Part 8

Apparently there has been scarcely any published study of the egg-laying or other habits of the adult _Oedemagena_ in relation to _Rangifer arcticus arcticus_, other than a few recent notes by Banfield (1951a: 31-32, fig. 17); but its behavior in relation to the Lapland Reindeer seems to be fairly well known, and it is summarized by Jacobi (1931: 245-246). In the case of the Reindeer, the fly's eggs are laid (during the summer) generally on the legs, belly, and tail region of the victim; the larvae, on hatching, bore through the skin, travel widely through the body, and finally (in the autumn) reach the place for further development--beneath the skin of the back on both sides of the vertebral column. Each one makes a breathing-hole through the skin, and uses this as an exit when leaving the host in the following June. Only the younger animals, from one to about four or five years old, are heavily infested; those still older are spared, possibly having learned to guard themselves better against the fly. Curiously enough, the fawns are said to escape this parasitism entirely.

My own observations on the larvae were restricted to a few Caribou specimens in June and in the autumn. As with the Reindeer, the Caribou fawns in their first autumn showed no visible infestation, as I noted in looking over some fresh hides on September 10, and as was noted again in a fawn of September 26. Fred Schweder, Jr., made the remark that larvae would be evident in the fawns by the following spring; this may indicate that the larvae have not, in the autumn, completed their journey to their final position on the Caribou's back. I learned of no immunity on the part of old adults.

Fullgrown larvae still remained in bucks secured on June 3 (fig. 17) and 18. According to Charles Schweder, they drop out in June. In the buck of June 3 there were perhaps several dozen warbles, each surrounded by a mass of repulsive tissue; in another buck of June 18, there were apparently more than 75. "It may be assumed," says Johansen (1921: 24), "that the pupae lie on the ground for about a month before the flies appear." He found (1921: 29) the adult flies abroad at Dolphin and Union Strait by July 14.

In a buck of August 17 the new warbles (or perhaps merely warble scars from the previous June--_cf._ Banfield, 1951a: 32) on the inside of the skin were not very numerous. Some were medium-sized, but most were so small that it was not deemed necessary to scrape them off; they had comparatively little fatty tissue about them and were merely allowed to dry up. The number of warbles (or warble scars) found in autumn specimens varied considerably, up to a maximum of roughly 200. They were situated mostly along the mid-dorsum, and more on the lower back or rump than farther forward. The number appeared to be approximately 130 in the skin of an adult doe that was nailed to the log wall of a cabin for drying on September 15 (fig. 18). A doe of September 21 seemed to have less than the usual number of warbles.

The nostril fly (_Cephenemyia_) is another serious dipterous parasite of the Caribou. The life history of the European _C. nasalis_ (L.) (or _C. trompe_ [L.]) and its effect on Reindeer are discussed by Bergman (1917), Natvig (1918), and Jacobi (1931: 245) as follows.

This fly attacks the host from June to September, depositing its viviparous larvae in the nostrils. The Reindeer attempts to fend off the fly, striking at it with its hoofs and keeping its nostrils closed as far as possible. Once deposited, the lively larvae crawl into the inner nasal passages and as far as the larynx, where they fasten themselves and live on the mucus. A Reindeer may harbor as many as 130 of these parasites. They range from 6 to 26 mm. in length. Their particular growth begins at the end of March, and they are ready for pupation up to May. The host assists their exit by continual sneezing and snuffling. In the last stages they are a great affliction for the host, and they sometimes cause its death. Pupation takes place in or on the ground, under some sort of cover, and it lasts for 15-19 days. The flies have been found emerging from July 12 to 31.

The corresponding parasite of the Barren Ground Caribou is a similar or perhaps identical species, with a parallel life history. Its chief activity as an adult doubtless occurs in July and August. A number of the mature larvae were found in the throat of the buck of June 3; two of them that were preserved measure approximately 7 mm. in diameter and 27 and 30 mm. in length. A large mass of such sizable parasites in the throat might easily become a serious obstacle to comfortable living or even to survival on the part of the host. Presumably the larvae drop to the ground at about the same period as those of _Oedemagena_. Fred Schweder, Jr., remarked concerning the buck of August 17 that these larvae are never found at that season, and Charles Schweder made the same remark concerning a doe specimen of September 21. It would appear either that they remain so small as to escape detection at this time or that they do not reach the throat on their short journey from the nostrils until some later period of the year. Johansen (1921: 24) records larvae only 2-3 mm. long in the nasal passage at the end of May.

