow, it goes prowling for larger prey and becomes a danger to reindeer calves in particular, and, in the mountains of northern Asia, to musk deer. While stalking a reindeer, the glutton usually waits until its victim starts digging for food in the snow, attacking it in the neck the very moment its head is bent to the ground. Reindeer calves are pur– sued through deep snow until they break down, as the glutton is one of the most persistent of all predators. Part of its kill is often cached for future use. It may divide a reindeer in two parts and on several occasions has been seen to make off with the head and antlers, to hide them high up in a tree. Incredibly powerful for its size, it may drag an entire sheep to a distant hiding place.
The glutton is equipped with anal glands which secrete a sharp-smelling fluid and in times of danger serve as an effective means of defense, even against large-sized attackers such as dogs, wolves, or bears. Eskimos from the [: ] Hudson Bay area maintain that, when a wolverine approaches, wolves will abandon their prey. Instances are known where a wolverine also drove off a bear and, in one case, two pumas in the Sierra Nevada of California. These stories seem all

Related to the American skunk? or to the marten family, see below.

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the more credible in view of the fact that the glutton, or wolverine, is re– lated to the North American skunk which uses its anal glands in similar effec– tive manner.
The glutton is an excellent climber of trees and when pursued by dogs often escapes into a treetop. It is also a good swimmer. Its gait is peculiar in that the animal never trots but either walks or gallops, a characteristic it shares with the rest of the marten tribe despite its longer legs. It is capable of wide leaps and may go bouncing along for days on end. When in a playful mood it cuts numerous capers or slides down a snow slope on its back, a trick it may repeat over and over again.
Despite the arctic or subarctic climate of its habitat, the female [: ] whelps as early as March or April, rarely before or after. Even in eastern Siberia, where spring sets in at a later date, the young are usually born in April. Their home is sometimes in a mountain cave or the like, but more often in a lengthy burrow dug into the snow. The litter consists of from 2 to 4 young which are unusually small and of a light gray or whitish color, very dissimilar from their parents. However, the smallness of their size does not necessarily point to a short gestation period (about 2 months) as was formerly assumed. There seems evidence now that the glutton, like other members of the marten family, mates in summer as well as in winter and that the period of gravidity may last from 7 to 10 months. The question whether or not this is due to delayed im– plantation is still open to debate. That gestation is delayed seems probable but has not been fully confirmed up to now.

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The Otter
The otter ( Lutra lutra ), too, is a member of the marten family, although it belongs to a different subfamily than those mentioned before. As with most aquatic [: ] mammals, the fur consists of two layers: a dense inner one of short wool which does not get wet and an outer coat of stiff glossy guard hairs. The otter is found in most parts of Europe, likewise in Asia except in certain parts of the south, and in northwestern Africa. To the northward, its range extends rarely beyond the tree limit, except along various Siberian rivers where fish are plentiful and where the otter may settle and breed.
When swimming, the otter propels itself forward with its tail and to a lesser extent with its hind legs, while the forelegs contribute little to its foreward motions, although they are sometimes used for steering purposes or may also serve as a brake. The base of the tail is generally thicker than that of the average mammal, very muscular, and flattened, not laterally like that of a fish, but [: ] dorso-ventrally. This corresponds to the swimming motions of the animals, for the tail moves upward and downward and not sideways like a fish. The hind legs, meanwhile, move in paddle-like fashion, backward and upward, and not rudder-like, to the right and left, as do those of the seal which are kept closely together. The ease with which the otter catches even such [: ] fast and strong fish as pike and salmon is nevertheless surprising and may largely be due to the animal’s endurance. A mammal can store far more oxygen in the haemoglobin of its red corpuscles and in the red myoglobin of its muscles than can the fish with its relative lesser blood content and its pale flesh — a fact of decisive importance where endurance is concerned.

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Besides fish, which constitute its main food, [: ] the otter also eats frogs, crabs, and even small rodents and the like.
The young are usually born in May or June, but there may be litters in any month and even in the northernmost parts of the otter’s range litters have occurred in December or January or else fully grown foetuses have been found inside females.
The Wolf
The original range of the wolf ( Canis lupus ), before it was limited by man, covered all of Europe including Great Britain, Ireland, and Sicily; all of northern and central Asia down to southern Arabia; northern India; Afghanistan; the western Himalayas; northern China, and northern Japan. The distribution area also in– cluded almost all of North America, from the Arctic down to Texas and Florida. Its range is therefore circumpolar. The wolf does not occur in Spitsbergen. Be– cause of variations in coloration, wolves of the various regions are often divided into subspecies and listed under different names ( lycaon , nubilus , etc.), but the majority of the races overlap and are too closely related to be considered inde– pendent species. Possibly an exception is the polar wolf, which is almost circum– polar in distribution on the barren grounds, and which is represented by a variety of subspecies from northern Greenland ( Canis lupus eogroenlandicus ), the arctic islands in Canada ( C. lupus arctos, bernardi , manningi , etc.), and from the northern part of mainland North America and eastern Siberia ( C. lupus tundrarum , mackenzii ). Although its skull is said to have some of the characteris– tics of the common wolf, it is definitely distinguished from the more typical varieties by its coloration which remains white throughout the seasons, except

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for a few markings of mixed gray and black hair. Hybridization forms are not known.
The common wolf, too, is a tundra inhabitant. However, it seems comfortable in all types of climate, and it is equally at home in the barren wastes of the desert and the steppe. In Great Britain it has long since been exterminated, and in central Europe it is now found only in the mountain regions of France and in the highlands of northern Scandinavia. In southern and central Sweden it has been extinct since the middl ^ e ^ of the nineteenth century. The Swedish variety, which is the variety referred to below unless otherwise indicated, is confined now to the northernmost highland areas. It is rare even here and would have dis– appeared altogether had its stock not been replenished repeatedly by [: ] arrivals from the east. For the past hundred years its young here have usually been born in dens located in the birch zone or in the lower parts of the mountain tundra zone and relatively seldom in the forest zone. Wolves often dig their own dens, usually in sandy soil, or else they make use of fox dens which they enlarge.
The wolf is strictly carnivorous, but its game varies with the seasons. In summer, its preferred food is small rodents, especially when the latter are abundant, but its prey also includes hares, reindeer calves, foxes, birds, etc., and even large-sized insects. In winter, when small rodents remain hidden under the snow, its chief game is the reindeer. At this season it usually hunts in families of from five to seven members, sometimes also in larger groups. The tactics of the group consists in a furious attack on a poorly watched herd in order to scatter it or at least to detach some of its members which are then singled out for further pursuit. At times such an attack results in a full-scale stampede and it has happened that several hundred reindeer were chased into an

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abyss where all of them perished. The wolves, however, rarely follow the domesticated reindeer far into the forest zone and, when winter comes, remain in the highlands to hunt the few specimens unintentionally left behind by the Lapps.
The wolf usually attacks the reindeer in the throat, or, if its victim at– tempts to escape, in the hind legs, so as to bring it down. When attacked by a lone wolf, a male reindeer or elk is sometimes able to defend itself and to ward off the killer with a few strokes of its powerful forelegs, but when faced by several wolves both reindeer and elk are usually helpless. Even bears will suc– cumb when attacked by several wolves. Three cases are known in Sweden where a bear was killed by packs numbering from 7 to 12 wolves. The North knows no greater scourge than a prowling group of famished wolves.
After killing a reindeer, the wolf first eats its tongue which it pulls out with the larynx. L. Munsterhjelm, in reporting on his hunting expeditions in northern Finland, tells of a case where 7 wolves killed 11 reindeer; only two carcasses were partly devoured, but the tongue of each reindeer had been pulled out and eaten.
The Eurasian Red Fox or Forest Fox
The red fox ( Vulpes vulpes ), much like the wolf, is at home in many climates, for its range includes North Africa, Mt. Sinai, Palestine, Arabia, and India as well as the areas of the North as far up as the tree ^ ^ line. It is common in the Sahara, in the interior of Arabia and other dry areas of southwest Asia, and in the vast steppes and deserts of Eurasia from southern Russian eastward to China.

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However, it is not commonly found in the tundra, at least not in the altogether treeless northernmost parts. The few specimens that have been observed, for instance, on Novaya Zemlya, are considered strays that have traveled up from the south. The species includes a number of geographic subspecies or races.
The habits of the common fox are too well known to necessitate a detailed description. In the following, therefore, only a few of its outstanding character traits are considered.
Like most predators, the fox often caches its food in the snow, under moss, or in the loose soil of a mole hill, etc., and there is sufficient proof to show that the animal is able to find it again at a later date. As a rule, the spoil consists of bird’s eggs which are carefully conveyed to a relatively distant hiding place and retrieved during the following winter. The eggs include those of capercaillies, ptarmigan, and sea divers. Such habits, although not exclusively those of the fox, seem to indicate not only a resourceful mind but a highly developed memory for places.
Equally interesting are the feeding methods of the female. When bringing food to the den for her young, she often carries an astonishing amount of small prey in her mouth and gullet. A female which was shot at the entrance of her den, was found to be holding 9 bird chicks in her mouth; another one carried 15 voles in her mouth; a third had returned with 3 young hares and 2 rats. On one occasion, a female was seen to arrange 12 dead rats in a row; she then picked them up in such manner that only the tails were still [: ] visible, six of them showing on either side of her mouth. The following even more astonishing inci– dent was observed in Norway where a dead female fox was found with a capercaillie

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in its mouth; wrapped inside the bird’s folded wings were 11 capercaillie ^ s ^ and black grouse chicks, together with a vole. Another had arrived at the den with a piece of meat, a rat, and a frog carefully wrapped in a folded lambskin. A third carried a large parcel of heather, dwarf birch and reindeer moss, tied to– gether with the long fine roots of the dwarf birch. When opened, the parcel was found to contain 26 voles and 2 small birds. The three cases seem to indi– cate that the fox is able to make use of tools, a faculty usually attributed only to two other (non-human) mammals, the monkey and the elephant.
The Arctic Fox
The arctic fox ( Alopex lagopus ) is definitely a tundra or mountain tundra denizen, far more so, at any rate, than any of the [: ] mammals described above. In Sweden, where a close study has been made of its habits and range, it is rarely found below the tree limit, except in winter, when it visits the nearby forests in search of game; usually it even avoids areas where a few birches may still be found. It is common in the lower parts of the mountain tundra zone, less common in the upper parts. In Asia, too, it ranges through the treeless zones. It is found here as far north as the shores of the Arctic Sea and withstands even more extreme climatic conditions than those encountered in the Scandinavian highlands. It is common also on the islands scattered over the northern seas, such as Bear Island, Jan Mayen, Franz Josef Land, Novaya Zemlya, etc., where it is more numerous than other land mammals. Its distribution in general is circumpolar.
The arctic fox exists in two varieties: one whose coat turns white in winter and another which assumes a blue-gray hue, shading faintly into brown.

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During the summer both are blue-gray or brown. The ability of the blue foxes to retain their blue-gray color in winter is determined by heredity, but is individual only, and it is possible for the same litter to contain both blue and white varieties. The “blues” are commercially more valuable than the “whites. The percentage of the blue foxes differs in the various regions. It is highest in certain coastal regions, for instance, in Iceland, where more than 50% of the foxes are blue. In other areas the proportionate number of blue foxes may be relatively small.
The arctic fox has two outstanding traits: curiosity and fearlessness. In areas where it is not generally pursued, it is easy to approach, and an observer stationed close to a den may see alternately one of the parents or one of the young peer out of the entrance. The food varies according to regions. In arctic coastal areas the fox feeds largely on invertebrates that have been swept ashore, while its diet in inland areas consists mainly of small rodents, ptarmigan, and the remains of spoil left by other predators. Foxes are usually more numerous following years when lemmings were abundant, as a rich diet results in larger litters. Although small, the fox may prove a menace also to larger animals, especially reindeer calves, as young cows which have calved for the first time cannot always defend their young. Usually, however, the fox is satisfied to feed on si x ^ c ^ k or dying calves found by the wayside, for the mortality among reindeer young is especially great when weather conditions are unfavorable during the breeding season. Like the red fox, the arctic fox also takes berries, such as blueberries and dwarf blackberries. Its voice is a hoarse bark, similar to that of a small dog.

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The female is said sometimes to whelp in a rocky cavern or under blocks of stone. Usually, however, it digs a burrow in sandy soil with the assistance of the male. The burrows have numerous entrances with the excavated soil form– ing slight elevations, from 10 to 35 meters long, above the gangways of the inner den. The largest of the burrows are inhabited by at least two families and may have from 50 to 100 entrances; on one occasion 172 entrances were counted. The “hills” are usualy visible from quite a distance as the foxes deposit their dung and the remains of [: ] spoil on their tops and a lush cover of grass is promoted by such constant fertilization. Up to about fifty years ago the highlands of northern Sweden abounded in such dens, of which H. Zetterberg has furnished a detailed description. However, the species is practically ex– tinct in Sweden nor due to persistent hunting, and the majority of the “hills” constitute nothing more than monuments of a vanished animal race. The dens of the arctic fox that are found in Spitsbergen and Novaya Zemlya are identical in type.
The Brown Bear
In Europe and Asia, the original range of the brown bear ( Ursus arctos ) comprised all forested regions from the timber line in the north to the Mediter– ranean, Lebanon Mountains, Persia, the Himalayas, southwest China, and Hokkaido Island of northern Japan in the south. Its former occurrence in northwest Africa is in doubt. Today its western limits are central Italy, the Carpathian Moun– tains, western Russia, and northern Scandinavia, with a wide gap isolating the Italian variety from the next eastward tribe in the Balkan mountains. In Norway bears are found only in five small isolated areas while the main range is in the

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highlands of northern Sweden where they occur in two regions, one located be– tween latitudes 61°30′ N., and the other between latitudes 64°30′ and 68° N. They are scarce even here, however, and inhabit only the coniferous forests and birch groves located at higher altitudes. Their range thus extends to the southern boundary of the mountain tundra zone, rarely beyond, although they may roam in the mountain tundra to feed on berries. The European bear is considered an independent species, as is the western Siberian one. A third nor– thern species inhabits Manchuria, the forests north of the Okhotsk Sea, and Kamchatka. Members of the last-named race are very tall and are closely re– lated to the North American grizzly bear. At the time of the Vega expedition Nordenskiöld observed brown bears on the northern shores of Chukotsk Peninsula. Most likely, the animals were attracted to the region by the presence here of the hoary marmot.
In view of the fact that extensive literature exists regarding the habits of the bear, only a few of its traits will be dealt with in this article. In winter, the bear of the Scandinavian highlands remains asleep in its den for a period of at least six months, the female and her young somewhat longer. A gravid female usually digs a den for herself and her future young on a sandy slope or underneath the root of a tree; she may also establish her den in a rocky cavern, under a fallen tree, or in a fir thicket. As a rule, the dens are in the forest zone, but cases are known where the bear, this genuine forest animal, has also hibernated in the mountain tundra zone. In northern Sweden, a large cave on a mountain top, 650 meters above the tree line, once served as a hibernation place to five or more bears at a time. A hunter who had killed 42 bears, secured 30 of them in this very cave or “bear hotel,”

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Opinions as to the mating habits of the bear have undergone certain re– visions in recent years. It was known for a long time that the young, usually two or three, are born in the den around the New Year or in January and that the cubs are very small at birth. On the basis of information furnished by hunters, it was assumed that mating took place in the late summer. However, according to observations made in animal parks or in the open, it is now certain that the sexes mate in May or June, which would mean that in the northern region, at least, fertilization takes place less than a month after the bears have left their dens. The period of gravidity is about 7-1/2 months which seems unusually long in view of the small size of the new-born cubs. Gestation may therefore be delayed as is the case with the marten.
The Arctic Hare
Although not a purely arctic and subarctic animal, the arctic hare ( Lepus timidus ) confines its range to the frigid and cold temperate zones. It is absent therefore in C ^ c ^ entral Europe but it inhabits the Pyrenees, the Alps, and the Carpathian Mountains, Scotland and Ireland, the Scandinavian Peninsula, and all of northern Asia as far east as the Bering Sea and northern Japan. The northern Siberian race tschuktorum is closely related to the Alaskan subspecies othus and may even be identical with it. The entire North American species Lepus arcticus is so closely related to the Eurasian Lepus timidus that the question may be raised whether or not both are geographic subspecies of a single circumpolar variety. Hares do not occur on Spitsbergen and Iceland. In Eurasia as well as in North America the species advances beyond and above the timber line (that is, its range includes both tundra and mountain tundra zones) and

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it is widely distributed even in the coldest parts of the Arctic. A seasonal change of pelage occurs due to molting, that is, the animals lose their brown summer coat and take on a white one in the fall. The change depends on the climate, [: ] however. Arctic specimens transplanted to a warmer climate, assume the coloration of their new environment.
Apart from its meat, the arctic hare is valued for its fur, although it is too thin to stand hard wear. Before World War I, two to three million white pelts were taken annually in Siberia, but the number has doubtless fallen off since then.
The Squirrel
The squirrel ( Sciurus vulgaris ) inhabits all of Europe and northern Asia, its range northward extending as far as the northern boundary of the coniferous forests and northeastward as far as longitude 180° E. (Bering Sea). About twenty different races have been distinguished, among them 16 in Russia and Siberia. The summer pelt of the northern Eurasian varieties is usually either reddish-brown or else darker with nearly black tail and ear tufts. The markings are individual, however, and not racial. Both types may occur in the same litter and hybrids are common. In central Siberia, the darker-hued type is said to occur primarily in forests where the Pinus cembra (stone pine) is the characteris– tic tree; the ligh ^ t ^ er-hued variety is more common in forests of Pinus silvestris (Scotch pine). In winter the reddish-brown coloration is replaced by a gray one while the under parts of the body remain white throughout the year. The purer the gray of the pelt, the higher is its commercial value. The finest are those of the northern Russian and Siberian variety which are a dark even gray with black

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tails and ear tufts. The light-gray squirrel does not turn a darker gray in winter and the tail and the turfs remain brown in the fall as these parts of the body are not affected by the molt.
The bulk of the squirrel’s diet consists of the seeds of conifers. Pine or fir cones are picked clear of their seeds in the following manner: The squirrel holds the cone in its forepaws with its top turned down and then tears off the scales until only the stem and uppermost scales are left. It is particularly fond of the fat seeds of the stone pine, but in the spring feeds also on the buds of trees. The ground underneath fir trees is often covered with sprouts gnawed off by squirrels. On one instance a squirrel was observed to drop 6 to 13 sprouts to the ground per minute.
Squirrels, too, have a habit of caching their food. When offered a nut or an almond in a park, the squirrel does not eat it on the spot but as a rule carried it off to buy it in a small hole which is then covered with a little moss or soil. Mushrooms or an apple (of which it eats only the seeds) are some– times fastened inside the crotch of a tree.
As its favorite abode is in trees, its nest, too, is built in tree tops, preferably in those of fir trees. Nests are fashioned of twigs, mosses, lichens, the fiber of bark, etc., and are usually built on two parallel branches close to the main trunk. In the northern regions, the gestation period averages 4 to 5 weeks. The litter consists of from 2 to 3 young which are born between March and May. As a rule there is only one litter a year, at least in the more northerly areas.
Squirrels have been hunted since time immemorial and they are still being

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eagerly pursued in many parts, especially in northern Russia and Siberia. At the beginning of the century the annual trade in squirrel pelts amounted to roughly 13 million pelts.
The Beaver
Beaver ( Castor fiber ) pelts were much in demand in former centuries, like– wise the secretions of the beaver’s anal glands, the so-called castoreum, which served as a remedy against various diseases. The demand became so great that the species was trapped or hunted to extinction within most parts of its original range. Its range in Europe today (roughly around [: ] 1930) includes the Rhone d ^ D ^ elta in southern France, possibly as far north as Avignon; the valley of the German river Elbe together with that of its tributaries between Wittenberg and Magdeburg: southern Norway where the European variety is still well repre– sented; and the vast swamp areas of western Russia and eastern Poland, along the tributa i ries of the Pripet (a confluent of the Dnieper) and the Niemen. In Siberia, where the beaver was common once as far east as the Lena b ^ B ^ asin its range is now said to be limited to the eastern slopes of the northern Ural Mountains and the Mongolian part of the Altai Mountains. In Sweden, it was exterminated during the nineteenth century, while the Norwegian beaver has been restocked since 1922 and is on the increase. In the highlands of northern Sweden beavers were occasionally found as high up as the birch zone, above the coniferous forest zone. Extermination of the species has doubtless been facilitated by the ease with which its houses and damen ^ dams ^ and other signs of its activities are spotted.
Today its trade value is negligible. Its one-time importance in international trade can be judged by the fact that in the sixteenth century thousands of beaver

EA-Zoo. Ekman: Eurasian Fur Bearers

pelts were annually shipped from Stockholm and that the pelts were divided into as many as seven categories according to quality.
Other Eurasian Fur Bearers
Among the northern Eurasian fur bearers three additional species deserve consideration here: the so-called Burunduk , i.e., the Asiatic chipmunk; the hoary marmot; and the Eurasian flying squirrel.
The Asiatic Chipmunk ( Eutamias asiaticus ) is from 15 to 16 cm. long; the tail has a length of about 10 cm. The pelage is marked by five horizontal stripes. The species occurs in northeastern Russia and throughout Siberia as far as the northern boundary of Kamchatka. The furs rank as export goods despite the small size of the pelts.
The Hoary Marmot ( Marmota caligata ) reaches a total length of 50 cm. or more. There are several races which are found in northwestern North America and in northeastern Asia. In northeastern Siberia its range extends to the shores of the Arctic Sea.
The Eurasian Flying Squirrel ( Pteromys volans , syn. Sciuropterus russicus ) is much smaller than the common squirrel but its pelt is very beautiful, a soft light gray in winter and tinted a yellow-brown in summer. Its range extends from the forests of northern Finland to the northern end of the Kamchatka Peninsula.
Other rodents of the Eurasian North, although not part of the category far bearers, include the Asiatic pika ( Ochotona hyperborea ) which inhabits the high mountains of northeastern Asia, Kamchatka, and the Chukotsk Peninsula, and some twenty varieties of lemmings and voles of which at least half are tundra inhabitants or include the tundra zone in their range.

EA-Zoo. Ekman: Eurasian Fur Bearers

Mention must also be made of two other species of mammals which, like the reindeer, are tundra inhabitants although their range is limited to definite localities. They are the musk deer ( Moschus moschiferus ) which follows the mountain ranges almost to the shores of the Arctic Sea, and the mountain sheep ( Ovis nivicola borealis ^ Ovis nivicola borealis ^ ) which persists as a relict species in the Verkhoyansk Mountains east of the Lena River and to the northwestward in the Byranga Range of Taimyr Peninsula north of 72° N.
Also to be included among the northern Eurasian animals are the reindeer and the polar bear which, however, are dealt with elsewhere in the Encyclopedia Arctica .
Sven Ekman

Wolf

EA-Zoology (A.W.F. Banfield)

WOLF

Throughout vast arctic regions of Asia and America the wolf ( Canis lupus ) dominates the natural scheme. It holds the apical position in the pyramid of numbers and natural food chains. Vegetation is transformed, through such intermediary stages as mice, ground squirrels, hares, and caribou, into wolves.
Considering its wide range, the wolf must be recognized as an efficient and successful organism. In the Northern Hemisphere it is circumpolar in distribution. It extends south in the Old World to central Europe and southern Asia, and in North America it extends south to central Mexico.
The wolf is a member of the family of carnivores Canidae (dogs). The species forms a compact group of closely related geographical races across ^^ the ^ its ^ entire range but there is intergradations along the margins of the various races. There is a close resemblance between the wolves of Alaska and those of eastern Siberia, suggesting the close relationship of these races and the probable exchange of individuals.
Goldman (11) recognized 23 subspecies of wolves in North America. Of these, eleven are recognized from the territory covered by this Encyclopedia. These subspecies and their ranges are outlined below: ^ G. N. F. B. Could you supply us with scimitar list for Old [: Nored] species? ^
  • 1. Canis lupus tundrarum - northern Alaska and Yukon.
  • 2. Canis lupus pambasileus - southern Alaska and Yukon.

    EA-Zoo. Banfield: Wolf

  • 2. Canis lupus mackenzii - northern Mackenzie District, Northwest Territories.
  • 3. ^^ Canis lupus occidentalis ^ Canis lupus occidentalis ^ - southern Mackenzie district, Northwest Territories.
  • 4. Canis lupus hudsonicus - Keewatin District, Northwest Territories.
  • 5. Canis lupus lycaon - southern Hudson Bay and James Bay areas.
  • 6. Canis lupus labradorius - Ungava Peninsula.
  • 7. Canis lupus manningi - Baffin Island.
  • 8. Canis lupus bernardi - Banks and Victoria Islands.
  • 9. Canis lupus arctos - High-Arctic Islands.
  • 10. Canis lupus orion - Greenland.
The larger subspecies are those occurring in Alaska, Yukon, and the Mackenzie District. Those inhabiting the Canadian Arctic Islands are smaller in size. Two of the largest wolves taken by the United States Fish and Wildlife Service, reported by Young (11), were: one at the Savage River drainage, Alaska, January 23, 1934, weighed 157 pounds, and one taken near the Seventymile River, Alaska, on July 12, 1939, weighed 175 pounds. Radforth reported a male killed on Shultz Lake, Northwest Territories, on January 4, 1912, which measured 67.7 inches in length and weighed 101 pounds. Another specimen reported by Young, taken at Mount Hayes, Alaska, on October 21, 1939, weighed 155 pounds and measured 69 inches in length and 38 inches at the shoulder. A group of wolves from Greenland averaged about 42 inches in length and weighed 45 to 63 pounds. Males are generally larger than females.
Color among wolves is of little scientific importance for racial descriptions. Wolves show great individual variation, and may be any color from pure white to jet black. They vary through every gradation of cream, gray, and rufous to dusky and black; gray is the most common color. In the arctic regions white

EA-Zoo. Banfield: Wolf

wolves are more common. The annual molt is variable in duration but occurs during the summer months.
The dental formula is: incisors <formula>(3-3)/(3-3)</formula>, canines <formula>(1-1)/(1-1)</formula>, ^premolars <formula>(4-4)/(4-4)</formula>,^ molars <formula>molars (2-2)/(3-3)</formula> = <formula>42.</formula> The ^^ carnassials teeth, used for tearing meat, are well developed. The wolf bolts its food rather than chews it.
The wolf is of rangy appearance, with short ^ stout ^ , long legs. The paws are ^^ large compared to those of a dog of equal proportions. Wolves run with a bound– ing gait that often appears clumsy; but they have been observed to run with a speed of 28 miles per hour for a distance of 200 yards. On a long run of a mile or two, the speed is reduced to about 22 to 24 miles per hour. The wolf seldom relies on speed alon g ^ e ^ to capture its prey, placing more reliance on ^^ surprise or, if necessary, endurance. The endurance of the wolf is legendary. MacFarlane (4) records a wolf which escaped with a steel trap at Ile a ^ à ^ la ^^ Crosses ^ , ^ Saskatchewan. It was killed at Green Lake a month later (approximately 100 miles south) with the trap and toggle attached to the hind leg.
The normal home range of a wolf covers a large territory. During hunting forays, they usually travel along well-marked game trails or geographical paths such as river bottoms or shore lines. Wolves have been known to use regular circuits of over 100 miles, the circuits being usually an irregular circle. During the winter months wolves may travel even greater distances, following migrating herds of caribou.
Wolves are good swimmers and do not shun taking to the water when their path lies across it. During the evening of August 3, 1948, the author watched a large black wolf following a well-worn caribou trail on the shores of Lake Clinton-Colden, Mackenzie District. When it reached the western shore s of Caribou Narrows, it entered the water without hesitation and swam rapidly

EA-Zoo. Banfield: Wolf

across the narrows, a distance of one-third of a mile.
The wolf can emit a variety of sounds which convey a wide range of ex– pression. Some are quite doglike such as the whines heard about the den, the quick bark on being disturbed or surprised, or the yelps of the chase. How– ever, the wolf cry most often heard — a long, deep guttural, quavering howl — is the wolf’s singular expression of his kind.
Wolves do not breed until they are two or three years old. Murie (6) mentions a captive Alaskan bitch which came into heat during her second year. The period lasted two weeks and she mated during the second week. She whelped on May 15 after a gestation period of 60 to 63 days. A male wolf, kept captive for two years at Prince Albert National Park, Saskatchewan, during 1945 and 1946, failed to mate with a female ^ dog ^ during its first two years of life.
The whelps are generally born during May or June in the Arctic. The litter may include up to fourteen in number; five to seven seems to be the average litter. In the Arctic, where the white color phase predominates, the young at birth are dull slate in color. They are born blind but their eyes are open after approximately one week. They nurse for six to eight weeks. By September they are learning to hunt for themselves and are about half grown.
The dens are usually situated in the center of the home territory. From the den the runways or circuits radiate. The den sites are usually chosen in dry sandy soil, on a promontory, to give a good view and to be near permanent water. North of the tree line, the sand eskers are favorite localities for ^^ dens. The den ^ s ^ may be dug by the wolves themselves, but more likely they are usurped and enlarged fox dens, beaver lodges, or natural crevasses in rockslides. ^^ The y ^ re ^ are generally several dens nearby to which the whelps may be moved as the season progresses. If the den is a remodeled fox den, there will probably be

EA-Zoo. Banfield: Wolf

several entrances which the growing whelps can use. The burrows may be up to thirty feet in length and lead to a chamber in which the young are born. This chamber is usually only two to five feet below the surface. A nearby knoll is used as a lookout from which the wolf can keep a careful watch over the sur– rounding country. There will also be depressions about the den month, used as beds by the adults.
The dens are usually occupied in April or May when the ground is still snow-covered. The same dens are generally used in successive years. The wolves in attendance are not only the mated pair but usually a small group which lives in harmony. These wolves are believed to be related, possibly the parents and the previous year’s litter l . All the members of the group assist in bringing food for the bitch and whelps. The dens are abandoned in September or October when the young wolves are being taught to hunt.
It is a common though disputed belief that wolves gather in packs for ^ AU: Ok? Ok FB ^ hunting purposes, especially in winter. These are thought to be temporary aggregations of families. The size of the pack would be limited by the law ^^ of d o ^ i ^ minishing returns.
In summer wolves appear to be evenly spread over the arctic plains. Some believe that pack territories are occupied. Clarke (2), from personal observa– tions and interviews with northern residents, estimated a population of about six wolves per hundred square miles. Considering the 600,000 square miles of caribou range in the northern Canadian mainland, this gave an estimated population of 36,000 wolves.
The population of wolves, like that of many other animals, fluctuates. During the past two decades (1930-49), there has been a high wolf population throughout northern North America. Periods of wolf scarcity have been noted

EA-Zoo. Banfield: Wolf

also. In his investigation of Mount McKinley National Park, Murie (6) stated that, according to information gathered, wolves were fairly abundant in 1880 and probably from 1900 to 1908. The population then declined and wolves were generally scarce from 1916 to 1925. Since 1927 they have been on the increase in the area.
The wolf has no predators except man. The grizzly bear ( Ursus sp.) and wolverine ( Gulo luscus ) are its most important competitors. Predation cannot be counted as a factor controlling the wolf population in arctic areas. It is generally believed that disease exercises an important role in controlling the number of wolves. Buffalo Jones believed firmly that the numbers of wolves are kept in check by disease. MacFarlane (4) given March 1868, as the date of a great epidemic among wolves which greatly reduced their numbers. Plummer (8) recently reported the identification of rabies in a wolf specimen from Baker Lake, Keewatin District. Other rabies identification has been made from ^^ specimen t s from Old Crow, Yukon Territory. Cowan (3) has reported the occurrence of sarcoptic mange in Rocky Mountain wolves. These diseases may exert a powerful influence on the numbers of wolves and foxes in the Arctic. [A recent view is ^ Ok FB? ^ ^^ that of trichinosis may account both for some direct fatalities and for deaths resulting from starvation brought on by inefficiency in hunting due to illness from this infection.]
The caribou is to the arctic wolf what the buffalo was to the plains wolf. During the summer months, when the wolf families are tied to the den locality by the whelps, caribou do not regularly form the major part of their food, for during this period the caribou herds migrate out of the territory of numerous wolves. Observations made at a den in southern Keewatin District indicated that small game was utilized under these conditions. Wolves were observed to

EA-Zoo. Banfield: Wolf

be capturing fish, voles, lemmings, ptarmigan, nesting waterfowl, and other birds.
During winter months wolves follow the migratory caribou herds. Clark ^ e ^ (2) ^^ has estimated, based on observations in the Thelon Game Sanctuary, that a wolf kills an average of twelve caribou per year. Wolves often follow trappers and rob trap lines. They may catch some arctic foxes under natural conditions. Beavers are favorite food item where available.
^^ Murie (6) examined 1,174 f a eces from the Mount McKinley area of Alaska and found that the remains consisted of 43% caribou ( Rangifer arcticus ), 25% Dall sheep ( Ovis dalli ), and 32% small game.
When trailing wolves, one frequently discovers that they are carrying meat or bones in their mouths. Young (11) reports that during late winter, excessive numbers of caribou and sheep are often killed in Alaska and the car– cases cached in the snow for food supply during the denning season. A den visited in May, when the snow was four feet deep, was surrounded by the remains of about four Dall sheep which were cached in the snow.
If food is plentiful, usually no special effort is made to cache the kill. On August 20, 1948, a wolf was seen to kill two calf caribou at Lake Clinton– Colden. One carcass was [: ] almost completely eaten, the other had only the tongue removed before the wolf departed. The following day the wolf returned to the carcass but departed when it sensed the presence of human beings. Although wolves usually make their own kills, they sometimes eat carrion.
There has been considerable controversy concerning the physical condition of the mountain sheep and caribou killed by wolves. Murie (6) examined the skulls of 221 Dall sheep which had perished during the previous two years from wolf predation and other lethal factors. He found that: 69% were in the

EA-Zoo. Banfield: Wolf ^ These are the figures given by Murie on pg 112. I suppose he left out decimals which would have raised to 100% ^

old-age class, 9% of prime age but diseased (actinomycosis), 4% prime, 3% ^ Au: should these add up to 100%? ^ diseased yearlings, 10% healthy yearlings, and 3% lambs. A further examination of 608 older skulls produced a similar grouping. From these observations it is indicated that despite the occasional observation of healthy animals succumbing, generally the older, weaker, and diseased stock are taken by wolf predation, these being the most easily captured.
Observations of hunting wolves by Murie and others have confirmed this fact. The wolves pursue caribou bands and finally ^ concentrate s on ^ the animal which drops behind, usually ^ Ok FB. ^ for one of two reasons: clumsiness due to age, or (in the case of bulls in season) because of the weight of the antlers.
Wildlife scientists ^ commonly ^ hold the opinion that a certain amount of predation is beneficial to a game species. The predators help to maintain a healthy stock by removing the aged, sick, and less wary individuals of the population. So Cowan (3) compared populations of elk ( Cervus canadensis ), moose ( Alces americana ), mountain sheep ( Ovis canadensis ), and mule deer ( Odocoileus hemionus ) in two areas of the Canadian Rocky Mountains. One area had a wolf population; no wolves were present in the second area. He found that there was no significant difference in the proportion of young animals in the populations of big game in the two areas. He concluded that factors such as available food, weather, and competition other than predation were critical in determining the annual increment.
Wolves have an insatiable curiosity and often trail hunters and trappers at a discreet distance to rob their traps, or pick up camp scraps. From many historical reports it seems the Old World wolves are more prone to attack humans than those of North America. However, occasionally, on the North American con– tinent, wolves have made unprovoked attacks upon man; such an attack was recently

EA-Zoo. Banfield: [: ] Wolf

reported by Peterson (7). The possibility of the wolf being rapid should always be considered in these cases. A recent attack upon a game warden and his dog team in the Mackenzie Delta during early 1948 was by a wolf presumably rapid, since the bitten dog died from a light would about two weeks later. Dogs belonging to Eskimos are frequently attacked and infected by rabies in the Canadian Arctic.
Since time immemorial the hand of man has been against the wolf. Some of the methods used to kill wolves by the Eskimos are very interesting. Richardson (9) reported the Eskimo use of an ice box trap, constructed of ice blocks with a heavy portcullis of ice which fell when the trigger, which was a peg baited with meat, was pulled out of position. One of the most unique devices is described by Mason (5) as a “piercer.” A small piece of whalebone is bent double to the appropriate size for wolves to swallow; the tension is held by a sinew; and the piece is wrapped with fat and blubber. After freezing the sinew may be cut, though this is not necessary. The baits are spread over the ground and, when swallowed, the fat and sinew are digested and the whalebone springs open and pierces the digestive tract. Another method is to encase a sharpened knife blade in frozen fat and stick the handle in a block of ice. The wolf, licking the blade, cuts its tongue; tasting blood, it becomes frenzied and inflicts greater injury. [Some authors consider this last to be folklore.]
Over large areas of the wolf’s former range, the increasing demands of civilization have necessitated the extermination of this predator because of its destruction to domestic stock. In the sparsely inhabited arctic regions, there are few areas where the investment in domestic stock is large enough to warrant the expenditure necessary to reduce materially the wolf population. In these arctic regions the wolf will probably remain a spectacular member of the native fauna for many years to come.

EA-Zoo. Banfield: Wolf

BIBLIOGRAPHY

1. Anderson, R.M. “Report on the natural history collections of the expedition,” Stefansson, Vilhjalmur. My Life with the Eskimo . N.Y. Macmillan, 1913, pp. 516-17.

2. Clarke, C.H.D. A Biological Investigation of the Thelon Game Sanctuary . Ottawa, Patenaude, 1940. Nat.Mus.Can. Bull . no.96.

3. Cowan, I.McT. “The timber wolf in the Rocky Mountain National Parks of Canada,” Canad.J.Res ., Sect.D, vol.25, no.5, pp.139-74, 1947.

4. MacFarlane, R.R. “Notes on mammals collected and observed in the northern Mackenzie River district, Northwest Territories of Canada,” U.S. Nat.Mus. Proc . vol.28, pp.673-764, 1905. (No.1405)

5. Mason, O.T. “Traps of the American Indians — a study in psychology and invention,” Smithsonian Inst. Annual Report for the Year Ending June 30, 1901 . Wash., G.P.O., 1902, pp.461-73.

6. Murie, Adolph. The Wolves of Mount McKinley . Wash., G.P.O., 1944. U.S. National Park Service. Fauna of the National Parks of the United States. Fauna series no.5.

7. Peterson, R.L. “A record of a timber wolf attacking a man,” J.Manual . vol.28, no.3, pp.294-95, 1947.

8. Plummer, P.J.G. “Further note on arctic dog disease and its relation to rabies,” Canad.J.Comp.Med . vol.11, pp.330-34, 1947.

