Figure 5.—Under unusually deep snow conditions, deer used lake shores heavily. (Photo courtesy of L. D. Mech.)Figure 5.—Under unusually deep snow conditions, deer used lake shores heavily. (Photo courtesy of L. D. Mech.)
In considering wolf mobility in snow, two types of movement must be recognized: the trot used during general travel, and the bounding used while chasing prey. The trot is an easy gait of about 5 m.p.h. on firm footing (Mech 1970), and can be continued for hours at a time. During periods of deep snow and high penetrability, most wolf travel is on frozen waterways, roads, snowmobile trails, and animal trails, including the wolves' own pathways, which become well packed with frequent use (fig. 6,7A, B). Such travel was observed during each of the three winters of this study.
Figure 6.—Wolves travel single file in deep snow. (Photo courtesy of L. D. Mech.)Figure 6.—Wolves travel single file in deep snow. (Photo courtesy of L. D. Mech.)
Figure 7.—(A) A single wolf must break his own trail through the snow. (Photo courtesy of L. D. Frenzel.) (B) Regular use by a pack keeps trails open. (Photo courtesy of L. D. Mech.)Figure 7.—(A) A single wolf must break his own trail through the snow. (Photo courtesy of L. D. Frenzel.) (B) Regular use by a pack keeps trails open. (Photo courtesy of L. D. Mech.)
The second type of wolf movement affectedby snow is the leaping and bounding associated with chasing prey. The shallower angle of the wolf's bound (fig. 8) (compared with that of the deer) often causes the wolf to flounder in snow that presents little hinderance to deer (Mech 1970). Such was the case in January and February 1967 in our study area. During 1967-68 no observations of wolves chasing deer were made by the authors, but reports by other field workers indicated that running conditions were similar to those of 1967.
Figure 8.—Wolves run at a shallow angle, thus hindering them in deep snow. (Photo courtesy of D. H. Pimlott.)Figure 8.—Wolves run at a shallow angle, thus hindering them in deep snow. (Photo courtesy of D. H. Pimlott.)
During the winter of 1968-69, wolves also bogged down a great deal in snow when chasing deer. However, after January 1969 the snow was so deep that deer were floundering even more than wolves in many cases. The fact that wolves could run in the trail broken by deer probably also gave the wolves an advantage under the conditions that severely restricted deer movements.
The above observations of snow conditions, deer movements, and wolf movements during the three winters of the study are in accord with observations made on the differences in the ability of the wolves to capture deer during the same period. Two indices support the conclusion that wolves had a much easier time catching deer during February and March 1969 than earlier in the winter and in the two previous winters: (1) the degree of utilization of wolf-killed deer, and (2) the kill rate of radiotagged wolves.
During the winters of 1966-67 and 1967-68, and in December and early January 1968-69, most wolf-killed deer found had been thoroughly eaten, and the bones—if present at all—were well chewed and scattered at each kill (fig. 9). All skin and flesh from the skull were eaten, and the mandible was usually separated from the skull. During late February and early March 1967, few fresh kills were even found, and wolves were returning several times to old kills that had been cleaned up many days before.
Figure 9.—Usually the remains of a wolf kill are well chewed and scattered before the wolves abandon them. (Photo courtesy of L. D. Mech.)Figure 9.—Usually the remains of a wolf kill are well chewed and scattered before the wolves abandon them. (Photo courtesy of L. D. Mech.)
However, in late January 1969 a substantial change began taking place. The skeletons of most kills found were almost intact, the flesh having been eaten from around the bones (fig. 10). Appreciably more skin was usually left on the carcass, especially on the side lying on the snow, and the neck and head were generally intact. This was true even of fawns, which in the past often were almost completely consumed.
Figure 10.—During a period of especially deep snow, wolves abandoned many kills before pulling apart the skeletons. (Photo courtesy of L. D. Mech.)Figure 10.—During a period of especially deep snow, wolves abandoned many kills before pulling apart the skeletons. (Photo courtesy of L. D. Mech.)