Since the bulk of the Caribou population has passed well to the northward of the Nueltin Lake region by the time the larvae of _Oedemagena_ and _Cephenemyia_ drop out of the bodies of their hosts to pupate briefly on or in the ground (say in the latter part of June), one is tempted to speculate on the possibility that the adult flies found here in August may have followed their prospective victims for many miles in their southward migration. However, Porsild remarks (1943: 386) that they "apparently do not travel very far."

Certain kinds of behavior exhibited by the Caribou in attempting to fend off the parasitic flies are discussed in the section on _Shaking off moisture and insects_.

The adults of _Oedemagena tarandi_ (L.) were determined by Mr. C. W. Sabrosky, of the Bureau of Entomology and Plant Quarantine; and the larvae of _Oedemagena_ and _Cephenemyia_ by Dr. W. W. Wirth, of the same bureau. The larvae of the latter genus are regarded as probably _C. trompe_ (L.); they were new to the collection of the United States National Museum.

_References._--Hearne, 1795: 197; Franklin, 1823: 241; Richardson, "1825": 328-330, and 1829: 242; Godman, 1831, +2+: 284; Murray, 1858: 210; B. R. Ross, 1861: 438; Pike, 1917 (1892): 58-59; J. B. Tyrrell, 1892: 128, and 1894: 442; Whitney, 1896: 239; Russell, 1898: 228-229; Jones, 1899: 411; Hanbury, 1900: 67, and 1904: 32, 137, 194; Preble, 1902: 41; R. M. Anderson, 1913b: 504; Stefánsson, 1913b: 204, 212-213, 333; Douglas, 1914: 191-192; Malloch, 1919: 55-56; Hewitt, 1921: 67; Johansen, 1921: 22-24, 29, 35, 37; Stefánsson, 1921: 247; Blanchet, 1925: 32, and 1926b: 47; Birket-Smith, 1929 (1): 56, 133; Seton, 1929, +3+: 109-111; Critchell-Bullock, 1930: 193; Hoare, 1930: 33, 37-38; Kitto, 1930: 89; Jacobi, 1931: 244-245; Munn, 1932: 58; Sutton and Hamilton, 1932: 84-86; Birket-Smith, 1933: 90, 92; Ingstad, 1933: 48, 135; Hornby, 1934: 105; Soper, 1936: 429; Henriksen, 1937: 25, 26; Hamilton, 1939: 247, 301; Murie, 1939: 245; Clarke, 1940: 70, 95; Downes, 1943: 226, 255; Manning, 1943a: 53; Porsild, 1943: 386; Gavin, 1945: 228; Harper, 1949: 228; Banfield, 1951a: 31-33; Barnett, 1954: 104.

_Ectoparasites_

It was in vain that I searched a number of fresh specimens for lice, mites, fleas, or ticks. The Schweder boys spoke of never having noticed any such parasites. Seton (1929) mentions none, and Jacobi (1931: 243) records only a louse (_Linognathus tarandi_) from the Reindeer. "Lice are not known from caribou according to Ferris (in conversation)" (Weber, 1950: 154).

_Relations to Reindeer_

Recent discussions of the possibility or advisability of introducing domesticated Reindeer to replace, or to augment the diminishing supply of, native American Caribou in various new localities prompt a brief review of the subject.

It may be remarked at the outset that acclimatization attempts in the Old World have generally been abortive. Wild Reindeer introduced from Finmark into Iceland in the eighteenth century flourished for a time, but by 1917 they were almost exterminated. A number of different introductions into Great Britain, Denmark, Germany, Austria, Switzerland, and Italy came to naught. On the other hand, the introduction of Lapland Reindeer on the subantarctic island of South Georgia in 1908 seems to have turned out successfully. (Jacobi, 1931: 158-165; Harper, 1945: 473-474.) A saving feature in each of the above-mentioned cases was the absence of any native Reindeer whose racial purity might have been destroyed by the newcomers.

Decrease of local American stocks of Caribou, and consequent suffering of native populations who had in past generations depended upon these animals for a major portion of their food supply, have led to introduction of foreign Reindeer in several regions of North America, from Newfoundland and Labrador in the east to Alaska in the west. The persons responsible were doubtless inspired by high humanitarian motives; but it is doubtful if they could have thoroughly considered or foreseen the serious biological consequences of their efforts.