9. Richardson, John. Fauna Boreali-Americana. Part 1. The Quadrupeds . London, Murray, 1829.

10. Seton, E.T. Lives of Game Animals . N.Y., Doubleday, 1925, Vol.1, pp.251-352.

11. Young, S.P., and Goldman, E.A. The Wolves of North America . Wash., American Wildlife Inst., 1944.

A. W. F. Banfield

Wolverine

(EA-Zoo. A. W. F. Banfield)

WOLVERINE

CONTENTS

Scroll Table to show more columns

Page
Physical Characteristics 1
Habitat 2
Economic Value 3
Habits 3
Bibliography 8

EA-Zoology (A. W. F. Banfield)

WOLVERINE
Probably no other northern mammal plays such an important role in camp– fire tales and folklore as does the wolverine ( Gulo luscus ). With few other mammals is there such a paucity of factual information of life history and eco [: ] logy.
Physical Characteristics . The wolverine or carcajou, as it is known to French Canadians, is the largest member of the weasel family (Mustelidae) of carnivorous mammals. It is a medium-sized mammal with a s ^ t ^ out badger-like body, a short, bushy tail, and large bearlike paws, each with five toes. The claws are partly retractile and are suitable for climbing; the soles of the feet are hairy. The ears are short and rounded. The dental formula (I = in– cisors, C = canine, P = premolar, M = molar) is: <formula>I (3-3)/(3-3), C (1-1)/(1-1), P (4-4)/(4-4), M (1-1)/(2-2) = 38</formula>
The following average measurements were reported by Seton (7): average of four mature males, length 41 5/8 inches, tail 8 1/2 inches, hind feet 7 7/8 inches. The females average smaller in size: average length 37 inches, tail 7 1/8 inches, hind feet 7 inches.
The weights of three adult males, as reported to Seton by Dr. R. M. Ander– son, were 35, 33, and 30 pounds; two females weighed 22 and 23 pounds each.

EA-Zoo. Banfield: Wolverine

Wolverines do not show great variation in color. The general color is a rich, dark brown, paler on the head and face. A band of light chestnut begins on each shoulder and passes back along the flanks to meet its fellow on the rump. On the flanks these bands are pale buff or creamy white. On the throat are irregular yellow splotches.
Habitat . Like many other northern animals, the wolverine is circumpolar in distribution. Its primitive range covered boreal America, Europe ^ , ^ and Asia. Over much of this vast area it has in recent years been greatly reduced in numbers in consequence of its value as a fur bearer. In North America it still occurs in Labrador and the Ungava Peninsula of Quebec and a few in– dividuals may yet exist in extreme northwestern Ontario and in northern Man– itoba and Saskatchewan. It is more numerous in the forested parts of the Northwest Territories, in Yukon Territory, and in central and northern Alaska. It has been recorded from several of the Canadian Arctic Islands, including ^ Ok FB ^ Ellesmere, Melville, Baffin, King William, and Victoria Islands. The wolverine also occurs frequently in the Canadian Rocky Mountains, south to the United States.
Two races of wolverine have been described for the area included in this Encyclopedia, namely, Gulo luscus luscus from the northern Canadian mainland and Gulo luscus hylaeus from Mount McKinley, Alaska.
The wolverine formerly inhabited the boreal forests of America . ^ in some numbers. ^ Today ^ ? ^ a few remain in the forests but a ^ the ^ greater population is found in Canada between tree line and the Arctic Sea. The animal is nowhere abundant and although one may be aware of its presence, it is possible to travel for many years without catching sight of it.
The wolverine seems to be adaptable in the choice of habitat. It

EA-Zoo. Banfield: Wolverine

occurs from the high alpine slopes above tree line of the Rocky Mountains to the sea coast and from dense coniferous forests and rocky shrub-clad hillsides to open, rolling arctic grasslands.
Little is known of the home range of the wolverine, but, judging from the animal’s habit of trailing trappers over long trap lines, it is probably a large area. This is in keeping with the wolverine’s carnivorous food habits. MacFarlane (4) writes of one trailing him for two days when he was traveling forty miles a day.
Economic Value . Throughout the arctic and subarctic regions, the fur of the wolverine is considered valuable. It is cut into strips and used as trim on hoods, moccasins, and mitts. It is a common belief of whites that in cold weather the moisture in the breath does not readily freeze on this ^ Au: OK? Ok FB ^ fur, when used as a trim for hoods; but anthropologists have pointed out that when whites first came it was chiefly the Eskimo women who used hoods trimmed with wolverine or wolf, the implication being that these fringes were used as Europeans use lace, chiefly as ornament. There is no doubt, however, that hoarfrost clings more readily to many other furs than it does to wolverine. Hardy (2) investigated this characteristic and compared the fur with that of coyote and wolf fur. It was found that the guard hairs of wolverine fur are straight and project an inch or more beyond the underfur, which is thick and even in length. This combination of factors results in the guard hairs being much less readily frosted than are other furs under the same conditions.
Habits . Wolverines are believed to mate during the last part of March. Hearne (3) writes that they make their dens in caves or clefts in rocks. Mac– Farlane (4) states that a discarded beaver lodge is sometimes used. Seton (7) recorded an Indian’s description of two beds, discovered near Lake Athabaska.

EA-Zoo. Banfield: Wolverine

They consisted of hollows in the ground under hanging spruce boughs and were lined with dry grass. The young were said to be born in these beds. The young are thought to be born in June after a gestation period of approximately 90 days. The litter may consist of two to five young. Two specimens in the American Museum of Natural History that were taken on May 1, 1901, in Siberia, were examined by Seton. They were three to four weeks old and were covered with dense, creamy, woolly fur. The color on the legs, back, and tail was grayish and the facial mask was brown. The females are brave in the defense of their young. Seton (7) reports that the Indian who discovered the beds previously described also stated that in each case the mother, with eyes blazing and teeth bared, attached the intruder. The young are suckled for eight or nine weeks. During the summer they remain in the den or bed and receive food brought by the mother. Late in the summer they follow their mothers and are taught to hunt. They continue to hunt with the dam all of their first winter. By the following spring they are full grown, drift, away, and establish their independence.
The wolverine is omnivorous. It consumes a wide range of edible roots, leaves, and berries as well as small game such as lemmings, mice, ground squirrels, hares, birds’ eggs and fledglings, and fish. In wooded country it is also known to kill beaver ( Castor canadensis ), which it secures on the trails between the feeding grounds and their ponds. It also seems to have a particular liking for porcupines ( Erethizon dorsatum ). However, it does not seem to have mastered the art of killing porcupines with impunity and is stated to succumb later, in some cases, from the quills. Other predators, such as the coyote ( Canis latrans ) and wolf ( Canis lupus ), kill porcupines by flip– ping them onto their backs and attacking their unprotected underparts.

EA-Zoo. Banfield: Wolverine

Carrion is also important in the diet of wolverines. They follow the migrating herds of caribou ( Rangifer sp.) in order to clean up the carcasses of caribou which were killed by wolves.
There is some evidence that wolverines may at times kill big game. Two observations that relate to this point are reported by Seton (7). Martin Hunter of the Hudson’s Bay Company saw a caribou sprint into a lake. When he observed it closely, he saw a black animal clinging to its neck. The caribou was pursued in a canoe and the attacking animal dropped off into the water. When shot, it turned out to be a wolverine. When the caribou was ap– proached it was found dead, with its jugular vein severed. During the winter of 1907-1908, J. Keele of the Canadian Geological Survey came upon a wounded young moose ( Alces americana ) helpless in a snowdrift. When it had been shot, it was found to have a large hole torn in its back. In the snow were tracks which indicated a running fight with a wolverine. The wolverine was later observed in the vicinity.
Like other members of the weasel family, the wolverine is pugnacious and strong. These characteristics seem to be developed in proportion to its size. It is said that a wolverine will defend a carcass against the attacks of wolves or even a g ir ^ ri ^ zzly bear ( Ursus sp.). it is not known whether these larger mammals are afraid to the wolverine or are merely treating it with the same respect they would give to a skunk ( Mephit e ^ i ^ s mephit e ^ i ^ s ), for the wolverine also possesses well-developed anal scent glands. It is possible that it puts its “mark” on the carcass, which further discourages intruders. In any event it is a courageous and resolute fighter and if cornered will fight to the last.
In movements the wolverine is generally slow and deliberate. If hunting or changing territory, it will trot tirelessly. It seems to have only mediocre

EA-Zoo. Banfield: Wolverine

eyesight and has the peculiar habit of sitting on its haunches and shading its eyes with one paw. Its senses of hearing and smell are acute and are used when hunting. The wolverine is generally solitary in nature except when accompanied by its young.
The short thick body of the wolverine is very muscular and it has prodigious strength and endurance. It can move stones that are as heavy as a man can roll, and can roll logs as heavy as a man can lift. It can swim and climb with ease.
An animal of this description would be expected to come into competition with humans sharing the same country. The wolverine in its tireless search for food frequently finds and follows a trap line. The traps are discovered, the trapped animals destroyed, and the traps themselves often hidden in the snow. The animals become very canny and remove the baits from traps with im– punity. They become wary of traps, snares, and other devices, and are dif– ficult to capture. If they follow the trapper persistently, it is sometimes necessary for him to abandon the trap line temporarily and search for a new route. On the other hand, some trappers and travelers report no difficulty in trapping wolverines when they appear on their trap lines or around their caches. It seems likely that young animals or other ^ s ^ that have had no experi– ence with humans may be less wary of traps than are experience individuals.
The wolverine also follows human tracks to get scraps of food which may be left about a camp site. Caches of furs, food, or camping goods are broken into when found. The furs are torn to shreds. The food is eaten or cached away in the snow and even the camping gear is sometimes hidden.
The writings of northern explorers and travelers contain many stories of the depredations of wolverines upon their caches and of the sagacity of

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these animals. Some writers, such as Richardson (6), state that the animals defile with glandular secretions what food stores they do not eat. Other writers, including Stefansson and Anderson (1), have found that their frozen meat had been gnawed and hidden in the snow, but what remained they found usable.
It is difficult to prepare a cache that is “wolverine-proof.” Seton made successful caches by placing supplies on raised platforms, the support– ing poles of which were stripped of bark and ringed by cod hooks. Warburton Pike (5) cached caribou meat by chopping a hole in the ice and freezing it in a lake. Anderson (1) reported a successful elevated cache with a platform that overhung the stripped poles by two to three feet, so that the animals were unable to climb over. Seton (7) writes of the wolverine: “It is not to be supposed that any part of the procedure of robbing caches is due to malice. Malice is a human weakness. Animals are not built that way.”
It is easy to understand how such a colorful animal as the wolverine has become a part of many of the legends of the North. Some trappers will repeat the common superstition that after death a trapper returns to earth as a wolverine to plague his fellow trappers.

EA-Zoo. Banfield: Wolverine

BIBLIOGRAPHY

1. Anderson, R.M. “Mammals,” Stefansson, Vilhjalmur. My Life with the Eskimo . N.Y., Macmillan, 1913, pp.494-527.

2. Hardy, Thora M.P. “Wolverine fur frosting,” J.Wildlife Mgmt. vol.12, no.3, pp.331-32, 1948.

3. Hearne, Samuel. A Journey from Prince of Wales’s Fort in Hudson’s Bay, to the Northern Ocean….in the Years 1769, 1770, 1771 & 1772 . London, Strahan and Cadell, 1795.

4. MacFarlane, R.R. “Notes on mammals collected and observed in the northern Mackenzie River district, Northwest Territories of Canada,” U.S. Nat.Mus. Proc . vol.28, pp.673-764, 1905. (No.1405)

5. Pike, Warburton. The Barren-Grounds of Northern Canada . London, Macmillan, 1892.

6. Richardson, John. Fauna Boreali-Americana. Part 1. Quadrupeds . London, Murray, 1829.

7. Seton, E.T. Lives of Game Animals . N.Y., Doubleday, 1926. Vol.2, pp.403-50.

A. W. F. Banfield

The Moose

(EA-Zoo. Belmore Browne)

THE MOOSE

CONTENTS

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Page
Physical Characteristics 1
Habits 3
Economic Importance 7
Enemies 8
Bibliography 10

EA-Zoology (Belmore Browne)

THE MOOSE
The largest member of the deer family reaches its greatest size in the American moose, Alces americana , and the Alaskan moose, Alces gigas , the latter attaining the more imposing proportions. They are related to the elk of the northern forests of Europe and Siberia, being differentiated from their Old World cousin by their larger size both of body and of antlers. The exact limits of the two North American species, that is to say the ge– ographical points of contact, have not been determined. Roughly one may say the American moose is found on the eastern side of the northern Rocky Moun– tains in the United States and Canada as far north as the arctic slope of northwestern Canada; the Alaskan moose, as the name indicates, is prin– cipally found in Alaska. Both occasionally occur beyond the Arctic Circle; sporadically they have been reported at the Alaska seacoast just southeast of Point Barrow and at the arctic coast of Canada on Coronation Gulf. The two are so alike in coloring and habits that they may be treated, in a brief study, as one.
Physical Characteristics . In color the moose varies from a mixed gray to black, the chest, shoulders, flanks and back being of the deepest tones. The legs are covered with short gray hairs that make them appear almost white.

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Individuals of a uniform mouse-gray color are occasionally seen and a few albinos have been reported.
An adult American bull moose will measure a total length of 102 to 108 inches and will stand at the shoulder 5 1/2 to 6 1/2 feet. The moose has a body short in proportion to its weight and height. It is much heavier through the shoulders than through the hindquarters. The powerful appear– ance of the shoulders is further accentuated by a distinct hump. When the size and importance of this animal is considered, it is surprising how rela– tively few accurate records of its weight exist. This is probably due to the fact that moose are usually shot in areas far removed from weighing facile– ties and the weight of entrails and lost blood has to be approximated. There is a recent record of 1,364 lb. for a bull shot in Quebec; many of the Alaskan specimens would weigh considerably more.
The head is massive. Beneath the lower jaw hangs a pouch or dewlap of hair-covered skin called by hunters the “bell.” Indeed, in younger bulls, the pouch is elongated into a slender appendage, suggestive of the clapper of a bell. As the males grow older this frequently disappears, leaving a shallower pouch. The bells of the cows usually retain the bell-clapper form. No definite function can be ascribed to the bell; but, as the moose habitually plunges its head beneath water to secure bottom grass and fre– quents marshy tickets where the brush is apt to be wet or snow-covered, it is possible that it serves the purpose of draining off some of the water that otherwise would keep the animal’s neck and chest wet.
The other distinctive features of the moose are the large p la ^ al ^ mated pro– jecting antlers possessed by both sexes but particularly large in the bull;

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its proportionally long legs; and the long fleshy muzzle, the hairless part of which is a small pad the size of a man’s thumbnail.
The moose is an excellent swimmer. It has been recorded that a bull moose swam across the entrance of Chugach Bay, Alaska, a distance of twelve miles. It walks with great rapidity and, when traveling a straight course, a moose can outdistance a hunter. The legs show a powerful muscular devel– opment. The cloven and lateral hoofs, called dewclaws, are well adapted to a marshy habitat. On the downward thrust they spread, contracting when lifted, thus enabling the animal to move with ease across bogs or marshy ground where a horse could pass, if at all, only with difficulty. This arrangement also enables them to move in such a way as to make almost no sound at all.
To those unfamiliar with the moose, its appearance is grotesque and lack– ing in the grace and beauty of the common deer; but in its natural habitat it presents an inspiring picture.
Habits . The moose is shy and elusive; but during the rutting season it becomes bolder, more inquisitive of noises or moving objects, and is, at times, easily approached by the hunter. If approached too closely, a moose will fight, but if there is an avenue of escape, it acts quickly and takes advantage of the available cover with great skill and judgment. The moose while under observation moves slowly for a time, then flees at top speed until cover is reached.
The rutting season begins when the antlers are at their prime, in Setpem– ber. The calves, usually one but not infrequently twins, are dropped eight months later.
Since the habits of all wild animals are influenced primarily by the

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type and location of their food supply, the Canadian moose could be listed as a lowland animal; for the green soft woods and aquatic plants on which it feeds are more abundant in the lowlands. But in the high mountains of northwestern Canada and throughout Alaska, there is sufficient moisture to sustain a heavy growth of moose provender, with the result that moose spend much of their time at, or even above, timber line. Even in the wintertime, moose will leave the snow-buried lowlands to feed on the wind-swept uplands, although flat valleys, well stocked with willows, spruce, tamarack, birch, aspen, and alder, are a favorite wintering ground.
By nature the moose is a solitary animal, but while “yarding” in deep snow, or in areas where they are very numerous, ten or a dozen may be seen close together. On the Kenai Peninsula, in Alaska, a hunter reported seeing nearly a hundred on one mountainside. Such gathering, however, would be accidental and the group would not remain together long, move compactly, or act in unison as a herd.
The seasonal migrations between the mountains and lowlands are caused by food and climatic conditions and insect pests. An understanding of local conditions is therefore necessary to the successful hunter. There are records of men dying of starvation in country where moose are plentiful simply because they mistook the “sign” — the hunter’s term for animal droppings — in the lower valleys for indications of an autumn habitat, whereas the moose were really gathered at timber line but a few miles away. As to moose sign, in the spring the summer when food is green and tender, it resembles the droppings of the domestic cow; but in fall and winter, when the food consists largely of twigs and bark, the sign consists of pellets about the size and shape of a pecan nut.

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In flat and forested country, moose frequent lakes and rivers in search of swamp vegetation and aquatic plants. For purposes of feeding as well as escaping insect pests, they spend much of their time in the water. When feed– ing on bottom vegetation in a lake, moose sometimes have not only the head but the larger part of the body submerged — some say occasionally the entire body. The sound of water sluicing from their antlers can be heard at a long distance. At such times the animals can be easily approached. In mountainous country, mouse feed along the willow-bordered streams and seek the wind-swept hilltops to escape the flies. Under such conditions they may be seen from afar and approached without much difficulty by the experienced hunter.
Moose frequent licks when available. The common belief is that all animal licks contain salt, but the deposits are often formed of a light clay and not infrequently salt is absent. Some of the well-known licks in the northern Rockies contain magnesium but no common salt. Salt, however, has a strong attraction for most or all the members of the deer family, as well as for wild sheep and goats. (It is reported by some observers that the arctic caribou are seldom or never seen putting their lips to sea water when they come down to the beach, while others claim they seek the beaches to lick salt.] Licks are easily recognized by the game trails radiating from them and the heavily tracked condition of the area. Although waiting for moose at a lick is universally condemned as unsportsmanlike, it would be excusable to the hunter in a serious emergency.
When forested country is burned over, a heavy growth of willows and other bushes spring up. These areas are frequented moose, as the fresh growth, tender and juicy, is desirable provender. That the burning of a forest increases the feed for moose is the reason why the forest Indians used to

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start forest fires deliberately; in some parts they do it still.
While the eyesight of moose is not particularly keen, their senses of small and hearing are extraordinarily acute. In an open stalk, the three prime requisites are the proper use of cover, silence, and a constant study of wind direction.
Hunting the moose in winter has certain advantages as well as disadvan– tages. The snow preserves the tracks of the animal and deadens the footfalls of the hunter. On the other hand, the slightest crust on the snow will produce a crunching sound and alert the moose. The snow curst is, however, a dis– advantage to the moose, as it cuts the legs of the animal, and the labor of plowing through deep drifts soon exhausts it. A hunter on snowshoes can, under such conditions, overtake and kill moose with case. This form of hunt– ing moose or any other animal is universally frowned upon by sportsmen except in rare cases where the securing of meat is a dire necessity. The forest Indian and the white trapper pursue this method constantly.
In any terrain and at any season, hunting the moose requires a high degree of skill and experience. Particularly is this so when the ground is covered with snow. The moose, along with other members of the deer family, would seem to realize that its trail is a source of danger. When preparing to lie down, the animals will, at times, move in a circle until a point is reached from which they can watch their back track. The result is that a hunter closely following the trail will be seen by his prey before he is aware of its presence. Experienced hunters travel on the downwind side of the trail, keeping a close watch ahead and locating the trail only at intervals in order to follow in the right direction.
In lower Canada “Moose calling” is practiced. A small horn made of birch

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or similar material is used and the long call of the cow or the grunting noise of the bull is imitated as conditions dictate. In western Canada the horn is rarely used and the hunter imitates the moose call through his cupped hands with equally successful results.
During the rutting period, or when wounded, a bull moose is dangerous. Even when apparently dead, a wounded animal should be approached with caution. A native of Cook Inlet, Alaska, was killed by a cow moose that he had mor– tally wounded. The cow, as is customary, struck with her front feet. Al– though the antlers of the male are formidable weapons and many men have been killed by them, the bull likewise uses his front feet as defensive weapons while the antlers are absent and during the period of their growth.
Economic Importance . The flesh of the moose resembles beef in flavor more than any other wild game of North America, with the exception of the bison and ovib u ^ o ^ s. Since time immemorial the moose has been the chief winter food of the forest-dwelling American Indian, and from the advent of the first white pioneer it has played an important part in the settlement of North America. Fresh, dried, smoked, or made into pemmican, it has furnished sustenance to wilderness travelers, from the lone trappers or prospector to the impressive brigades of the Hudson’s Bay Company. At present its meat is of vital importance to many individuals and communities through the forested North. When in prime condition, prior to the rut or mating season, the moose carries a plentiful supply of fat, the most important food item to those who live in remote areas. The Canadian voyageurs esteemed above all other parts of the animal the large gelatinous muzzle, called moufle by them; it is still considered a delicacy, and is eaten boiled or sometimes roasted.

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Moose hide has likewise played an important part in the lives of Indians and the white pioneers. When tanned, it is soft, pliable, and warm. In the North it is still used for countless purposes — for coats, leggings, mittens, moccasins, and sleeping robes. Abraham Lincoln as a boy slept beneath an elkskin; had he lived farther north, his bedcover would have been moose. Untanned mooseskin or rawhide has innumerable uses: winter covering for cabin floors, mattresses for sleeping bunks, and, cut up in strips, for ropes and string. The hide of the hock, in shape, bears a rough resemblance to the human foot. When case-skinned, the small end can be sewed with a sinew and the large end cut of admit the foot, thus forming an excellent emergency moccasin.
The long sinews which follow the backbone are composed of countless threadlike fibers. When the sinew has dried, a thread of any requisite size can be secured by splitting the end of the sinew with a sharp knife. It is then immersed in water until soft. An inch or two of the end is kept dry and hard and is used as a needle in forcing the thread through the holes per– forated by an awl in the material to be sewn. Clothing sewed with sinew lasts longer and is stronger than when sewed with cotton thread.
Enemies . Man, the wolf, and disease constitute the moose’s natural enemies. The sportsman is interested primarily in securing a fine trophy and seeks only the bulls carrying the largest antlers, which are, in most instances, approaching the end of their reporudctive ^ reproductive ^ value. The steady consumption of moose meat, summer and winter, by Indians, trappers, and frontier communities is a more serious matter, for the strain on the moose population is heavy and constant and both sexes are killed indiscriminately. The final result of man’s presence on the frontier is the growth of ranches and stock ranges. At this

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point the moose becomes a problem. A bull moose can walk through a barbed wire or rail fence with ease, the horses or cattle passing through the break may scatter for miles, causing many days or hard riding by the stockmen if not the actual loss of valuable animals. The advent of the rancher and stockman presages the end or retreat of the moose unless government sanctuaries be made for their protection.
The wolf is the only North American animal that habitually preys on the moose. The grizzly bear is strong enough to kill a moose and, on rare occa– sions, does so, but there are few if any records of black bears having killed moose. There is little doubt, however, that they kill calves when the oppor– tunity presents itself. The killing of calves by the smaller predatory animals is only barely possible, as the cow moose is a powerful and dangerous adversary. The fact that wolves sometimes hunt in families accounts for their ability to kill so large and formidable an animal as the moose. Most of the wolf killings occur during the wintertime, when the moose is seriously hampered by deep snow. At the present time (1950), there has been a considerable increase in the number of wolves throughout the North and the moose may have suffered seriously because of it. In spite of local fluctuations, the past ten years have seen a steady decrease in the moose population of western Canada and Alaska.
Under normal conditions, the moose is a healthy animal and in large areas, where they can move at will during their seasonal change of feeding grounds, cases of disease are apparently rare. On ranges where overcrowding has occurred, such as the Kenai Peninsula, Alaska, cases of diseased moose have been reported.
Fortunately the moose, if protected, adjusts himself to the presence of man, and a realistic conservation plan, including strategically placed sanc– tuaries and well-enforced game laws should secure the future of this noble animal.

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BIBLIOGRAPHY

1. Dixon, J.S. Fauna of the National Parks of the United States. Birds & Mammals of Mount McKinley National Park, Alaska . Wash., G.P.O.; 1938. U. S. National Park Service. Fauna series no.3.

2. Osgood, W.H. A Biological Reconnaissance of the Base of the Alaska Peninsula . Wash., G.P.O., 1904. U. S. Biological Survey. North American Fauna no.24.

3. ----. The Game Resources of Alaska . Wash., G.P.O., 1908. U. S. Dept. of Agriculture. Yearbook 1907, pp.467-82.

4. Sheldon, George. The Wilderness of Denali . N.Y., Soribner, 1930.

Belmore Browne

Mountain Sheep of Northern North America

EA-Zoology (Belmore Browne)

MOUNTAIN SHEEP OF NORTHERN NORTH AMERICA

CONTENTS

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Page
Habitat 2
Physical Characteristics 2
Habits 4
Natural Enemies 7
Status of Geographical Varieties 8
Stone or Black Sheep 8
Fannin or Saddleback Sheep 9
Dall or White Sheep 9
Kenai Sheep 10
Summary 10
Bibliography 12

EA-Zoology (Belmore Browne)

MOUNTAIN SHEEP OF NORTHERN NORTH AMERICA
Until recent years wild mountain sheep occurred in all the western mountain regions of North America, from the arctic coast to Mexico. Fossil– ized remains show that they inhabited the Yukon region in the Pleistocene period. It is though that mountain sheep entered North America from Asia by way of the Bering Sea land bridge. In the region between Mexico and British Columbia and Alberta, ten subspecies are listed, all of which are loosely referred to as the bighorn. The northern limit of the bighorn’s range is at some unidentified spot north of the Smoky River area on the British Columbia-Alberta boundary. North of that, for several hundred miles, a break occurs where no wild sheep have been reported. From 55° N., then, to the Arctic Circle is the mountainous territory over which range the northern mountain sheep, of which four subspecies have been described. The northern mountain sheep differ from the bighorn in three important characteristics: their size is similar; their horns are less massive and tend to curl farther outward from the head; and they are lighter in color.
Although sometimes attributed to Alaska, both the Rocky Mountain sheep, Ovis Canadensis , and the black or Stone sheep, O. dalli stonei , may be said to have their limits within Canadian territory and to portions

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of northwestern United States. The distinctive Alaskan sheep is the Dell’s sheep, O. dalli dalli and O. dalli kenaiensis . There is also the fannin or saddleback sheep, O. fannini , but the description of it as a distinct subspecies is now questioned.
Habitat . All mountain sheep prefer the wild, exposed mountaintops, but they do not hesitate to descend into timber. Their range is confined to a narrow altitudinal strip between 2,000 feet above sea level and the highest grass lien. Other northern animals, such as the caribou, have a much wider range, including all territory where vegetation is found, from sea level upward, and sometimes overlapping the habitat of the mountain sheep.
Physical Characteristics . The form of the mountain sheep is compact and powerful yet full of grace. The horns are not shed but continue to grow throughout the life of the individual, the rings of growth being an indication of the animal’s age. For this reason, as well as because of their beauty and comparative rarity, the horns of mountain sheep have always been highly prized as trophies of the big-game hunter. The horn consists of a tough outer sheath that surrounds a core of bone which is a part of the skull. The period of fastest growth is during the summer. During the winter, little or no horn is added and a crack of depression between the periods of growth form “year rings” on the surface. The horns of the ewes are much smaller than those of the rams, averaging only about nine inches.
Apparently the main function of sheep ^^ s horns is aggressive action during the mating season, when rams savagely fight each other for possession of the ewes. Instance of their using the horns as protective weapons against predatory animals (including man) care rarely reported.

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In the extreme northern ranges ^ , ^ sheep rarely live longer than thirteen years, whereas the more southern species have a greater life span.
Mountain sheep are among the surest footed and most agile of animals. The hoof is cloven, each half being capable of freedom of action, a valu– able asset on rough rocky formations. Each half, moreover, is composed of a hard toe of horn and a softer center which, wearing away more rapidly than the shell, preserves a concave form with a sharp cutting edge. To this is added a soft rounded heel which possesses a suction-like quality, capable of adhering to smooth surfaces.
The almost incredible climbing ability of the wild sheep is well known. Its poise and fearless balance on steep slopes and the abandon with which it plunges downward on treacherous cliffs are constant sources of amaze– ment and admiration to mankind. This great agility and lightness of move– ment is particularly remarkable when one remembers that the rams carry a burden of curling horns which sometimes weights thirty pounds or more.
They are keen of vision and, unlike most game animals, depend little upon scent for warning of danger. It is exceedingly hard for a hunter to approach these alert, agile, and farsighted animals on an open mountainside.
Very little reliable material has been published on the relative weights of the different members of the wild sheep family. They are usually killed in remote and rugged areas and hunters rarely posses the equipment or leisure to secure accurate results. A rough approximation, based solely on experience in back-packing the carcasses, would place the weight of, for instance, an adult Stone sheep at about 200 pounds. A Dall or white ram would weigh about 30 pounds less, with the saddleback somewhere in between. These figures will suggest the quantitative meat value of the

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northern sheep. As food, most sheep hunters state that between the months of August and November sheep meat is superior to that of any other American game. It is fat, tender and nutritious, and the liver, brains, and kidneys are esteemed as delicacies. The large intestine, when turned inside out, washed, and roasted on a green sapling, is prized by northern forest Indians and rare white initiates.
Their close-packed pelage of brittle hair is practically impervious to cold. A noticeable shedding or molting takes place in early spring. The brittleness of sheep hair, which breaks or separates easily, renders the hides of little use. Skins collected in August, however, when the hair is still growing, are used by some northern forest Indians and Alaska Eskimos for coats and bedding. They are ideal for sleeping bags, being as light in weight as eiderdown, yet shedding frost and snow as no eider– down bag can do.
Despite its strength and hardihood, the wild sheep does not adjust itself to the encroachment of civilization. It is susceptible to diseases introduced by domestic animals and requires low-altitude wintering grounds well removed from the presence of man.
Habits ^ Habits ^ . In North America the mountain sheep rarely use the western or humid side of the many ranges forming the western cordillera. Their preference seems to be for the rolling grasslands on the eastern or dry side of the western highlands. The main exceptions to this rule occur at the northern and southern extremities of their habitat: on the arctic coast west of the Mackenzie River, western Alaska, and on the northwestern coast of Mexico.
Snow is their constant neighbor and, while it may threaten their

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existence during hard winters, it waters the mountain pastures, cools the air, and thus preserves the sheep from insect pests in summertime. During heavy blizzards they may seek the ice of cliffs or descend to the protection of the timber line but, in such case, as soon as the storm has passed, they return to their high grass slopes where the wind blows the snow from the winter pastures and the winter sun warm their coats.
The sheep winter on grass-grown foothills, still above timber line and, where possible, facing southwest. It will be found that ridges connect their feeding grounds with higher outcrops of rock; these form a safe line of escape during attacks from predatory animals.
Early in the morning, the sheep begin feeding downward. They usually speed the noon hour at rest on some exposed shoulder of the mountain. During the afternoon they feed upward to the high ground where they spend the night. Their beds consist of small level areas which they hollow out in crushed rock or gravel — this type of bed providing better drainage in times of rain or snow than would grass or moss.
With the coming of spring, the sheep leave their wintering grounds and follow the green grass upward into the higher ranges. The mass movements of sheep are not migrations in the accepted sense but merely journeys to fresh feeding grounds. Great animation is evident on the spring journey. Different bands converging from the lower spurs may, in a wild and well– populated range, form a band of several hundred individuals. The presence of newborn lambs adds to the confusion and to the trepidation of the ewes.
The mating season occurs in late October and November; the period of gestation is about seven months. The young, usually one, sometimes twins, are born in May, in sheltered nooks under protecting cliffs. Though

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soon able to follow their mother, the lambs spend the first few weeks of their lives close to easy concealment in the rocks. By June they are scam– pering about on the grassy slopes in little bands of ^ from ^ four to ten. A lamb can make a vertical jump of six feet, so that when the “migration” takes place in June the young are able to take care of themselves, while remaining under the watchful eyes of the ewes.
The avalanche of plunging, leaping bodies moves from crag to crag, an unerring instinct seeming to lead them along the safest routes. Year after year the same general routes are used, resulting in clearly defined trails across the cliffs, rockslides, and grassy slopes, Some of these trails can be seen several miles away, particularly when they become etched with melting snow. No other mountain animal equals the sheep as a trail maker.
During these spring journeys the sheep gather the mineral deposits known as licks. These are not, as formerly thought, salt deposits, since recent examination has disclosed that the alkaline ingredient attractive to the sheep is magne is ^ si ^ um oxide. This mild laxative, combined with clay, brings about a scouring of the animal’s intestinal tract. Trails always converge from all directions to these licks.
While the rams follow the trails used by the ewes, they descend to the wintering grounds later than do the ewes and leave earlier in the spring. Ewes and lambs habitually range lower than the rams. The reason for this is not clear, the ewes being as capable as the rams of reaching the higher pas– tures and having the added incentive of removing their young from the haunts of predatory animals. A possible reason is that ewes and lambs form larger bands than do the rams and, therefore, require more extensive and less broken pasture land.

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Young rams of from three to six years of age exhibit an indecision concerning their associates and spend much of their time wandering between the bands of ewes and the groups of older rams. They have outlived the need to be protected by the former and are not yet accustomed to the monotonous and detached lives of the latter. As they grow older they gradually become associated with a band of rams and adapt themselves to new ways.
As the summer advances, the separate bands gather into larger groups, influenced, perhaps, by the prevalence of food in certain areas. Such a band, composed of rams of medium ages, may number as high as thirty-five head. A band of this size, moving across some steep mountainside, their powerful bodies and massive horns standing out in bold relief against a backdrop of snow-capped peaks, makes an impressive pictures. The oldest rams are seldom found in a band of this kind but, instead, are apt to segregate themselves, apparently preferring solitude. There are seldom more than six rams in a band of patriarchs. These, because of the extraordinary size of their horns, are the ones most sought after by the persistent sportsman. Mountain-wise as they are, these older rams seem to become indifferent to their surroundings in extreme old age, being caught sometimes far back in a mountain “pocket” when the heavy snows fall, cut off by drifts from the grass of the lower foothills. As weakness overcomes them, the weight of their horns forces them inexorably downward into some snow-filled gully where escape is impossible.
Natural Enemies . Mountain sheep are hunted by men and have always been preyed upon by wolves, wolverines, coyotes, gold ^ en ^ eagles, and lynx. In the past fifty years, there has been a rapid decline in number, and the

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wild sheep is faced with extermination. Occurring now chiefly in national parks and game preserves, the mountain sheep has become one of the major problems of game conservation. Since in game preserves and national parks the predatory animals are protected as well as their victim, special legislation against such animals, particularly the wolf, has been and is being advocated.
^ Status of Geographical Varieties ^
Stone or Black Sheep ( Ovis dalli stonei ). This sheep occurs chiefly within the confines of British Columbia, but has been reported in the area north of 55° to the south-central Yukon Basin. On the west it does not penetrate very far into the humid, snow-covered mountains of the Alaskan boundary, but on the eastern side it has been found along the tributaries of the Peace and Liard rivers. On the headwaters of the Taku River, the Stone sheep seems to blend with the white Dall sheep, the blending becoming more pronounced in the more northern areas. Osgood (3) failed to find signs of Stone sheep in the northern regions and merely notes that “both white and gray sheep are reported, though all are said to be white in winter.”
The Stone sheep is the largest of the northern sheep and the darkest in color. Its color pattern resembles that of the bighorn, but the dark shades are deeper, running to blackish brown and dark bluish black. The light markings consist of a distinctive rump path and a ribbon of white on the back of each leg, with light shading on the belly. The horns are slender and curl outward from the head. They are seldom blunted or “brushed” on the tips as is frequently the case with the close curled and more massive horns of the bighorn.

EA-Zoo. Browne: Mountain Sheep

Fannin or Saddleback Sheep ( Ovis fannini ). When this sheep was first discovered and described, it was accepted as a distinct form or subspecies. Today, after careful study of its range and physical characteristics, it is recognized as an intergrade between the Dall and Stone sheep. Its range ex– tends from the headwaters of the Yukon to the headwaters of the Peel River and westward to the Tanana Hills.
The color of this intergrading species in darker in the southern part of its range where it meets the black sheep, and lighter in the north and west where it meets the white sheep. The average trend of color is more toward the light or Dall coloring than toward the darker shades of Stone sheep. Throughout all its color variations the pattern, however dim, is suggestive of the Stone or bighorn pattern.
Dall or White sheep ( Ovis dalli dalli ). These sheep were unknown to science until specimens were obtained for Edward William Nelson from the upper Yukon region. He first described the species in 1882, naming it Ovis dalli in honor of the early Alaskan explorer and naturalist, William Healey Dall. The Dall sheep occur in two distinct regions. The southern range, roughly describing an arc, runs from west of Whitehorse, Yukon Territory, through Nutzotin and Chugach Mountains and on to the Alaska Range. In its northern range, the Dall sheep occurs from the vicinity of the Liard River on the south to the arctic coast along the Mackenzie Divide in the north, and thence westward to the Baird Mountains which lie northeast of Kotzebue Sound.
The Dall sheep is in general appearance pure white, although individuals have been reported with a few dark hairs in the tail. It is an animal of grace and beauty. In the past it was of great economic value, its meat pro– viding food for the laborers who built the Alaska Railroad and other government

EZ-Zoo. Browne: Mountain Sheep

projects. In former times carcasses were on sale in most of the mining camps from Yukon Territory to western Alaska. Despite the heavy slaughter and the unnecessary waste that always accompanies commercial killing, the sheep held up numerically very well. However, they have decreased alarmingly in recent years; which, in the author’s opinion, is due to the increase of wolves.
Kenai Sheep ( ovis dalli kenaiensis ). According to Osgood, the most noticeable difference in this sheep as compared with the true dalli is that the upper side of the tail is white, whereas in the dalli there are dusky or brownish hairs. Also, the horns of kenaiensis average thicker at the base, particularly on the lower side, and are less divergent at the tips than in dalli .
Summary
The wild mountain sheep everywhere in North America is in danger of ex– tinction, a fact to be deplored since this is one of our most beautiful animals. It now occurs almost exclusively in game preserves and national parks. The national parks, however, also extend their protection to animals which prey upon the sheep, and such predatory animals are by way of increase– ing numerically as a result. Chief of these predatory animals is the wolf; and its rank have been strengthened to late by its related species, the coyote, which is steadily increasing its range northward, having been reported even farther north than the most northerly sheep.
Basing his arguments on a hypothetical beneficent “balance of nature” and on the theory of the “survival of the fittest,” Murie (2) has opposed all legislation aimed at checking the wolf population of Mount McKinley

EA-Zoo. Browne: Mountain Sheep

National Park, or other game preserves. He denies that the sheep population has decreased due to the depredation by wolves, and asserts that only old and feeble sheep can fall victim to the wolf. Dixon (1) also seems to favor the continued protection of wolves but advocates the extermination of the newcomer, the coyote, from Mount McKinley National Park. He admits, how– ever, that one wolf den containing four pups, which he observed in 1932, showed that several mountain sheep had been eaten by this one litter. But, as Sheldon (4) and others have pointed out, when a wolf visits sheep country all the sheep become terrified and run well back in the mountains, not resuming their normal ranging habits for several days. It is clear that such a flight from the pasture lands might well results in the diminution of a band of sheep through starvation.
The fact must certainly be emphasized that in the four years from 1941 to 1945, according to the author’s investigations, the sheep population in Mount McKinley National Park dropped from around 1,000 to not over 500, at which figure it has since, presumably, remained.
[The effects of predators, other than man, upon animals like the moun– tain sheep and caribou, are at present hotly disputed. Views like those of Mr. Belmore Browne are generally held by sportsmen-naturalists; many profess– sionals naturalists oppose them. The Editors]

EA-Zoo. Browne: Mountain Sheep

BIBLIOGRAPHY

1. Dixon, J.S. Birds & Mammals of Mount McKinley National Park, Alaska . Wash., G.P.O., 1938. U. S. National Park Service. Fauna Series no.3.