In several cases, only about half of the flesh had been eaten from the carcasses. On February 2, 1969, four deer recently killed by wolves were found along a 1½-mile stretch of Birch Lakeand nearby Polaris Lake (Minnesota-Ontario border). One large doe was completely uneaten and remained so for at least 24 hours after discovery from the air. Further, one fawn had only a few pounds of flesh eaten, a yearling doe was half eaten, and another fawn was about 75 percent eaten. Hazardous landing conditions during this period severely limited the number of carcasses that could be examined from the ground, but on February 6 a yearling doe was discovered that had only about 5 to 10 pounds of flesh eaten, and on February 8 an adult doe was found that was completely intact except for wounds.
In past winters some kills had been located that had been only partly eaten, but in each case the carcasses were soon revisited and cleaned up (Mech 1970). This was often not the case in 1969. For the rest of the winter most of the deer killed by wolves in our study area were not as completely consumed as in previous winters. Pimlottet al.(1969) found a similar relationship between the severity of the winter and the degree to which wolf-killed deer were utilized.
Correlated with the above information was the kill history of our radiotagged wolves (Mechet al., p. 1). From December 1968 through January 1969 No. 1051 had killed three or possibly four deer, and generally had spent 6 or 7 days feeding on each. However, throughout most of February this animal visited a new deer carcass (which presumably he killed) every 3 days, and he spent only 1 or 2 days at each. In two cases two new carcasses were found in the immediate vicinity of this animal during the same day, and in each case the wolf spent only 1 day in the area. A second wolf (1053) which had spent most of December and January scavenging on the remains of both deer and moose (Alces alces) that had died long before, made her first known kill of a deer on January 31, 1969. The kill rate of the other three radiotagged wolves also increased, although the data for them are less complete.The average kill rate for all radiotagged wolves and their associates was one deer per wolf per 16 to 20 days before February 1, and one per 8 to 12 days after February 1 (see Mechet al.,p. 1).
FOOTNOTES:[36]M. H. Stenlund. Personal correspondence to L. D. Mech, Oct. 10, 1969.
[36]M. H. Stenlund. Personal correspondence to L. D. Mech, Oct. 10, 1969.
[36]M. H. Stenlund. Personal correspondence to L. D. Mech, Oct. 10, 1969.
Under usual snow conditions throughout most of the range of the white-tailed deer, healthy vigorous individuals can probably escape most attacks by wolves. Observations by Mech (1966), Rutter and Pimlott (1968), and Mechet al.(p. 1) indicate that a high percentage of attempts by wolves to kill deer during winter are unsuccessful. This is further implied by the figures of Pimlottet al.(1969) and Mech and Frenzel (p. 35) showing that at least during winter wolves tend to kill a disproportionate number of old deer as well as those with various abnormalities and pathological conditions.
However, during a winter with extremely deep snow, the usual relationships seem to change somewhat. Fewer deer are able to escape wolves, and a surplus is killed. This means that some individuals not vulnerable under the usual snow conditions become vulnerable during extreme conditions. There are two main possible reasons for this, the effect of the extreme weather conditions on the health and vigor of the deer, and the physical effect of the snow on the escapability of the deer.
In regard to the first possibility, there was limited evidence that during February and March 1969 some fawns and yearlings in our study area were losing their fat stores. Two of three yearlings, and both fawns intact enough for examination during this period lacked back fat, and the marrow in one of six fawn femurs was partly fat depleted. Nevertheless, the third yearling inspected still had back fat, and a 3½-year-old doe had heavy omental, renal, heart, and back fat during the same period. Thus, although an abnormal decline in the physical condition of some deer in the late winter might partly account for the increased kill by wolves during February and March 1969, the effect of snow on the escapability of the deer probably was also involved.