In Alaska, importation of domesticated Siberian Reindeer began in 1892. By the 1930's the herds had increased to an estimated total of 600,000. For various reasons, however, the industry has so far declined that by 1949 the total number of Alaskan Reindeer had become reduced to about 28,000 head. Disinclination of Eskimos for reindeer-herding and mixture of their stock with wild Caribou were important reasons for this decline. (Lantis, 1950.) From the biologist's point of view, the most unfortunate result was the large-scale interbreeding with the native Grant's Caribou (_Rangifer arcticus granti_) and the progressive extermination of that fine animal in a pure form by dilution with inferior alien blood. Among Alaskan Reindeer, "constant inbreeding has led to a noticeable reduction in the prolificness of the females, and degeneration is to be observed in many herds" (Hewitt, 1921: 323).

In 1908 Dr. Wilfred T. Grenfell brought 300 Lapland Reindeer to Newfoundland. After some years they were transferred to the north shore of the Gulf of St. Lawrence and finally to the island of Anticosti. (Hewitt, 1921: 324-328; Seton, 1929, +3+: 92.) In 1911, 50 of these Reindeer were shipped from Newfoundland to the Slave River region. Most of them escaped (probably to contaminate the local stock of Caribou), and by 1916 the last survivor of this band in captivity had succumbed (Hewitt, 1921: 329-330).

"A large part of the reindeer in Alaska are south of the Arctic Circle on the comparatively mild shores of Bering Sea, where there are several months of open tidewater navigation; vegetation is more luxuriant [than in Arctic Canada] and conditions easier in general. There the reindeer were introduced into a country where the wild caribou had been virtually exterminated, and a large native population were anxious to take up a new mode of support. The percentage of profits has appeared unduly large in Alaska because statisticians have been unable to take into account the value of the services of a large body of devoted missionaries, government teachers, and other unselfish persons who put their best efforts into years of unpaid extra work to make the reindeer successful and beneficial to their charges.

"Canada has a large area of Arctic and sub-arctic lands beyond the reach of possible cultivation, still occupied by large numbers of wild caribou and remnants of musk-oxen, with native inhabitants who derive a living from them and add to the national wealth by fur production. These Indians and Eskimos are still far from being either able or willing to enter upon a pastoral stage of existence, and moreover, they are now enjoying an era of prosperity from the fur industry which may be temporary, but which they will not relinquish for the slower and less profitable prospects of the herder." (R. M. Anderson, 1924: 330-331.)

In 1921 some Norwegian Reindeer were landed at Amadjuak, Baffin Island (Seton, 1929, +3+: 92). The lack of further reference to the Baffin Island animals by such subsequent investigators as Manning and Soper would seem to indicate that the reindeer have not survived, unless through mixture with the native Caribou. An attempt in 1922 at acclimatization in Michigan "ended in total failure" (Seton, 1929, +3+: 93).

"The Barren Grounds . . . still feed enormous herds of caribou. . . . The greatest danger to this industry [reindeer-raising] is just these wild herds, which would be very apt to absorb the tame animals. This problem may perhaps become a fatal one to the Eskimos, for there might very easily come a most difficult transitional period, when the caribou would be too few in numbers to form a definite basis for the existence of the people, but on the other hand numerous enough to make reindeer breeding difficult." (Birket-Smith, 1933: 121.)

In northwestern Alaska "large numbers of reindeer are constantly escaping the herders and joining the wild caribou. It seems that it will be but a short time until there will be no pure bred caribou along that part of the coast. . . . As the reindeer are protected, and the caribou are killed at every opportunity, the former will doubtless prove the dominant animal and in time overcome the caribou, with hybridization the inevitable result." (Bailey and Hendee, 1926: 22.)

"The caribou's greatest menace is not the wolf, nor the hunter, but man's economic developments, principally the raising of reindeer. Wherever reindeer herds are introduced, caribou must of course disappear, for both cannot occupy the same range. The disappearance of the caribou along the Bering Sea and Arctic coasts, while regrettable, was unavoidable in view of the development of reindeer herding in this section, which is ideal for the purpose. . . .

"The mingling of reindeer with the main caribou herds should be avoided. Reindeer herds maintained in close contact with migrating caribou suffer frequent losses through strays. Already the domestic reindeer are mingling with the caribou herd of Mount McKinley National Park. . . . [Hybridization] would be regrettable in interior Alaska, which has produced a splendid type of wild caribou, coming near at least to being the largest on the continent." (Murie, 1935: 7.)