2. Murie, Adolf. The Wolves of Mount McKinley . Wash., G.P.O., 1944. U. S. National Park Service. Fauna Series no.5.

3. Osgood, W.H. “Mammals of the Yukon Basin,” U. S. Biological Survey. Results of a Biological Reconnaissance of the Yukon River Region . Wash., G.P.O., 1900, pp.21-45. The Survey. North American Fauna no.19.

4. Sheldon, Charles. The Wilderness of Denali . N.Y., Scribner, 1930.

Belmore Browne

Animal Population Cycles

(EA-Zoo. Leonard Butler)

ANIMAL POPULATION CYCLES

CONTENTS

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Page
Lemmings 3
Arctic Fox 9
Red Fox 12
Snowy owl 14
Ptarmigan 15
Mice 16
Marten 17
Lynx 18
Mink 18
Muskrat 18
Other Animals 19
Caribou 19
Arctic Hares 19
Polar Bear 19
^ Ermine ^ 20
Bibliography 21

EA-Zoo. Butler: Animal Population Cycles

LIST OF FIGURES

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Page
Fig 1 Arctic fox returns and lemming reports for Canada 9a
Fig. 2 Colored fox returns showing four-year and ten-year cycle 13a

EA-Zoology (Leonard Butler)

ANIMAL POPULATION CYCLES
The number of animals in a certain region does not stay constant from year to year but fluctuates between abundance and scarcity. Sometimes these fluctua– tions are gradual and at other times they are sudden or eruptive. Many of these fluctuations have a rhythmic periodicity, the time interval between two consecu– tive periods of abundance being the same. The periods of abundance are spoken of as peaks, and when these peaks are easily recognized and occur with sufficient regularity the population is said to be cyclic. The average time between peaks gives the length of the cycle, so that it is customary to speak of four-year cycles (i.e., four years between each peak) and so on. Whether an animal popula– tion exhibits regular cycles or irregular fluctuations depends upon both the species of animal and the locality. The beaver population does not show cyclic behavior, whereas foxes have a ten-year cycle. Marten in bush country have a ten-year cycle, whereas in the Subarctic they have a four-year cycle.
In a country where the natives depend in the main upon wildlife resources for their living, fluctuations in animal population assume serious importance. The most important and striking fluctuations in the Arctic are found in the populations of arctic fox, lemming, snowy owl, and ptarmigan. The fox is impor– tant because of the value of its fur and the fact that it apparently transmits disease to sledge dogs, which are necessary to the normal mode of travel in the

EA-Zoology. Butler: Animal Population Cycles

North. The lemming is important because in many sections it is the main food of the arctic fox. The snowy owl is useful as an index of lemming abundance; when the owls are plentiful the lemmings are also plentiful, and when the lemmings become scarce the owls move away, often undertaking long flights to the south. Ptarmigan provide food for both man and fox.
The periodic fluctuations in the members of lemming, ptarmigan, snowy owl, and fox, all appear to ebb and flow together. The period of the fluctuations or the length of time from one peak to the succeeding peak averages four years, varying sometimes to three or five. The regularity of the cycle and the fact that it affects several species were noted as early as 1780 in the writing of Gmelin. P ^ L ^ ater Pennant (1784), Schrenk (1854), Collett (1895 ^ ) ^ , and many others ^^ referred to this p eh ^ he ^ nomenon. Wrangel (1839) and several others suggested that the cycle is due to the permanent wandering of all foxes around the North Pole. The migration was assumed to take four years, so that the animals would be abundant in any particular place only once in four years. This theory was disproved as soon as large-scale date were observed because it was noted that there was no succession of peak years occurring at approximate intervals around the Pole.
The simplest theory of the ca su ^ us ^ al relationships between the cycles in lemming, snowy owl, white fox, and ptarmigan is as follows: the lemmings, for some reason at present unknown, increase in numbers and become abundant. This increase makes the area attractive to the snowy owl, which feeds on lemmings, and the snowy owl moves in. The young owls have a better chance to survive and snowy owls become numerous in the area. The food supply furnished by the increased number of lemmings also affects the resident fox. Its numbers increase either because of larger litters or greater survival of the young cubs. The ptarmigan

EA-Zoology. Butler: Animal Population Cycles

becomes abundant either because the favorable factors affecting the lemming also cause it to increase, or else the presence of abundant lemmings reduces the predatory pressure on ptarmigan and allo ^ w ^ s them to increase. Later, the lemmings decrease in number either because of lack of food, looses from preda– tion, migration, or some other cause. The fox population also decreases, through starvation or through being more easily trapped when in a weakened condition; or their numbers may be reduced through disease, for at the peak of fox abundance and epidemic can spread rapidly throughout an area. The snowy owl, because of luck of food, moves to other areas which still have lemmings, or else flies south in search of alternatives food. The ptarmigan, now preyed upon more heavily, also decreases.
Lemmings
The lemming cycle has attracted much attention because of the spectacular migration which often occurs at the peak of the cycle. Most of these migration reports come from Norway, where the lemmings live in the subalpine regions. When they become excessively abundant they begin their migration, vast numbers of them, regardless of obstacles, always advancing in the same direction in which they originally started but following more or less the course of the great valleys. Down the mountainsides and valleys they come, through the tree belt, and out over the lowlands, crossing fields, swimming lakes and rivers, and even passing through busy towns. Often they become so numerous around a town that dogs and cats become exhausted from killing them and will eat no more. The lemmings that survive move steadily onward until they reach the coast where they plunge into the water and swim out until they drown or reach an island. These creatures travel across powerful physical barriers and without fear of man, yet they die easily of shook from the slightest injury. In the Arctic the migra– tions are not as pronounced, although they have been recorded for several regions.

EA-Zoology. Butler: Animal Population Cycles

The lemming, a stout rodent which resembles a very large mouse, weighs from 75 to 100 grams as compared with 20 to 40 grams for a mouse. The lemming is variously colored: black, brown, yellow, or white. The short stumpy tail and bustling gait make it an odd little creature. It is remarkable for its voracity and its powers of reproduction. The lemming is mainly nocturnal and lives in burrows from two to three feet long, ending in a ne x ^ s ^ t chambers, four or five inches in diameter, lined with grass and moss. Near the nest there is often a branch burrow which is used for sanitary purposes and as a place of refuge when the main burrow is invaded. In the nests, during the early summer, the litter of from two to eight young is born. There are several litters a year so that when conditions are favorable the population can increase very rapidly from dearth to abundance.
Lemmings are active throughout the winter, making many tunnels along the surface of the ground under the snow which are revealed when the snow melts in the spring. These tunnels are their foraging road, safe from the storms of winter. At times when gales blow the snowy covering off, the lemming wander aimlessly about on top of the snow until they perish from cold or are caught by some enemy.
Lemmings feed entirely on vegetation, and many theories regarding this food supply have been advanced to explain the fluctuations in lemming population ^ s ^ . It has been suggested that at certain times the food is richer in vitamins and minerals than at other times and that it is this extra richness which causes the lemming to breed and survive better. It has been suggested that the food in certain years contains a vitamin so potent that it leads to an orgy of reproduction and finally to mass migration and self-destruction. ^ Another ^ S ^ s ^ upposition has it that when the lemmings become overabundant they eat up all the available food and then have either to move on or starve to death. None of these theories has any experimental proof to substantiate it.

EA-Zoology. Butler: Animal Population Cycles

That the lemming has sufficient voracity to eat up its food supply is shown in the writings of Elton and Freuchen. Elton ^ (6) ^ records an observation in [: ] ^ w ^ hich two Dicrostonyx (collared lemming) were released in a pen with a surface area of 25 square yards. The ground in the pen had a lush covering of grass and other vegetation about eight inches thick when the lemmings were introduced on March 30. On about April 14, five young were born. By this time there were numerous roads make in the grass and some of it had been nibbled and eaten. By May 28, the lemming family of seven had, in eight weeks, destroyed every vestige of vegetation in the pen. One factor contributing to the destruct– tion was the lemmings’ habit of eating roots as well as stems and leaves.
There are two main types of lemmings: brown lemmings and collared lemmings. The brown lemming has the same brown coat both winter the summer, while the collard lemming changes from its brown summer cost to a white one in winter. Another main difference between the collared and brown lemming is that with the change to the white winter coat, the collard lemming develops a thick, horny pedlike growth on the underside of the two middle claws of the front feet. These pads re molted in the spring when the winter coat is shed. The collar lemming inhabits the high, dry ground, while the brown lemming inhabits the low, moist spots.
There are three main species of brown lemming, all belonging to the genus Lemmus. The genus is circumpolar in distribution and fossil remains of lemming are found as far south as Portugal, indicating that these rodents moved southward ahead of the icecap. The northward retreat of the ice past the southern part of the White Sea split the lemming population into two parts, at the same time pushing them higher and higher up the mountainsides. These mountain lemmings to not form one continuous connected population but live in normal years as

EA-Zoology. Butler: Animal Population Cycles

separate breeding colonies isolated from one another by deep valleys. The population to the west of the White Sea is confined chiefly to Lapland and the mountains of Scandinavia; it constitutes the species Lemmus lemmus . East of the White Sea, as far as Siberia, and in Novaya Zemlya, the species L. obensis is found. The third species, L. trimucronatus , which closely resembles the Siberian one, ranges from Alaska to Hudson Bay. The last-mentioned species has three subspecies commonly known as Back’s lemming, Alaska brown lemming, and tawn o y lemming. ^^
The collared or varying lemming ( Dicrostonyx ) is represented in Canada by two species and five subspecies. D. hudsonius , the Labrador varying lemming, covers the barren ground of the Ungava Peninsula, extending down the coast of Labrador and the east coast of Hudson Bay. The four subspecies of D. groenlandicus cover the rest of Canada from Baffin Island to Alaska; the common names for the subspecies are: Greenland varying lemming, Alaska varying lemming, Mackenzie collared lemming, and Richardson’s varying lemming. The main point about this distribution is that the brown and collared lemmings co-exist throughout their Canadian range except in Quebec and Labrador.
Most of the data on lemming cycles and their relationship to the predator cycles are given in Elton’s book Voles, Mice and Lemmings (6).
Of the many accounts by arctic travelers and naturalists on the increase in lemming population and its subsequent decrease, the following taken from Peter Freuchen’s observations is typical (7). As a member of Rasmussen’s Fifth Thule Expedition, Freuchen spent several years in various parts of the Melville Peninsula. He noted that during 1921 lemmings were not very numerous; foxes and predatory birds were also scarce. “In 1922, however, matters were quite different. There were lemmings everywhere, and there were traces of many more foxes, rough-legged buzzards and owls, more peregrine falcons, and more ravens.

EA-Zoology. Butler: Animal Population Cycles

The Eskimo brought large numbers of ermine skins to the trading posts, but their catch of foxes was nothing extraordinary.” The natives claim that often when lemmings are abundant, the foxes are hard to catch as they are not attracted by baits. In this particular year, Freuchen notice s ^ d ^ that both caribou ^^ and hares were left entirely in peace by the wolves because there were sufficient lemmings. In 1923, the lemmings swarmed in still greater numbers, even invading the tents of the expedition’s living quarters.
Then suddenly the lemmings seemed to disappear and, by December 1923, Freuchen was able to notice a remarkable difference. The foxes caught were not gorged with lemmings as had been the case in the spring of 1923, and in the previous year. Predatory birds were very scarce and their stomach contents contained on lemmings. This “crash” was also noted on Baffin Island the same year.
The best-documented source of information on the rhythmic ups and downs in lemming populations in furnished by the Canadian Arctic Wildlife Enquiry. Residents and travelers in the North are asked to record whether lemmings are more abundant, less abundant, or no change as compared with the previous season. These reports are assembled and analyzed by the Bureau of Animal Population at Oxford (3). From these reports it is possible to get both regional and general changes in the lemming population of Canada. The lemming population has had peaks in the following years:

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Year 1933-34 1936-37 1940-41 1944-45
Per cent observers reported increase 85 65 79 68
It will be seen that adjacent peaks are 3 years, 4 years, and 4 years apart, which is close to the orthodox four-year interval expected. In recent years the country has not been treated as a whole, but grouped into two sections: the

EA-Zoology. Butler: Animal Population Cycles

Eastern Arctic where the cycle is very pronounced, and the Western Arctic where the cycle is not quite so definite. That there are differences between these two regions is seen in the following comparison for the year 1942-43”

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Eastern Arctic Western Arctic
Per cent observers reported increase 17 47
Per cent observers reported decrease 62 18
Per cent observers reported no change 21 35
In the Eastern Arctic, 1942-43 was the second year of scarcity after the 1940-41 peak. The majority of places in northern Quebec and Baffin Island re– ported decrease, while at Padlei and Baker Lake, on the edge of the Eastern Arctic section, increase was reported. The third year of increase of lemmings on the islands of the Western Arctic was in 1942-43, and increase was also re– ported at Bathurst Inlet. At Cambridge Bay this was the peak year; numerous lemmings in their fall coats were found dead on the ice in the spring.
These observations indicate that while there is general synchronism in the cycles there is no absolute synchronism. The agreement between the Canadian figures and those for Lapland and Norway is as close as between various sections or Canada. The last three peaks for these countries recorded by Elton (6) are 1930-31, 1933-34, and 1938, all of which correspond closely with the figures given above for Canada.
It is well established that there is a rhyth e m in the population cycle of the lemming and that peak populations do not occur at the same time throughout the circumpolar region, but there is a synchronization of these peaks which, in the whole region, often occur the same year. Whether the similarity in timing is accidental or because the same [: ] casual agent is exerting the same influence throughout the region cannot be determined until more is known about the causes

EA-Zoology. Butler: Animal Population Cycles

of cycles. The five factors that may cause or control the lemming cycle are: ( 1 ) climatic factors affecting the food or rate of reproduction of the lemming, ( 2 ) predators taking too large a toll of the lemming population, ( 3 ) epidemics of disease, ( 4 ) migration, and ( 5 ) the removal through some unknown cause of one generation of young. This last cause is a factor only when the animal has a short life span.
Arctic Fox
The life history of the arctic, or white fox (genus Alopex ) is dealt with fully in the article “Fur Bearers of North America.” The figures for the number of pelts collected at various trading posts give an index of the number of animals present in that particular section in any year. This index is only a relative one as there is no evidence to indicate that the percentage of the population trapped each year is constant. In fact, there is a good deal of evidence to indicate that at, or just after, the peak of the cycle a much larger percentage of the foxes is trapped than at the bottom of the cycle. This is borne out by the fact that at the bottom of the cycle it takes an expert trapper to catch a fox whereas when they become abundant it is possible for a rank amateur to trap them. As far as the cycles are concerned, it means that the peaks are probably exaggerated but that their periodicity is the same as the population periodicity.
Figure 1 shows the total Canadian white fox catch from 1915 to 1946. This illustrates the magnitude of the increases (for instance, from 19,000 in 1929 to 68,000 in 1931), and the regular periodicity of peaks being 3, 5, 4, 4, 3, 4, and 4 years apart. Most of these pelts are obtained from the Eastern Arctic, so the graph shows the cycle which is in operation in the area east of Fort Ross on the Boothi s ^ a ^ Peninsula. West of this point the cycle is not as regular but it ^^ ^ followed by 9a ^
Fig. 1. Arctic fox returns and lemming reports for Canada.

EA-Zoology. Butler: Animal Population Cycles

does show violent up-and-down movements. The Alaskan data show no violent fluctuations nor any sign of a cycle. In Greenland, there is also no sign of a cycle, the pelts fluctuating, in general, from 1,000 to 3 0 ,000, with a few exceptional highs of 5,000 pelts.
The main causal factor in the white fox cycle is presumed to be the availa– bility of the lemming food supply. An index of the lemming population in east– ern Canada is plotted on the same graph along with the white fox data. This shows a marked correlation between the two cycles, the peak of the white fox cycle usually coming one year after the peak of the lemming cycle.
If the lemming is the main causal factor, then in areas where the fox does not feed to any extent on lemming, or where an alternative food supply is available, the fox cycle should not be as definite. In general, this is true; on small islands or other areas where a good deal of marine life is available for fox food, there are no rhythmic population changes. In Greenland, Braestrup (1) divides the foxes into two types: the lemming foxes of the north and the coast foxes of the south. There is not a sufficiently large take of foxes in the north to indi– cate whether or not these “lemming foxes” are cyclic or not. There is, however, a cyclic tendency in the percentage of whites. Braestrup interprets this as being the result of migrations from Canada where the white phase is more common. The peak percentages of the whites seem to agree with the population peaks in Canada.
Within the Eastern Arctic of Canada where the white fox cycle is well synchronized, there are minor abnormalities. The peaks in northern Baffin Island and on the western shore of Hudson Bay occurred the same year until 1940-41. In that year there was a peak catch at Repulse Bay, Baker Lake, and Chesterfield Inlet on the west shore of Hudson Bay, while at Arctic Bay, Clyde, and Pond Inlet in

EA-Zoology. Butler: Animal Population Cycles

northern Baffin Island, there was the lowest catch of the cycle. The following year there was a decrease on the west shore of Hudson Bay and a peak collection in northern Baffin Island. By 1945-46, both sections were showing similar fluctuations in their white fox populations. During the same years, Sugluk ^ , ^ on Hudson Strait, exhibited the same accelerated peak. It had its highest collection at the same time as the west coast of Hudson Bay and one year ahead of the peak in Baffin Island. By the next peak, it was back in step with the general cycle.
There are several cases known of this speeding-up or retarding of peak years in certain sections but none has been completely analyzed so the possible cause is still unknown. It is interesting to note that the cycles usually synchronize again at the next peak. It has been suggested that a local abundance of lemming or other food may have encouraged migration into the area, or faster reproduction of the indigenous stock, but the whole question of ptarmigan and marine sources of supplementary food needs more investigation. All that can be said with certainty is that, where lemming is the chief source of food, the arctic fox exhibits a clear-cut cycle of extreme fluctuations in pelt numbers and that this cycle runs one year behind the similar cycle for lemmings.
Disease may also play an important part in the crash of the arctic fox population. There have been numerous reports of disease in sledge dogs occurring just at, or after, a peak year in foxes. These epidemics have at times been very severe and have decimated the whole dog population of the area. Isolation of dogs and restrictions on dog teams have not been enough to stop the epidemic which seemed to break out simultaneously at several points and spread rapidly through the country. The fact that these epidemics occurred at approximately four-year intervals and that there appeared to be some connection between fox peaks and dog epidemic led Elton to institute an inquiry. This showed that

EA-Zoology. Butler: Animal Population Cycles

occasionally dead foxes were found and also sick ones were seen around the settlement at the time of dog epidemics.
The symptoms were extreme boldness, running or frothing at the mouth and nose, glazing of the eyes, and the nonavoidance of obstacles in the fox’s path. The disease was assumed to be of a virus nature and was classified variously as distemper, encephalitis, pseudorabies, and crazy disease. Plummer (10) has shown recently that the disease is true rabies and that it is present in both dogs and foxes in the Canadian Arctic. It has been reported previously from the Soviet Arctic population of arctic fox.
Red Fox
The red fox (genus Vulpes ) is a comparative newcomer to the Canadian Arctic (see “Fur Bearers of North America”). In strictly arctic regions, the red fox has a four-year cycle which is very definite and in every way comparable to the cycle of the arctic fox. In the Subarctic, the red fox has a ten-year cycle similar to that of many of the bush animals.
In Labrador, Elton has shown, the red fox has a definite four-year cycle. In 92 years there were 23 cycles, and only once was the interval between peaks as long as 6 years. In 92% of the cycles, the time interval varied from 3 to 5 years, with 4 years the most common interval. Various points on the Labrador coast showed the same periodicity with identical peak years, indicating that the total population was controlled by the same factors. When a peak at any point gets out of step, the subsequent peak tends to fall back in line again. The peak years for red fox were identical with those for the arctic fox.
Prior to 1930, the fur statistics for Canadian arctic posts other than Labrador showed t a ten-year cycle for red fox, indicating that the reds probably did not breed to any extent north of the bush line but migrated to the Arctic in

EA-Zoology. Butler: Animal Population Cycles

times of population peaks in the south. Gradually both the northern range of the red fox and its numbers increased, as pointed out by Butler in his paper on color phases in the wild red fox (2). With this increase in numbers there is a suggestion s of minor peaks at four-year intervals imposed on a ten-year pattern, ^^ and, finally, interaction between four- and ten-year cycles. In Figure 2 the red fox collection at Great Whale River, which is in the tundra, is plotted for comparison with the collection at bush posts, three hundred miles to the south. The dotted line on the graph gives the southern collection taken at Eastmain and Rupert’s House and indicates a typical ten-year cycle. The peaks in this ten-year cycle came in 1917-18, 1925-26, 1935-36, and 1944-45. The solid line on the graph shows the Great Whale River collection which has the same characteristics as the collections from Ungava Bay and other points to the north. The cyclic picture here is much more complicated. It starts off with the typical ten-year cycle with minor peaks due to the four-year cycle superimposed in 1921-22 and 1930-31. By 1934-35, the population had begun to breed in the Arctic and the population fluctuations changed into the typical four-year cycle. The white fox ^ peaks ^ for the ^^ Eastern Arctic are marked on the same graph and it will be seen that in all cases the red fox peaks agree with the white fox. In 1925-26, a population peak was reached by the southern red fox and they went north. Food and other conditions were favorable, and the migrants were so prolific that the following year there was a real peak in the north which, although one year later than the southern peak, synchronized with the arctic fox peak. Similar conditions prevailed in 1945 and 1946. In 1935-36, the crash in the northern four-year cycle occurred at the same time as the peak population of the southern red fox so that the only influence s of the southern migrants was to make the catch larger than in previous ^^ or succeeding lows. ^ followed by 9a ^
Fig. 2. Colored fox returns showing four-year and ten-year cycles.

EA-Zoology. Butler: Animal Population Cycles

The colored fox is a predator of the white fox as well as a competitor in their common search for food. Because of this relationship, the magnitude of the cycle of an arctic fox population may be changed by the influx of red foxes. Since, however, the length of the cycle and the timing of the cycle are the same in both species, such predation and competition should not affect the interval between peaks.
Snowy Owl
The snowy owl, Nyctea nyctea , breeds in the tundra. When lemmings are plentiful the owl population increases rapidly and observers who may scarcely have been a snowy owl the previous season will report them as abundant. These owls, when lemmings or other food become scarce, quickly move to where there is a more plentiful food supply. Since most of the lemming populations diminish simultaneously, the snowy owl is often forced to migrate south in search of food. Many of these owls stay in southern Canada and the northern United States but some go as far as the southern United States and even to Bermuda. These migration years have been charted by A. O. Gross (8) and more recently by L. L. Snyder. The years of migration show the same periodicity as was present in the arctic fox and lemming populations. The year of migration corresponds to the peak year in the pelt collection of the arctic fox. Most of these southward movements have taken place in November and December; the last migration recorded was in the winter of 1945-46, following a peak lemming population in the years 1944-45.
Several other predatory birds have similar but less well-marked cycles. The northern shrike, Lanius borealis borealis , which breeds immediately south of the timber line, has a four-year cycle in the numbers that migrate southward in winter, the years of maximum abundance coinciding with those of the snowy owl and

EA-Zoology. Butler: Animal Population Cycles

arctic fox. Long-tailed skuas and rough-legged buzzards also seem to show the same type of cycle.
Ptarmigan
The willow ptarmigan, Lagopus lagopus , and the smaller rock ptarmigan, L. rupestris , are the birds of the greatest importance in the economy of the North. Widely distributed to the most northerly islands, and prolific breeders, they form an important part of the food of the fur-bearing carnivores. While not killed by the natives in any l large numbers, they are a great asset as they are virtually the only feathered game available during the severe months of winter. In winter the ptarmigan migrate southward to the bush country where their food of willow buds is available. Such migrations are not complete, some remaining behind all winter. These southern migrations show a period rhythm which is the same as the lemming’s. The peak migrations do not always occur all over the north country the same year, nor do all parts of the country witness peak migrations. There may be a peak migration in a certain section one year and then two peak migrations may miss this section s , so that there may not be a large migration in this section for about twelve years.
It is not certain that a large southward migration indicates a peak population of ptarmigan. It may indicate only that feeding conditions are bad in the Arctic so that a greater proportion of the population has migrated. There is some evidence, however, that the decrease in predation by arctic foxes, which accompanies an increase in the lemming population, does allow the ptarmigan to increase.
The ptarmigan population figures for Greenland give a slight indication of a cycle but there are much larger fluctuations occurring at about fifteen-year intervals which have obscured the short-term cycle.
In Norway the willow ptarmigan is called the rype. Its populations show a

EA-Zoology. Butler: Animal Population Cycles

definite four-year cycle in central Norway. The peak years and times of scarcity from 1869 to 1927 were tabulated by Kloster. The ^^ rype ^^ cycle became less regular after 1905, and several reasons have been advanced to account for this. Brinkman noted that disease was frequent in rype population and sought to explain it by the disappearance of predators, which might be expected to maintain standards of rype fitness by weeding out the diseased and weaker birds. Nordhagen suggested that there was a connection between the berry crop and the winter survival of the birds. The chief berries eaten are the crowberry, bilberry, bearberry, and c ^ l ^ oudberry, which fluctuate a great deal in abundance from year to ^^ year. unfortunately the data on the yearly fluctuations of the berry crop were not sufficient to correlate with those on the fluctuations of rype.
Disease appears to play a large part in the crash of the Lagopus cycle. The chief disease is coccidiosis, which is especially severe in the young birds. Since the outbreaks occur in widely separated parts of the Arctic at more or less the same time, and since the crash usually occurs at the same time as the crash in the lemming cycle, there must be some background influence which is common to all.
Mice
There are many species of mice in the arctic and subarctic regions. The most common genera are: Microtus , Clethrionomys , Peromyscus , Synaptomys , and Phenacomys . Most of these mice are not easily identified by the ordinary observer, so that when mice are reported as being abundant in any particular region it is not possible to tell with certainty which species is being reported. The interrelations between the various species are little known, but it has been shown that one species of mouse can be increasing while another is decreasing or very scarce. Because of the lack of trained observers in the region, little

EA-Zoology. Butler: Animal Population Cycles

is known about the fluctuations in the mouse populations. There appears to be a cycle in the mouse populations which corresponds with that of lemmings but it is not as definite nor as widespread. Certain areas will be reporting in– crease in both lemmings and mice, while adjacent areas will report increase in lemmings with no change or decrease in mice. Part of this lack of agreement may result from the fact that adjacent areas are often occupied by different species and that conditions which result in the increase in one species will not cause a corresponding increase in another species.
Marten
The marten (genus Martes ) usually exhibits the typical ten-year cycle of the Canadian bush country, but in Labrador, it has been shown by Elton, there is a four-year cycle. The great majority of good marten years either coincide with or come one year after the corresponding peak in foxes.
Some of the best marten country in Canada is found in the subarctic regions, especially in the Mackenzie River section. In 1850, this section was producing an annual catch of more than 30,000 pelts, but by 1930 the catch had declined to less than 5,000. Simultaneously with the decline in catch, the regular ten-year cycle, evident in the earlier figures, became less marked and harder to decipher.
The marten is a shy breeder and the young do not breed until they are two years old. This low [: ] reproductive potential makes it hard to account for the four-year cycle in Labrador by any reason except the supposition that in good mouse and fox years more young survive to maturity. Such a theory is borne out by the data which show that the magnitude of the fluctuations is not as large in the case of the other short-cycle animals.

EA-Zoology. Butler: Animal Population Cycles

Lynx
The Canada lynx (genus Lynx ) is the classi x ^ c ^ example of the ten-year cycle because its fluctuations are great in magnitude and the peaks occur at very regular intervals. The population changes are closely allied to those of the snowshoe rabbit which is its main prey. In fact, it is often cited as a typical example of the predator-prey relationship. An increase in the prey leads to an increase in the predator because of better feeding conditions. A crash in the prey population leads to a rapid decrease in the predator because their large population is confronted with a scarcity of food.
The lynx is confined to the bus y ^ h ^ country but it does occur in fair ^^ quantities in the Mackenzie River valley. There is definite evidence that the lynx is being overtrapped so that the cycle is now hard to perceive. There were well-marked peaks in 1916-17, 1926-27, 1935-36, and a less definite one in 1943-44. The cycle interval over the past 120 years is 9.7 years between peaks.
Mink
Mink (genus Mustela ) are found throughout the Subarctic and in favorable locations in the Arctic. They have a definite ten-year cycle and so far there seems to be little evidence for assuming that they have a four-year cycle in some localities. In northern Quebec there is often a minor peak four years after the major one, which may indicate the possibility of the short-term cycle. The peak years were in 1929-30 and 1940-41 in most of the Canadian Subarctic.
Muskrat
In the bush country, the muskrat (genus Ondatra ) has a well-marked ten-year cycle. There are some exceptions to the general cycle in any area and these are often found in a delta region where the fluctuating water conditions often modify

EA-Zoology. Butler: Animal Population Cycles

the normal cyclic picture. Trapping pressure also seems to play a more important part in the muskrat cycle than it does in the cycle of the other fur bearers. This may be due to the fact that a greater percentage of the total population is trapped than with other species, or it may be connected with the fact that one of the main limiting factors in a muskrat population is intraspecific strife.
The cycle in northern areas is not as definite as it is in southern ones possibly because most northern muskrats come from delta areas, or it may be the influence of the short-term arctic cycle. In the leading muskrat-producing area of the Mackenzie Delta, Arctic Red River had peaks in 1929-30, 1939-40, and 1945-46. At Fort MacPherson in the same locality, there were peaks the same years and other peaks in-between, indicating a modified type of four-year cycle. At Aklavik, the farthest point north on the delta, there were only two peaks during this period; they occurred in 1932-33 and 1940-41. Until more data are available it is impossible to say whether there is, or is not, a muskrat cycle in northern areas.
Other Animals
Caribou (genus Rangifer ) are in much better condition some years than others and they periodically change their migration routes. The cause of these changes is unknown and it has not been ascertained whether the changes occur regularly enough to be classed as cycles.
Arctic hares (genus P ^ L ^ epus ) fluctuate to a large extent, but there is not enough information to indicate a periodicity. They may have a four-year cycle like the lemming and fox, or else a ten-year cycle like the allied snowshoe rabbits in the forests of the south.
Polar bear (genus Thalarctos ) fluctuate locally, but this is thought to be due to the erratic movements of the sea ice rather than to over-all changes in the population.

EA-Zoology. Butler: Animal Population Cycles

E ^ r ^ mine (genus Mustela ) appear to have the same short-term cycle as the fox, but the peak years may not occur at the same time. The ermine population of arctic regions is so small and the information on population changes so meager that nothing definite can be said at this time.

EA-Zoo. Butler: Animal Population Cycles

BIBLIOGRAPHY

1. Braestrup, F.W. “A study on the arctic fox in Greenland,” Med.Grønland vol.131, no.4, 1941.

2. Butler, Leonard. “The genetics of the colour phases of the red fox in the Mackenzie River locality,” Canad.J.Res . vol.25, pp.190-215, 1947.

3. Chitty, Helen, and Chitty, Dennis. “Canadian Arctic Wildlife Enquiry 1942-43,” J.Animal Ecol . Vol.14, no.1, pp.37-45, May, 1945.

4. Collett, Robert. Norges Pattedyr . Kristiania, Ascheho n ^ u ^ g, 1911-12.

5. Elton, C.S. “Epidemics among sledge dogs in the Canadian Arctic and their relation to disease in the arctic fox,” Canad.J.Res . vol.5, pp.673-92, 1931.

6. ----. Voles, Mice and Lemmings . Oxford, Clarendon, 1942.

7. Freuchen, Peter. “Field notes and biological observations,” Degerb o ^ ø ^ l, Magnus and Freuchen, Peter. Mammals . Copenhagen, Glydendalske Boghandel, 1935, pp.68-278. Thule Expedition, 5th, 1921-24. Report vol.2, no.4-5.

8. Gross, A.O. “Snowy owl migration 1930-31,” Auk , vol.48, pp.501-11, 1931.

9. Pennant, Thomas. Arctic Zoology . London, 1784, vol.1.

10. Plummer, P.J.G. “Further note on arctic dog disease and its relation to rabies,” Canad.J.Comp.Med . vol.11, pp.330-34, 1947.

Leonard Butler

The Ten-Year Cycle in the Subarctic

EA-Zoology (William Rowan)

Controversial-needs editorial note before publication. See Dunber letter of april 19-1951

EA-Zoology (William Rowan)

THE TEN-YEAR CYCLE IN THE SUBARCTIC
LIST OF FIGURES

Scroll Table to show more columns

Page
Fig. 1 Lynx fur returns of the Northern Department, Hudson’s Bay Co., 1821-1913, and of equivalent area 1915-34 3a

EA-Zoology (William Rowan)

THE TEN-YEAR CYCLE IN THE SUBARCTIC
One of the most inexplicable phenomena of subarctic regions is what has come to be known as the ten-year cycle, a periodic oscillation of marked regu– larity in the numbers of many nonmigratory birds and various species of the smaller mammals. As its name indicates, the interval is approximately ten years between peaks of abundance which are conspicuous not only through the enormous numbers of animals attained but in the precipitate decline — the “crash” — that immediately follows. Thus juxtaposition of superabundance and scarcity creates such a striking contrast in the resident fauna in the brief period of only a a ^^ year or two that it cannot escape attention. The phenomenon is universally familiar to country dwellers of the Canadian northland.
The periodicity of the cycle has been worked out from fur returns, chiefly of the Hudson’s Bay Company, over the past 200 years. Fur records are subject to so many irregularities owing to weather and transport difficulties, market fluc– tuations, trapping activities, etc., that in their crude state they fail to reveal the underlying accuracy of the ten-year interval. Elton and Nicholson (2), how– ever, have sifted out these and other complicating factors from ^ the ^ Hudson’s Bay Co . ^ mpany ^ ^^ lynx returns over a period of 100 years, from 1830 to 1930; the true regularity of the cycle is consequently revealed and shows itself clearly in their graph (see Fig. 1). The long-term average periodicity works out at 9.6 years. The

EA-Zoology. Rowan: The Ten-Year Cycle in the Subarctic

relative constancy of the interval makes the ten-year cycle almost unique. Whereas the sunspot cycle, for instance, averages around 11 years, with the individual intervals between sunspot maxima varying from 7 to 15 years, the interval between consecutive rabbit and grouse peaks varies little from the ten-year span that gives the cycle its name. it may be 11 years on one occasion and 9 the next; an d ^ y ^ given district may be a year or even two in advance of another, ^^ or possibly behind, but the over-all picture for the Dominion of Canada is a cycle of extraordinary regularity of 9 or 10 years, with the long-term average somewhere in between.
Another significant aspect of the cycle is that it is confined to the northern regions of the world and is apparently synchronous around the globe. Since resident mammal life gets scarcer toward the pole and includes chiefly marine forms, such as seal and walrus, or large land mammals, such as wolf, polar bear, reindeer, and musk ox, which show no recognizable cycle as far as is presently known, or lemmings and voles with a four-year cycle, the ten-year cycle is not as conspicuous in the extreme north as farther south. It also gets less and less apparent with decreasing latitude, becoming difficult to detect south of the Canadian border. It still remains detectable, but shows relatively low amplitudes.
The most conspicuous member of the cycling fraternity, the snowshoe hare or rabbit ( Lepus americanus ), does not occur on the arctic islands and is scarce along the coastal regions, being replaced by the arctic or polar hare ( L. arcticus ). MacFarlane (5) was familiar with the ten-year cycle in the North but does not credit this latter species with the vast fluctuations of the snowshoe.
Through the vast Canadian forests and muskegs lying north of about latitude 52° N., the cycle reaches its most impressive proportions. Here, at ten-year intervals, incredible numbers are attained by the snowshoe hare, various species of

EA-Zoology. Rowan: The Ten-Year Cycle in the Subarctic

grouse, and many fur bearers. The amplitude varies from decade to decade with all of them as it does with the lynx (see Fig. 1). Lynx skins procured by the Hudson’s Bay Company, for instance, have run during the past hundred years from something below 10,000 at one peak to more than 60,000 at another. It is a striking fact that the introduced Hungarian partridge and ring-necked pheasant now show cycles synchronous with those of the native grouse.
A complete cycle, commencing at the peak year with prodigious numbers, shows the same general picture in each decade. In the case of the snowshoe rabbit, Seton (7) estimates that this species may attain a density of several thousand to the square mile in favored districts and a possible total population in an area the size of the Province of Alberta of perhaps 100,000,000. Because rabbits are normally of no commercial value, recorded figures are rare, but some exist in the Hudson’s Bay Company’s archives. Only a few posts on the shores of Hudson Bay apparently bothered with rabbit skins; the highest figure for any one year appeared to be 143,930, in 1867.
During World War II, however, owing to lack of shipping facilities from Australia, an unexpected demand for rabbit skins from Canada developed in the United States. From Alberta alone, mainly from the Peace River country, nearly 6,000,000 skins were shipped out in 1942, a peak year. contracts for the follow– ing season, however, when the cra c ^ s ^ h hit, had to be cancelled for lack of rabbits; ^^ there were not enough left to make collections worth while (6). The episode well illustrates the precipitate nature of the crash. A percentage generally survives the first year, but two years after the peak it may be impossible to find a single winter rabbit track over miles of territory. Slowly the animals recover, within ten years, to reach another stupendous peak, when the story repeats itself. In rough outline, this is the fate of all the cycling fraternity-
Fig. 1. —Lynx fur returns of the Northern Department, Hudson’s Bay Co., 1821-1913, and of equivalent area 1915-34

EA-Zoology. Rowan: The Ten-Year Cycle in the Subarctic

upland game birds, fur-bearing mammals, and such birds as magpies.
Grouse are probably never as abundant as rabbits, but they nevertheless exist in millions when the peak has developed. The greatest movement on record occurred in 1932, when a heavy influx of grouse, supposedly from the Hudson Bay region, flooded southern Ontario and Quebec (8). In spite of their vast numbers, the grouse never established themselves, for the subsequent years were crash years when a hoodoo seemed to pervade the North, and none of the cyclic species appeared to hold its own.
The outstanding feature of the ten-year cycle is its synchronous nature across Canada, affecting the whole Dominion approximately simultaneously. It is also synchronous as to certain species. The snowshoe rabbit and all species of grouse, for instance, come and go together. Many of the fur bearers lag behind by a year, or two years, or several years, but their peaks occupy the same relative position in the cycle ^ decade ^ after decade. The regularity is such as to make forecasts of oncoming peaks at least a reasonable p r robability.
Certain species offer peculiar complications, among the foremost being the muskrat and beaver. Both are dependent on water levels for subsistence. While the rat has to take things as they come, the beaver can, up to a point, conserve its own water supply and is less directly affected. Yet it is obvious that this dependence on water must modify the cycle in both species. The large mammals seem to be entirely immune, while mice, as already mentioned, have a much briefer periodicity of approximately four years. Foxes, and probably coyotes, which indulge in a mixed diet running from insects and mice to carrion, also show com– plications, depending possibly on their feeding habits. Possibly there is only one fur bearer that is wholly dependent on rabbit, the lynx, which has been described as “merely an animated rabbit.” The lynx chart (Fig. 1) is consequently

EA-Zoology. Rowan: The Ten-Year Cycle in the Subarctic

a close duplicate of the rabbit curve. Next to the lynx, the various species of grouse and the Hungarian partridge, and possibly the magpie, show an almost equally precise coincidence with rabbit numbers, and it is in this apparent correlation that the chief scientific interest of the ten-year cycle centers. What controls this remarkable periodicity? What is the common factor that keeps the cycles of so many species, including certain insects (e.g., grasshoppers), birds, and mammals, in incessant phase decade after decade?
The answer is unknown, but there seems to be two possibilities: ( 1 ) weather, and ( 2 ) some cosmic factor. The possibility that numbers alone are responsible for synchrony can be dismissed. In the first place, there is no reason to suppose that the rate of the increase of numbers would be the same in different areas or in different species, and secondly there is a clear disparity in peak populations from period to period. If 1,000 rabbits to the square mile form a sufficient crowd to permit disease to destroy the hordes, why should three or four times that number survive during some subsequent and much greater peak? It as least suggests the probability that the number does not lie in numbers alone. The impression is heightened by the fact that numerous diseases (not just one) affect rabbits and grouse at the time of crash, suggesting that re– sistance may then be at ^ a ^ low ebb, due possibly to vitamin or other deficiencies. The investigations of ^ Dr. ^ R. G. Green of Minneapolis on the snowshoe rabbit strongly support this view(3).
As to the possibility of weather as the common factor, the probabilities, are almost wholly against this explanation, for it is impossible to believe that the marked discrepancies of weather across the 3,000 miles of Canada’s breadth, with drought frequently prevailing in the west simultaneously with floods in the east (or vice versa), could do anything but disrupt the synchronization that is so typical of the cycle. It could hardly induce it.
A cosmic factor of some sort could theoretically provide the perfect key

EA-Zoology. Rowan: The Ten-Year Cycle in the Subarctic

to the ten-year cycle. The chief difficulty is that no such factor is known to exist. There are, however, some suggestive scraps of information, such as the records of atmospheric ozone, made in London and Paris during the years 1880 to 1910 (4). They show very marked peaks that coincide almost precisely with those of Canada’s rabbit population. Unfortunately, they are the only records of their kind in existence. Some authorities might say that the ozone records are European while the rabbit records are Canadian, and that there is probably no connection between them. When, however, one recalls that the grouse maxima and minima — as far as they are known — of northern Europe appear to coincide with those of Canada, there may after all be a correlation. If one adds to this the known association of the terrestrial ultraviolet supply with the ozone layers of the upper atmosphere, one can perceive a possible biological factor of great importance that could conceivably provide the key to the entire ten-year cycle. All of this must be considered speculative at present, but it suggests an approach that might be most remunerative.
The main facts of the ten-year cycle are now sufficiently well known for general acceptance. It is clear that the cycle is a phenomenon of the North with basic repercussions on human as well as animal life. When grouse, rabbits, and other smaller mammals in the Subarctic die off in millions at the time of crash, a vacuum is left in the fauna that cannot help but have a destructive effect on all predators (except marine forms), and man is a predator. The explorers and surveyors who today fly into the northern wilderness amply supplied with canned foods may be entirely immune, but such is not the case with the native population of nomadic Indians who depend neither on the sea (like many Eskimos) nor on the amenities of civilization for survival. They, like the rabbits, show a ten-year cycle of welfare: periodic starvation and disease are

EA-Zoology. Rowan: The Ten-Year Cycle in the Subarctic

equally their lot. From that viewpoint alone, the ten-year cycle becomes a matter of direct human concern.