The key difference in snow conditions between the two periods—(1) the winters of 1966-67, 1967-68, and December-January 1968-69, and (2) February and March 1969—was the heavy, persisting accumulation of snow during the latter period, combined with the increasing density of the snow. As our observations show, this greatly hindered the movements of deer fleeing from wolves.
Under more usual conditions, a running deer might sink through the snow to the ground and thus obtain a firm footing from which to spring again. In discussing wolf-caribou relations in snow, Kelsall (1968, p. 249) stated the following: "While caribou (Rangifer tarandus) will sink into snow even deeper than wolves, their longer legs permit them to run efficiently where a wolf will bog down. Nasimovich (1955) considered that roe deer and sika deer could be taken by wolves when snow was not more than 30 cm. (11.8 inches) in depth. At depths above that their pursuit becomes difficult or fruitless."
However, it appears that when snow becomes extremely deep, wolves then gain the advantage. With 22 to 48 inches or more of snow to plow through, a deer would have trouble even touching a firm foundation. According to Kelsall (1969), deer measure only 20 to 24 inches from hoof tip to chest, with legs extended.
It is true that wolves stand even shorter than deer and so might be expected to flounder even more. However, this is where another factor becomes important, the "weight-load-on-track" or total weight per area of track. As Kelsall (1969) has pointed out, the mean weight-load-on-track for deer is extremely difficult to measure directly, because the actual under-surface of the deer's foot slants vertically, and a much greater area may be used to support an animal in snow than on a hard surface. This probably explains the discrepancy between Kelsall's measurements and work done by Verme (1968) in Michigan. According to Kelsall, deer weight-load-on-track (hoof only) varies between 431 and 1,124 gm./cm.2. However, Verme stated that his compaction gauge (with a weight load of about 211 gm./cm.2, described earlier in this paper) sank in virtually the same amount in snow as did deer. Under the snow conditions in our study area, we found that the same type of compaction gauge generally penetrated to a depth within a half inch of that to which deer were sinking. On this basis, it seems reasonable to suggest that a deer in snow is supported by more of its foot thanjust the hoof, and that the actual weight-load-on-track of deer in snow is about 211 gm./cm.2.
For wolves, this measure varies from 89 to 103 gm./cm.2(Foromozov 1946). This means that for the same amount of force applied during running, a wolf would have twice as much support as a deer. It also means that in deep snow a walking wolf generally is much less restricted than a walking deer. Late in February 1969, for example, when deer were seriously limited in their ability to travel, wolves were able to travel widely (Mechet al.,p. 1).
Even though wolves have much greater support than deer, when running they still sink into the snow almost as much as deer under most conditions, probably because both run with such force that snow usually offers little support. Nevertheless, with extremely deep snow, the difference in support factor between wolves and deer could become critical, and this is probably what happened during February and March 1969. With deer seriously restrained by the deep snow, even a slight advantage in favor of the wolf could increase hunting success. A high snow density during that period would accentuate this advantage. This is because until the snow becomes dense enough to hold a running deer, each increase in density would further the advantage of the wolf, which would require only half the density to support it, while it would hinder the deer.
One result of the extreme snow conditions of early 1969 was that deer tended to gravitate to lakes, where snow was shallow and footing was firm. Initially upon disturbance by human beings, and probably by wolves, these deer usually headed inland, but it is apparent from a number of kills examined that when pressed hard by wolves inland, deer headed out onto lakes where possible. Apparently they could run there with better footing. However, frozen lakes also provide wolves with good running conditions, and even seem to give them an advantage (Rutter and Pimlott 1968, Mech 1970), so many of these deer were killed (fig. 11).
Figure 11.—On frozen lakes, wolves often seem to have the advantage over deer, such as in this case where the wolf (center) has just killed a deer and is trying to discourage a raven from joining him in the feed. (Photo courtesy of L. D. Frenzel.)Figure 11.—On frozen lakes, wolves often seem to have the advantage over deer, such as in this case where the wolf (center) has just killed a deer and is trying to discourage a raven from joining him in the feed. (Photo courtesy of L. D. Frenzel.)