Murie's extensive experience with these animals in Alaska has led him to remark further (1939: 245):

"The greatest hazard to the Caribou is the possible occupation of the range by man's agricultural activities. . . . The most serious danger is introduction of domesticated Reindeer on wild Caribou range, for the wild herds must be removed in order to make possible the safe herding of the domestic animals. . . . There is not room for both of these animals on the same or closely adjacent ranges."

Porsild points out (1943: 386, 389) that sparsely covered grazing areas are suitable for Caribou but not for Reindeer; and that the former disappear before expanding Reindeer culture.

"Perhaps the worst threat of all to the caribou has been the introduction of reindeer culture along the arctic coast. This has resulted in interbreeding between the wild caribou and their inferior domesticated relatives. When and if this mixture extends to all the herds of the Barren Grounds, the caribou may be written off the record as a pure species; the animal will have become extinct through dilution, as the biologists express it." (Harper, 1949: 239.)

The American Society of Mammalogists, at its annual meeting in 1950, passed the following resolution (_Jour. Mammalogy_ +31+ (4): 483, 1950):

"That the American Society of Mammalogists urges that the Canadian Government not undertake the introduction of reindeer into Ungava. Before any introduction even is seriously considered, those persons involved in any planning are urged to make a thorough study beforehand of the problems of integrating lichen ecology, reindeer biology, and native culture--serious problems that have not been solved to date on any workable scale on the North American continent. It would be particularly deplorable if an introduction, to aid the natives, led to early successes and high hopes, then eventual failure."

Porsild, who knows the Reindeer thoroughly at first hand, has made (1951: 53) the following observation:

"Thus far these experiments [at introduction into America] have met with only partial or indifferent success, because reindeer nomadism is incompatible with present trends of cultural development and because the North American Arctic is too thinly populated to provide a ready market for reindeer products."

Referring to the region of the Brooks Range in northern Alaska, Rausch says (1951: 190):

"The mixture of inferior reindeer bloodlines with the native caribou is serious. This has already occurred to a considerable degree, and it is hoped that proper control will be exercised if the reindeer industry is revived in Alaska. Ear-notched animals have been killed in the Anaktuvuk Pass country, and white reindeer have been seen running with the caribou. The number of unrecognized reindeer passing through could be great."

At present the Barren Ground Caribou is apparently the third most abundant member of the deer family on our continent, being exceeded by the White-tailed Deer and the Mule Deer (_cf._ Jackson, 1944: 7-8). No other member of this family could be expected to be so eminently and thoroughly adapted to its Arctic environment or to thrive so well on the very ground where nature has been molding and perfecting its characters for thousands of years. No naturally occurring relative--Moose, Deer, or Woodland Caribou--undertakes to compete with it on its own particular range. It requires practically nothing for the maintenance--and increase--of its present numbers, other than an enlightened policy of conservation. (As indicated on a previous page, the feminine wearers of Arctic Fox furs must bear a heavy share of responsibility for the decline of the Barren Ground Caribou in recent decades.) Our highest authorities have pointed out the impracticability of Caribou and Reindeer occupying the same range.

Would it not be the part of wisdom to exclude the inferior domesticated alien, with its difficult and generally unsuccessful culture in North America, and thereby to give the wonderful wild Caribou of the Barrens its best chance for survival?

_References._--Chambers, 1914: 350-351; Hornaday, 1914, +2+: 105-108; Hewitt, 1921: 323, 329-330; R. M. Anderson, 1924: 330; Kindle, 1928: 74; Seton, 1929, +3+: 92-93; Blanchet, 1930: 53-54; Birket-Smith, 1933: 121; Godsell, 1934: 276; Murie, 1935: 7, 1939: 245-246, and 1941: 435; Porsild, 1943: 386, 389; Rousseau, 1948: 96; Harper, 1949: 239; Polunin, 1949: 24; Lantis, 1950; Hustich, 1951; Porsild, 1951: 53; Rausch, 1951: 190; Scheffer, 1951.