EA-Zoology. Rowan: The Ten-Year Cycle in the Subarctic

BIBLIOGRAPHY

1. Christian, E.V. Unflinching . London., Murray, 1937.

2. Elton, Charles, and Nicholson, Mary. “The ten-year cycle in numbers of of the lynx in Canada,” J.Animal Ecol . Vol.11, no.2, pp.215-44, Nov. 1942.

3. Green, R.G. “Shock disease and the snowshoe hare cycle s ,” Science , vol.87, ^^ pp.298-99, 1938.

4. Huntington, Ellsworth. Mainspring ^ s ^ of civilization . N.Y., Wiley, 1945.

5. Mair, Charles, and Macfarlane, R.R. Through the Mackenzie Basin , Toronto, Briggs, 1908.

6. Rowan, William. The Ten-Year Cycle. 2nd ed. Enl. And new ^ (rev) ^ . Edmonton, ^^ University of Alberta, Dept. of Extension, 1948.

7. Seton, E.T. The Arctic Prairies . Lond., Constable, 1912.

8. Snyder, L.L. A Study of the Sharp-Tailed Grouse . Toronto, Univ. of Toronto Press, 1935. Toronto. Univ. Studies Biol.Ser . No.40.

9. Stefansson, Vilhjalmur. The Friendly Arctic . New ed. N.Y., Macmillan, 1943.

William Rowan

Fossil Mammals of the North

EA-Zoology (Edwin H. Colbert)

FOSSIL MAMMALS OF THE NORTH

At the present time the Arctic is the abode of a considerable mammalian fauna. In the north circumpolar region are found various mammals well adapted for life in these high latitudes. On land there is the usual ecological balance between herbivorous mammals and the carnivores that prey upon them, here exemplified on the one hand by such animals as reindeer or caribou, moose, musk oxen, mountain goats, and various rodents, especially lemmings; and on the other hand by wolves, foxes, and wolverines. In the oceans and along the shores are polar bears, seals, and various cetaceans, preying in part upon one another and in part upon fishes and other denizens of the waters. The great pageant of life is enacted here as it is in more southerly regions, but on a re– stricted scale.
There is reason to think that similar mammalian associations were characteristic of the arctic region in the immediate geologic past, while in more distant geologic epochs the mammalian life of the Far North was even more abundant and varied than it is now. This last statement is based upon two considerations. In the first place it is known from geologic evidence that climates were much milder during most of Cenozoic times than they are today. Until the beginning of the Pleistocene period,

EA-Zoo. Colbert: Fossil Mammals

when there was a widespread refrigeration of the Northern Hemisphere consequent upon the advance of the first great continental ice sheet, zonal climates would seem to have been less sharply established than they now are, and temperate conditions would seem to have extended to very high lati– tudes. Thus it can be supposed that characteristic temperate-zone mammals then lived in the Far North; that the adaptations to an arctic environment had not yet taken place.
Secondly, the fossil record shows that there were extensive intercom– tinental movements of mammals by way of a trans-Bering land bridge between Eurasia and North America during Cenozoic times. Such a northerly migration route would have led to the northern spread of mammalian faunas during the time the intercontinental land connection was in existence.
Unfortunately, however, there is virtually no fossil record of mammalian life in the Far North during the pre-Pleistocene phases of the Cenozoic era of earth history. Perhaps future explorations, especially in Siberia, may bring to light some truly northern pre-Pleistocene mammalian faunas, but at the present time we must face the fact that our knowledge of this aspect of mammalian evolutionary history remains a blank. In most northern regions the Pleistocene glaciations effectively removed any records of pre-Pleistocene mammalian life, if such existed. Therefore, our knowledge of former mammalian life in the North, as based upon an adequate fossil record, is confined to the Pleistocene period. Here the record is well known and rather widely spread. Pleistocene mammals are known from Alaska, parts and rather widely spread. Pleistocene mammals are known from Alaska, parts of northern Canada, Siberia, and the Scandinavian Peninsula.
The Pleistocene fossils of the north circumpolar region show that mammalian life during the great ice age was rather similar to what it is now,

EA-Zoo. Colbert: Fossil Mammals

but more extensive. In other words, there were various mammals then inhabiting the North which no longer exist; life in that region at the present time is impoverished as compared with the life of the Pleistocene. This is the result of a taxonomic reduction in the mammalian faunas of the North during the transition from Pleistocene to Recent conditions.
Ecologically, the Pleistocene period was a time of fluctuating environ– mental conditions in the arctic region. During the times when the great continental glaciers advanced southward, extreme arctic environments pre– vailed in the northern part of the earth. During the interglacial phases, when the glaciers retreated to the north, temperate environments extended into high latitudes. These alternations are reflected in the distribution of Pleistocene faunas by the extension of arctic mammals to the south during glacial stages, and to a lesser extent by the advance of temperate mammals to the north during interglacial stages.
Zoogeographically, the Pleistocene was a period of broad circumpolar mammalian distributions, a condition that has been continued into Recent times. Of course there were certain differences between the Pleistocene faunas of Eurasia and those of North America, as there are today, but generally speaking the resemblance far outweighed the differences. Con– sequently, in Pleistocene times as today there was a great holarctic fauna that inhabited the land masses around the North Pole. Similar or even identical species extended from northern Europe through northern Asia, across to Alaska, and through northern Canada.
In Pleistocene times various large herbivores were prominent in the faunas of the northern latitudes. Among the artiodactyls, or even-toed hoofed mammals, it is evident that large herds of reindeer or caribou

EA-Zoo. Colbert: Fossil Mammals

( Rangifer ) roamed the arctic tundras through northern Eurasia and North America. The musk ox, Ovibos , also had a wide circumpolar range in Pleistocene times; and during glacial stages this animal moved as far south as latitude 40° N., as did [: ] so many arctic mammals. In addition to Ovibos , there were other Pleistocene musk oxen belonging to several distinct genera distributed over the northern part of North America.
Bison were varied and numerous during the Pleistocene period, and the fossil record shows that at times they extended into northern latitudes. For instance, abundant remains of bison have been found in Alaska. These for the most part belong to a species now extinct, a very large bison with enormously long, curved horns. It should be emphasized here that the bison was of Eurasiatic origin and that the migration of this animal into North America (of which region it is commonly regarded as typical) took place across the trans-Bering land bridge during early Pleistocene times. As the bison spread through North America in the Pleistocene period it divided into a number of species, of which only one, Bison bison , has survived into Recent times. The sol d ^ e ^ surviving Old World bison, which was widely spread in northern Eurasia in the Pleistocene period, is the aurochs, Bison bonasus .
In this connection it should be said that other large artiodactyls in addition to bison were of Old World origin and cross ^ ed ^ the trans-Bering land bridge into the New World in early Pleistocene times. The reindeer and the musk ox, already discussed, are examples of such a distributional history. Another example is Oreamnos , the mountain “goat” (actually a rupicaprine antelope), found in Pleistocene deposits in Alaska. Another is the moose, Alces . Still others are the bighorn sheep, Ovis , and the wapiti, Cervus , these two not so generally arctic as the forms previously considered.

EA-Zoo. Colbert: Fossil Mammals

Certain artiodactyls crossed from northern Asia into Alaska in Pleistocene times, but never extended beyond the Alaskan area. Thus their distribution was dominantly palearctic or Old World, rather than holarctic or circumpolar. Prominent examples of artiodactyls that reached Alaska but got no farther in the Pleistocene period are the saiga antelope, Saiga , and the takin, Budorcas .
The perissodactyls, or odd-toe s ^ d ^ mammals, are represented in the arctic region durin f ^ g ^ the Pleistocene period by a horse, Equus , and by two rhinocero s ses, the woolly rhinoceros, Coelodonta , and the elasmothere, Elasmotherium .
The Pleistocene horse, Equus , originated in North America, from which region it migrated into Eurasia by way of the trans-Bering land bridge at the beginning of the Pleistocene period. Thus its origin and distributional history constitute a record just opposite to the records for the various artiodactyls which have been discussed. The horse is and probably always has been a subtropical to temperate-zone mammal, so that presence of fossil horses in Alaska is an indication of relatively warm conditions there at the time the horses inhabited that area. Perhaps hoses lived in arctic latitudes during interglacial stages, as probably did some of the artiodactyls such as the bison, the moose, and the takin. It is interesting to see that the horse eventually became extinct in the land of its origin, but continued successfully in Eurasia.
One of the characteristic Pleistocene mammals of the Old World was the woolly rhinoceros. This animal, related to the modern white rhinoceros of Africa, was clothed with a heavy covering of long hair, an obvious adaptation to arctic conditions. It lived in northern Eurasia, where fossils

EA-Zoo. Colbert: Fossil Mammals

have been found, including some mummified remains, and where excellent cave drawings made by early men have been preserved.
The woolly rhinoceros seems to have been perfectly adapted for wide wanderings across the circumpolar continents, yet this animal never crossed the Pleistocene land bridge into North America. The woolly mammoth, closely associated ecologically with the woolly rhinoceros in Eurasia, readily made the crossing into Alaska, from whence it spread widely across North America. If the mammoth could do this, why could not the rhinoceros? The failure of the woolly rhinoceros to enter the New World is one of the many puzzles of Pleistocene paleontology.
The other northern rhinoceros, Elasmotherium , a large and highly specialized rhinoceros characterized by a gigantic single horn on the forehead, was even more limited in its north Eurasiatic distribution than was the woolly rhinoceros.
Perhaps no mammal is more characteristic [: ] of the Pleistocene period in the Northern Hemisphere than the woolly mammoth, Elephas primigenius . This long-haired elephant was well known to early man in Europe, who has left us many drawings of the animal on the walls of caves. Moreover, frozen carcasses of the woolly mammoth have been found in Siberia and in Alaska, so we have a rather full knowledge of the soft anatomy of this beast, and even of the food it ate.
The woolly mammoth was closely related to the m modern Asiatic elephant, and was similar to the modern elephant in size and general proportions. It was characterized, however, by its heavy coat of reddish-brown hair, which enabled it to live in arctic climates, and by its unusually long, curved tusks, especially large in the males. The tusks of the woolly mammoth have

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been found in such great abundance in Siberia as to constitute an important source of ivory in past years. Evidently these animals were abundant during Pleistocene times; probably they traveled in great herds as do the Asiatic and African elephants of the present time.
As mentioned above, the woolly mammoth crossed from Eurasia into North America in early Pleistocene times and rapidly spread through the northern part of the New World. Consequently this animal had a wide distribution through much of the Pleistocene period.
[: ] Another proboscidean that reached northern latitudes in the Pleistocene period was the American mastodon, Mastodon americanus . This was a heavier animal than the mammoth, not so tall at the shoulder, and with gently cur b ^ v ^ ed tusks. Like the mammoth, the American mastodon had a heavy covering of hair, which enabled it to live in cold climes.
Remains of the mastodon have been found in Alaska (they are very abundant throughout the United States and southern Canada). There is some indication that the mastodon may have crossed into Siberia, but, if so, it did not become firmly established in northern Asia. It was essentially a North American mammal.
So far we have been concerned with the large herbivores that lived in the arctic or subarctic region during Pleistocene times. These animals certainly constitute the bulk of the northern Pleistocene faunas, partly because they were large. Their bones were less easily destroyed than were the remains of smaller mammals, and therefore were more li ^ k ^ ely to be preserved as fossils.
However, there obviously were many small mammals, especially rodents and hares, that lived in the arctic region during the Pleistocene period. For

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instance, lemmings are especially charactistic of the Arctic and the present time, and such was probably the case during the Pleistocene period. Beavers, Castor , have been found in the Pleistocene of Alaska, and it is reasonable to think that hares were abundant in the arctic in Pleistocene times, as they are today.
The record of the rodents is obviously incomplete, as it usually is in fossil faunas. Perhaps careful collecting in the future will add greatly to our knowedge of these small mammals as they lived in far northern regions during the Pleistocene period.
In a normal mammalian fauna the presence of large herbivores, such as perissodactyls and artiodactyls, is balanced by the presence of large and highly predaceous carnivores, while the presence of small herbivorous and omnivorous mammals like rodents and hares is balanced by the presence of small carnivores. Consequently, there were large and small carnivorous mammals in the arctic region similar to the carnivores that still live in this area.
Perhaps the most ubiquitous of the large carnivores was the wolf, Canis , which ranged through the arctic portions of Eurasia and North America, as it does today. This predator harried the large mammals, with the possible exception of such giants as the rhinoceroses, the mammoth, and the mastodon. Also present as an aggressive predator was the wolverine, Gulo . Foxes belonging to the genera Vulpes ^ Vulpes ^ and especially Alopex (the arctic fox) preyed upon the rodents and hares.
Bears, Ursus , were prominent in the northern faunas of Pleistocene times. These large carnivores were obviously as omnivorous in their feeding habits in Pleistocene times as they are today; consequently they must have

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lived upon berries, fish, small mammals, and occasional large mammals, as do their modern descendants. In Europe the giant cave bear, Ursus spelaeus , was a contemporary of Pleistocene man. It ranged into northern Europe, and it quite possibly extended into the arctic region, although the evidence for this is not definitive.
The extension of the ranges of large Pleistocene felines into arctic areas cannot be definitely decided, upon the basis of present knowledge. In Europe the cave lion, Felis spelaea , was found in England and comparable latitudes, but whether this large ca r ^ t ^ ever reached areas north of latitude 60° N. is at present unknown. The same might be said for the large cats in North America. However, it is probable that some small cats, especially the lynxes, inhabited the subarctic and arctic regions in Pleistocene times, as they do today.
The discussion up to this point has been concerned with the Pleistocene terrestrial mammals of the North. These are the forms for which the fossil record is most satisfactory. Something should be said now of the aquatic mammals, of which scattered fossils are known.
The evidence would seem to indicate that the aquatic mammals of Pleistocene times were essentially the same as they are today. In the deeper oceanic water were various cetaceans, while along the shores and the ice borders were several genera of seals, the walrus, Odobenus , and the polar bear, Tha r ^ l ^ a ^ r ^ ctos . The ecological relationships of these mammals — with each other and with their environment — are well known, and need not be elaborated at this place. One element in the Pleistocene and Recent aquatic fauna of the Arctic is now missing. This is the northern or Steller sea co s ^ w ^ , Rhytina , which became extinct a scant two hundred years ago.

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The end of the Pleistocene period in the Northern Hemisphere is generally defined by the retreat of the last great continental ice sheet – the Wisconsin glaciations in North America and the Wűrm glaciation in Europe. This glacial retreat evidently began about thirty or forty thousand years ago in the lower latitudes that mark the extreme advance of the ice. Naturally from south to north the disappearance of the con– tinental glacier occurred at progressively later dates, so that in the extreme northern latitudes this even took place only a few thousand years ago. Consequently, it is impossible to assign a precise date for the end of Pleistocene times.
However that may be, the fact is that the extinctions of certain large mammals, commonly regarded as taking place at the end of the Pleistocene period, actually did not occur until after the beginning of the glacial retreat. In fact, from the fresh condition of many Pleistocene fossils (with the frequent preservation in arctic regions of hair, muscle tissue, and other perishable details of soft anatomy) there is good reason to think that some mammals, now extinct, were living a very few thousand years ago. Therefore, properly speaking, their extinction was a post-Pleistocene event.
Whether the extinctions occurred slightly before or slightly after the close of the Pleistocene period is a matter of academic importance. The crucial fact is that various mammals did disappear from the earth during the gradual transition from Pleistocene to Recent times, and it is the disappearance of these mammals that marks the dictinction between Pleistocene and Recent faunas.
In the arctic regions of the northern continents the notable extinctions that took place near the close of the Pleistocene period were as follows:

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( 1 ) The woolly mammoth, widely distributed and very numerous in arctic Eurasia and North America became extinc e ^ t ^ everywhere. ( 2 ) :Likewise the American mastodon disappeared completely. ( 3 ) The woolly rhinoceros of northern Eurasia became extinct. ( 4 ) The elasmothere also disappeared; this extinction may have taken place well before the end of the Pleistocene period. ( 5 ) The horse disappeared from the New World, but it continued in Eurasia. ( 6 ) All musk oxen except the genus Ovibos became extinct. ( 7 ) Various species of bison disappeared, leaving only Bison bonasus in Eurasia and Bison bison in North America. (Farther to the south, still other large mammals became extinct, but they are outside the scope of the present discussion.)
Such extinctions as those listed above were indeed of great import, and since they affected large and numerous elements of the northern faunas they emphasize the differences between the faunas containing them and subsequent ones. They point up the fact that the modern arctic faunas are indeed impoverished as compared with those of Pleistocene times.
Why should there have been such broad extinctions at the end of the Pleistocene period? This is a question to which there is no satisfactory answer. The problem of extinction is a baffling one, and as yet we have but little insight to help us solve this problem. So far as we can see, many of the large mammals so typical of the Pleistocene period should have continued into Recent times, yet they became extinct completely and rather suddenly.
It is interesting that early man was associated with these now extinct mammals in the Old World, and that he certainly entered the New World before the Pleistocene or post-Pleistocene extinction took place. Was man a

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significant factor in any of these extinctions? This is something to speculate about. Whether or not man had anything to do with the extinction of mammals at the end of Pleistocene times, the sad fact is that he is a potent force that threatens to blot out various contemporanesou species of mammals.
BIBLIOGRAPHY
This list of references is not intended to be complete. Boule and Piveteau, and Romer, are general works on fossil vertebrates that include the genera of fossil mammals characteristic of the arctic Pleistocene. In Osborn, 1910, there is a long chapter on the Pleistocene, in which some attention is given to the Pleistocene of the far north. The large monograph by Osborn on the Proboscidea discusses the distribution of mammoths and other proboscideans in the circum– polar regions. The monograph by Tolmachoff, while concerned primarily with mammoths and rhinoceroses, does contain and extensive bibliography in which there are included works of a more general nature that have to do with the Pleistocene mammals of the arctic region. The same is true of the monograph by Skinner and Kaisen.

1. Boule, M., and Piveteau, J. Les Fossiles: Elements de Paleontologie . Paris, 1935.

2. Buckland, William. “On the occurrence of the remains of elephants, and other quadrupeds, in the cliffs of frozen mud, in Escholtz Bay, and in other distant parts of the shores of the arctic seas,” Beechey, F. W., Narrative of a Voyage to the Pacific and Beering’s Strait . London, 1831, vol.2.

3. Digby, George Bassett. The Mammoth and Mammoth-Hunting in Northeast Siberia . London, Witherby, 1926.

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[: ]

4. Fischer de Waldheim, G. “Notice sur les boeufs fossiles de Siberie,” Soc.Imp.Nat.Moscou, Bull ., seconde annee, art.6, 1830.

5. Gilmore, C.W. “Smithsonian exploration in Alaska in 1907 in search of Pleistocene fossil vertebrates,” Smithson.Misc.Coll . vol.51, pp. 1-38, 1908.

6. Hay, O.P. “The pleistocene of North America and its vertebrated animals from the States east of the Mississippi River and from the Cana– dian Provinces east of longitude 95°,” Carneg.Inst.Wash. Publ ., no.322, 1923.

7. ----. “The Pleistocene of the middl ^ e ^ region of North America and its verte– brated animals,” Ibid ., no.322A, 1924.

8. ----. “The Pleistocene of the western region of North America and its vertebrated animals,” Ibid ., no.322B, 1927.

9. Osborn, Henry Fairfield. The Age of Mammals in Europe, Asia and North America . New York, 1910.

10. ----. Proboscidea: a Monograph of the Discovery, Evolution, Migration and Extinction of the Mastodonts and Elephants of the World . N.Y., Amer.Mus.Nat.Hist., 1936-42, 2 vols.

11. Pallas, Peter Simon. “De ossibus Sibirias fossilibus,” Acad.Sci.Imp. Petropolitanse, Novi Comment ., vol.13, 1769.

12. Pavlova, Marie. “Description of fossil mammals collected by the Russian Arctic Expedition, 1900-1903,” Akad.Nauk.Classe Phys.Mat., Mem . Zapiski , ser.8, vol.21, 1906. (In Russian.)

13. Pfizenmayer, E.W. Mammutleichen und Urwaldmenschen in Nordost-Sibirien . Leipzig, 1926.

14. Quackenbush, L.S. “Notes on Alaskan expeditions of 1907 and 1908,” Amer. Mus.Nat.Hist., Bull . vol.26, pp.87-130. 1909.

15. Romer, A.S. Vertebrate Paleontology . Chicago, 1945.

16. Skinner, Morris, F., and Kaisen, Ove C. “The fossil Bison of Alaska and preliminary revision of the genus,” Amer.Mus.Nat.Hist. Bull ., vol.89, art.3, pp.123-256, 1947.

17. Tolmachoff, I.P. “The carcasses of the mammoth and rhinoceros found in the frozen ground of Siberia,” Amer.Phil.Soc. Trans ., vol.23, pt.1, art.1, pp. 11-74, 1929.

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18. Tscherski, I.D. “Beschreibung der Semmlung posttertiärer Säugethiere. (Die wissenschaftliche Resultate von der Kaiserlichen Akademie der Wissenschaften zur Erforschung des Janalandes und der Neusi– birischen Inseln in den Jahren 1885 und 1886 susgesandten Ex– pedition),” Akad.Nauk. Mem . ser.7, Vol.40, 1890.

19. Wilkerson, Albert S. “Some frozen deposits in the goldfields of interior Alaska. A study of the Pleistocene deposits of Alaska,” Amer . Mus.Novit . no.525, 1932.

Edwin H. Colbert

Siberian Mammoth

EA-Zoology (Eugene A. Golomshtok)

SIBERIAN MAMMOTH

CONTENTS

Scroll Table to show more columns

Page
Distribution 2
Preservation of Carcasses 3
Siberian Discoveries 4
Adams’ Mammoth 5
The Berezovka Mammoth 7
Other Discoveries 9
The Taimyr Mammoth 11
Physical Characteristics 12
Feeding Habits 14
Mammoth and Prehistoric Man 15
Early Description and Folklore 17
Ivory Industry 20
Bibliography 22

EA-Zoology (Eugene A. Golomshtok)

SIBERIAN MAMMOTH
The Siberian or woolly mammoth ( Elephas primigenius ) is an extinct circumpolar form of elephant which lived during Quarternary times. It is considered to be one of the final members of a long series of special and ascending mutations extending back through the entire Pleistocene period, and first recognized in the upper Pliocene of Italy as Elephas primigenius mut. astensia . It was described by Deperet and Mayer in 1923. More speci– fically, the north Siberian mammoth developed from the south Siberian or European animal in the same way that the latter had originated from Elephas trogontherii and E. antiques , i.e., through decreases of the dental plates and the thickness of layers of enamel.
The name “mammoth” corresponds to the Russian mamont or mamut , a word of uncertain origin. Besides numerous popular and semipopular designations, the scienti s ^ f ^ ic name for the northern mammoth underwent many changes. One of the earliest was the term Elephas mammonteus (Cuvier, 1796). Although the current designation E. primigenius was assigned by Blumenbach in 1806, other names ( E. primaevus , E. mammouth , E. jubatus , E. paniscus , E. giganteus and Mammut sibiricus ) were used for awhile. In 1924 the great American specialist H. F. Osborn insisted that the fundamental cranial and dental characteristics and proportions clearly distinguish the mammoth from the modern genus Elephas ,

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as typified by the Indian elephant. Consequently he introduced the generic designation Mammoteus , typified by Mammonteus primigenius — the Siberian mammoth which includes Mammonteus primigenius americanus , Mammonteus primigenius compressus , and other subspecies of the northern steppes and tundras. Another species, apparently existing at the same time as Elephas primigenius in European Russian, is a dwarf form E. stenotoechus .
Distribution
The mammoth, which appeared at the end of the Pliocene era, is definitely associated with cold and arctic conditions. During the inter– glacial periods, it migrated to the north, following the receding ice fields.
The Siberian mammoth (including American and Alaskan subspecies) appears to have ranged almost exclusively a north of latitude 40° N. from the British Isles across northern Europe (except the Scandinavian Peninsula), northern Siberia, Alaska, and northern Canada. In western Europe the mammoth followed the path of the glaciers and it is known that at one time it migrated as far south as the Mediterranean, where its remains have been found. In France mammoth remains have been found in archaeological sites from the Acheullean period to the upper Paleolithic period, after which it disappeared. In Siberia, especially on the New Siberian Islands, the mammoth existed during the last glacial and postglacial periods.
Conditions during the last glacial age were evidently favorable for the existence of the mammoth, and its disappearance coincides with the retreat of the glaciers toward the Baltic. Here the finds are much rarer, and farther northwest they disappear completely. Climatic and geological changes which subsequently took place are the main causes for the disappearance of the mammoth. The change in climate was accompanied by severe snowstorms which

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caused great quantities of snow to accumulate in the valleys, and the transformation of this snow into fossil ice prevented the mammoth from obtaining sufficient food. The extreme degree of specialization (incurving form of the tusks) and general clumsiness, which made the mammoth easy prey for Paleolithic hunters, may be considered as contributory factors in its extinction. Some writers, including George B. Digby, consider the large number of mammoth finds in Siberia to be the result of a migration of the animals fleeing from European hunters.
Preservation of Carcasses
Most of the remains of the mammoth have been found in Siberia, including complete frozen carcasses which were often in a surprisingly good state of preservation. Carcasses are usually found on top of bluffs buried near the surface in the layer of ever-frozen ground. The mammoth-bearing deposits sometimes reach a thickness of several tens of feet, but sometimes they are very thin. In northeastern Siberia layers of fossil ice underlie these deposits. Contrary to the assertion of some writers, mammoth e remains are never found in the fossil ice. Thus both stratigraphically and structurally the mammoth belongs to the youngest generation of the fossil elephant family.
There are many theories explaining the excellent state of preservation of mammoth carcasses in Siberia. One of the most widely accepted is that the animal may have fallen into the soft mud bed of a stream caused by the thawing of frozen ground and fossil ice. Such streams usually contain extremely soft and sticky mud, which is the product of loam layers covering the fossil ice. These mud beds are absolutely impassable. One trapped in a moving mud stream, the bulky body of the mammoth formed a sort of dam against which mud piled up

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until it covered the whole animal and suffocated him. Several recorded cases of death from asphyxia support this theory. Such must have been the fate of the Berezovka mammoth and one found near Moscow.
Another possibility is that the mammoths had drowned in rivers and lakes, especially during winter or early spring, and were either buried on the spot, or drifted downstream to be buried somewhere in the lower parts of the rivers within their deltas or embouchure sediments. Such was the case of Adams’ mammoth.
The widespread theory that mammoth flesh may be eaten by human beings has been discarded by many authors. The meat usually has a strong smell of putrefaction, the result of decay which set in after it was exposed. In addition, a slow process of decay was taking place in the carcass even when’ it was frozen. Soon after the animal’s death, by alternate exposure to weak sunlight and cold air during the day, and freezing at night, a slow drying-up process began, thus insuring preservation, but a sufficient degree of decay took place to make the flesh unpalatable to men, although dogs are reported to eat it readily.
Siberian Discoveries
The first western European mention of the mammoth is presumably that of Josias Logan, who in 1611 brought to London and “elephant” tusk which had been found by the Samoyeds in the region of the Pechora River. Later, in 1644, Mikhail Stadukhin reported that according to the Yakuts the islands now called New Siberian Islands were rich in “great elephant” bones.
The first written acount which introduced the knowledge of preserved mammoth bodies to western Europe was apparently that of Nicolas Witsen, in Noord en Oost Tartaryen , published in Amsterdam in 1692. A more detailed

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description was given by Evert Y a ^ s ^ brants Ides in Driejaarige Reize Naar China s (Amsterdam, 1704). He described the finding of mammoth remains in the Lena, Yenisei, and other rivers, when the ice broke up in spring and the floods, washing away the banks, revealed bones, tusks, and sometimes preserved flesh. He reported that his companion, whose business was the collection of mammoth ivory, “found the head of a mammoth uncovered by a fall of frozen earth. When they dug it out they found most of the flesh was putrefied…” According to Ides, the foreleg of the mammoth was found somewhere On the Yenisei and was taken to Turukhansk.
Subsequently, a number of other travelers (D.S. Messerchmidt in 1719. Khariton Laptev in 1739, and the Gmelin brothers in 1765) reported mammoth finds. The descriptions by Witsen, Ides, and others were generally dismissed as fantastic and incredible, and little attention was paid to them. A scientific work on the subject was published in St. Petersburg in 1771 with the title D e reliquiis animalium exoticorum per Asiam borealem repertio complementum .
In 1787 the body of a mammoth washed out by spring waters was discovered in an upright position near the village of Alazeia, 65 miles from the town of Sredne-Kolymsk. It retained its skin, and, in some places, its hair. Unfor– tunately, Lieutenant Sarychev, who received the news of the discovery, could not reach the place before the carcass had disappeared.
Adams’ Mammoth . The famous Adams’ mammoth, the basis for the first scientific description of the animal, was found by a Tungus hunter in 1799 in the delta of the Lena River. The body was exposed gradually and was reported to be nearly complete. In the fifth year after the discovery the tusks became exposed and the Tungus hunter chopped them off and sold them.

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It was not until 1806 that the zoologist M. F. Adams, a member of the Russian Academy of Sciences, arrived at the site of the discovery, only to find that the body had suffered a good deal from the ravages of wolves, bears, and dogs. It was a long-maned male animal. The trunk and tail, which had been seen and sketched by an eyewitness, Boltunov, were gone. One of the ears, measuring over ten inches in length, was found, and one eye, which still retained its color. The color was lost, however, during the drying process later on. The skin on the head, on one foot, and on the side of the animal which was lying on the ground were the only soft parts which survived. It is said that the Yakuts had fed their dogs with mammoth flesh during famines.
Adams brought with him almost a complete skeleton, with ligaments, about 35 pounds of hair, and some entrails. He purchased two other tusks to replace the missing ones, which were reported to have been ten feet long and to have weighed 360 pounds. Samples of the hair of this ^ m ^ ammoth have found their way to various museums in Europe, and the skeleton is in the Museum of the Russian Academy of Sciences. Between the teeth and in the stomach, needles of coniferous plants and young shoots of leaf-bearing shrubs were found. The most important results of Adams’ expedition were published in the Journal du Nord, vol. 32, in St. Petersburg, 1807. It was established that the mammoth had been found in permanently frozen ground.
In 1805 Captain Potapov, who was bringing supplies to the Kruzenstern hydrographical expedition, saw the body of a mammoth on the shores of the Arctic Sea. He collected some hair and sent it to Blumenbach. About 1820 a mammoth was found in the Yakutsk region, and a well-preserved leg was seen by Schrenk. In 1839, on the steep shore of a tundra lake south of the Yenisei,

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a body, less preserved than Adams’ mammoth, was found. It was brought to the Moscow University Museum where it was studied by Gliebov. A. F. Middendorf, in 1843-44, found a fairly complete skeleton in a poor state of preservation on the Taimyr Peninsula. Other discoveries before 1910 are noted on a map in Les Mammouths de Siberie , by E. W. Pfizenmeyer (Paris, 1939).
The Berezovka Mammoth . The most important discovery, up to 1947, was that of the well-preserved body of a mammoth found in northeastern Siberian by a Lamut hunter, Tarabykin, in 1900. It was located 200 miles northeast of Sredne-Kolymsk on the Berezovka River, the right tributary of the Kolyma.
At first Tarabykin found a tusk which weighed 166 pounds, and then nearby he found the head of a smaller specimen with a smaller tusk. He cut the tusk off and sold it to a Russian Cossack who reported the find to the authorities. As a result of the report, an expedition of the Academy of Sciences set out from St. Petersburg to Yakutsk in 1901, whence they traveled over 1,500 miles to the site of the discovery, a three months’ journey. It took a month to complete a skilful excavation.
The carcass was found in the best imaginable state of preservation and was almost untouched by wild animals. The mammoth’s pose, with the forelegs thrust out, suggested that it had made great efforts to free itself from some trap into which it had fallen. It was thought that perhaps it had broken through a crevice or plunged into soft ground and died from the resulting injuries. This theory is supported by the fact that the pelvis, a forefoot and a few ribs were found broken. The possibility of death by suffocation in the mud still exists.
This specimen was a young male animal with a small tusk (the other tusk had been removed before the expedition arrived.) The trunk and the skin on

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the head had been eaten away by animals, but most of the skin was preserved and was suitable for tanning. Most of the fur had fallen out but lay near the body and was gathered up. The flesh was so well preserved that it was soft and its outward appearance reminded one of boiled beef. The dogs ate it willingly. The remains of partially masticated food were found between the teeth. The tongue was preserved, but it had dried out. The brain, liver, and alimentary canal had decayed. The analysis of dried clots of blood showed that the red blood corpuscles and hemoglobin had completely disappeared. The stomach with its contents was completely preserved.
The skeleton was complete except for the missing tusk. The Zoological Museum of the Academy of Sciences undertook a series of studies of the remains, which were brought there in a frozen state. They also prepared the specimen for mounting in the original pose in which it had been found. The results of the study of the frozen portions of the tissues were published in Transactions of the Academy of Sciences.
A number of [: ] scientists participated in the study of the Berezovka mammoth. The general report was made by the head of the expedition, O. F. G ^ H ^ er g ^ t ^ z. Osteology and tooth structure were described by B. Zalenski, geology and stratigraphy of the find by I. P. Tolmachev, anatomy by N. M. Maliev, microscopic investigation and histology by F. A. Bialinitski-Birulia, and the contemporary flora by V. N. Sukachev. A good over-all description of the discovery and excavation appeared in E. F. Ffizenmayer’s Les Mamouths de Siberie .
The results of the histological and microscopic studies of the material brought to St. Petersburg demonstrated that one cannot speak of the complete preservation of tissues, at least not in the case of the Berezovka mammoth.