Stenlund (1955, p. 44) reported as follows on years of low snowfall, the opposite condition, which demonstrated the same relationship between snow depth and kills on lakes: "The winters of 1951-52 and 1952-53 were abnormally mild with little early snow. As a result, few wolf-killed deer appeared on the lakes and most deer attempted to outrun wolves in the woods."
Thus it appears that extreme snow conditions in our study area increase the vulnerability of deer to wolf predation in three ways: (1) by causing a decline in the health and nutritional state of some members of the deer population; (2) by hindering the escapability of the deer; and (3) by causing deer to congregate on frozen lakes where wolves have the advantage in running.
During the winters of 1966-67, 1967-68, and 1968-69, the interactions of wolves (Canis lupus) and white-tailed deer (Odocoileus virginianus) were observed in northeastern Minnesota from aircraft. Snow depth and supporting ability were also measured during these winters, and the ability of wolves to capture deer was compared for a period of usual snow conditions versus a period of extreme snow conditions.
It was found that during February and March 1969, when snow remained from 2.5 to 3.9 feet deep and failed to support running deer, wolves were able to capture deer more easily. This was evidenced by kills that were left partly or completely uneaten, and by a higher rate of predation by radiotagged wolves and their associates.
Although both wolves and deer floundered in the extremely deep snow, the relatively lighter weight-load-on-track of wolves evidently gavethem a greater advantage than under the usual snow conditions, when wolves were observed floundering more than deer. This factor, plus a decline in the health and vigor of some segments of the deer population and a tendency for deer to congregate on frozen lakes, where wolves have an advantage, help explain the increased vulnerability of deer to wolf predation during the winters of deep snow.
This study was supported by Macalester College, the Minnesota Department of Conservation, the USDA Forest Service, the U.S. Bureau of Sport Fisheries and Wildlife, and the New York Zoological Society. Pilots John Winship, Pat Magie, Jack Burgess, and Don Murray flew the observation planes during radiotracking. Miss Elizabeth Dayton, Mr. Wallace C. Dayton, and the Quetico-Superior Foundation, all of Minneapolis, financed Mech during the writing of this report.
Thanks are also due L. J. Verme, J. P. Kelsall, and J. M. Peek for their helpful reviews.
Foromozov, A. N. 1946. The snow cover as an environment factor and its importance in the life of mammals and birds. (Moskovskoe obshchestvo ispytatelei priroda) Materialy k poznaniyu fauny i flory SSSR, Otdel. Zool. n. 5 (XX). (Translation from Russian published by Boreal Institute, Univ. Alberta, Edmonton, Alberta.)
Kelsall, J. P. 1968. The caribou. Can. Wildl. Serv. Monog. 3, 340 p.
Kelsall, J. P. 1969. Structural adaptations of moose and deer for snow. J. Mammal. 50: 302-310.
Mech, L. D. 1966. Hunting behavior of wolves in Minnesota. J. Mammal. 47: 347-348.
Mech, L. D. 1970. The wolf: the ecology and behavior of an endangered species. 389 p. New York: Natural History Press, Doubleday.
Nasimovich, A. A. 1955. The role of the regime of snow cover in the life of ungulates in the U.S.S.R. Moskva, Akademiya Nauk SSSR. 403 p.
Pimlott, D. H., Shannon, J. A., and Kolenosky, G. B. 1969. The ecology of the timber wolf in Algonquin Provincial Park. Out. Dep. Lands and Forests Res. Rep. (Wildl.) 87, 92 p.
Rutter, R. J., and Pimlott, D. H. 1968. The world of the wolf. 202 p. Philadelphia and New York: J. B. Lippincott Co.