NUMERICAL STATUS

There seems to be a general impression, among those who have known the Barren Ground Caribou at first hand for a considerable period, that the population has been reduced by something like a half during the past generation. "Recent preliminary aerial survey has indicated that their numbers, although less than the previous estimates of 3,000,000 (R. M. Anderson, 1938; Clarke, 1940), which were based upon the carrying capacity of the Arctic tundra, are probably comparable to their primitive numbers in the central portions of the range" (Banfield, 1949: 478). A definite reduction is indicated along the Arctic coast and on the Arctic islands (R. M. Anderson, 1937: 103, and 1938: 400; Banfield, 1949: 478, 481, and 1951a: 13-14). While large numbers still remain in southwestern Keewatin, there are no reports of any such mass occurrence as was witnessed by the Tyrrell brothers on the upper Dubawnt River on July 29, 1893; that throng was estimated at 100,000 to 200,000 animals (J. B. Tyrrell, 1897: 165).

During the big movement of the last week of August, 1947, I may have seen as many as 500 Caribou on one or two days, in herds numbering up to 150 individuals. A striking proportion of those observed seemed to occur in bands of roughly 25 animals. On August 25 Fred Schweder, Jr., reported about a thousand crossing Little River, in bands of as many as 100 individuals. On October 11 Charles Schweder observed a thousand Caribou resting on a hill 3 miles long in the vicinity of Four-hill Creek. In November he found thousands, in herds up to 300 strong, moving south from the upper Kazan River. These figures may give a faintly approximate idea of the numbers occurring in the general region of Nueltin Lake in a year considered less good than an average one. On the other hand, toward the coast of Hudson Bay, there were reports of a greater number of autumn migrants than in ordinary years.

In October, about 1944, tracks indicated that 2,000 or 3,000 animals had crossed Windy River in the vicinity of Four-hill Creek in the night (_fide_ Charles Schweder). About October 10, 1946 (a year of unusual abundance), Fred Schweder, Jr., witnessed the passage of thousands in one day in this vicinity; he got the impression of "the hills moving with Deer." (Yet this was the season when the Caribou passed mainly to one side of the upper Kazan River, so that nearly one-third of the local band of Eskimos starved to death.) In the first part of May, about 1942 or 1943, John Ingebrigtsen came to a nameless lake, about half a mile by a mile and a half in extent, somewhere east of Duck Lake, Manitoba. It appeared "absolutely full of Caribou," and he estimated their number at not less than 20,000. This would mean a density of no more than about 50 per acre.

_References._--Jones, 1899: 368, 374; J. B. Tyrrell, 1894: 442, and 1897: 10, 49-50, 165; Whitney, 1896: 240; Seton, 1911: 220, 258-260; R. M. Anderson, 1913b: 502; Hornaday, 1914, +2+: 225-226; Nelson, 1916: 460; Thompson, 1916: 100-101; Kindle, 1917: 108-109; Buchanan, 1920: 130-131; Hewitt, 1921: 56, 64-66; Stefánsson, 1921: 255; R. M. Anderson, 1924: 329; Blanchet, 1926b: 48, and 1930: 52; Kindle, 1928: 72-73; Seton, 1929, +3+: 131-134; Critchell-Bullock, 1930: 159-160; R. M. Anderson, in Hoare, 1930: 52-53; Kitto, 1930: 87; Jacobi, 1931: 201-202; Munn, 1932: 58; Birket-Smith, 1933: 89; Ingstad, 1933: 160; R. M. Anderson, 1938: 400; Clarke, 1940: 65, 84-91, 101-104; Downes, 1943: 258-260; Wright, 1944: 185-188, 191, 193; Yule, 1948: 287-288; Banfield, 1949: 478, 481, and 1951a: 9, 13-14; Harper, 1949: 231, 239; Anonymous, 1952: 261; Barnett, 1954: 96.

GENERAL HABITS

_Daily periods of activity and rest_

According to Charles Schweder, the Caribou do not move about much at night; that seems to be their principal time for sleep. They exhibit a definite tendency to pause and rest also toward the middle of the day. Several instances have already been given of the animals resting at such a time on frozen lakes and rivers: lakes southwest of Reindeer Lake, March 18; lakes south of Lake Athabaska, April 16; Seal River, May 31; Windy Bay, June 6 (mid-morning). Open hilltops are evidently sought likewise for both nocturnal and mid-day rests: knoll by Windy River, June 3; Josie's Hill, June 20; ridge by Little River, August 24 (about 9 a.m.). (For details, see sections on _Winter range_, _Spring migration_, and _Fall migration_.)