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It was established that some changes took place due to decay, though at t a much slower tampo than normally, perhaps soon after the death of the animal. Some of these changes are of the type which can be expected when the process of decay occurs at a low temperature with considerable moisture present and without a sufficient amount of air. These would correspond to the conditions to which the body was exposed as it lay for a long time in cold running water. It is probable that the temperature of the surrounding ground was periodically raised and lowered, allowing a certain amount of decay. Later, in the permafrost condition, the thawing stopped altogether and the body remained frozen until historic times, when the shores of the lakes and rivers, undercut by water, fell and exposed the Berezovka mammoth as well as other specimens.
On the basis of Zelenitski’s description, O. P. Hay in his Observations on Some Extinct Elephants , Washington, D. C., 1912. Identified the Berezovka find as a new species, Elephas beresovkius , a term which was not adopted by the Russians.
Other Discoveries . Subsequently a number of finds of carcasses in a good state of preservation were reported, but either they proved to be false, or the investigating group arrived too late to gain any valuable knowledge. At present more than forty localities are known where parts of frozen mammoth carcasses have been found. In addition to these frozen specimens, a large number of mammoth bones and more or less complete skeletons are known to have been found in Siberia and the European part of the U.S.S.R.
In 1924 an unknown Tungus hunter found the well-preserved and of a mammoth’s trunk in the permanently frozen soil on the shore of the Bolshaia Bazaikha River in the Kolyma district. The trunk passed from hand to hand,

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and someone cut off the tip and kept it, throwing the rest away. This tip, the tissues dried and hardened, was accidently discovered in 1929 among the curios of a housewife in the town of Sredne-Kolymsk, and was brought to the Zoological Museum of the Academy of Sciences.
The specimen measured 28 centimeters in length, and was dark brown in color. The inner side was darker than the outside. While it was similar in general to the trunk of the modern elephant, it had certain significant anatomical differences, which may be considered adaptations to a different mode of living. The “lip” at the extreme and tapers gradually to form a finger-like appendage without any abrupt contraction. This appendage is much longer and wider than that of any modern elephant species, and gives the entire tip a two-lobed structure, which enabled the mammoth to pluck large bunches of grass and moss with greater ease.
The original hairy covering had long since disappeared b y ^ u ^ t microscopic examination showed where two sizes of hair were once rooted. There was no hair on the inner or outer surface of the lips.
In addition to its paleontological interest, the trunk tip confirmed anew the keen powers of observation of the ancient cave men, as well as their ability as accurate draftsmen. The bilobal trunk shown in drawings of memmoths in Paleolithic caves of Europe had always puzzled archaeologists, who considered these representations to be inaccurate, in conformity with current paleontological concepts which assumed the shape of the tip to be like that of the modern elephant.
On the America a n continent frozen remains of the mammoth have been found, either washed out by rivers or uncovered by mining operations. One of the most interesting finds was the skin of the face, the trunk, and the

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foreleg of a baby mammoth found in Fairbanks Creek, Alaska, on August 28, 1948. This is described by Harold E. Anthony in Natural History , September 1949.
The Taimyr Mammoth . In 1948 the Leningrad Academy of Sciences was informed that the skeleton of a mammoth had been discovered on the Taimyr Peninsula, and an expedition was sent in May 1949 to make a thorough investigation.
The skeleton was found to be complete except for the tail vertebrae and a few ribs, which may have been torn away by carnivorous animals soon after the mammoth died. The bones were almost entirely devoid of flesh; some tufts of hair, fragments of hide, and a few soft parts were found, but there was no trace of the stomach.
A permanently frozen peat bog and a deposit of gravel surrounded the skeleton, and layers of peat adhering to the bones indicated that at the time when the animal was buried the development of peat bog in the tundra was much more rapid than at the present time; therefore the climate must have been warmer. Further proof of this was shown by the presence of large trunks of willow trees in the deposits of the first terrace of the Mamontova River. The investigators came to the conclusion that the gravel in which the skeleton was found did not represent the spot where the animal had originally lai a ^ n ^ . It must have died on a river terrace and fallen when the river changed its course and washed the bank away.
For the first time the discovery of mammoth remains was accompanied by a detailed study of conditions of stratification in the Quaternary period and a comparison of vegetation with that of the past. According to these investigations, it appears that the last habitat of the mammoth was charac– terized by a type of tundra and forested tundra which is now found in more southerly regions. Scientists hope that the material collected by this

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expedition will help to clarify the hitherto unsolved problems of the conditions of life of the mammoth and the reasons for its disappearance.
Physical Characteristics
The mammoth was an elephant-like animal about 10 feet in height. Its body was built in different [: ] proportions from that of the present-day Indian elephant; the head constituted about one-third of the body. The head was pointed, with a bulbous or well-rounded forehead rising like a sloping dome at the top. The entire length of the skeleton of Adams’ mammoth, from the fore head ^ ward ^ ar e ^ c ^ of the tusks to the tuberosities of the ischium was 15 feet, the vertebral column was 9 to 10 feet long. Other measurements were as follows: height ^ , ^ 9 feet 3 inches; tusk, 9 feet; humerus, 3 feet 4 inches; ulna, 2 feet 11 inches; femur, 3 feet 10 inches; tibia, 2 feet 4 inches. The cranium exhibits fore and aft compression resulting in bathyc ^ e ^ phaly and hypsicephaly.
The body covering consisted of two types: ( 1 ) soft, wooly hair about an inch long, varying from faded yellow to h ^ y ^ ellow-brown; ( 2 ) much coarser hair, dark rust-brown in color, often reaching a length of 20 inches. This longer hair was especially well developed on the sides of the lower part of the body and formed a fringe, similar to that of the yak, which covered the cheeks, shoulders, rump, and legs down to the horny ends of the toes.
The skin was unusually thick, with a 3-inch layer of fat just below its surface.
The tusks were from 9 to 10 feet in length and were curved to form the shape of a crescent moon. In adults and ends of the tusks curved inward and in many cases they crossed in the middle line. They could not used

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for digging, fighting, or uprooting, because of their extreme curvature. These tusks weighed from 180 to 200 pounds, on the average.
The trunk was extremely well developed, and its tip was different from that of the Indian elephant, with the two-lobed structured which has been described.
The ear was somewhat smaller than that of the Indian elephant; it measured 15 inches in length and 6 1/2 inches in breadth, and was covered with a thick coat of short wool and long hair.
The tail was conical in form and about 23 inches long. It was about 14 inches in diameter at the root and ended in a sharply pointed bunch of stiff dark-colored bristles about 8 to 14 inches in length.
The chewing apparatus consisted of massive teeth weighing about 18 pounds each. These teeth had thin layers of enamel on the chewing surface which formed from 14 to 16, and 18 to 28 sharp corrugated plates on the second and third molars, respectively. These were like millstones and were used to masticate coarse food like twigs and grasses. The worn mid– coronal surface is sometimes fully horizontal, i.e., at right angles to the perpendicular ridge plates. The typical ridge formula of the third molar is 24/24, or near that of the Jeffersonian mammoth’s 25/25, but the third molar is shorter and deeper. As compared with Elephas columbii and Archidiskodon imperator the 10 ridge plates of Elephas primigenius are compressed into a line 100 millimeters in length with a minimum of 8 plates in 100 millimeters and a maximum of 13 plates in 100 millimeters.
The E. primigenius differs from Parelephas jeffersonii , the more southerly species, which was, perhaps, also of a hairy type but lacked the heavy undercover of wool, and Archidiskodon imperator , the still more southerly

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species, which was hairless and had great incurved tusks attaining gigantic size.
In a period marked by frequent torrents of sleet, rain, and snow, the sloping hindquarters of the mammoth were well adapted to shedding water. The bulbous accumulation of fat on the forehead and the layer of fat under the skin served as a supply of energy for the winter months. During the summer these portions were extremely full in appearance, but by spring they assumed their normal shape.
Feeding Habits
Valuable information about the feeding habits of the mammoth was obtained when a large wad of food was discovered between the upper the lower teeth of the Berezovka mammoth. Apparently the animal met its death quite suddenly and did not have time to swallow the food. In addition, the entire stomach, containing about 12 kilograms of undigested food, was found. An analysis of this material permitted identification of the contemporaneous flora, which consisted of foxtail grass, sweet grass, several types of sedge, wild thyme, field oxytrope and alpine poppy, also Hypnum fluitans , Aulacomniun turgidum , Alopecurus alpines , Beckmannia cruciformis , Agropyron cristatum , Hordeum violaceum , Carex lagopina , Ranunculus scros , and Oxytropis sordida . All of these species are typical representatives or the meadow flora of northern Siberia at the present day.
In winter the animals probably ate the bark and twigs of birches, fir trees, and other types of low-arctic vegetation. They grazed in large herds, mostly on lowlands near watering places.

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Mammoth and Prehistorin Man
The mammoth must have attracted prehistoric man by the great mass of flesh, fat, brain, and bone marrow which its carcass could supply. It was hunted extensively, as can be seen from the numerous remains found in Paleolithic sites of Europe and Siberia.
Paleolithic man hunted the mammoth with club, da ^ r ^ t, and spear, and probably most often stalked young, sick, or trapped animals. The unwieldy bulk of the mammoth, and the fact that its tusks were useless for defense, made hunting relatively simple, Use was made of pits dug on paths leading to watering places, with sharp-pointed poles at the bottom. Sometimes the animals were driven toward narrow gorges where they would fall easy prey to the hunters. The use of vegetable prison has been suggested by some writers, but cannot be proven.
Hunting had to be done by a group, perhaps members of the same camp site; otherwise it would have been impossible to transport portions of mammoth bodies weighing up to one ton to the sites where, on a number of occasions, articulated bones of the animal have been found. Sometimes the animal was killed too far away from the settlement and a feast took place on the spot. Remains of such a temporary camp were found near Tomsk, with an almost complete skeleton, traces of fireplaces, and numerous flint implements.
A fondness for brain and marrow is the only explanation for the presence near camp sites of the long bones and skulls of mammoths, extremely heavy and difficult to transport, which were always found split to extract the marrow. Often large stones which had been used to split these bones were found nearby.
In the Aurignacian and Solutrean sites of eastern and central Europe

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and Siberia large piles of mammoth bones, representing dozens or even hundreds of animals, were found. These accumulations of bones are too large to be merely refuse piles; it is believed that they represent a supply of bone used for some definite purpose. On some occasions (for example, on the site known as Kostenki I, Central R.S.F.S.R.) such supplies were kept in a specially dug-out pit.
The mammoth was a source of building material and fuel, as well as food, for Paleolithic man. The use of bones as material for walls of semisubterranean huts, and as fuel for fireplaces, was demonstrated in a number of sites. We may assume that mammoth skin and hair were also used in primitive economy.
Mammoth tusks were highly valued because they furnished excellent material for bone implements, the manufacture of which began to develop in the upper Paleolithic period. Piles of tusks stored separately from other bones were found in western and eastern European loess sites. The use of mammoth ivory for implements and carvings was widespread. Large adzes (presumably for splitting wood), spear points, needles, and a variety of decorations such as pendants, plaques, bracelets, and earrings have been found. A large number of animal and human figuriess carved out of mammoth ivory appear in archaeological sites all the way from western Europe to eastern Siberia: several such female representations were discovered at Malta, near Irkutsk.
Paleolithic man was well-acquainted with the mammoth and he pictures it in carvings, engravings, and paintings. The caves of southern France furnish excellent examples of mammoth representation in the form of line engraving on the soft limestone walls of caves, monochrome outline drawings, and even polychrome paintings with gradations of color and shading. The figures outlined in black are the earliest. Next in age are drawings washed

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in red, which are followed by incised drawings. Finally there are poly– chromes, boldly sketched in three colors and representing the finest example of Paleolithic art. The most famous of these are from French caves at Combarelles and Font-de-Gaume.
Early Descriptions and Folklore
Early concepts of the mammoth visualize it as a gigantic subterranean animal. Thus K’ang-his (1654-1722) emperor of the Manchu dynasty in China, quoting older sources, called the mammoth fen-shu or [: ] yu-mu (the hidden mouse) or shu-mu (mother of mice). He said that the Manchu call it ice rat and adds that “in the north in the land of the Russians, these rats, as large as elephants, live in the ground. When air, sunlight or moonlight touches them they die immediately. The flesh of this animal is ice-cold and very efficaciou x ^ s ^ in fevers. There are fen-shu which weigh up to 10,000 pounds. Their teeth are like those of elephants and the people of the north make vessels, combs, knife-handles and other articles of them.”
Another Chinese book (1771) says: “The ice-rat, or the mountain stream rat lives in the ground under the thick ice of the northern regions. Its flesh is edible. It hair is several feet long and is used to make a woven material which keeps out dampness.”
Early European travelers such as Witsen (about 1692) concluded, on the basis of native information, that “the great puzzle of g the earthquake is easily explained by the movement of this gigantic rat.” He quotes the Muscovites as believing that the mammoth is a southern animal, and considers it proof that the earth made a complete turn.
Russians believed that the mammoth lived underground, split the waters when crossing them, and brought misfortune to anyone who saw it. Some of

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their traditions ^ ^ describe it as the animal brought by the Deluge from India to Siberia, or as the vestige of the “march of the Israeli a tes,” who “passing through the warm lands brought along a great many elephants who perished from the bitter cold of Siberia.” It is also said that the mammoth was [: ] so large that Noah’s ark could not harbor it and it perished during the Deluge. Another story was that the mammoth was saved by Noah in the ark, but perished later, after it left the ark because the soaked soil could x not bear such a ponderous animal and it sank into the underground where it has been found. Others saw in the mammoth the remains of the elephants which ran away from Hannibal.
Numerous Siberian tales of the mammoth were recorded by Tatshchev, one of the first and best investigators of this animal, in 1730. He cites native beliefs that the mammoth is an underground animal who feeds on the earth, lives far from human habitations, and dies when exposed to daylight.
Siberian Ostiaks believe that the mammoth exists now, lives in the lakes, and can pass underground from one lake into another. The Yakuts call it “water bull,” the Tungus call it the giant animal, and Yukaghirs consider mammoth remains to be bones of a gigantic bird.
Like most of the other Siberian natives, the Chukchi are afraid of mammoth remains and consider the animal to be a kind of evil spirit. They believe that if mammoth tusks are seen above the ground it is a bad omen, and if incantation is not uttered something unpleasant will happen to the observer. In former times the search for mammoth ivory was taboo. Later the finder of a mammoth tusk was required to make various sacrifices to the spirit of the place in which the tusk was found, in order to appease it. According to one of their folk tales, some Chukchi men found two mammoth tusks

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protruding from the earth. They began to beat the drum and performed several incantations, and the whole carcass of the mammoth came into sight. The people ate the meat, which was very nutritious, and they lived on it all winter. When the bones were stripped of all meat they put them together again, and in the morning they were again covered with meat.
The widespread belief in the medicinal value of mammoth tusks was not confined to North America, Siberia, and the European part of Russia, but was also found among Romans, Venetians, Mexicans, Indians, Spaniards, French, Italians, Germans, and peoples of the Arabic world. Until the seventeenth century, [: ] European medical men believed in the [: ] curative value of the tusks. The Roman emperor Augustus collected them for his Museum on the Island of Capri.
Mammoth tusks were venerated either as the remains of kings or saints or as freaks of nature, and as such were placed at the entrance of churches in western Europe (Helmstaadt, Haudersheim, Brunswick, and Vienna.)
Fantastic reconstructions of mammoths were made both by scientists and charlatans. Leibniz recreated it as the one-tusk animal Unicorne fossil . The physician Masurie in 1613 showed mammoth bones for money as the bones of the Tevtobokh, the kin d ^ g ^ of the Kimvrs.
Even when the difference between the elephant and the mammoth was established by Cuvier, a number of theories suggested the original home of the mammoth to be in Central Asia – the convenient home of many puzzling things — and it was supposed that mammoth bodies “floated down” Siberian rivers to the northern area, where no one could imagine that any such animal could have lived.

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Ivory Industry
While the fossil ivory (called in Russian mamontova kost - mammoth bone) was long known in Siberia, it did not become a regular object of trade there until about 1582, the date of the conquest of Siberia by Yermak, and it first reached western Europe in 1611, being brought by Logan who purchased it from Samoyeds on the Pechora River.
The use of mammoth tusks a material for manufacturing objects is quite old. It was known to the Greeks and Arabs who bought some of it on the Volga in the ninth and tenth centuries. Another route was through Archangel and Novgorod, and in the eighteenth century Khiva and China became the centers of trade. Collection of mammoth ivory was very widespread and the Tears tried to establish a monopoly over this trade.
A great hunting ground for mammoth ivory was discovered in the New Siberian Islands in 1712. In 1809, the Cossack Sannikov brought 9,000 pounds of ivory, representing 80 to 100 animals, from there. Another collector in 1821 brought 18,000 pounds from the same locality. During the first half of the nineteenth century about 36,000 pounds of ivory were sold yearly in Yakutsk, and between 1825 and 1831 this amount reached 72,000 pounds. In addition about 3,000 pounds each came from Obdorsk, Tobolsk and Turukhansk. Middendorf estimated that in the course of eighteenth and nineteenth centuries the tusks of more than 20,000 mammoths were sold, and Nordenskiöld considers this figure too low. Taking the latest figures into consideration, Tolmachev estimates that since the time of the Russian conquest at least 46,750 animals must have been discovered.
Siberian natives have used mammoth ivory as material for various objects. The Yakuts make rings, combs, boxes of fine fretwork, powder horns, match

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boxes, knife handles, parts of harnesses for horse, reindeer, and dogs, and many other articles from ivory. Also wall executed are their models of summer huts and of sledge with either an ox or a reindeer harnessed to them. The Yakut combs, of two types, are especially well made. The smaller one 2 1/2 inches long and 1 1/2 inches wide is usually cut from the “bark” of the tusk, and the side edges in many cases consist of a gray– brown, bluish or red-brown strip of the actual surface. The longer ones measure 6 by 4 inches and are decorated in the middle with fretwork designs, initials of the carver or animal representations. Tobacco pipes have knife-carved ivory handles and barrel-shaped bowls lined with native steel beaten very thin. Ivory hunting-knife handles have grooves cut in them and are filled with rings of blackish horn to prevent slipping.
Koriaks make fishhooks, harpoons, and arrowheads out of ivory. The Chukchi make an ivory coat of mail out of plates sewn together with leather thongs, and arranged in three rows. [: ] Each plate measures 12 centimeters long, 4 centimeters wide, and about 1 centimeter thick. They also make buckles and hooks.
Eugene A. Golomshtok

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BIBLIOGRAPHY

1. Adams, M. “Relation d’un voyage a le mer glaciale et decouverte des restes d’un mammouth,” Jour.du Nord , vol.32, 1807.

2. Anthony, H.E. “Nature’s deep freeze,” Nat.Hist . 1949 (A separate.)

3. Bialynitsky-Birulia, F.A. Histological and Microscopical Observations on the Tissues of the Berezovka Mammoth . St. Petersburg, 1907. (In Russian.)

4. Digby, George Bassett. Mammoth and Mammoth-Hunting in Northeast Siberia . London, Witherby, 1926.

5. Efimenko, P.P. Primitive Society . Leningrad, 1940. (In Russian.)

6. Golomshtok, E.A. “La trompe du mammouth Siberien,” L’Anthropologie , vol.42, no.5-6, 1932.

7. ----. “The old stone age in European Russia,” Amer.Phil.Soc. Trans ., vol.29, part 2, 1938.

8. Hay, Oliver P. Observations on Some Extinct Elephants . Wash., D.C., 1922.

9. Illarionov, V.T. Mammoth . Gorkii, 1940. (In Russian)

10. Osborn, Henry Fairfield. Procboscidea . Amer.Mus.Nat.Hist., N.Y., 1942, vol.2.

11. Pevlova, Mary. “Description of fossil mammals collected by the Russian Arctic Expedition, 1900-1903,” Akad. [: ] Nauk. S.S.S.R., Classe Phys.Math., Mem.Zapiski , ser.8, vol.21, no.1, 1906. (In Russian.)

12. Pfizenmayar, E.W. Les Mammouths de Siberie . Paris, Payot, 1939.

13. Tikhomirov, B.A. “Ne Taimyr za momontom.” (To Taimyr after the mammoth.), Vokrug Sveta , no.8, p.42, 1950.

14. Tolmachoff, I.P. “The carcasses of the mammoth and rhinoceros found in the frozen ground of Siberia,” Am.Phil.Soc., Trans ., vol.23, pt.1, pp. 12-14, 1929.

15. Zenzinov, V.M. On the trade in the North of the Territory of Yekutsk . Moscow, 1916. (In Russian.)

Eugene A. Golomshtok

Invertebrates

Arctic Insects

EA-Zoology (Marie Hammer)

ARCTIC INSECTS

CONTENTS

Scroll Table to show more columns

Page
Plecoptera, Neuroptera, and Trichoptera 6
Orthoptera 7
Hemiptera 7
Coleoptera 8
Lapid e ^ o ^ ptera 9
Hymenoptera 10
Diptera 12
Bibliography 14

EA-Zoology (Marie Hammer)

ARCTIC INSECTS
Little is known as yet concerning insect life in Arctic Canada, Alaska, Greenland, Spitsbergen, northern Scandinavia, and Siberia. The best-explored of these areas are Greenland and northern Scandinavia, from which many expeditions in the course of years have brought back extensive material, giving a relatively good knowledge of the insects of these countries. Apart from this, nothing, or almost nothing, is known about the insects’ wintering conditions, their nourishment, copulation, egg– laying, etc., and there is even less information from the remaining arctic regions. The Canadian Arctic Expedition (1913-18) returned with relatively comprehensive material, although it represented only a fraction of what one cou p ^ l ^ d expect to find there. Only 2 species of mosquitoes were found by the above-mentioned expedition, whereas a thorough examination in later years of mosquitoes at Churchill (which, although not actually in the Arctic regions, is situated on Hudson Bay) revealed about 20 species. In the case of collemboles, of which the Canadian Arctic Expedition brought back 12 species, some collections from near the Arctic Sea obtained the summer of 1948, with apparatus specially suited for the purpose, brought the number up to 70, perhaps 100. It would presumably be the same with all groups, which actually means that only a small proportion of the insects living in

EA-Zoology. Hammer: Arctic Insects

the northern regions are known today. In Spitsbergen the insect fauna is poor, which is doubtless due equally to its position isolated, from all [: ] other large land masses, and to its being situated so far north; only about 130 species, mostly Diptera, have been brought back from numerous expeditions. As far as Siberia is concerned, little is at present available, especially as theses on the subject are for the most part written in Russian. Therefore, this article will be confined to the available matter in European languages.
To understand insect life in the Arctic, one must first realize how it has come into existence. The arctic regions were thousands of years ago (and still are partially) covered by immense ice masses. As, during the ice age, these ice masses spread southward across the continents, they forced all animal life which did not perish to move southward in fron ^ t ^ of the ice border; later, as the ice slowly retreated, a fauna which had acclimatized itself to the prevailing conditions wandered northward, closely following the receding ice. The glacial period is usually imagined as having been intensely cold, but its conditions may not have differed greatly from those in modern Greenland, where an ice age still exists. There even large animals like the musk ox pasture near the ice edge and warmth-loving thermophile butterflies flutter in the sun not far from the inland ice. In Iceland , also there is abundant insect life in sheltered valleys right up to the edge of the great glaciers, so it is easy to understand that a rich fauna cold live under similar conditions during the glacial period.
Some insects immigrated as soon as the land was accessible, many others came later, and presumably some even survived the glacial period on the highlands or other areas which were not covered with ice. It is

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known that certain areas in the Urals in Siberia, in the Scandinavian mountains, and in Greenland and Alaska must have been free from ice, judging by the rugged and pointed appearance of the mountains and the presence of certain plants, and presumably a more or less scanty insect life has persisted here. If these areas resembled the stretch of coastal border in Greenland which is free from ice, with a climate not particularly colder, as is believed then the entire fauna might have been able to survive the ice period. On the other hand, if there were only a few scattered open patches in the ice, only the most hardy would survive, for instance, such insects as collemboles, scale insects, be t ^ e ^ tles, and gnats.
A third element in the insect population appeared later, as before men– tioned, th o ^ r ^ ough immigration or rather through transportation by winds, rivers, sea currents, birds, and ships; a great number of the insects found in Greenland today owe their presence to these different means of transportation. One need only think of the many insects often found in and around human dwell– ings in the North ^ , ^ such as lice, fleas, bedbugs, carpet beetles, larder beetles, beetles, in timber, and the like. Insects carried by wind, water, and birds have on the whole no great importance; most of them have a rather poor chance of finding foothold and surviving as the necessary conditions for existence are not present; presumably, therefore, in most cases they die after a few days, perhaps after a summer.
Finally there is a constant active immigration of insects, the species depending on the climatic conditions. During hot summers, it is thus possible to discover a few foreign insects such as dragonflies, butterflies, and grasshoppers, although they will quickly die out. Lindroth mentions that the hawk moth, Herse convolvuli , has been found in Iceland time and again,

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and it can only have come to the island by itself.
Now that we have noted the elements which make up the fauna, we will consider the extreme conditions prevailing in the arctic regions, which limit the insect life. There are almost no woods or large trees, only low scrub or stunted trees and bushes to replace the forests of the regions farther south. This means that many insects would not be able to exist there; in fact, all insects connected with the forest, which either live directly on its products of leaves, bark, wood, etc., or are indirectly associated with it as parasites or inhabitants of the forest ground, living on rotting leaves and branches. Without forests, insect life in greatly restricted, and, although many insects may arrive in the arctic regions, they will not find suitable conditions for existence and will quickly perish.
This leaves the insects which can adapt themselves to conditions on swampy tundra stretches, stony mountain fields with only scattered plant covering and more or less fertile valleys and mountain slopes. Also, con– sidering that the earth is covered with snow or ice the greater part of the year, so that insects have only two or three months or less in which to take nourishment and to multiply, it is no wonder that insect life is poor in the arctic regions.
Keeping all these restricting factors in mind, we must also realize that arctic insects can be observed only in the short summer. On a good warm day, bees can be heard buzzing among the flowers, while butterflies flutter across the scattered flowery banks and slender crane flies dance on the swampy meadows, laying their eggs. If one sits down for a short rest, he is immediately sur– rounded by the tormentors of these regions, mosquitoes and buffalo goats, which can make even grown men cry. They are specially troublesome in still, damp days and after sundown.

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Only the possessor of infinite patience should think of collecting insects in the arctic regions. Apart from gnats and flies, which can be found everywhere, the ^ and ^ collemboles, which are difficult to catch without special means, practically no insects can be found except by very thorough searching. Weeks spent in turning over stones to find beetles, scale insects, etc., and netting in the low vegetation, give apparently a very poor result; but when all material is assembled it is often better than expected as practically all groups of insects are represented. Not all insects are visible during the whole summer, however, and the time at which one happens to come across certain species is often more dependent on the presence of the right vegetation, pollen, or honey , than on the temperature. This of course only applies to insects which suck nectar, such as butterflies, bee ,s ^ s, ^ and certain flies, while the many others which live on rotting vegetation, on dead animals, or as insects of prey or parasites, can be found during a longer period. It is typical of arctic insect life that a great number life on rotting animals and plants, of which the y supply is always abundant, some on pollen and nectar, and others as parasites and insects of prey, while the actual plant eaters, like the phytophagous ^ phytophagous ^ beetles (Chrysomelidae) are few. A closer examination of the different groups will illustrate this. The Apterygota, the most primitive of all insect groups, are richly repre– sented by numerous collemboles. It seems that the short arctic summer brings the insects to a climax of development, where the number of individuals even exceeds that in regions farther south. On distinctly dry lichen plains at Angmagssalik in East Greenland in June, about 900,000 individuals per square meter found,. A colossal number considering that the vegetation is very low, so that the insects have little room to spread upward and have

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no possibility of finding nourishment underneath where there is only a sand or stone. Not only is the member of individuals great, but also the number of species. Some 40 species are known from Greenland, about 20 from Arctic Canada and Probably a similar number from Siberia. Throughout all of the Arctic there is a high percentage of primitive forms such as Hypogas–trura species (of which especially H. Armata is found everywhere), Anurida granaria, Onychiurus sibiricus , O. Groenlandicus , and O. armatus, all of which have short springing organs and live more or less in the ground. The more highly specialized forms, such as Enthomobrya, Tomoceras, and Sminthuridae species, which have well-developed springing organs and are more associated with trees or flowers, are only poorly represented. Collemboles are better suited than any other insect group for life in the Arctic, as they can stand extreme cold and will come forth when the temperature is 5° below freezing. A few degrees of warmth in the middle of the day will bring them out in spring and they will lay their eggs even a temperature of 7°C. They live on all sorts of plants, such as lichens, moss, fungi, and rotten vegetation. The majority doubtless spend the winter as fully f ^ g ^ rown insects; in any case, chiefly fully grown specimens are found in winter, and in early spring, whereas mostly small ones are found late in the summer. Collemboles ^ need ^ only a little food, so they are to be found wherever there is the slightest vegetation. They are the only insects found on the Antarctic Continent.
Plecoptera, Neuroptera, and Trichoptera . Only a few species of Plecoptera (stone flies) and Neuroptera ( ;acowomts ^ lacewings ^ ) are known from the arctic regions, a perlid having been caught in Alaska and not less than three Hemerobius species found in Greenland. Trichoptera or caddis flies

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are common, and numerous species thrive in these regions; in Lapland more than a hundred, while Greenland and C ^ a ^ nada can claim only a few species, among others some belonging to the genus Limnophilus. The larvae of Plecoptora, Neuroptera and Trichoptera live in water, where they winter.
Orthoptera. The warmth-loving Orthoptera are also known [: ] far to the north, but they are seldom seen and are possibly only carried there accidentally. Information from Greenland stating that the dragonfly Calopteryx virgo was seen there, is doubtless erroneous; in the summer of 1948 the author saw a large dragonfly fluttering across the cliffs at Coppermine in Arctic Canada, but it was impossible to catch. Grasshoppers have not been met with in Greenland, apart from the greenhouse grasshopper Tachycines asynamorus , which like Forficula auricularia and Blatta orientalis , both found in Greenland, belongs to the group of insects which definitely keep near human dwellings. In Canada and Siberia, on the other hand, grass– hoppers have been found several times. Melanoplus frigidus has been found, amon t ^ g ^ other places, at Franklin Bay and in Alaska near latitude 70° N.; the author has found it at Coppermine, N.W.T., where it was previously known. It seems to have a wise circumpolar distribution. Also M. borealis is well known in arctic regions, although its radius stretches somewhat farther south. Strangely enough, neither dragonflies nor grasshoppers are found in Iceland, a fact which doubtless relates to the difficulties of crossing the wide open seas. In Lapland there are not only dragonflies and grasshoppers, but even a species of cockroach.
Hemiptera . Hemiptera are well known throughout the northern regions, although the number of species and individuals is not great. As they are plant suckers, conditions for their existence are rather unfavorable, but

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they have nevertheless adapted themselves to [: ] circumstances, and they can be found under the ground sucking roots, as well as on the low, scanty vegetation. Several sorts of Aphis can be found on grass and willow in Greenland, Iceland, Canada, and Lapland. The jumping plant lice Psylla ambigua and P. alni live in Greenland, and in all probability elsewhere in the A arctic regions. Nysius groenlandicus can be found up to latitude 74° N., and Ligyrocoris constrictus in common all over Canada. The best known of all Hemiptera is doubtless the scale insect Orthezia cataphracta , which can be found among leaves, under stones and branches, and in grass, where it sucks roots. Like many other scale insects it carries its young in a bag which protrudes from its back and is formed of long, broad wax threads. Also the well-known bedbug Cimex lectularius thrives in the Arctic, and is often a great plague to the population. The majority of these insects winter as larvae or nymphs hidden under leaves, in cracks and crevices.
Coleoptera . The order of Coleoptera or beetles, which contains the greatest variety of types of all insect groups, is rather poorly repre– sented in arctic regions. In Greenland 44 species have been found, in Canada up to 70, while Lapland can claim about 420 different types. Many of the Canadian species originate from the forest districts, where they live in bark and wood. In Greenland, where there are no forests, there are, nevertheless, a number of beetles which originally came from forests, such as Pityogenes chalcographus, as well as several Cerambycidee ( Tetropium castaneum , Callidium violaceum , C. variable , Molorchus minor , Pogonochaerus fasciculatus, and many others). None of these are actually natives of Greenland, but have been transported there with timber. The many beetles

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living in and around dwellings have presumably also been introduced, such as the larder beetle Dermestes Laradarius , the carpet beetle Attagenus Pelio , Anthrenus museorum , Ptinus fur , and others. The most common arctic beetles are Carabidae, Dytiscidae, Gyrinidae, as well as Staphylinidae, Byrrhidae and Curculionidae, which are nearly all small and insignificant. They can be found in water, under stones, and among grass and leaves. Most of them live on other insects or on vegetation. The actual phytophag e ous leaf beetle is, on the other hand, seldom found in arctic regions. Occa– sionally it is brought in; in Canada there are various types, such as Chrysomela , Lina scripts , and Haltica. These beetles seem to be able to winter in all stages.
Lepid e ^ o ^ ptera . The Lepidoptera belong mainly to warmer climates than usually found in arctic regions, and this is especially true of the Rhopal ^ o ^ cera or butterflies. The fact that butterflies nevertheless have been found far to the north in Greenland is due, according to the opinion of some research workers, to their having immigrated from North America during a warmer period. We find them in sheltered valleys where they may have retreated when climatic conditions became worse; for instance Argynnis polarisv, A. charicles and Colias hecla have been noted. These sheltered valleys are situated particularly in East and north Greenland, where there is little rain and the climate is somewhat continental, with much sunshine. Colias hecla is circumpolar and found in three geographical subspecies, of which two live in the Soviet Arctic.
In Iceland there are no indigenous butterflies; according to the opinion f Lindroth, this is because there are too few sunny days for them to feed and copulate. A few species, Pieris rapae , Pyrameis ( Vanessa )

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atalante and P. cardui , have, however, been found, but these must have been introduced. In Canada, butterflies are more common than [: ] in Greenland; many speci d ^ e ^ s could be mentioned here, such as Argynnis bischoffi , various Colias species, the colorful swallowtails ( Papilio machaon, P. glaucus ), the closely related Parnassuis sminthus , as well as some cabbage butter– flies ( Pieris occidentalis, P. napi ) and the Euchloe species related to them, ( E. creusa, E. ausonides ), as well as many others. Until now only one species of hawk moth has been found in Greenland, a larva of Hyphoraia lapponica from East Greenland. Several types of the same genus ( H. parthenos , H. alpine and H. festiva ) live in Canada. There are no hawk moths in Iceland, although Herse convolvuli has been found several times - probably introduced like the butterflies. The lesser Lepidoptera, such moths as Noctuidae, Geometridae, Tortricidae, and Tineidae are common everywhere in the North; 18 Noctuidae, 10 Geometridae, 7 Tortricidae and 2 Tineidae are known from Greenland; from Canada a somewhat greater number, and in Iceland they [: ] are also frequent. Special mention should be made of the widely distributed Agrotis occulta , whose larvae are cutworms, because their destruction of grass has been suggested as one of the major reasons for the disappearance of the Norsemen from Greenland. As butterflies suck nectar, they are chiefly found at the height of summer, vanishing early in autumn. They spend the winter mostly as larvae of pupae among roots and leaves in the earth, in crevices in tree stumps, or in other protected places under stones or branches.
Hymenoptera . The Hymenoptera are a group with a great variety of species in arctic regions, including some of the insects which penetrate farthest to the north. Bumblebees have been found in Greenland as far north as latitude 81°50' N. They seem to be present everywhere, both over land and

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sea, and often they can be seen several miles from the nearest coast. Bumblebees, of which two species live in Greenland, winter as queens, coming forth ^ ^ early in the spring and finding their first nourishment in the numerous willow blossoms, which are soon replaced by many others ( Dryas, Cassiope , Papl Papilionaceae, etc.); later in the summer fireweed ( Epilobium ) is a special attraction. In arctic North America there are no fewer than ten species of bumblebees, including the two Greenland species, which are also found in the Soviet Arctic. Iceland is represented by only one species of bumblebee, Bombus jonellus , which is also common in the northern parts of Scandinavia with ten other species. Wasps are rare, although, Vespa marginata has been found in Alaska. Wood waspa (Sirex) are seldom seen, but they can stray far from their domain, or be introduced. The author caught a well-grown specimen by the Arctic Sea in the summer of 1948. Presumably it had come from the wooded areas farther south; it has also been found near Hudson Bay. In Iceland Sirex (species gigas ) has been observed several times. Sawflies (Tenthredinidae) are quite common. Six species are found in Greenland, where their larvae live in grass or on willow and poplar leaves; in Canada about twenty species have been found, mostly on different kinds of willows, where the larvae often produce galls. Not less than eighty species of T ^ I ^ chneumonidae and other small wasps live in Greenland, whereas the figure for arctic North America is much smaller.
Unlike most of the other Hymenoptera, ants are not common in the North; they have not been reported from Greenland or Iceland. Several species are known from Alaska, and during the summer of 1948, the author found these ^ three ^ species in the Mackenzie Delta near latitude 69° N. One of these, Camponotus hercu lanus, lives in rotting tree trunks; another, Leptothoras

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canadensis , in smaller branches; while the third, Formica fusca , makes its home in the earth under leaves and roots. Most Hymenoptera winter as larvae or pupae, hidden away in cracks and crevices in wood, or buried under dead leaves.
Diptera. Diptera are the most abundant in both species and individuals of all insect orders in arctic regions. About 275 species are known from Greenland. It is common knowledge that the millions of gnats and buffalo gnats can be a torment to all living creatures. They live beside the sea, by lakes and rivers, everywhere in the tundra. What these enormous swarms live on is hard to say, but presumably they suck nectar from the numerous flowers. It is believed that the female must eat blood in order to lay mature eggs, but where it finds enough blood is a problem, as most of the regions full of gnats are rather thinly inhabited by ^ v ^ bertebrates. Only one species of mosquito (Culicidae) is found in Greenland, ( Aedes nigripes ), while in Arctic Canada two species are known, Aedes nearcticus and Aedes sp .) They winter as imagoes. Few species of buffalo gnats are known, five from Greenland and three from Arctic Canada. They are, like the mosquitoes, an insufferable torment, not so much because of their stings, but because they fly headlong into eyes, no es ^ se ^ and mouth. The larvae live in running water, especially where there are rapid streams or small waterfalls; they winter as larvae.
Many other Nematocera could be mentioned, of which Chironomidae, Tipulidae, Sciaridae and Mycetophilidae are common everywhere in the arctic regions. There are also numerous species of Brachycera, for instance, the big, beautiful Syrphidae, and the genus Calliphora , (blow-fly) which can trace the slightest smell of meat from great distances.

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Even the well-known housefly, Musca domestica , and the lesser housefly, Fannia canicularis , are found. Members of the Scatophagidae, whose larvae live in manure and rotting vegetation, are often met with; while Hypoderma tarandi , is a great torment to the caribou in the tundra. Finally, there are large members of other small dipterous flies, which find nourishment chiefly in the many arctic flowers.
From the above it will be seen that the insect fauna of the Arctic is not restricted to that region alone, but is rather a branch of the southern faunal element, of which the hardiest species have been able to spread over the northerly regions, where they find ^ some what ^ unfavorable conditions for existence.

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BIBLIOGRAPHY

1. Canadian Arctic Expedition, 1913-1918. “Insects,” The Expedition. Report , vol.3. Ottawa, Acland, 1922.

2. Degerbøl, Magnus. “A contribution to the investigation of fauna of the Blosseville coast, East Greenland, Medd.Grønland, vol.104. no.19, 1937.

3. Hammer, Marie. “Studies on the oribatids and collemboles of Greenland,” Ibid . vol.141, 1944.

4. Henriksen, K.L. “Insects collected on the Fifth Thule Expedition,” Thule Expedition, 5th, 1921-1924. Report , vol.2, no.8, 1937.

5. ----. “A revised index of the insects of Grønland,” Medd.Grønland, vol.119, no.10, 1939.

6. ----, and Lundbeck, Will. “Landarthropoder (Insecta et Arachnida),” Ibid. vol.22, pp.481-821, 1917.

7. Jansson, A., and Sjöberg, O. “Bidrag till Kännedomen om Insektfaunanii Hamra Nationalpark,” Svenska Vetenskapsakad. Skr. no.20, 1932.

8. Jensen, Ad.S. Grønlands Fauna, et Frosøg pea en Oversight . København, Bianco Lunos Bogtrykkeri, 1928.

9. Johansen, Frits. “Insect life on the western arctic coast of America,” Canadian Arctic Expedition, 1913-1918. Report , vol.3, pt.K, 1921.

10. Lindroth, C. “Die Insektenfauna Islands und ihre Probleme,” Zoologiska Bidr. Uppsala, vol.13, 1931.

11. Sjöstedt, Y. “Insektfaunan inom Abisko Nationalpark I-II,” Svenska Vetenskapsakad. Skr . no.16-17, 1931.

12. Thor, Sig. “Beiträge zur Kenntnis der invertebrathen Fauna von Svalbard,” Norsk Polarinstitutt. Skr. no.27, 1930.

13. Twinn, C.R. “Studies of the biology and control of biting flies in northern Canada,” Arctic , vol.3, no.1, pp.14-26, Apr. 1950.

Marie Hammer

Land and Freshwater Invertebrates

EA-Zoology (Elisabeth Deichmann)

LAND AND FRESHWATER INVERTEBRATES

CONTENTS

Scroll Table to show more columns

Page
Mollusks 1
Habitat 4
Dispersal 6
Variation 7
Geographic Distribution 8
Crustaceans 11
Phyllopeda 16
Cladocera 18
Copepoda 19
Ostracoda 19
Habitat 20
Other Aquatic Invertebrates 22
Sponges 23
Bryozoans 23
Hydra 23
Flatworms 24
Oligochaetes, Nematodes ^ , ^ and Tardigrades 24
Rotifers 25
Terrestrial Arthropods 25
Spiders 26
Mites 31
Other Terrestrial Invertebrates 36
Oligochaetes ^ 36 ^
Lumbricidae 36
Enchytraeidae 38
Nematodes 40
Rotifers 41
Tardigrades 42
Bibliography 44

EA-Zoology (Elisabeth Deichmann)

ARCTIC LAND AND FRESHWATER INVERTEBRATES
I. MOLLUSKS
Our knowledge of arctic nonmarine mollusk fauna can probably be said to have begun in 1843, when Middnedorff reported a freshwater snail form a small po i nd on the Taimyr Peninsula is Siberia (73°30' N. lat.) and enthusi– astically wrote: “undoubtedly the most northern, almost unbelievably high northern locality for a fresh-water mollusk.” With that suspicious beginning, the study of arctic land and freshwater mollusks was inaugurated on a large scale. During the next hundred years a colossal amount of data was collected by amateurs and professionals, some merely records of occurrence, some def– initely erroneous, but nevertheless so much that when the ecological and zo– ogeographical studies began there were many sources from which to draw. In a great number of instances the old shells had been preserved in various museums and were available for the new generation of ecologists and zoogeo– graphers, and advances in geological exploration have made it possible to connect the recent fauna with those which existed in earlier periods.
Russia n , with its territory extending from the Baltic Sea to the Pacific Ocean, has been active in the study of northern nonmarine mollusks; first under the aegis of the Tear, and later under the Soviet government. Often this research dealt with purely practical economic problems, such as fisheries, and additional information came from the geological discoveries which were ^ made in ^

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large areas in connection with the vast Soviet engineering projects. North America in many respects offers a striking parallel to northern Asia, and its fauna is actually more rich in species but the northernmost region is not nearly so well explored. Particularly in Canada very little work has been done, and the ecologists can start with a fresh slate beginning with modern methods such as those Thorson (1946) recommends for [: ] future studies in the arctic region.
Europe has, of course, been most carefully investigated from the beginning of systematic zoology, but only a very small part of northernmost Scandinavia is of interest in connection with the study of arctic mollusks, and even that region needs to be reinvestigated by persons familiar with ecological technique.
It must be made clear that, as far as land and freshwater mollusks are concerned, it is almost impossible to define the special arctic zone. What exists in the highest latitudes is merely a fraction of the mollusk fauna which occupies the land farther south. Modern writers have chiefly considered the entire subarctic region, roughly the zone which lies between 50° N. lati– tude and the Arctic Circle, and then briefly, amost causally, taken in the fauna which extends farther north. The old definition of the A ^ a ^ rctic zone, used for example in Fauna Arctica ^ Fauna Arctica ^ , as everything which lies north of the Arctic Circle, is as artificial as it can possibl e ^ y ^ be. The subarctic zone, intelli– gently interpreted, with allowance for the irregularities which are produced by climatic and other factors, represents a natural region. As its southern border there is a sudden wholesale disappearance of a large number of land and freshwater forms, while within its border, and stretching up into the northernmost part of the Frigid Zone, one finds a sudden blossoming out of certain genera which appear with a wealth of species and vari ^ e ^ ties. Also one finds in the

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subarcti x ^ c ^ zone a few forms, mostly quite primitive, which occupy a very lim– ited area, confined to the coldest part. If they are known from the Temper– ate Zone at all, it is from isolated spots in high altitudes. These forms represent the relicts of an earlier fauna, while the remaining forms are the hardy outposts ^ - ^ tolerant, less specialized in their demands, and [: ] often extremely successful in the North.
Our present knowledge of the composition of the subarctic land and fresh– water mollusks was summarized in 1938 by Mozley, who not only has done a considerable amount of field work in northern Siberia, Canada, northern Scan– dinavia, and Finland, but also has had occasion to study a large number of the older collections in European museums, and has thus been able to eliminate a number of doubtful forms and synonyms. He lists 223 species and vari ^ e ^ ties for the entire region, of which 55 are terrestrial. The remainder are aquatic forms, divided between 63 species of clams and mussels and 105 species of snails. Some of the snails are slightly amphibian in character, and there– fore presumably able to migrate from pond to pond when necessary. A large number of the mussels and clams belong to the Unionidae, and the subarctic species represent a rather heterogeneous selection of that family — some fairly large species — while the rest belong to the Sphaeriidae, a family of small species which seem particularly well suited to the conditions pre– vailing in the North. Among the aquatic snails one finds a similar division; a number of representatives of the most diverse groups, which as a whole are much more abundant in the Temperate Zone, and also two large families, the Lymnaeidae and Planorbidae, fairly closely related, which make up the bulk of the subarctic and arctic species.

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With the uncertainty which besets the definition of the arctic region, with nobody willing to commit himself, one can only guess that about 10 to 15% of the subar a ctic mollusk fauna ranges into the most frigid northern zone. The terrestrial forms on the whole are poorly represented in the Arctic. Not many of them are found in the subarctic either, as Mozley’s figures clearly show. Land mollusks are more or less dependent on deciduous trees, and conifers have little attraction for them; therefore living conditions are unfavorable in the tundra of the North, where only a few deformed trees stray from the deciduous forests into a region where they do not belong. Mozley mentions one land snail from the Hudson Bay region and two from the northern part of Siberia; in Greenland only two land forms appear to be indigenous. Iceland, in contrast, has about eighteen terrestrial snails, and no less than six slugs. The only northern regions having a greater number of terrestrial forms are those where forests are present, such as the mountains of western Alaska and the highlands of Kamchatka.
^ Habitat . ^ Thus it is the freshwater forms which dominate in the North. With the abundance of wet localities, including rivers, lakes, ponds (temporary and permanent), and extensive systems of bogs and marshes, there seem to be un– limited possibilities for the aquatic fauna. The factors which restrain the mollusks from filling up the entire zone are: ( 1 ) the climate, ( 2 ) the scarcity of food in large parts of the region, and ( 3 ) poor adaptation by mollusks for rapid dispersal. The low temperature in itself is not an ab– solute hindrance, but in the higher latitudes, the exceedingly short summer makes it difficult or even impossible for many species to complete their devel– opment. Also, plant life is comparatively poor in many different types of water bod ^ i ^ es, particularly so in regions farther north.

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Best suited for mollusks are the smaller lakes and the ponds, which often have an abundant plant life. An interesting type is the temporary pond, formed by melting snow, which Mozley was the first to describe, both in Canada and Siberia, as an exceptionally good habitat for mollusks. In these ponds he found, besides the typical arctic communities of Entomostraca, such as Apus , Lepidurus , and Branchipus , up to eight species of mollusks in Canada and five in Siberia. Curiously enough, such ponds seem to be totally lacking in Finland, where Branchipus and related forms do not occur, and they are rare in Scandinavia. The snails hibernate in the adult stage. As soon as the snow begins to melt, they start mating and lay their eggs which develop rapidly. When the pond dries up, after one or two months, the snails hide in the mud and begin their summer sleep, which passes without interruption into the winter sleep, lasting until the spring thaw of the following year. In the ponds which contain water the year round, a similar fauna is usually present whe ^ n ^ plant life is rich enough to support the mollusks, and when unfav– [: ] orable factors, such as sphagnum, are absent. Both in Siberia and in Canada such ponds are common, and characteristically they harbor decidedly few species than the temporary ponds; the present census reads, three snails, representing three genera, and one clam, in both regions. In Greenland an even smaller community is known to exist, with only two species, both of the genus Lymnaea , while in the ponds of Ko p ^ l ^ guev Island in the Barents Sea only a single variety of the same ubiquitous genus is present. The larger lakes of the North have not been well explored, but it is probable that they are not favorable habitats for mollusks, partly on account of their scarce vege– tation; the same holds true of the stream beds of the rivers. In Siberia many mollusks have succeeded in migrating down to the mouths of the large

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rivers, but as they are mostly adapted to quiet water, they are rarely found in the main channels, where the rapid current prevents the growth of plant life. In Canada little work has been done on the deltas of the arctic rivers, but the chances are that similar conditions obtain here and that possibly quite a number of species of mollusks have found their way far northward.
Dispersal . The Entomostraca are eminently adapted for rapid dispersal, with their hardy eggs which can withstand desiccation and freezing and be carried away by the wind when the temporary ponds are dried up. In contrast to them, the mollusks are about as ill-fitted for migration as any inverte– [: ] ^ br ^ ate group can be, with the possible exception of the earthworms. They can fill a certain habitat to perfection, and the hardier forms can tolerate ex– tremely unfavorable conditions and withstand great variations in temperature; but they cannot move into near l ^ b ^ y localities if there is the least obstacle in the way, and their eggs and young are not suited to endure changes in moisture for any great length of time. In both North America and Si x ^ b ^ eria where the original fauna was completely driven out by the glacial period, the mo [: ] llusks have had to work their way northward, as Dall expresses it, “step by step.” The terrestrial forms have been bound to follow the northward trek of the plant communities with which they were connected, while the aquatic forms have had to move from water body to water body as conditions permitted. To some extent they have utilized the rivers; many of them were crushed in the rapids or smothered by the silt, but in the course of time some have succeeded in reaching the mouths of the rivers in the north and gradually populating suitable side waters; however, dispersion from one river system to another has usually been beyond their powers. To a large extent the mollusks have also been distributed through the air, when their eggs, or even adult animals

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of the smaller species, stick to bits of plants on the feet of water birds. This transportation has been particularly possible for the inhabitants of small ponds with a rich plant life, while the forms which live in larger lakes or in the quiet parts of rivers where there are few plants usually have had to remain ^ ^ inside the limits of their own river system. The Unionidae, which have a par ^ a ^ sitic larval stage, are dependent on freshwater fish for their distribution, while the smaller forms, such ^ as ^ [: ] ^ Pisidium ^ , which lack such such a stage, have to depend on birds, and have occasionally been known, in the adult stage, to clamp onto the legs of aquatic insects and let them– selves be carried to some other locality.
Variation . For the geneticist and the zoogeographer the study of the variation of the mollusk shell has always been of the greatest interest, and the arctic fauna, living under extreme conditions, offers a particular challenge. As a whole, it can probably be said that variation is less pro– nounced in the s ^ S ^ ubarctic and Arctic Zone than farther south, possibly because the low temperature influences the tendency to variation, or possibly because most of the region has become inhabited so recently that there has not been time for the animals to become isolated and divided up into definite races or varieties. In spite of the large amount of material which has been studied, our knowledge is still incomplete, and scientists are rather blindly groping their way forward. As far as the outer conditions and their influence are concerned, the tendency is here, as in many other groups, for the more recent immigrants from the south to become dwarfish as they reach farther north. This seems to have been demonstrated satisfactorily in a number of Unionidae. On the other hand, such typical ^ ^ northern cold-water forms as Aplexa hypnorum , which is at home on the Taimyr Peninsula, grow larger here than farther south,

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and when, in rare cases, specimens are found in the Temperate Zone, they are the ones which have become dwarfish. But when one tries to work out how the animals react within areas having reasonably uniform life conditions, one finds that there are no rules for their behavior. Thus, according to Mozley, the Siberian members of Valvata can easily be recognized when examined in– dividually; but when large series are compared, there is such an intergrada– tion that one is inclined to merge them all into one variable species. And while the different forms seldom occur together, there seems to be no reason, ecological or zoogeographical, which explains why in one locality they develop into one form, and in a similar environment into another. In the large family Lymnaeidae, which is one of the most successful and widespread groups in the subaractic zone, it was found that one species, taken from a number of ponds which were lying close together, showed no tendency to form local races but varied widely inside each locality. Another species, taken from rock-filled ponds which were separated by distances up to a hundred miles, showed a def– inite tendency to split up into local races. In the case of the well-known widespread form Lymnaea e stagnalis , Mozley found in the whole of northern Siberia only one variable form, while in Switzerland, which represents an older, more stable region, with less extreme temperatures, twelve distinct local races or varieties were reported.
Geographic Distribution . Geographically the nonmarine mollusk fauna can be divided into two main provinces, the Eurasian and the North American, and, as a special subdivision, the arctic islands, with their impoverished fauna consisting of the most hardy species and representing the last part of the Arctic Zone to be re-populated after the ice age. Aside from a few circumpolar species, the two main areas have a separate fauna derived from the richer fauna

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of the Temperate Zone farther south. Only in the mountains around Bering Sea do the faunas overlap. Between Europe and North America there seems to be no faunistic relationship, except for the peculiar case of Helix ^ ^ hortensis . This is supposed to have been found as a fo l ssil in Maine, and has been reported from various localities in Labrador and even in Greenland (three records — one from the stomach of an eider duck. Scientists should investigate such puzzling problems as this before going too deeply into the question of former land bridges between Europe and North America, or considering the validity of Wegener’s theory of continental drift.
The circumpolar portion of the Soviet Union offers several interesting features which Shadin has briefly summarized, though only for the aquatic mollusks. The region can be divided into a smaller western province in which the man-made canals have helped to spread the European forms up into the White Sea region and a larger eastern province. Around Pechora a considerable number of European forms are still present, but farther east the characteristic lit– tle form Ancylus fluviatilis drops out. Then the Siberian fauna takes over, with the conditions reversed — a decrease in n number of species from the eastern part, with its richer fauna, toward the west. A particularly inter– esting subregion is the mountainous Kamchatka with its almost insular character. Its fauna seems to have been impoverished by the great natural catastrophes which it has undergone, its ice age and volcanic eruptions. It has some af– finities with the Amur region as well as with North America, as Dall pointed out in 1905, and possesses, in addition, a few endemic species and some of circumboreal distribution. In its hot springs a number of species of Lymnaea , have been found, and in its cool, rock-filled rivers the river pearl mussel re– appears, after having dropped out in the westernmost part of the circumpolar zone, near the Finnish border.
(Page 9a follows)

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Although the northern fauna of Siberia has been well explored, especially by the Vega expedition, the total number of species and varie– ties remains remarkably low. The reasons are to be found in the turbulent past of the Asiatic continent, with the originally rich fauna of the Ter– tiary and interglacial epochs being more or less wiped out by the ice, in combination with the changes which central Asia has undergone, resulting in a desert and salt steppe zone which now cuts off all possibilities for a repopulation from the south.
In North America the conditions are far more suitable for a migration northward from the Temperate Zone. Both the central part of con– tinent and the western mountain ranges favor a penetration from the south, and it is easy to understand how so many species have been able to reach Alaska and large parts of Canada. The northernmost region, how– ever, is still very incompletely studied. Most of the available informa– tion about the fauna of the mountainous part of Alaska and its islands was gathered by the Harriman Expedition (1899), and very little has been added to Dall’s observations made at that time. The Canadian Arctic Expedition (1913-18) was likewise undertaken before the methods of intensive ecological work had been developed; few freshwater and terrestrial mollusks were brought back from the North and few observations have been added in later years. Probably it will be found that quite a number of mollusks have become well established in the deltas of the arctic rivers, and that the fauna here compares favorably with that of northernmost Siberia.

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Island faunas are notoriously poor, and the mollusk faunas of the arctic islands, that is, Greenland , and the islands north of Europe and Asia, are no exception. These localities, with their extremely select fauna, deserve to be discussed separately. They are the only portions of the Arctic Zone in which nonmarine mollusks were considered at all in Fauna Arctica , that invaluable though some hw ^ wh ^ at unwieldy depository for much of our earlier know– ledge of the fauna of the Far North.
In spite of the fact that Greenland has been unusually th ^ o ^ roughly explored during the last 150 years, the list of land and freshwater mollusks still re– mains stationary and is exceedingly small. Leaving out the utterly unreliable records, and a few cases of obvious introduction through man (such as the lone slug, Arion fuscus , which came in with a load of cabbage from Denmark, and was promptly shipped back to the zoological museum) there are only about ten species present, so far all from the west coast, which is by far the most inhabitable. [: ] The species consist of a single small clam, two terrestrial forms, and a few aquatic snails. Except for one apparently endemic species, they all belong to widespread circumpolar forms. It is hard to form any opinion about their origin. Most probably they came from North America during the favorable periods when several American species of different groups were able to migrate into Greenland. When northern Canada becomes better known in regard to fossil and recent forms, the answer may appear quite obvious. It must, however, be men– tioned that all the species, except the one which is endemic, also seem to live in Iceland, which has about three times as many mollusks as Greenland. Since most of the species found in Iceland have been taken near the places where the Vikings settled (besides other localities farther north), there is the possibil– ity that Greenland actually received most of its nonmarine mollusks in historic times from Iceland.

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Until quite recently the other arctic islands were considered totally uninhabited by mollusks, but with the intensified colle [: ] cting and better tech– nique which is characteristic of modern expeditions, this picture has changed and may well change ever more. Shadin, ^ in ^ 1935, reported from one explorer’s collection not less than seven species from the Solovetski Islands north of Siberia. One is the ubiquitous Lymnaea stagnalis and the rest are also wide– ranging northern species. From Kolguev Island in the Barents Sea has been reported a vari ^ e ^ ty of another Lymnaea ^ Lymnaea ^ , and finally Novaya Zemlya has been found to harbor a clam, a very primitive Pisidium , one of the typical cold-water forms, which grows larger in the north than farther south. It seeks a deeper level in Novaya Zemlya than in northern Norway and Siberia, where it is always found in shallow water. This clam has recently been discovered also on one of the Aleutian Islands, so its dis [: ] tribution has proved to be much wider than was originally assumed.
II. CRUSTACEANS
It has been estimated that 15% of the ice-free land in the Arctic is covered by water bodies, ranging from huge rivers and large lakes down to shallow ponds and bogs of which some are filled with water only during part of the short summer. In fairly early times explorers began trying to find out what lived in these locations by means of dip nets or dragnets; and they round and astounding number of different forms, even though one might say from casual observati ^ o ^ n that the water looked very dead as contrasted with similar situations in the Temperate Zone, where the vegetation in most cases is far richer. For many divisions of aquatic invertebrates our knowledge is still in the exploratory state in the Arctic, but in one group a tremendous amount of work has been done — namely, among the crustaceans. Those

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which have been studied in fresh ^ ^ water are chiefly the more primitive smaller forms, the Entomostraca, or “small crustaceans” as the Scandinavian writers usually call them, in contrast to such higher forms as lobsters, crabs, and shrimps which are referred to as “big crustaceans.” One of the reasons why so much work has been done on these animals in the Arctic is that they are easy to collect, particularly those which live free in [: ] the water, where they occur during the summer in large numbers and where even during the winter months it is sometimes possible, through the ice, to observe those few forms which normally continue to stay active until the next summer. Vanhőffen, one of the earliest explorers to spend a winter in Greenland on one of the nunataks, noticed how life continued under the ice of the l ka ^ ak ^ e; later the Scandinavian expeditions to the east coast of Greenland made extensive studies of the conditions found in the larger lakes during the winter months. In the second place, no [: ] other group has been studies so intensely in the Temperate Zone as have the Entomostraca. This is true particularly in Germany, Switzerland, and Scandinavia, and more recently also in the United States and the Soviet Union. Hence a whole school and tradition of limnology have been developed in the twentieth century, and the [: ] technique of study is well established. In mountainous regions a complete series of zones has been noted, roughly corresponding with those of the higher latitudes, showing a similar flora and fauna and a similar decrease in number of species. The background is therefore complete, and even where the arctic material is not always as plentiful as one could desire, it is often possible to draw analogies from the biology of the same species, or very similar forms, under alpine conditions.
The least explored regions in respect to arctic freshwater crustaceans are Siberia and North America. In the first-mentioned area we have as the

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one large contribution Vereshch ^ a ^ gin’s work (in Russian) on the Cladocera of the Iamal Peninsula, and in the second the exploratory reports of the Canadian Arctic Expedition; for both regions there is in addition a number of smaller, scattered papers, dealing with isolated areas. No doubt the freshwater fauna in these regions will be found to be quite rich, though with the countries running in an east-west direction there will be great uniformity. The situa– tion is presented by Haberbosch, who has compared the en ^ o ^ rmous number of Cla– docera, which Vereshchagin reports from one relatively small spot of Siberia, with the few species known from the whole of well-explored Greenland.
Haberbosch notes that on Iamal one finds a coast region which has been raised up from the sea during the last few thousand years, a tundra with a continuous, still undefined reticulum of water bodies, connected through the large Siberian rivers with the hinterland of Asia. In Greenland is a large coastal area, only recently freed from ice, completely cut off by the sea from other continents and i ^ s ^ lands, with water bodies mostly lying on archaic rocks or on basalt, entirely independent of each other. These differences have undoubtedly exercised a great influence on the composition of the fauna of these two arctic regions and may still be actively at work. For just as the water bodies at the foot of the Alps and the Scandinavian mountains repre– sent a reservoir from which a new faunal element can migrate continuously into the higher altitudes, so can the Iamal Peninsula depend on new inhabitants coming incessantly from the rich reservoi [: ] rs of central Asia to settle down in the less favorable arctic regions in the North. But in Greenland condi– tions are entirely different; here the sources from which new additions to the fauna should come lie hundreds of miles away on other continents [: ] and are separated from them by the sea. From our limited knowledge of these two

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large regions, Siberia and Arctic North America, it appears that the crustacean fauna offers striking similarities although the individual species in many cases are not the same.
The situation is quite different elsewhere in the North, namely, in Greenland, the Canadian Arctic Islands, Spitsbergen (or Svalbard), Franz Josef Land, Bear Island, Jan Mayen, and the northernmost part of Scandinavia. Here the faunas are small but extremely well analyzed, and the knowledge gained by European scientists in Middle Europe and the Scandinavian countries has been tu ^ ut ^ ilized in the North. Nevertheless, as Thorson has outlined, there is still much to be done, particularly during the winter, and in the line of long-range studies continued over a period of many years. Our two chief cources of in– formation are Haberbosch’s work on the Greenland Entomostraca and Olofsson’s studies on Spitsbergen fauna. The findings of Haberbosch and Olofsson show great similarity, and both men went to their task equipped with the same thor– ough training in the subject: one having acquired it in the Alps under the leading freshwater biologists of that region; the other with a similar back– ground received from the most prominent workers in Sweden.
Haberbosch’s work is based exclusively on a study of collections made by other persons, often with very scant information as to the ^ ^ type of water body from which the animals were taken and ^ not ^ too many temperature observations, al– though the author has been able to piece the existing knowledge together to make a reasonably coherent picture. Olofsson’s study is based on a single person’s observations during a summer visit which was extensive enough so that he was able to cover most of the ground reasonably well, and to visit most of his stations at least twice during the summer. These obse [: ] rvations were sup– plemented by a large amount of data obtained from the weather station on

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water ^ winter ^ conditions such as temperature, thickness of ice, and dates of disappear– ance of snow and ice. Spitsbergen had a much smaller fauna than Greenland, and Olofsson therefore found time to supplement his studies of the Entomos– traca with work on freshwater rotifers (rotatori a ^ n ^ s) which in many respects formed an interesting parallel group to the crustaceans.
The Greenland fauna comprises 46 species divided among 3 Phyllopoda, 24 Cladocera, 12 Copepoda, and 10 Ostracoda. Except for the last group, which has been less well explored until recently when better methods for collecting bottom samples were devised, it is not likely that many more species can be expected. From the scarcity of certain species it appears that these are less well established, although the existing records themselves are beyond doubt, and when more data are gathered it may be possible to say why these forms are ill-suited for the particular conditions found in this part of the world. In Spitsbergen only 27 species are known and most of them are the same as in Greenland. In the other arctic islands the fauna is even smaller, with only the most widespread and hardiest species left; last on the list comes Jan Mayen with only one freshwater crustacean. It seems certain that all the crustaceans in Greenland and Spitsbergen perished during the ice age, although some authorities claim that possibly a few of the moss-living forms which only recently have become better known, may have survived. In both Greenland and Spitsbergen it seems certain that birds are responsible for the presence of all the species, except the few which are of marine origin, while in Siberia and North America they may have spread by way of rivers. All the Entomo ^ s ^ traca are hardy forms, with eggs which can resist desiccation and are well fitted to be carried by the wind for short stretches or by birds for longer distances. Some species may have been transported by the ice with soil from the shore,

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accidentally washed down on the ice while it was still attached to the land. In the case of Greenland, the composition of the fauna is so typically hol– arctic -- , almost all the species occur everywhere in the Arctic — that it is not possible to state whether it originally came from the Old World or from America. The chances are that most of the species are of American ori– gin, just as Spitsbergen probably received most of its fauna from Europe through migratory birds or by the ice. Only two species are known in Green– land which have a more limited distribution — as far as our present know– ledge goes; a few more collections may well throw this distribution on the scrap heap. One is a palearctic form which seems to have its western limit of distribution in Greenland, while the other is a well-known American species which has reached eastward to Greenland and Iceland. In addition one might mention a brackish-water form, a primitive phyllopod which occurs in ^ on ^ Green– land’s west coast, both in salt water and in a few lakes. It was originally considered identical with the widespread brine shrimp, but now it is believed to be a Greenland variety of a species hitherto known only from the northern part of Canada. It constitutes, therefore, a further link with America.
[: ] Phyllopoda . A few words may be said about the different groups. The phyllopods are largest in size, reaching a length of several centimeters, which is colossal for the “small crustaceans.” The common fairly shrimp, which occurs in water pools in the Temperate Zone during the early spring and then disappears when the pools dry up, may be familiar to many. In the arctic region there is one species ( Branchinecta ) which ranges about as far north as any freshwater animal ca, to 81° N. latitude, in Polaris Bay, Green– land, and, interestingly enough, it has also been discovered in the Carpathian Mountains in Europe, where it must have settled down after the ice age. Less

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known to most people is the dark brown Lepidurus arcticus , for which there is no popular name, although related forms occur intermittently in the Temperate Zone. It looks like a diminutive horseshoe crab, or, as a little Indian boy explained to a Stanford professor on an expedition to Alaska, “like a turtle.” The largest specimens measure 2 to 3 centimeters. It is usually found in shallow-water pools but may also occur in larger lakes where there is enough plant food. Its occurrence or nonoccurrence in different water bodies may be cited as an example of the peculiar fastidiousness which these arctic animals have toward certain external conditions, , a characteristic which, considering their extreme hardiness in most respects, is rather baffling and hard to understand for a mere warm-blooded animal such as man. In the Arctic these animals live mostly in shallow pools where the summer temperature often goes up to 15°C., and if they live in larger lakes they never go down into the real cold water. Olofsson, therefore, considers it “not too arctic” in its taste. In the higest mountains of Norway and Sweden the animal completely avoids the shallow pools and occurs exclusively in the larger lakes at a moderate depth; hence, Ekman notes it typical “high-arctic” behavior. It is not found in the Alps, although it is well adapted for wide dispersal and has been found in fossil form in various parts of Europe. Olofsson finally concludes that it is not the ab ^ s ^ olute temperature but the wide daily variations which it avoids. Since in the Arctic, particularly in Spitsbergen, the variations are small, it can live in pools there; in Norway the conditions are sufficiently stable in lakes only; and in the Alps, where the sun’s rays strike at a much higher angle , and the temperature varies widely in the upper strata of the lakes, it cannot live at all, since the conditions deeper down are not suitable for other reasons, including perhaps the lack of food.

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Caldocera . The Cladocera are one of the most successful groups of the Entomostraca, and occur mostly in fresh water which they have comp ^ l ^ etely con– quered. They are all small — a ^ ^ length of 4 to 5 millimeters is considered gigantic. Many people have seen them in aquaria, with their compact bodies covered by a fold of chitin like a little jacket, their large complex eyes, and their dorsal spine, moving along with small jerky jumps. In the Temper– ate Zone they produce a number of broods during the summer, some developing from unfertilized eggs, while other generations consist of males and females which produce fertilized eggs, usually called “winter eggs” because the last brood which carries the species through the unfavorable season is of this type. The winter eggs are more thick-shelled and are also protected by part of the skinfold of the female, which is thrown off and forms a peculiar case (ephip^ephip^ pium^pium^) around the eggs. In the arctic region the species have cut down their productivity and there is usually only one cycle; that is, only once a year does a winter-egg-producing generation appear. The resistant, well-protected eggs of these forms are par ^ t ^ icularly suited for distribution by birds, much better than the eggs of the next group, the copepods, which, generally speaking, are most successful where they can wander from one body of water to another without going on land, as in Asia and North America. In Greenland there are twice as many cladocerans as copepods, and the remarkably low number of species known from Spitsbergen (five [: ] clad ^ oc ^ erans against t h en species of copepods) is a riddle which we have not yet been able to sovle. The majority of the arctic species are widespread forms. In Greenland, Daphnia pulex (up to 5 millimeters long) seems invariably to occur in all the same localities as Lepidurus arcticus as well as in a number of other types of water. It is also known from large

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parts of Europe and America. Like the other Cladocera, it varies much accord– ing to external conditions, and Olofsson has made a large series of observations on its variation and its relation to the environment — drying up of pools, lack of food, temperature conditions, etc.
Copepoda . The third group, the Copepoda, have their main distribution in the ocean, where they constitute one of the most important food elements for the whalebone whales. In fresh water they are quite successful and closely parallel the Cladocera, though they have not developed generations which con– sist exclusively of females, and their eggs are less well adapted for desiccation and transport. The majority are panktonic forms with an oval body and large an– tennae which assist them in floating and swimming. Recently a large number of species have been discovered which live chiefly in wet moss and are almost never seen in the open plankton zone. In Greenland about a dozen species are known, the dominant forms being two widespread cosmopolitan species which are steno– thermal (demanding cold water), in contrast to many eurythermal species which tolerate a wider range of temperature.
Ostracoda . The fourth [: ] group are the Ostracoda which live exclusively in the bottom of the water bodies, either in the mud or among the plants. [: ] They are very small and look like diminutive mussel shells; hence, they are often called “mussel crustaceans.” As far as is known they produce on ^ e ^ generation of eggs per year, and the eggs remain mostly cn ^ con ^ cealed in the shell of the female. As in the Cladocera, the eggs can withstand extreme conditions, and there is no doubt that far more species will be discovered when the technique of bottom sampling improves and a careful examination of the aquatic vegetation becomes part of the usual procedure.

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Habitat
An extensive literature exists on the different types of water bodies which are found in the Arctic. The rivers are in a class by themselves, for except in the parts where they flow slowly and contain a rich vegetation and thus simulate the conditions of a lake, they are not suitable for a permanent population of planktonic or near-planktonic animals. They merely act as high– ways along which animals can be transferred into other regions. In general, arctic water bodes may be divided into the following types: warm or cold, shallow or deep, containing much or little plant life. The shallow water bodies consist of bogs, ponds, and lakes with little depth, and many of these habitats are temporary and exist only during a few spring and summer months. Nevertheless it is here that the richest populations of crustaceans are found, including such species as Lepidurus and Daphnia pulex and other forms which can complete their growth and produce their eggs during the sum– mer months and afterward die. The number of species is low. In Greenland one can at the most find 15 species in one locality, while in Finland up to 56 different species may be found in one lake; but the paucity of species is offset by the large number of individuals. In the larger and deeper lakes, the crustaceans are rarely found in large numbers, their presence depending upon the quantity of plant life in the lake. It is a general rule that the deeper the lake, the colder the water and the fewer the plants, both along the edges and in the open, and with that goes a decrease in the number of species and individuals of crustaceans. The water of the lake can be divided into three zones — the shallow water zone, covered by plants or barren; the [: ] deeper bottom zone with its varying thickness of more or less rich mud; and

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and the open water in the middle. Most zoologists find that there is no real planktonic zone in the arctic lakes, that is, there is no community which spends its whole life floating freely in the open water. Some species from the shore zone visit it frequently: the young of Lepidurus have been ob– served as nauplius and metanauplis in the open water, but in the later stages this genus returns to the plant-filled edges of the lake and stays there for the rest of its life.
A common belief is that conditions in the Arctic are very uniform, and, when contrasted with the sudden thaws and frosts of the more temperate zones, it is true that the climate of the Arctic is more dependable. There are, how– ever, seasonal and annual variations and there may be considerable difference in the length of the summer season, which may cause some variation in the number of broods produced among copepods and cladocerans. Usually the peak of the season is in July or the beginning of August, when the majority of the crustaceans lay their eggs. For some reason those which hibernate in the egg stage lay their eggs quite early, while those few forms which continue their development under the ice or hibernate as adults lay them much later; some even breed during the cold season and utilize the warm period for growth.
Water bodies are influenced not only by varying weather conditions but by location. Even in Spitsbergen, a comparatively small areas, it was discovered that the conditions are more favorable on the west coast, where the effect of a small arm from the Gulf Stream is felt. Within a single fjord, Olofsson found pronounced differences between the water bodies situated close to the opening of the fjord and those near the inner end. In so huge a country as Greenland there are, of course, even greater differences. As a whole it appears that the conditions are less favorable on the east coast. Although probably all

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the Entomostraca known from Greenland will be found to occur there, it is [: ] ^ l ^ ikely that the number of individuals will be far smaller than in similar areas on the west coast. Along the southern part of the west coast, certain forms which might be expected do not occur. Particularly striking is the complete absence of Lepidurus everywhere south of Holsteinsborg, which lies nearly on the Arctic Circle. This may be caused by the pack-ice barrier which drifts in around Cape Farewell from the Arctic Sea, making the condi– tions less s suitable in some particular respects. What these respects are we do not yet known, and so far not enough data have been gathered; but the problem illustrates how much detailed study is necessary before our under– standing of these animals can be said to be fairly complete.
The higher crustaceans play a negligible role in the arctic freshwater fauna. Of the Isopoda only the common Asellus aquaticus has been reported from Greenland and Siberia, and the few amphipods which are known are all de– rived from salt water. The little shrimplike Mysis relicta , famous as a relict form in many fresh waters, occurs here and there in the Arctic, in northern Canada and also in Spitsbergen. In the latter locality, Olofsson was able to follow the development of the marine M. oculata which now and then became trapped in fresh water, and he found that it gradually acquired the character– istics of M. relicta . Hence, the modifications are not inherited but are due to the effects of the environment, something which is worth remembering, par– ticularly where one is dealing with such a comparatively recent fauna as that of the arctic regions, where isolation has not yet had time to do its work.
III. OTHER AQUATIC INVERTEBRATES
Among the other invertebrates some groups are evidently not suited for

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life in the North, [: ] while others may prove to be more widespread there than they appear at present, if better equipment is used and more time is spent in study.
Sponges . It seems certain that the freshwater sponges are out of place in the North in spite of their gemmulae, which permit parts of the colonies to survive the unfavorable season as small unites, and later to become widely dispersed as efficiently as if spread by eggs. The few records we have of sponges are almost all from Siberia, where the chances are that reinforcements may be received from farther south even if the whole population of the Arctic Zone is temporarily wiped out in unfavorable years.
Bryozoans . The freshwater B ^ b ^ ryozoans, with their statoblasts which repre– sent a similar adaptation for survival and dispersal, are likewise a group almost nonexistent in the Arctic. A few statoblasts have been found here and there, indicating that the species have been able to reach the North but , that the conditions for survival have not been suitable. An exception is the discovery of two widespread species in a stony mountain stream in northwest Greenland, where apparently the colonies found the conditions much to their taste. Possibly it will be found that these species are not uncommon in this type of habitat.
Hydra . The H ^ h ^ ydra, a characteristic element of freshwater bodies in the Temperate Zone, has been reported from a few places in northern Canada and on the west coast of Greenland. The material which has been identified belongs to well-known, widely spread species. Roman Kenk ( in litt .) reports that a large form of Hydra is common in some lakes and streams in southern Alaska, where it is found in shallow water attached to the undersides of stones. Fur– ther investigation may show that Hydra is more widespread than had been assumed.

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Flatworms . Until fairly recent years the freshwater flatworms had been reported from only a few places in the Arctic, mostly in Siberia and North America. In the 1920’s two collectors visited Greenland, and in one summer raised the number of species known there from near zero to almost sixty. Most of the flatworms were of American origin, inhabiting the shallow plant-filled ponds characteristic of the A ^ a ^ rctic tundra. Their eggs, always small and resis– tant to drought and cold, might easily be carried by the wind for long dis– tances, even over the ice-covered sea. While most of the species have crossed to Greenland from arctic North America in this way, others were probably car– ried by water birds. The one species having large, thin-shelled eggs was not nearctic in origin, but was undoubtedly brought in by the Vikings, in the wet moss which clings to the roots of the herb angelica. Another species, likewise of non-American origin, has proved to be an offshoot of the marine fauna which had previously existed in some warm springs where the animals are now found; its close ^ s ^ t affinities are with species now occurring in the North Sea.
According to Roman Kenk ( in litt .): “Subarctic Alaska has a comparatively rich fauna of freshwater triclad Turbellaria living in streams and lakes. No triclads have so far been found in Alaska north of the Arctic Circle, though Rhabdocoela are encountered as far north as Point Barrow.”
Oligochaetes, Nematodes, and Tardigrades . The oligochaetes have a number of representatives in fresh water, mostly primitive forms which are also found in brackish water, but they have not been thoroughly studied, and the same is true of the aquatic representatives of the nematodes and tardigrades. The two last-named groups, however, like the rotifers, belong to the select microfauna which has been found in the lakes of the nunataks, and they may possibly claim to represent the few original inhabitants of the arctic regions which have sur– vived the ice age. To prove that they did so is difficult, for they all belong to widespread forms which could easily have been brought in from the outside,

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with stop-overs in the temporary lakes of melting ice on the inland ice itself, in which these animals have often been taken.
Rotifers . The rotifers (or rotatorians) have not been thoroughly studied even in the Temperate Zone. They rival the crustaceans and [: ] nema– todes in number of species, but at present not all of them have been described and classified. The members of the group are extremely widespread but at the same time very selective in their choice of habitat. In Spitsbergen, Olofsson has made the first attempt to study their biology as they have been studied in the Temperate Zone by Wesenberg-Lund and others, and his results have been most promising. The Rotifera follow in many respects the pattern of the Entomostraca to which they have a curious superficial resem– blance. The chief difficulties are the problem of preserving the animals well enough for identification, and the necessity for long-range studies extending over an entire summer season and, if possible, over periods of years. With the modern trend of extending biological research over decades instead of weeks, it seems likely that these microscopic animals may become recognized as one of the most important elements in the ecology of the arctic freshwater communities, comparable in importance to the mites in the soil.
IV. TERRESTRIAL ARTHROPODS
The terrestrial arthropod fauna in the Arctic is, aside from the in– sects, mostly composed of spiders and their allies. There are only a few higher crustaceans in the Temperate Zone which have become adapted for life on the dry land, namely, the well-known sow bugs. Of these only two species have been reported frequently from the arctic region; they give the impression of having been accidentally introduced, and it is not known whether they have

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become permanently established in these localities. Thus, in Greenland, so reliable an observer as Fabricius stated that he had seen one kind, but it has never been found again since his time, around 1780. A number of myriapods and millepedes have also been reported from the arctic region, but so little is known about these forms that they are hardly worth discussing here. They do not appear to be a group of animals particularly suited to life in the Arctic, and most of the records are from Siberia and North America, where it has been comparatively easy for many of them to intrude into the arctic region. In Greenland, there has so far never been any record of them.
Spiders
Except for a few moss scorpions, a few daddy longlegs, and similar aberrant forms, the arachnids in the Arctic are divided among the spiders and the mites, the hunters and the hunted. The spiders, being conspicuous animals, usually found running actively on the ground, have early attracted attention here as everywhere [: ] else, and in the Temperate Zone an immense number of observations have been made of their interesting habits. The mites, being smaller and usually living quietly in the soil, were little noticed before the twentieth century. With the advent of the Berlese funnel they were discovered to be one of the most interesting and important groups, both from an ecological and a zoogeographic standpoint.
About 400 species of spiders were listed in Fauna Arctica around the begin– ning of the twentieth century. This was an exaggerated number, as the boundary lines for the Arctic Zone had been drawn rather generously, but in a way that made it easier for a beginner to estimate what species might be expected to ex– tend their range into the Far North. Probably 250 to 300 sp i ecies would be a

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closer estimate. Although a number of new species have been added, and a number of forms will be merged, and although the distribution area for the individual species will be greatly modified when the enormous area becomes better explored, nevertheless it may be said that the composition of the fauna is reasonably well known. The arctic spiders are all of the most specialized types, the primitive forms being all restricted to the tr po ^ op ^ ical and subtropical zone. The fauna includes most of the larger groups, with a preponderance of those which make elaborate webs — of the sedentary species, to use an old fashioned term. Especially well represented are the members of the subfamily Linyphiinae, and of the free-living wolf spiders, while there is a notable scarcity of crab spiders and jumping spiders. The species are all of moderate size or small, even extremely small, which would be expected in a region where the life conditions are rather severe. Looking over the larger geographic divisions, Siberia, Alaska, Canada, etc., one finds a remarkable similarity in the proportions of genera of the fauna from each region. On the arctic islands there is a sharp drop in the number of species found, though possibly a number of the smaller species may still be reported. From Spitsbergen, for example, about a dozen are known. On Akpatok Island in 1931 an English expedition discovered 19 species; strangely enough, 11 of these were known from the palear [: ] ctic region and not less than 9 were known from England, which exemplifies how many of the species have a wide dis– tribution. Roughly one can divide the species into the widespread holarctic forms, and those which belong either in the northern part of the Old World or in North America. Greenland, which lies at the crossroads, so to speak, has a fauna of about 50 species, composed of elements from both east and west. The history of the nomenclature of its spider fauna shows an interesting pattern.

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In the beginning, almost all the species found were considered identical with species known in Europe. Later the majority were considered new and endemic to Greenland, except those which obviously represented very well-known Euro– pean species. But as time went on, it was found that the majority of the endemic species were merely outposts from American species, and the list of endemic species has been gradually decreasing, so that in the opinion of most arachnologists there probably are no endemic species in Greenland. This may seem a trivial matter, except for the fact that the presence of endemic species indicates that a region has been i os ^ so ^ lated for a very long time.
It is natural to pay particular attention to Greenland, because it oc– cupies a strategically important position where the nearctic and palearctic faunas overlap, and because its limited fauna has been comparatively well ex– plored by numerous expeditions. It is the one place in the Arctic where some attempts have been made to study the ecology of arctic spiders throughout the year.
So much is written about the spiders and their habits in general that little need be repeated here. In the arctic region the animals lay their eggs in midsummer so that the young spiders can creep into the ground about the first of September. When they reappear in the spring, about the first of May, they have either to mature in an extremely short time or to stretch their life cycle into another year. Spiders which take more than one year to complete their development are known in the Temperate Zone, though they are uncommon; but in the North the finding of mature femal ^ e ^ s emerging from the winter sleep, and the presence of three or four size classes among immature spiders caught simultaneously during the summer, suggest that the longer life cycle may be

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fairly common there. It is quite possible that the amount of food available is so small that spiders cannot find enough to enable them to complete their development in one summer, even if they make use of the whole 24-hour-long day, as for example the bumblebees are known to do. Food is not plentiful in the Arctic, and spiders must, to a great degree, depend on what they find in the ground, mostly mites and collemboles; even the web-builders cannot rely on an abundant supply of insects.
The distribution of spiders has been shown to a great extent to fol ^ l ^ ow climatic zones, and Greenland, with its ice-free land strips running approx– imately north and south, shows different climatic types which coincide with the habitats of the majority of spiders. A few forms are completely indif– ferent to the amount of moisture present in the air, and to the abundance or lack of sunshine; these animals have a wide range, and are found almost everywhere except where food is scarce. Brandegaard has found that a number of other species are distributed only in the northern part of East Greenland, where the air is dry and the climate pronouncedly sunny; while south of Scoresby Sound, where the sky is cloudy, he fou [: ] nd exclusively forms which enjoy a humid atmosphere. In only one instance did he find that a species from the humid region had wandered up into the arid zone, and in that case it was invariably found hidden in the damp ground, thus inhabiting a situation resembling that to which it was accustomed in the humid south, where it is always found in the open. On the west coast, which has been less thoroughly investigated, he found that in the southern part the ice-free land strip is wide enough to permit the existence of two different climatic zones, a humid, foggy, sunless coastal zone

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and a sunny warm hinterland in the deep fjords. In the first are found some of the forms known from the humid part of the east coast, while in the other, a small species, Dictyna borealis , known from the arid zone of the east coast, suddenly appears widespread, though it is totally absent in the southern part of the east coast.
How the spiders migrate from one locality to another in the Arctic is not known in all cases. They are active wanderers, at least during part of their lives, and there is even a record of a wolf spider being found on a nunatak, one of the isolated mountain peaks which never were ^ was ^ covered by ice, and still lie ^ s ^ more or less completely surrounded by glaciers. In the Tem– perate Zone small spiders travel extensively by air, not only the newborn ones but in many species the adults, floating in the upward currents of the air attached to a long silk thread. Only in a few cases has this method been ac– tually observed in Greenland, but as far as size is concerned a large number of the Greenland spiders are well suited for this form of transportation; the Linyphiines in particular are good “balloonists,” or “aeronatus” as the modern term has it.
Where the spider fauna of Greenland came from is a question s which has always interested students of geographic distribution. It is certain that no spiders survived the ice age, as some of the mites possibly may have, and being dependent on other animals for food they cannot have become established until there was something for them to eat. Of the usual transportation methods, driftwood from Siberia, with its lengthy sojourn in the arctic water, is not probable, though it may be that Jan Mayen received some of its spiders in that way. Birds are out of the question, as spiders instinctively try to detach themselves from anything which moves. Man can probably be counted out

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in this case, as not even the cosmopolitan house spider has ever been reported from Greenland. A few Greenland species are found also in Den– mark, where one of them is extremely rare; the others were taken on the east coast of Greenland long before that region had begun to have any con– tact with the outside world. From Iceland the Vikings might have brought a few species, but those which are likely to have been introduced from there , happen also to be common in North America. The wind may account for some distribution of species; it is considered certain that one of the northeast coast species has been carried by warm winds across the inland ice from the west coast. But for the majority the distances to the nearest land areas are too large, and in case of land which lies reasonably near, as Iceland, the faunas differ considerably. Moreover, there are a few heavyweights which definitely are not able to utilize the wind as a method of distribution. So it appears that in the case of spiders, as in other groups, it becomes neces– sary to resort to a theory of former land bridges, or to Wegener’s drift theory, in order to explain their presence in Greenland.
Mites
The terrest ir ^ ri ^ al mites, particularly the smaller ones, were almost unknown at the beginning of the twentieth century, when various workers began to study them; it was not until the Berlese funnel began to be generally used, in con– nection with soil studies, that the microfauna came into its own. The o ^ r ^ iginal studies of the soil were undertaken in the Temperate Zone, and proved to be of the greatest practical value for agricultural and forestry research; but it was in the arctic region, where the scarcity of other inhabitants of the soil simplified the picture (and incidentally permitted a far larger number of

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individuals to live per square unit), that the study of these mites opened up a whole new world in the realm in ^ of ^ ecology. Except for some work done on Svalbard most of the studies have been undertaken in Greenland, where particularly the east coast has been surveyed extensively and throughout the greater part of the year. One of the most important long-range studies was unfortunately left unfinished on account of World War II, which forced the worker to abandon his arctic station without having carried his studies over into the summer as had been planned. Nevertheless, his observations have supplied a large amount of much-needed data on the composition of the microfauna during the winter, when the mites are found either as eggs, larvae, or adults.
The entire known arctic mite fauna was tabulated in Fauna Arctica in 1905 by Träga ^ a ^ rdh, who also reviewed the history of the group and emphasized the tremendous gaps which exist in our knowledge. The Oribatidae or moss– mites were at that time imperfectly known, although it was already evident that they were one of the most successful forms of mites in the Arctic. From Greenland, in which the arthropod fauna had been systematically explored for years, only a few Oribatidae had been listed at that time.
The te ^ c ^ hnique of analysis of the microfauna of the soil is fa ri ^ ir ^ ly simple. Standard samples of soil are collected from different types of biotopes, which are characterized chiefly by the vegetation: bogs, meadows, rocky hillside or scarp, lichen fields, etc. All the small animals which are contained in the sample are driven out by slow desiccation, or by speeding up desiccation by means of a hot water bath. A funnel beneath the container which encloses the sample permits the animals to escape and fall down into a glass tube filled

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with preservative fluid. Afterward the animals are sorted out, identified, and counted. In this work the systematic lines are generally ignored and the mites are usually treated with the collemboles, small primitive wingless insects which, like the mites, occur in large numbers, and therefore are also well suited for a statistical treatment. The samples are taken from as many different types of soil as po ^ s ^ sible, and the sampling is repeated at intervals of about fourteen days, so the worker can follow the development, egg-laying, hatching, and other processes of the mites. During the winter frozen samples are collected so as to learn how the animals spend this time of the year. Eggs of course give a negative result, but by having a continuous series of samples, one can find out whether a species in fact occurs in a given type of ground even if it is lacking in some samples. The animals occur in the loose debris which forms a layer of varying thi e ckness over the soil, and down to a depth of about two centimeters in the soil itself; deeper down only a few scattered animals are found. The advantage of hibernating so close to the surface is obvious. It is just as cold here as farther down, but the animals are “on the spot” as soon as the snow melts and the sun warms the up– permost layer of the ground. The moss mites live, as the collemboles do, on decaying parts of lic ^ h ^ ens, moss, and similar substances which are present al– most everywhere except in the most barren areas. The range of population density in different biotopes is very wide: 1/100 square meter in the poor– est biotope, the rocky hillside, may contain 30 individuals (mites and collem– boles) while the richest, the lichen field, may have up to 7,800.
By tabulating all the species from each type of soil, it has become pos– sible to give a clear account of what the ecological conditions are, and

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hence what other kinds of animals one may expect to find. The main divi– sion is between wet and dry biotopes, with a number of small subdivisions, which are characterized by their “index species.” In some cases one finds a kind of double-decker arrangement, as in the “glumiferous meadow” where the large grass tufts may support a wet community in the lower part near the roots, while higher up in the tuft a pronounced dry community is present.
Although the Greenland mite fauna is still incompletely known, in spite of the advances our knowledge has made in the last decades, it has become clear that the mites are chiefly related to those of the Old World — a striking contrast to most of the other more mobile forms of animals, from the musk ox downward, which are largely of American origi a n. As the mites of arctic North America become better explored this difference seems to be more pronounced. It seems, therefore, that we may have in the mites an actual remnant of the original fauna which has been able to survive the ice age, hidden away in the isolated mountain peaks, the nunataks, which have been found to harbor a small select flora and fauna of various hardy forms. It is to be hoped that in the future the nunataks will be investigated in much greater detail than the earlier explorers were able to do. Later addi– tions to the mite fauna may possibly be very slight.
Some mites may have been carried by birds; extremely few may have been able to use the wind, judging from the scarcity of these animals in air-samples taken at high altitudes. Driftwood seems not very suitable as a carrier, since the mites under consideration are never connected with trees, and man as an agent has definitely been most unimportant. Whether the entire mite population (of the more immobile types) is of preglacial origin, or whether certain types have wandered in later, at a time when

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Greenland may have been connected with Eurasia, cannot be decided at this moment. In one other relatively well explored area, Svalbard, it seems as if the entire fauna consists of preglacial species.
V. OTHER TERRESTRIAL INVERTEERATES
Oligochaetes
The remaining groups of terrestrial invertebrates which play a role in the arctic region are the O ^ o ^ ligochaetes, which consist of the earthworms and their allies, and all the smaller forms, such as roundworms, tardigrades and rotifers, which are microscopical or near-microscopical and still income– pletely known as to distribution and ecological importance.
Less than a hundred species of O ^ o ^ ligochaetes have so far been reported from the arctic region. Aside from a few primitive forms which are exclu– sively aquatic, and some of which are found in brackish water ^ or ^ on the seashore, they can be divided into two distinct families: the Lumbricidae, which repre– sent the highest developed groups, the well-known earthworks; and the more primitive forms, the Enchytraeidae, usually called “white worms,” familiar only to people who know them as pests in their flowerpots or those who have aquaria and raise them as food for their fishes.
Lumbricidae . The Lumbricidae are a group which is not well represented in the Arctic. Only about a dozen arctic species are known, most of them having a very wide distribution and belonging to the so-called peregrine species which have been spread passively, mainly through the activities of man. The family has been well studied by Michelson and is represented by a large number of genera and endemic species in the Temperate Zone. There it forms almost a continuous belt around the world, but farther north, in the

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region which once was covered by ice, the number of genera and species is small, and it drops quickly as one enters the Arctic Zone. How well the few species have been able to establish themselves in the Arctic is not known in most cases. Often they are abundant in the spots around settlements where there is plenty of leafmold and rich garden soil, and in the few places where such conditions are naturally present they are also likely to be found. Our knowledge of the earthworms in Greenland co ^ m ^ es chiefly from observations made by botanists and other nonspecialists. In the southern part of West Greenland “common ea ^ r ^ thworms” have been reported, occurring in large numbers on the surface after rain in the sheltered sunny valleys, where the moisture of the soil is sufficient to permit the decaying leaves of deciduous plants such as willows to form a considerable layer of black mold. A little farther north, near Godthaab, Warming observed them in large numbers in similar local– ities and commented on their absence in the adjoining areas where the typical arctic tundra condition prevails, with its lack of rich soil. Vanhöffen noticed that no earthworms seemed to occur north of Egedesminde, where there is also a sharp drop in the number of insects; that seems to be the extent of our knowledge in one of the regions which has been energetically explored for most animal groups. One must, however, remember that much of the work done in the Arctic is still in the purely exploratory stages, and that individual collectors, having one or two days at their disposal, have tried to find the most typically arctic spots instead of concentrating on the local garden plots and kitchen middens. Actually very little recent work has been done on the earthworms in Greenland, and much of it has been con– cerned with studying the older records rather than with new field work. Three species are known with certainty; all are widespread in Eurasia, but

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their exact area of distribution in Greenland is unknown. How they came to Greenland is not known; possibly they were introduced by the Vikings, as they occur in the places where those men settled. In the more northern localities whalers may have brought them in accidentally. In the remaining parts of the Arctic little is known except that certain species have been collected there; they may or may not have become permanently settled.
Enchytraeidae . Quite a different picture is presented by the family Enchytraeidae, a group which is well developed in the arctic region. Instead of a scant dozen species, about eighty forms have already been listed, and a much larger number can be expected when the soil has been more carefully inves– tigated. In Greenland very few species have been found, presumably because they have not been looked for. The majority of species which have been col– lected are from Siberia and northern Canada, and as a large number were orig– inally described by the same persons the incidence of wrong identifications is unusually small. In almost all cases the species have their chief distribution in the arctic regions, some in both the New and the Old World. Only a few species are found which have an erratic but wide distribution and for these man is undoubtedly responsible. The enchytraeids are mostly small species, less than two centimeters ^ long, ^ and threadlike, though a few species may be stouter. An unusual giant measuring six centimeters is known from Alaska to California. Enchytraeids usually occur in large numbers in mud, under decaying matter, and sometimes along the seashore under dead seaweeds; some are almost aquatic, While the life histor y ies of the common species in the Temperate Zone are fairly well worked out, little is known of the habits of the arctic forms: how resistant they are to unsuitable conditions, how hardy their egg capsules are, or how they migrate. They are more primitive forms than the earthworms

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and apparently less sensitive to salt water, therefore they are probably far better suited for transportation along the coastal zones than are the earthworms; and as they are often found in muddy localities, water birds may be active in distributing them. It may be also that most of the species were uniformly distributed over the entire northern zone and simply migrated northward after the ice age. Perhaps our present division into nearctic and palearctic species is simply caused by our incomplete knowledge and will gradually be eliminated as larger and larger areas become adequately explored.
Most spectacular are the so-called glacier worms of the genus Mesenchy traeus , which has about twenty representatives in the Arctic Zone, many living in mountainous areas under peculiar life conditions. The first observations were made by Nordenskiold who in 1886 saw some small dark-colored worms on the inland ice of Greenland. He concluded that here was actually a true in– habitant of the ice itself and guessed, rightly, that they subsisted on the microscopic algae which were found there. Twelve years later a new species was described from an Alaska ^ n ^ glacier, and in the following years two more species were described from the same part of the world. Some were discovered to pass part of their life in the soil next to the glaciers, but it was cer– tain that others spent their entire life on the ice of the glacier, with the eggs being hatched in the little pools which are found there, and the adult worms usually hiding under the snow when the sun was too warm and coming out later in the afternoon. The food seems in all cases to consist of micro– scopic algae. In contrast to most other members of this group, the glacier worms are dark-colored, but it is not yet proven that this is an adaptation to their life on the glacier, as dark-colored forms have also been found in other types of habitats.

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Nematodes
The N ^ n ^ ematodes or roundworms occur in large numbers in the soil. In the United States they have been thoroughly studied, because certain species are harmful to crops, and it has therefore become necessary to distinguish between the harmful and the harmless ones. In the Arctic they represent one of the last groups to be considered, and particularly those which are found in the soil have escaped notice much longer than the marine and fresh– water forms. They are usually present in enormous numbers and as soon as they have been well observed they should not be difficult to understand, though their uniform streamlined body and their few distinctive characters usually make students steer away from them as rather [: ] ^ u ^ ninteresting animals. Our incomplete knowledge of these forms is evident from descriptions of the origin of the material. Almost all the species known from Novaya Zemlya, for example, were by-products from some samples of moss, collected for the sake of the tradigrades which they might contain, and no less than 27 species were secured from one locality. In Greenland no free land and freshwater nematodes were known before 192 5 ^ 7 ^ , aside from casual remarks by earlier ex– plorers. Then in 1921 some collectors were requested to look for these ani– mals, and the result was 22 species of which 15 came from a single day’s ex– cursion into one locality. With little known about the nematode fauna of the larger part of the Arctic, with most of Europe and the northern part of North America still inperfectly mapped out, it is impossible to say much about the significance of these animals from an ecological standpoint. Both the adults and the eggs are hardy, they resist low temperature as well as desiccation, and they are well suited for passive transportation by water or wind. As far as we know they are widely distributed, but within small areas they may well

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prove to be useful indicators for certain conditions, just as some of the cosmopolitan rotifers are considered to be.
Rotifers
The arctic rotifers, or rotatorians, are on the whole insufficiently know ^ n ^ ; particularly the terrestrial forms have attracted little attention. Few scientists have realized that they are far easi re ^ er ^ to study than the aquatic forms, since they can be shipped in a tight container with a sample of moss and then, months after they have been collected, studied at leisure in the comfort of the laboratory. By means of a little water they can be kept alive for a long time, while workers who were attracted by the aquatic species had to depend on preserved material which was later not always found to be in good condition. Spitsbergen is the only region where the terrestrial forms have been reasonably well studied. One of the first species known from the Arctic was obtained by Ehrenberg in 1869, in a sample of moss from Spits– bergen, and since that time it has been customary to submit samples of moss from that region to specialists. Bryce, as early as 1897, reported 26 species from the few samples he received from Spitsbergen; from one sample no less than 15 different species were picked out. The species which live in moss almost all belong to a single family, and they differ considerably from the numerous, often fantastically shaped, species which are found in fresh water. All the species taken from Spitsbergen are known also from other parts of the world, and it is quite possible that other species which may be found to occur in the Arctic will prove to be cosmopolitans, for there is probably no group better suited to world-wide dispersal than the rotatorians. It remains to be seen whether the terrestrial forms will prove useful as indicators of

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certain ecological conditions, as the aquatic forms appear to be; for exam– ple, a lake in the Arctic Zone was found to contain exactly the same species of rotatorians as a similar lake in the Alps, no more, no less. A thorough study of these forms of life is evidently needed.
Tardigrades
Finally there remains a small terrestrial group in which a great deal of work is still to be done, the tardigrades. Like the other minute inhabi– tants of moss, they have so far escaped notice in most places, while their aquatic counterparts have long been known; for example, the rotifers were listed among the few animals which were found to inhabit the lakes of the nunataks in Greenland. Our knowledge of the occurrence of tardigrades in the Arctic is still in the most elementary exploratory stage. Only ten species so far, aquatic and terrestrial, have been reported from Greenland, while Spitsbergen leads with more than forty terrestrial forms, thanks to the moss-collecting tradition which belongs to that island. Although some efforts have been made to find them in Novaya Zemlya the results have been rather meager, and apparently there is a definite paucity of species in Bear Island; Franz Josef Land harbors several species. These differences may prove to be only variation in the ability of collectors to find the right kind of material, but they may possibly indicate something more. Under all circumstances the tardigrades are well worth studying. There are only a limited number to consider; the entire world fauna consists of less than two hundred species, a striking contrast to the other groups, the rotifers and roundworms, where the number of species is far larger. They are well characterized, and with their tremendous ability to survive unfavorable

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situations and to withstand desiccation for years on end, they may be studied under all sorts of laboratory conditions.

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49. Murray, J. “Arctic Tardigrade, collected by Dr. W. S. Bryce,” Roy. Soc. Edinb. Trans . vol.45, pp.669-81, 1907.

50. Olofsson, O. “Beitrag zur Kenntniss der Harpactiden, Familien Echinosomidae, Cantocamptidea (Gen. Maraenobitus) und Tachidiidae, nebst Be– schreibungen neuen und wening bekannten Arktischen Brackwasser und Süsswasser Arten,” Usala. ^ Uppsala ^ Univ. Zoologiska. Bidr. Uppsala , vol.6, pp.1-40, 1918.

51. - - - -. “Studien ueber die Süsswasserfauna Spitzbergens. Beitrag zur Systematic, Biologie und Tiergeographie der Crustaceen und Rotatorien,” Up ^ p ^ sala . Univ. Zoologiska Bidr. Uppsala , vol.6, pp.183-646, 1918.

52. Posselt, H.J. “Grønlands Brachiopoder og Bløddyr,” Medd. Grønland , vol.23, no. 1, 1899.

53. Randell, J.A. “Araneae. Results of the Oxford University Expedition to Akpatok in 1931,” Zool. Soc. Lond. Proc . 1933, pp.145-60.

54. Remane, A. “Die Rotatorien,” Fauna Arct. Jena , vol.6, pp.93-114, 1932.

55. Richters, F. “Arktische Tardigraden,” Fauna Arct. Jena , vol.3, pp.493-508, ^ 1904. ^

56. Rosen, O.V. “Mollusques terrestres et d’eaux douces, recueillis au Kamtchatka par 1 ‘expedition de M. T. Riabusinsky en 1908-1909,” Akad. Nauk. Zool. Muz. Ezhegodnik , 1926, pp.262-74.

57. Sars, G.O. “Freshwater Ostracoda from Canada and Alaska,” Report . Canadian Arctic Exp. 1913-1918 , vol.7, Part I, 1926.

58. Schaudin, F. “Die Tardigraden,” Fauna Arct. Jena , vol.2, pp.18 6 ^ 5 ^ -96, 1901.

59. Schlesch, H. “Note sur la faune malacologique du Groenland et de l’Islande,” J. Conch . vol.74, no.1, pp.115-27, 19 4 ^ 3 ^ 0.

60. Schultze, P. “Hydriden des Arktisches Gebietes,” Fauna Arct. Jena , vol.6, pp.55-58, 1931.

61. Shadin, W.I. “Uber die ökologische und geographische Verbreitung der Süss– wassermollusken in der USSR,” Zoogeographica , vol.2, pp.495-554, 1935.

62. Smith, Frank. “Oligochaeta collected by the Canadian Arctic Expedition 1913-1918. 1. - The Lumbriculidae,” Canadian Arctic Expedition, 1913-1918. Report . vol.9, pt. A, pp.3A-8A, 1919.

63. Steinböck, O. “Die Turbellarien des Arktisches Gebiet,” Fauna Arct. Jena , vol.6, pp.295-342, 1932.

67a. Thor, Sig. “Neue Beiträge zur Kenntniss der Investebraten Fauna von Svalbard,” Zoolischer Anz. vol.107, pp.114-39, 1934.

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64. Steiner, G. “Freilebende Nematoden von Nowaja-Seml y ^ j ^ a,” Zool. Anz . vol.47. pp.50-74, 1916.

65. Strand, E. “Die Arktischen Araneae,” Fauna Arct. Jena , vol.4, pp.431-78, 1906.

66. Thiele, J. “Ar c ^ k ^ tische Loricaten, Gastropoden, Scaphopoden und Bivalven,” Fauna Arct. Jena , vol.5, pt.2, pp.561-632, 1928.

67. Thor, Sig. “Beiträge zur Kenntnis der Invertebraten Fauna von Svalbard,” Norsk Polarinstitutt. Skr . no.27, 1930.

67a. Thor, Sig. “Neue Beiträge zur Kenntniss der Investebraten Fauna von Svalbard,” Zoolischer Anz. vol.107, pp.114-39, 1934.

68. Thorson, Gunnar. “Technique and future work in Arctic animal ecology.” Medd. Grønland , vol.144, no4, pp.1-40, Copenhagen, 1946.

69. Trädgaardh, J. “Monographic der Arktischen Akariden,” Fauna Arct. Jena , vol.4, pp.1-78, 1906.

70. Ude, H. “Die arktischen Enchyträeiden und Lumbriciden, sowie die geo– graphische Verbreitung dieser Familien,” Fauna Arct. Jena , vol.2, pt. 1, pp. 1 3 -34, 1901.

71. - - - -. “Die Oligochaeten der Arktis und ihre geographische Verbreitung,” Fauna Arct. Jena , vol.6, pp.41-54, 1931.

72. Vanhöffen, E. “Die Fauna und Flora Grønlands.” Ges. F. Erdkunde , 1891-93 ^ 1891-93 ^ , vol.2, pp.137-76, 1897. [Part of Drygalski’s Greenland Expedi– tion, vol.2.]

73. Verescagin, G. J. (Vereshchagin) “Sur la plancton de basin de la presqu’ile de Yamal.” St. Petersburg Ann . Mus. Zool. Acad. Sci., vol.18, pp.169-220. (In Russian)

74. Warming, Eng. “Om Grønlands vegetation,” Medd. Grønland , vol.12, p p.32, 1886-87.

75. Welch, P.S. “Alaskan Glacier Worms (Oligochaeta).” Montreal, McGill Univer– sity. Its Bionomial Leaflets no.2, 1917.

76. - - - -. “Oligochaeta collected by the Canadian Arctic Expedition, 1913-1918. II. - The Enchytraeidea,” Canadian Arctic Expedition, 1913-1918. Report . vol.9, pp.9A-19A, 1919.

77. - - - -. “Snow-field and glacier oligochaeta from Mt. Rainier, Washington,” Amer. Micr. Soc. Trans . vol.35, pp.85-124, 1916.

78. Wesenberg-Lund, Elise. “Igler og Oligochaeter,” Medd. Grønland , vol.23, Suppl. no.18, 1926.

Elisabeth Deichmann

Parasitology

(EA-Zoo. Thomas W. M. Cameron)

PARASITOLOGY

CONTENTS

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Parasites of Man 2
Parasites of Food Animals 9
Para ^ s ^ ti ^ it ^ es of Dogs and Related Carnivores 16 ^^
Conclusion 18
Bibliography 20

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ERRATA
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“We have records of this parasite in dogs in the Arctic, although its main hosts are wild carnivores (q.v.).”
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“The Arctic fox is also infected with the adult tapeworm but the source of its infection may well be small rodents, such as the Tundra vole and related species. The vole has been found infected with the cystic stage in Alaska.
“All these carnivores may be infected with the Trichina worm.” [continue with paragraph beginning: “It is obvious . . .”

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LIST OF FIGURES

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Fig. 1 Natural history of the parasite 4 ^ a ^

EA-Zoology (Thomas W. M. Cameron)

PARASITOLOGY
Our knowledge of parasitolgoy in the Arctic is extremely inadequate, and although we have accumulated during the past two decades (1930-50) a certain amount of information concerning the parasites of man and of dogs, there are still great gas to be filled. We may assume that these parasites are circumpolar, but there are still too few data on the relative importance of the various species. The main factor in the distribution of animal para– sites is climate; the arctic climate with its short, warm summer and its long, cold winter is not favorable to the development of species which require a considerable portion of the life cycle to be spent on the ground. Accord– ingly, the most common parasites are those which are carried by contact or which develop in intermediate hosts. This is true not only for man, his food animals, and his dogs, but for wild animals as well. The intermediate host is usually an animal which is eaten by the definitive host; however, a few parasites do exist which pass from host to host directly. The most im– portant of these occur in carnivores, and, as the infective season is short and limited to summer, they seldom reach the importan t ce that related forms do in warmer climates. This article evaluates our present situation in regard to man, his food animals, and his dogs. Much of the information is based on the survey initiated in 1932 by the Institute of Parasitology, Macdonald College, Ste. Anne de Bellevue, Quebec. Additional more recent reports from North America and northwestern Europe have also been consulted;

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recent information from the U.S.S.R. is not available to the writer.
Parasites of Man
These include the trichina worm, the oxyurid worm, the ascarid worm, various fish-carried tapeworms, the intestinal protozoa, and lice.
Trichina . By far the most important parasite is the trichina worm ( Trichinella spiralis ). It probably existed in the Arctic before the advent- of man, although it is only in recent years that its presence has become known. It is a small roundworm (the female is only about one-eighth of an inch long and the male half this size) which lives in the small intestine of flesh-eating animals. The femal,e lying in close contact with the mucous membrane, produces some six hundred minute sausage-shaped embryos which are passed directly into the mucosa. Those which enter the lumen of the bowel die; T those in the mucosa, however, are carried passively by the blood stream to all parts of the body and finally come to rest in the capillaries. Those which arrive at organs also die, but in dying cause some toxic changes. Those in the skeletal muscles, and those alone, are able to survive. They enter the muscle substance and grow at its expense, finally becoming so large that they have to coil in [: ] spirals — the fact which is responsible for their speci– fic name. By this time they are almost as large as their parents, and an en– closing fibrous cyst is formed around each. This cyst is not at first visible to the naked eye but, as time goes on, chalk granules become deposited in it and it hardens and becomes apparent as a small whitish poin about the size of a pinhead. The young worm within the cyst is often dead, although it may sur– vive for a time within the chalky wall. If it is to develop further, the cyst must be eaten by some other animal. In this second animal, the cyst is digested

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away and the young worm emerges, crawls into the folds of the lining of the small intestine and develops to maturity in about a week. The entire life cycle may be completed in four weeks, or it may take as long as six or seven years.
It is obvious from this life history that the worm was originally a parasite of flesh-eating animals, particularly of those which eat other carnivorous mammals. It is legitimate to assume, therefore, that it was originally a parasite of subarctic and arctic regions. The author, like other workers, has found it frequently in polar bears ( Ursus ( Thalarctos ) maritimus ), arctic foxes ( Alopex lagopus ), and red foxes ( Vulpes fulva ) in the Canadian Arctic. It has been found commonly in dogs in the Arctic, while Roth et al. have found it once in the bearded seal ( Erignathus barba–tus) and the walrus ( Odobenus rosmarus ) in Greenland. However, the author has examined a considerable number of seals for this parasite with negative results; it is probably not common in this host. In Alaska it has been found in white whales, wolves, and wolverines, as well as in foxes, dogs, and Alaska brown bears ( Ursus arctos dalli ). It has also been recorded as occurring in the polar bear in Siberia.
The trichina worm has little host specificity for mammals, and it will probably parasitize any animal which will eat meat. While certain herbivore– ous animals can be artificially infected, this does not happen in nature; thus bovine animals and deer may be regarded as free from infection. It spread to man in temperate climates and is perpetuated in him mainly because man infected his domestic pigs with the same parasite, and, by his system of animal management, has maintained it in them. From the pigs it spread to rats, cats, and other mammals which would eat raw meat scraps. Man can be infected.

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from any source. In temperate climates, undoubtedly, his main source of in– fection is uncooked pork; in cold regions it is probably the polar bear, al– though other animals cannot be ruled out, and the evidence points to seals and walrus as playing at least a minor part.
The disease has only recently been recognized as a human one in the Arctic, although probably it has been of long standing there. The reason for this is the complex clinical picture, which follows closely the natural history of the parasite (Fig. 1).
The disease is a generalized one, the severity of which depends on the number of infective larvae which are ingested. It commences with a gastro– intestinal disturbance, with abdominal pain, nausea, diarrhea, and general weak– ness; this period corresponds to the development of the worm in the intestine. In ten to fourteen days, the larvae are distributed throughout the body by the blood stream and are entering the muscle fibers, causing pains, swelling (es– pecially about the face), difficulty respiration, and some degree of fever. This goes on for about a month, when the symptoms gradually subside. These symptoms vary greatly, not only depending on the number of viable larvae present in the meat, but on the individual’s response. So great is the difficulty of inter– preting them without laboratory aid, that in an outbreak involving a dozen people, a dozen different diagnoses have been made by the same physician. Typhoid fever and influenza are the commonest misdiagnoses.
There is no doubt whatever that clinical trichinosis does occur in the Arctic, and that a number of sharp and even fatal outbreaks which were not diag– nosed at the time were due to this cause.
Diagnosis is difficult. However, certain signs are strongly suggestive, e.g., a history of having eaten uncooked bear or other meat a short time
^ followed by 4 a ^
Fig. 1.

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earlier and of illness in others who partook of the same meal, the acute onset of the disease, the fever (often 104°F.), the swollen face and muscular pain, and the difficult breathing, owing to involvement of the muscles of respira– tion — not the lungs.
Confirmation can be made by demonstrating an eosinophilia in the blood, by an intradermal test or a serological test — the latter being possible only in a laboratory. Biopsy, frequently suggested in textbooks, is of little value, and fecal examination is useless.
Prevention is simple. Meat must be cooked until the proteins are coagu– lated, or it must be refrigerated until the young worms are killed, or it must be treated chemicaly in some way. In passing, it should be noted that only the skeletal muscles are potentially infected and that the heart, liver, and other viscera are safe.
The cysts are not morally ^ normally ^ visible to the naked eye but can be seen with a low-power microscope. However, it is safer to regard all [: ] raw meat in the Arctic (except that from ruminants) as potentially parasitized and to treat it accordingly.
The temperatures required to kill the larvae are not extreme, but it is important that all cysts be exposed to these temperatures, not only those in the outer portions of the meat. The common arctic habit of bringing pieces of meat to a boil in water, cooling, and eating will not sterilize the larger pieces. The temperature must be sufficiently high to change the color of the center of the piece of meat.
The larvae can be killed by cold, although our data here are less well understood. A temperature of −35°F. will kill them almost at once, while −15°F.

EA-Zoo. Cameron: Parasitology

will take a day and a half; but these temperatures must reach the center of the meat, and that may take time. Cooking is the safer procedure.
Pinworm . A more common roundworm, which probably entered the Arctic with the earliest human beings, is the pinworm or seat worm ( Oxyuris ( Entero bius ) vermicularis ). This worm is exclusively human and the infection can come only from man. It is very widespread in its distribution, but is probably most common in temperate climates. However, in the 1930’s the author found it common in all stations in the Eastern Arctic at which the summer supply ship called, while more recently Brown et al ., using more modern techniques on Southampton Island, found about a quarter of those examined harbored this parasite. It is probably common in all arctic regions, the habits of the Eskimos being suitable for its continuance.
The female pinworm is about half an inch long; the male is about one– third this size. Both sexes live in the caecum and appendix, where the female becomes fertilized and gradually moves to the lower bowel. When the gravid period is complete, the female migrates down the rectum and out onto the perianal skin where she ruptures and deposits all her eggs. These eggs are sticky and irritate the skin to which they are attached. This causes scratch– ing, and the microscopic egg may stick to the fingers and be carried to the mouth. Alternatively, the eggs may become detached from the person and become part of the [: ] ^ house ^ dust; some of this reaches the mouth and is swallowed. In either event, the egg becomes swallowed and hatches in the intestine. The young worm which emerges becomes mature in about two to three weeks.
The biology of this worm is such that it tends to infect all the members of a household, and, although it is more common in children, it has been found in Eskimos of all ages. While the pinworm is not a serious parasite, it causes

EA-Zoo. Cameron: Parasitology

a considerable amount of distress. It has been suspected as a cause of ap– pendicitis; while it may cause a chronic type of wriggling appendicitis, its relationship to the fulminating type is more obscure. Diagnosis of the presence of the parasite is by finding the worm in the stool, or the eggs on the skin or under the fingernails. The eggs can be seen only with the aid of a microscope.
Roundworm infections caused by Ascaris lumbricoides have been recorded from the Old World Arctic and Greenland, but they are relatively rare and may have been accidental introductions from the south.
Tapeworm . Tapeworm infections are common in the Arctic, and our records show that these worms occur in all regions there, although we have no figures which can determine their actual incidence. All the specimens we have seen are fish-carried tapeworms of the genus Diphyllobothrium (synonym Dibothrio–cephalus ), but the species involved is uncertain, and there may be more than one. The distribution down the Mackenzie River is continuous with Diphyllo–^i^um latum (or what is usually regarded as this species) in temperate North America, but a second species may exist along the arctic littoral and in the islands. However, we have no indication that any clinical sympton follows in– fection with these worms in North America, and the question of moen ^ nomen ^ clature is at present academic. In the Old World Arctic the species is definitely D. latum , and in certain regions this worm has been associated with symptons of pernicious anemia, although convincing proof that the parasite is the cause of these symp– toms is still not available. No such symptoms have been seen in North America, where the parasite also occurs in dogs, foxes, and bears, these animals prob– ably acting as a reservoir from which man becomes infected.
These tapeworms are large (up to 28 feet long) and live in the small intes– tine where they have to coil in o ^ r ^ der to accommodate themselves to the space

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available. They have no adequate organs of attachment, and so have to keep in continual movement to maintain their position; this movement can often be felt by the patient. They have no digestive system, and they nourish themselves at the expense of the contents of the bowel, digesting foodstuff and intestinal secretions. Their food requirements are relatively large, because a fully developed individual produces several million eggs daily. These are minute structures passed to the exterior in the patient’s excrement. They are not embryonated when laid and develop only if they reach water (fresh water in the case of Diphyllobothrium latum ). In due course they hatch, and the larva passes first of all through a small crustacean and secondly through one or more fish. Man is infected through eating raw fish containing the immature tapeworm, or plerocercoid.
There are numerous related tapeworms in nature with a similar life cycle and numerous species of fish carry these plerocercoids, only some of which are infective to man. They are often several inches long, resembling pieces of fibrous tissue lying in the viscera or muscles. They have no obvious diagnostic features and there is no way of being sure of their infectivity; consequently, all fish should be cooked before being eaten. As the adult tapeworms occur also in wild animals, no effective system of control is avail– able.
Intestinal Protozoa . Probably all the usual intestinal protozoa occur throughout the Arctic in about the same proportions as in Canada. Only a few surveys have been made and accurate figures are not available; however, all the common species have been recorded. Only one, however, is potentially pathogenic, namely, Entamoeba histolytica , which occurs in the large bowel of man in all parts of the world. It was first discovered in Archangle and since

EA-Zoo. Cameron: Parasitology

Hydatid cysts (which are discussed more fully under the parasites of rein– deer) are the bladder-worm stages of a small tapeworm which lives in the intes– tine of carnivores, such as dogs, foxes, and wolves. The bladder stage can live in man and it has occasionally been reported from Indians in the Subarctic. Until recently, however, it [: ] has not been regarded as of real public health importance as its distribution was mainly in wild animals. Recently, however, serological tests in Alaska have shown it to be not uncommon in certain localities among Eskimos, who may receive their infection from arctic foxes or local dogs. The extent or seriousness of the infection is not yet fully appreciated. Else– where, however, the larval parasite which has a very long potential life, is an extremely serious and often fatal parasite, usually infecting the liver of the lungs but capable of developing in almost any part of the body. When fully grown it may be divided into a large number of smaller cysts varying in size from a pea to an orange. There is no medicinal treatment available for the bladder-worm stage in man but it can often be removed surgically. Its presence can be confirmed by serological tests. Infection can result only from the swallowing of the tapeworm egg as a contamination of food or in dust, and the tapeworm egg can come only from the droppings of dogs, foxes, wolves, and related carnivores. The egg, which is microscopic, has considerable resistance to dry– ing and can be blown about in the dust. Prevention, when the tapeworm infection is mainly sylvatic, is difficult, but when dogs alone are infected some good can be done by periodic worm treatment with an anthelminitic such as aerocline ^ arecoline ^ .
Intestinal Protozoa . Probably all the usual intestinal protozoa occur throughout the Arctic in about the same proportions as in Canada. Only a few sur– veys have been made and accurate figures are not available; however, all the common species have been recorded. Only one, however, is po r tentially pathogenic, namely, Entamoeba histolytica , which occurs in the large bowel of man in all parts of the world. It was first discovered in Archangel and since

EA-Zoo. Cameron: Parasitology

has been found both in the Canadian Arctic and in Alaska.
We have no actual knowledge of its pathogenicity in these regions. It is commonly believed, but by no means proved, that not all strains of Enta–moeba histolytica can cause disease. Those which can do not always produce clinical symptoms and may be pathogenic only when some still unknown factor is present. Until we are sure, it is safer to treat all infections as poten– tially dangerous. When it does produce disease, it does so by invading the lining of the large bowel, causing ulcers and some degree of diarrhea or actual dysentery. However, it may also invade the blood stream and cause abscesses in various parts of the body, especially the liver. Such cases do actually occur in the inhabited section of Canada and may result in death, but they are comparatively rare.
Diagnosis is difficult since there are many other causes of dysentery. The diagnosis depends on the microscopic demonstration of the parasite during a clinical attack or of its cyst between attacks. The parasite is carried from person to person by this cyst, which is passed in the stool and subse– qently ingested as a contaminant of food. Efficient sanitation is the most satisfactory m method of breaking this cycle, just as it is with most of the other diarrheic diseases. Treatment is reasonably efficient and specific, but should be carried out only under medical supervision.
Lice . Head lice have been recorded from the Eskimos, as they have from all races of mankind, and their presence in the Arctic calls for no special comment.
Parasites of Food Animals
Whales and Seals . Although whales and seals are intensively infected with parasites of one kind or another, little other than appearance is known about most of these parasites.

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The bearded seal ( Erignathus barbatus ) and the Atlantic walrus ( Odobenus rosmarus ) are much more frequently infected with intestinal worms than are the ringed seal ( Phoca hispida ) and the harp seal ( P. groenlandica ). Both of these species also have been found by Roth to be infected occasionally with the trichina worm, and ther [: ] ^ e ^ is strong circumstantial evidence pointing to walrus meat as the source of several human epidemics in the Arctic. However, the normal feeding habits of these animals suggest that the infection rate must be low. Nevertheless, seal and especially walrus meat should not be eaten raw.
Seals are infected with several species of ascarid worms, that are related to but distinct from those occurring in man and land animals. In at least one case, the ascarid worms have fish of the cod family as intermediate hosts, and in certain areas the musculature of these fish is frequently found to contain young worms an inch or so long. These worms are harmless to human beings and will develop only after being eaten by a seal.
Seals also harbor tapeworms of the genus Diphylobothrium and are sus– pected of being the reservoir for the human tapeworms of this genus found in the arctic islands.
Seals are infected with small thorn-headed worms, and at least one species, Corynosoma semerme , can also live in dogs.
The white whale ( Delphinapterus leucas ) is frequently infected with ascarid worms ( Anisakis spp.) and thorn-headed worms ( Corynosoma strumosum ). These are not transmissible to man and appear to do little damage to the whale. However, the trichina worm has been recorded from this host in Alaska and has been suspected elsewhere.

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Reindeer . The larval stages of three canine tapeworms occur in reindeer. Cysticercus tenuicollis , a bladder worm about the size of an egg, occurs in the liver and other viscera. When eaten by a dog, wolf, or coyote, it develops in the intestine to a tapeworm called Taenia hydatigena . While the tapeworm will develop only in Canidae, the bladder worm will live in most herbivorous animals. Except when very heavy infections occur, the cysts are harmless.
Cysticercus krabbei is a small bladder worm about the size of a pea. It resembles superficially the human bladder worms, taenias, which, however, are absent from the Arctic. It is capable of developing only in Canidae, where it becomes Taenia krabbei . This parasite is common in reindeer (and in Barren Ground caribou, Rangifer arcticus ), and is found in the musculature, being especially common in the deep muscles of the quarters, causing a condition analogous to “measles” in beef or pork. It is harmless to man, and, in any event, is destroyed by cooking.
Hydatid cysts occur in the lungs of reindeer (as well as other Cervidae in the North) as bladder worms of irregular shape, filled with fluid and con– taining small secondary bladders. They are usually no larger than an orange, but as their growth is slow, they may be much larger in older animals. Multiple infections occur and may cause so extensive a destruction of lung [: ] tissue as to destroy the animal. The hydatid cyst is the larval stage of an almost microscopic tapeworm, Echinococcus granulosus , which lives exclusively in the small bowel of wolves, coyotes, and dogs. As in the previous two cases, the ruminant becomes infected by the microscopic tapeworm eggs voided in the excrement of the dog. These eggs are resistant to drying and are swallowed as a contamination of food. Each egg gives rise to a single hydatid cyst. Although man can harbor the cyst but not the tapeworm, he can be infected

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only from the egg passed in the faces of the carnivore; the cysts in the reindeer are harmless to men, but they are the source of infection of car– nivores and should be destroyed by cooking or disposal in such a way as to be inaccessible to dogs or wild Canidae.
There is no treatment for the cystic stages in reindeer, but dogs locally should have their tapeworms removed by medicinal treatment. As these worms also occur in wolves and coyotes, this procedure may not com– pletely eliminate the parasites in the stock, but it will reduce their number. As dogs can be reinfected from caribou offal, treatment should be at regular intervals.
Young reindeer harbor two species of adult tapeworms in their small in– testine. These are not related to the cysts described above, are harmless to man, and nearly harmless to the reindeer.
Roundworms in reindeer occur in the digestive tract and in the lungs. Those living in the lungs, Dictyocaulus viviparous , are whitish creatures, two to three inches long, and are found in the main air tubes, sometimes in con– siderable numbers, causing bronchopneumonia and severe coughing. These round– worms lay minute eggs which are passed in the animal’s droppings. They hatch on the grass and the resulting larvae are in due course swallowed by the deer when grazing. There is no satisfactory treatment known for this condition.
Various small, hairlike roundworms ( Ostertagia sp . and Nematodirus (s.l.) spp.), occur in the true stomach and small intestine of reindeer. These worms are related to species found in other domestic ruminants and those found in the true stomach can be serious parasites. The worms lay minute eggs which pass to the exterior in the droppings where they hatch. The larvae which emerge are ingested with the grass and develop to maturity in three or four weeks.

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Each egg produces only one larva and ultimately only one adult, but each adult lays several hundred eggs. Consequently, continual grazing over the same area results in heavy, gradually increasing infections. While a few worms cause no symptoms, numerous worms cause serious illness and even death, especially in fawns. Infection takes place only during summer, and rota– tional grazing will do much to keep the numbers of parasites within reason– able limits. Where grazing areas are limited, preference for clean areas should be given to the fawns. The larvae of these worms will probably be able to survive over winter, provided they are covered with snow before any severe freezing takes place.
The reindeer warble fly ( Oedemagena tarandi ), which is active during the summer, is a beelike insect with, however, no functional mouth parts, no sting, and only two wings. It lays eggs on the underdown of the reindeer, especially such parts as touch the ground when the animal is lying down; but it also attacks standing animals, causing much gadding among the herd by its buzzing action. The eggs hatch in about a week; the young larvae bore into the skin and migrate gradually to the back of the animal. There they puncture the hide to make a small breathing hole and remain there from October until June. At that time, the maggots make a relatively large hole out of which they crawl, to fall to the ground. There the outer skin hardens and they remain quiescent for about four weeks, at the end of which time the tip of the hardened skin is pushed off and the fully formed fly emerges and commences egg-laying within a few days.
This fly is quite closely related to the warbles of Bovidae farther south, but it produces a rather small warble. It is especially prevalent in yearlings which may have as many as a thousand maggots on their backs; however, infection

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becomes progressively lighter as animals become older. The direct damage done to the living animal is not very severe, but the warble openings are ex– posed to screwworm flies of which the commonest is the black blowfly ( Phormia terrae-novae ). The damage to the hide is more serious than in cattle, because of the thinness of reindeer skins.
The maggots can be removed mechanically in April or May or treated with derris washes. If this is done regularly, not only will the warbles be re– duced in number, but screwworm damage will also be reduced.
The reindeer nostril fly ( Cephenomyia trompe ) is a blackish, two-winged fly which causes great annoyance from June to September. The fly deposits its living larvae directly at the entrance to the nose, and the larvae crawl up the nasal passage and attach themselves at the rear, where they stay until the end of May. The larvae give considerable irritation to their hosts, causing a nasal discharge, staggers, and coughing. In summer the larva falls to the ground, its outer skin hardens in a few hours, and it remains quiescent for two to eight weeks; then the adult fly emerges. There is no available treat– ment, but the pori ^ pro ^ vision of long, dark sheds has been suggested as a relief for reindeer during the fly season.
The maggots of various blowflies are able to develop in living, diseased tissues as well as in decaying animal refuse such as stale meat or dead animals. Some of these flies — especially the black blowfly ( Phormia terrae-novae ) — are serious pests around settlements in the Arctic, attacking wounds on rein– deer and other animals. They attack old warbles, sores on the skin, broken antlers, and so on. Simpl y ^ e ^ surgical methods adopted early will prevent any serious lesions, but if no treatment is given, great destruction of tissue and even death may result. Blowfly control is a matter of hygiene and

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enforcement of sanitary regulations as to disposal of offal.
Caribou . The Barren Ground caribou ( Rangifer arcticus ) has been less studied than the reindeer. Our records show that its parasites are the same as those of the domesticated types, but we have no knowledge of their rela– tive importance. Although their migrations probably reduce infections caused by roundworms, they may increase exposure to larval tapeworm infection because of accompanying wolves and other predators.
Moose . The moose ( Alces americana ) is really a stray into the Arctic. It is often sever ^ e ^ ly infected withhydatid cyst of the lungs (in the Subarctic) and this may result in its death. In Canada it is often severely infected with the winter tick, Dermacentor albipictus , but nothing is known of its parasites in the Arctic.
Musk Ox . There is no information on the parasites of the musk ox ( Ovibos moschatus ) in the arctic regions.
Bear . The important parasites of bears in the Arctic are the trichina worm and the fish tapeworm. The trichina worm has been found in polar bears in both the Old and New Worlds and a considerable percentage (about 50%) appears to be infected. This parasite also occurs in Alaskan brown bears, and probably will be found in the grizzlies and other north-country species. As bears are undoubtedly the most important source of human trichinosis in the Arctic, all bears should be regarded as infected and no bear flesh eaten with– out previous cooking. Human infections have been traced to dried bear “jerky” which was so hard that it had to be chipped with an ax. It should be noted that the liver is not infected with trichina worms, and sickness following the eating of uncooked polar bear liver is not trichinosis.
The bears can be infected only by eating flesh of other mammals which in

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turn have fed on mammalian flesh. While it is not yet clear what is the normal source from which they become infected, it may well be dogs or foxes, which in turn become infected from bear, dog, or fox carcasses. Seals and walrus prob– ably play a minor part in this cycle.
Bears are also a reservoir of infection — probably the main reservoir — of the North American freshwater fish tapeworm. They also are occasionally infected with ascarid worms related to those in dogs but not transmissible to human beings.
Parasites of Dogs and Related Carnivores
Dogs . Sledge dogs throughout the Arctic are infected with numerous in– ternal parasites, of which the most important are roundworms and hookworms. Roundworms ( Toxascaris leonine and Toxocara canis ) are common and important species of particular importance to pups and one species may infect pups in utero. ^ utero. ^ They are cosmopolitan in their distribution. [: ]
Hookworms ( Uncinaria stenocephala ) are tissue-feeding worms related to but quite distinct from the hookworms of man in warm climates; this species is not transmissible to man. As the worm during its feeding processes causes a considerable loss of blood, a heavy ^ ^ infection can result in a fatal anemia, particularly in young animals on a diet deficient in iron. The adult worms live in the small intestine, where the female lays microscopic eggs which pass to the exterior with the droppings. Under summer conditions these hatch, and the resulting larva, after about a week’s growth in the open, is able to pen– etrate the skin of the dog, to return to the intestine, and in a month or so commence egg laying. Dogs confined to corrals or limited spaces in summer are accordingly liable to become heavily infected. Treatment is by the

EA-Zoo. Cameron: Parasitology

administration of tetrachlorethylene to eliminate the worms, and feeding the animal foods high in iron and protein.
The trichina worm is common in arctic dogs but normally causes no symp– toms. Its importance lies in the fact that dead, infected dogs may form a source of infection for bear, which in turn may be eaten by man. Infected dog flesh, of course, is also infective to man unless cooked.
Whipworms ( Trichuris vulpis ) are are common in dogs in temperate cli– mates have not been recorded from the Arctic.
While dogs may be infected with all three taenia-like tapeworms, the larvae of which occur in reindeer and caribou, we have records of Taenia hydatigena and T. tarandi only. It is doubtful if these tapeworms cause any symptoms in the dog, but they should be removed to help prevent infections in herbivores. The larval stage of the third tapeworm, Echinococcus granu–losus , is the hydatid cyst which can develop in man, causing a serious disease; and infected dogs should be destroyed and burned. However , w ^ W ^ e have at records of this parasite in dogs in the Arctic, where ^ although ^ its main hosts are wild carni[][].^vores^ ^ (q.v.). ^
Dogs in the Canadian Arctic act as hosts for two species of fish tapeworms ( Diphyllobothrium spp.), which correspond in their distribution to the two forms from man and are probably identical with them. These worms are spread by feeding on raw fish, and are not directly infective to human beings. More– over, dogs do not appear to be suitable hosts for them and the eggs which they pass are often sterile. The bear is probably the better reservoir for the freshwater species, and seals are suspected as reservoirs for the salt-water species in the arctic islands.

EA-Zoo. Cameron: Parasitology

Liver flukes are common in dogs in the Subarctic, and scanty records suggest that they may occur in the Arctic also. These creatures are small leaflike structures in the bile ducts of the liver, and they cause a serious disease when present in numbers. Infection is contracted by eating the larval stage in fish, which in turn are infected by previous stages develop– ing in snails feeding on the eggs in dog feces. In Canada, the only fish known to carry this fluke is the common sucker ( Catostomus commersonii ). In Old World subarctic species, which differ from the American form, other species of freshwater fish are involved. All these flukes are transmissible to man and to most carnivores, if the infected fish are eaten raw. Cooking completely dest [: ] ^ ro ^ ys all parasites in fish.
Wolves, Coyotes, and Foxes . Wild carnivores have basically the same parasites as sledge dogs, but, in America at least, wolves and coyotes are much more frequently infected with the hydatid tapeworm ( Echinococcus granu–losus ) than are dogs. They are the main source of lung infections of reindeer and caribou, as well as moose and elk in the Subarcitc. These tapeworms are very small creatures, less than half [: ] an inch long, living in the small intes– tine of carnivores; infections are usually multiple, and thousands of the small worms can be found in a single host. As each worm produces several hundred eggs, and as each egg can cause one hydatid cyst, a single infected wolf can infect a large number of gregarious grazing animals in an area. Epidemics even among the comparatively solitary moose have been traced to this cause. All wild carnivores may be infected with the trichina worm. ^ See Reff. in Errata to be inserted here ^
Conclusion
It is evident that a great deal has still to be learned about parasites

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in the Arctic, not only concerning the species present but their distribution and importance. There is an obvious need for surveys such as that commenced by the Institute of Parasitology in 1932 and which is still in progress (1950). The technique adopted was simple. Large garbage cans were taken north by supply ships during summer, and with each was sent a supply of concentrated formalin and glycerin in quart cans. These materials were left at various stations, with instructions to dilute the formalin with water to a 1:20 solu– tion in the garbage can. Formalin is an excellent general pres s ervative, while the glycerin i , being hydroscopic, prevents complete drying of speci– mens. The entrails of animals taken at the various stations were placed in this solution (with a label attached giving the animal’s name and other in– formation), and the material remained there until the next year’s supply ship arrived. Then the liquid was discarded, some moss placed over the entrails as packing, the lid tied on the garbage can, and the material returned to Montreal. There the dissections were completed, and parasites were collected and examined. While far from ideal, this technique is practicable and enables a considerable amount of material to be collected.

EA-Zoo. Cameron: Parasitology

BIBLIOGRAPHY

1. Brandly, P.J., and Rausch, R. “A preliminary note on trichinosis investigations in Alaska,” Arctic, vol.3, pp.105-7, 1950.

2. Brown, M., Sinclair, R.G., Cronk, L.B., Clark, G.C., and Kuitunen-Ekbaum, E. “Intestinal parasites of Eskimos on Southampton Island, Northwest Territories,” Canad.J.Pub.Health vol.39, pp.451-4, 1948.

3. Brown, M., Cronk, B., Desinner, F., Green, J.E., Gibbons, J.E., and Kuitunen-Ekbaum, E. “A note on trichinosis in animals of the Canadian Northwest Territories,” Ibid. vol.40, pp.20-21. 1949.

4. Cameron, T.W.M. “Trichinosis,” Canad.J.Comp.Med . 1937.

5. ----., Parnell, I.W., and Lyster, L.L. “The helminth parasites of sledge-dogs in Northern Canada and Newfoundland,” Canad.J.Res . D, vol.18, pp.325-32, 1940.

6. Connell, F.H. “Trichinosis in the Arctic: a review,” Arctic , vol.2, pp.98-107, 1949.

7. Hadwen, S. and Palmer, L.J. Reindeer in Alaska . Washington, Govt. Print. Off., U.S. Dept. of Agriculture, Bull . No.1089.

8. Lyster, L.L. “Parasites of some Canadian sea mammals,” Canad.J.Res ., vol.18, pp.395-409, 1940.

9. Parnell, I.W. “Animal parasites of north-east Canada,” Canad.Field Nat . vol.48, pp.111-15, 1934.

10. Roth, H. “Trichinosis in arctic animals,” Nature , vol.163, p.805, 1949.

11. Thorborg, N.B., Tulinius, S. and Roth, H. “Trichinosis in Greenland,” Acta Pathologica at Microbiologica Scandinavica , vol.25, pp.778-94, 1948.

12. ----., ----., and ----. “Trikinose paa Grønland,” Ugeskrift for Laeger vol.110, pp.595-602, 1948.

13. Wardle, R.A. “Fish-tapeworm,” Canada. Biological Board. Bull . no.45, 1935.

T. W. M. Cameron

Dog Diseases

EA-Zoology (Thomas W. N ^ M ^ . Cameron)

DOG DISEASES

CONTENTS

Scroll Table to show more columns

Page
Distemper 3
True distemper 3
Paradistemper 4
Contagious canine hepatitis 5
Rabies, 6
Nutritional Diseases 10
Hypervitaminosis A. 10
Avitaminosis B 1 11
Bibliography 13

EA-Zoology (Thomas W. M. Cameron)

DOG DISEASES
Comparatively little is yet known concerning diseases of dogs in the Arctic. Those caused by animal parasites have been referred to in the article “Parasitology.” Such diseases are usually endemic or sporadic. However, in addition to these, epidemics affecting dogs ^ ^ have been recorded periodically while similar diseases have been reported from arctic foxes and wolves. In 1931 Elton (1) collected accounts of many of these out– breaks and, while the symptomatology (accounts of which were based on local reports) varied, emphasis was placed on the frequency of nervous symptoms. An analysis of his data is strongly suggestive of the existence of more than a single cause. It has recently been proved by Plummer (5; 6) that ra v ^ b ^ ies occurs in foxes, wolves, and dogs in the Arctic and has existed there for a long time. It is now firmly established that, once clinical symptoms of rabies appear, the animal invariably dies. Many of the dogs mentioned in Elton’s survey recovered; it follows that these cannot have been suffering from rabies and that the so-called arctic dog disease must be more than a single condition.
Disease syndromes in dogs, including nervous symptoms, are numerous. Some are nutritional in their origin, others are functional, while still others are caused by parasitic worms. However, when the condition is epidemic

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and numerous animals are affected, the disease is usually caused by a virus. The ultravisible viruses are a heterogeneous group of organisms which are invisible to the highest powers of the visual microscope and are so small that they can pass filters which can hold back even the smallest bacteria. In consequence, they are known only by their effects, and while some undoubtedly can and do exist without causing disease, their presence can only be demonstrated by their ability to cause disease in some suitable animal. In other words, it is extremely difficult to demonstrate a carrier state in any virus disease.
There are two known groups of virus disease affecting dogs and related carnivores, and the available evidence suggests that both may be present in the Arctic. The first of these groups includes distemper, paradistemper, and contagious hepatitis (or fox encephalitis), and is apparently carried mainly by droplets spread by coughing or sneezing; the second includes rabies and is mainly carried by inoculation of infective saliva. The first is confined to certain species of carnivores, the second probably will infect any species of mammal. As these viruses are easily destroyed by physical and chemical agencies outside the body, infection can come only from other infected animals. Infectious diseases can never arise spontaneously.
While the distemper group of diseases is most commonly seen in young animals, this is not invariably so, and in isolated communities, such as are found in the Arctic, they might be expected to occur in minor epidemics, affecting all dogs which had not previously had the specific disease. There is no age incidence in rabies but the disease will occur only in animals bitten by a rabid dog, fox, or wolf.

EA-Zoo. Cameron: Dog Diseases

Distemper Group
“Distemper” has long been known as a serious disease of dogs, but the word as used in the earlier days included a variety of conditions. However, in 1926, Carr e ^ é ^ ’s earlier work was confirmed by Laidlaw and Dunkin and dis– temper was believed to be finally and specifically recognized. Since then, however, at least two other related conditions have been identified in dogs and our concept of distemper has to be revised. All three diseases are caused by viruses, all three occur mainly (but by ^ ^ no means exclusively) in young dogs, and all may produce symptoms involving the nervous system. These three diseases are: true distemper, paradistemper, and contagious hepatitis.
Tru [: ] ^ e ^ distemper is a disease of dogs which is transmissible to red foxes and wolves and to ferrets, weasels, ermine, marten, and mink. The virus at first occurs in the blood, but is later concentrated in the upper respiratory tract. In experimental infections, there is an incubation period of three to six (usually four) days followed by comparatively mild symptoms of lassitude, inappetence, fever, and a discharge from the eyes and nose. However, almost always, secondary bacterial invasions take place and there is a patchy pneumonia, digestive disturbances, and nervous symptoms such as muscular spasms, epileptiform seizures, or convulsions; such cases are often but not necessarily fatal.
Infection is by contact or by droplets of saliva, and an animal which recovers is usually immune for life. Most pups are resistan c ^ t ^ for the first four or five weeks of their life but thereafter become susceptible unless artificially immunized.
Vaccines, virus, and serum are available for prophylactic purposes

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and are usually employed either by injecting vaccine (that is, inactivated virus) followed in 10 to 21 days by dried living virus reconstituted just before use, or by ^ ^ injecting virus followed by antiserum in a few hours later. The antiserum can also be used in treatment, which involves treatment of the secondary infections by specific antisera, sulfonamides, and antibiotics, in addition to symptomatic treatment, nursing, and dieting.
Paradistemper or “hard pad” disease is closely related to and often confused with true distemper. [: ] It is caused, however, by a distinct virus which affects not only dogs (including husky dogs) but ferrets as well; it is not known whether it affects foxes or wolves. It is closely related to true distemper, and a partial protection against it is given by distemper inoculation or by a previous attack of distemper; under these conditions dogs are less susceptible and the disease is milder. In dogs the disease runs a distemper-like course and the initial symptoms may be mild with only a slight fever. In some cases the animal may recover and be immune; in others the animal may die in convulsions within a week. The popular name for the disease, hard pad, refers to the frequent hardening of the pads of the feet and sometimes the skin of the nose, at a time when the animal appears to be recovering. This is followed by the onset of encephalitis with symptoms of chores and local muscular twitching, chattering of the teeth, convulsions, continuous hysterical yelping, and movement in circles. This condition either slowly improves or becomes general, accompanied by complete paralysis and death. The disease has been transmitted to ferrets, but these animals, while contracting hard pads, did not develop an encephalitis.
There is no vaccine at present but an antiserum is available in limited

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quantities for treatment. Otherwise the symptoms are treated as they arise, including the use of small doses of aspirin and barbiturates; sulfonamides are, however, contraindicated.
It is still not clear what is the exact relationship between these two disease, whether encephalitis is produced only by paradistemper or whether both are caused by the same virus — one becoming neurotropic, the other enterotropic, as is known to occur with certain other virus infections.
Contagious canine r ^ h ^ epatitis has only recently been recognized as a clinical entity in northern Europe and Great Britain, but it appears to be identical with fox encephalitis d i ^ e ^ scribed by Green and his colleagues from North America (2). It affects dogs, red foxes (and their color varieties), and wolves, but not ferrets and mink. The natural method of transmission is unknown. The virus occurs in the nerve substance, frequently in the spleen, and sometimes in the blood, and persists for some days after death in the absence of putrefaction.
In foxes the disease appears suddenly and may kill the animals without obvious symptoms. More usually, however, there is a loss of appetite for a day or so followed by violent convulsions; this in turn is followed by a deep lethargy sometimes interrupted by convulsions. The animal usually dies in a coma within two days after the onset of symptoms; toward the end paralysis of the whole body or of the hindquarters is common, caused by hemorrhages into the brain or spinal cord, both of which are inflamed.
In dogs the disease has a variable incubation period and often a very short course, the animal dying in a coma in 12 to 24 hours. However, if the dog lives longer than two days, there is a fever accompanied by serious digestive disturbances, abdominal pain, and tonsilitis (including

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great difficulty in swallowing); encephalitis may also develop, especially in pups with hysteria. With encephalitis complications, violent convulsions and death may occur within a week to ten days. It would seem, however, that quite a number of dogs show only transient symptoms and recover, while others seem to show an inherited resistance. “Worming” may cause the onset of serious symptoms in a latent case.
The virus of this condition appears to be longer-lived than that of the other distemper-like diseases and healthy carriers are suspected to exist. In the fox an attenuated virus has been used for protective vaccina– tion with satisfactory results. There is no cross immunity with distemper.
It seems probable that some of the arctic outbreaks are due to this disease although no laboratory confirmation has been attempted. However, it is impossible to rule out true distemper or paradistemper or even some still unknown virus. It is known, for example, that some virus s es which live normally in an inapparent condition in birds are transmitted to certain mammals by mosquitoes, causing symptoms of encephalitis. Both birds and mosquitoes are present in abundance in the Arctic in summer and there is ample opportunity for infection to take place.
Rabies
Rabies is an ancient disease and is widely distributed throughout the world. It exists in several epidemiological forms, such as urban or “street” rabies, which is confined to South and Central America and is primarily a disease of vampire bats. Sylvatic rabies occurs in numerous parts of the world, including South Africa, East Africa, India, and the United States, and Plummer (5; 6) has recently demonstrated conclusively

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its presence in the Arctic in foxes and wolves. It is otherwise absent from Canada. The virus of rabies affects the brain and nervous system, and, while there is some evidence that a number of variants exist, all appear to be strains of the same virus rather than different species. Infection is by ^ ^ inoculation, the virus slowly passing up the nerves to the brain, establishing itself in that organ, and then descending other nerves. In this way it may reach the mouth and pass into the salvia which is the infective agent. The virus is carried by the teeth when an animal bites. It is occasionally, but rarely, carried by the tongue and an existing wound can be infected in this way. It is important to note that in most rabid animals saliv e ^ a ^ is infective only after the development of clinical symptoms, and then only in a proportion of cases; in other words, the bite of an infected animal is infective only after symptoms have appeared and not always then. (This is not necessarily the case with bat rabies, when apparently normal vampire bats can carry the infection. ^ ) ^
The incubation period varies from about ten days to more than six months. In general, the closer the bite is to the head, the sooner symptoms develop. The amount of virus injected may also be of importance. In most cases the incubation period is between three and nine weeks.
Probably all species of mammals are susceptible to this disease, including man, ruminants, and rodents, and once symptoms have developed it is invariably fatal.
The earliest symptoms in dogs may be merely a change in disposition; a dog often becomes more friendly, but is usually irascible and will bite if restrained. It tends to become restless, snapping at moving objects, and showing a depraved appetite. It may lose its appetite, show accelerated

EA-Zoo. Cameron: Dog Diseases

breathing, and run a fever. These generalized symptoms may be absent but if present may last for about three days. Then the animal usually becomes vicious, will bite furiously at anything near it, and often bites off its tail. It no longer knows its master and will bite him as readily as a stranger. It shows a tendency to roam away from home and develops a peculiar howl. Its eyes are often gla x ^ z ^ ed and wide open and it may froth and bleed at the mouth. Subsequently paralysis sets in, usually in the hindquarters, and progresses over the entire body. The dog usually dies in a coma, occasionally in convulsions. These symptoms do not last longer than ten days, and death may occur even sooner. Sometimes there is no furious stage. The dog is sleepy, hides itself, shows no tendency to bite, but becomes paralyzed and dies in three days.
In foxes and wolves the symptoms are somewhat similar but the furious stage is ^ often ^ more severe. They show no fear of man or dog; they will invade settlements and are not only insensible to pai d ^ n ^ , but cannot be frightened by blows or gunshot.
Plummer (5; 6) has demonstrated conclusively the widespread nature of this disease in the Arctic and proved its presence in both arctic foxes and wolves. There is some evidence that the virus is slightly modified and the incubation period shorter in the wolf than in the fox and the dog. It is highly probable that the fox is the reservoir from which dogs are infected, but it is not known how foxes become infected or whether there is another reservoir in such animals as the lemmings. The lemming form a considerable proportion of the food of the arctic fox, and there is little doubt that they could be infected with the virus. It has been suggested that a disease in these rodents, manifested by lassitude, coma,

EA-Zoo. Cameron: Dog Diseases

and death may be rabies. Although there is no proof of this, the lemmings cannot be ruled cut as possible reservoirs.
Rabies was apparently present in bats in South America before the advent of Columbus. It is more than likely that it was introduced into the New World in the early days of human settlement and came to South America with man and his dogs via Alaska. Arctic rabies is also somewhat aberrant, and it too may belong to the same original stock as did the bat virus. On the other hand, it may have been introduced subsequently in wild Canidae from Siberia, or in dogs from Europe or temperate North America. In any event the disease is now well established in the Arctic. It is rather surprising that it has not spread south to temperate Canada, where rabies is absent except for an occasional inroad from the United States.
There is at present no known way of preventing rabies in wild animals. As the disease is only carried by the bite of an animal showing symptoms, considerable protection can be given by confining dogs at night. However, there is an efficient vaccine, available for dogs which gives good protection for a year. Accordingly, animal vaccination with approved vaccine will virtually eliminate the disease in dogs. It must be repeated yearly for an indefinite time, and be administered to dogs of all ages. There is no treatment for rabies.
The diagnosis of rabies in the Arctic is one of some difficulty, but the presence of “crazy” foxes in the vicinity of settlements is strong presumptive evidence of the local existence of the disease. In dogs there may be a history of a bite from such an animal — although not all animals bitten by mad dogs, foxes, or wolves develop rabies. It is important to remember that any carnivore capable of transmitting rabies

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will itself die within a fortnight (usually sooner) and so, if a biting animal is confined instead of immediately destroyed, and is alive a fortnight later, it did not suffer from rabies. If it dies, the evidence that it did have rabies is not conclusive but it is strongly presumptive. The diagnosis can be confirmed only by a laboratory examination of the brain, but pending this examination it is well to assume that the animal died from rabies and to take steps accordingly.
To secure a laboratory diagnosis, the head should be removed, pre– served in ice, and shipped under continuous icing conditions to a convenient central laboratory. (In Canada this should be preferably the Animal Diseases Research Institute, Hull, Quebec, the central veterinary laboratory of the Dominion Government.)
Nutritional Diseases
Symptoms involving the nervous system ^ (for example, canine hysteria) ^ can be caused by roundworms and by improper feeding, (for example, canine hysteria) but there are two nutritional conditions of special importance in the Arctic which may be involved — hypervitaminosis A and avitaminosis B 1 .
Hypervitaminosis A. it is well known that the livers of polar bears, white foxes, bearded seals, and huskies are sometimes toxic to dogs and to human beings, and it has been [: ] shown recently by Rodahl (8) that this toxicity is due to an enormous excess of vitamin A. in bears this is apparently due to recent eating of seals; in the other animals it is probably caused by eating sea fish. The symptoms, which appear suddenly, are those of an acute intoxication, that is, general malaise, drowsiness, muscular weakness, reduced activity, and sometimes death. Small quantities

EA-Zoo. Cameron: Dog Diseases

of these same livers eaten over a longer period produce a picture typical of scurvy, with scattered small hemorrhages all over the body. A similar clinical picture might be expected to occur in animals fed on whole salt– water fish, but this has not as yet been recorded. The disease has so far been seen only in animals fed on livers of mammals which habitually feed on seals or salt-water fish. An excess of vitamin A does not occur in walrus or arctic hares, and presumably does not occur in the livers of other herbivorous animals such as caribou.
Certain workers have divided vitamin A into two fractions: A 1 , common in freshwater fish, and A 2 , in slat-water fish. If this is correct, then this arctic disease is almost certainly due to a great excess of vitamin A 2 . It should be noted that the disease is not trichinosis, which occurs in the same animals. The trichina larva is present only in muscle, never in the liver, and trichinosis cannot be contracted by eating any part of the viscera.
Avitaminosis B 1 occurs in the Subarctic, probably also in the Arctic, and is known as Chastek paralysis. It is actually canine beriberi, but it is caused by an enzyme present in the viscera of numerous species of fish, both in fresh and salt waters, as well as certain shellfish. This enzyme destroyed vitamin B 1 (thiamine or aneurin) and can destroy it in other foodstuffs if left in contact with them. When the diet contains more than 10% of these fish the disease may occur.
The symptoms vary with the severity of the disease and include muscular weakness, leading to failure to eat and loss of voice, as well as rheumatism, especially after heavy work; nervous symptoms including psychic disturbances, cramps, unsteady gait, excessive sensitivity of the

EA-Zoo. Cameron: Dog Diseases

nerves; peripheral neuritis, the animal licking and biting its paws; and paralysis in which the muscles are rigid and the tendon reflexes increased. In severe cases the animal dies, sometimes following convulsions.
These symptoms, if they have not progressed too far, may be relieved by injections of vitamin B 1 or by feeding a diet containing vitamin B 1 , but separate from the raw fish diet, so that the thiamine is not exposed in the stomach to the effect of the enzyme. Cooking of fish destroys the enzyme, so that the vitamin B 1 is not affected.
The relationship between nutritional and virus diseases is not clear, but both involve nervous symptoms. Undoubtedly the existence of a subclinical dietary disturbance could influence the course of a virus infection, and might even be responsible for the symptoms of encephalitis.
The only virus infection definitely proved to exist in the Arctic is rabies, but there is strong presumptive evidence that others do exist, and a high probability that their course is modified by the specialized diets of arctic dogs.

EA-Zoo. Cameron: Dog Diseases

BIBLIOGRAPHY

1. Elton, C. “Epidemics among sledge dogs in the Canadian Arctic and their relation to disease in the arctic fox,” Can.J.Res . vol.5, pp.673-92, 1931.

2. Green, R.G., Ziegler, N.R., Green, B.B., and Dewey, E.T. “Epizootic fox encephalitis. I. General description,” Amer.J.Hyg . vol.12, pp.109-29, 1930.

3. MacIntyre, A.B., Trevan, D.J., and Montgomerie, R.F. “Observations on canine encephalitis,” Vet.Rec . vol.60, p.635, 1948.

4. Pay, T.W.F. “Infectious canine hepatitis ( Hepatitis contagiosa canis (Rubarth)), Ibid . vol.62, pp.551-55.

5. Plummer, P.J.G. “Further note on arctic dog disease and its relationship to rabies,” Ibid . vol.11, pp.330-34, 1947.

6. ----. “Preliminary note on arctic dog disease and its relationship to rabies,” Canad.J.Comp.Med . vol.11, pp.154-60, 1947.

7. Rodahl, K. “Hypervitaminosis A and scurvy,” Ibid . vol.164, pp531, 1949.

8. ----. “Toxicity of polar bear liver,” Nature , vol.164, p.530, 1949.

Thomas W. W ^ M ^ . Cameron
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