Stenlund, M. H. 1955. A field study of the timber wolf (Canis lupus) on the Superior National Forest, Minnesota. Minn. Conserv. Dep. Tech. Bull. 4, 55 p.
Verme, L. J. 1968. An index of winter severity for northern deer. J. Wildl. Manage. 32: 566-574.
L. David Mech and L. D. Frenzel, Jr.
The timber wolf (Canis lupus) of northeastern Minnesota occupies an area within the range given by Goldman (1944) for the eastern timber wolf (C. l. lycaonSchreber). However, this area is within 150 miles of the eastern edge of the former range of the Great Plains wolf (C. l. nubilusSay), and there is some question as to whether the Minnesota wolf is really an intergrade between these two subspecies. Writing ofnubilus, Goldman (1944, p. 444) stated: "Specimens from eastern Minnesota and Michigan seem more properly referable tolycaon, but relationship tonubilusis shown in somewhat intermediate characters."
In describinglycaonas basically a gray wolf, Goldman made no mention of the occurrence of black or white color phases in that subspecies. However, in discussingnubilus, Goldman (1944, p. 442) wrote the following: "Many color variations are presented. Individuals may be nearly white at any season, except for a sprinkling of black hairs over the back, a small, narrow, but conspicuous, black patch over the tail gland, and a more or less distinctly black tip. Black individuals may occur in the same litter with those normally colored." Goldman also referred tonubilusas "now probably extinct."
Figure 1.—A few wolves observed in the study area were jet black. (Photo courtesy of L. D. Mech.)Figure 1.—A few wolves observed in the study area were jet black. (Photo courtesy of L. D. Mech.)
In the eastern part of the range oflycaon, color phases other than gray appear to be rare as Rutter and Pimlott (1969, p. 188) attest: "The uniformity of the color of timber wolves in many areas is evidenced by the work in Algonquin Park, in Ontario. There, over the past eight years, dozens of packs have been observed from the air. However, we have never been able to discriminate between any of them on the basis of the color variation of individual animals."
Thus it seems significant to report on incidences of black and white color phases in wolves that we have observed in northeastern Minnesota during some 480 hours of flying associated with wolf research (Mechet al.,p. 1). The observations took place in the Superior National Forest, in northern Cook, Lake, and St. Louis Counties during the winters of 1966-67, 1967-68, and 1968-69. A total of 309 sightings were made of wolves that could be classified by color; of these, 11 (3.6 percent) were jet black (fig. 1) and two (0.6 percent) were creamish white, with the cream color the most intense on the back. No doubt some of the grays, and perhaps the blacks and whites, were repeated observations, but the figures should provide a reasonable approximation of the incidence of these color phases in this area. All black or white animals except one were observed with gray wolves (table 1andfig. 2).
A number of black wolves, and a few white wolves, have been seen by other observers, all in the three counties listed earlier. To gain some idea of the past incidence of these color phases in the same general area, we asked Conservation Officers Robert Hodge, Robert Jacobsen, and Frank Baltich of the Ely, Minnesota, area about the numbers of each phase that they took before 1960. They reported killing an approximate total of 580 wolves, of which four were black and three were white or creamish white.
Table 1.—Observations of wolves of black and white color phases
DateLocationColor combinationswithin each packFeb. 24, 1967T64N-R8W-S1Vera Lake3 grays; 1 black; 1 whiteMar. 4, 1967T63N-R9W-S27Lake Two3 grays; 2 blacksDec. 18, 1968T63N-R8W-S35Lake Insula2 grays; 2 blacks[37]Jan. 17, 1969T65N-R8W-S27Carp Lake1 gray; 1 whiteFeb. 1, 1969T63N-R8W-S13Lake Insula4 blacks; 2 grays[38]Feb. 5, 1969T63N-R8W-S8Benezie Lake1 blackFeb. 6, 1969T63N-R10W-S33Clear Lake3 grays; 1 black
FOOTNOTES:[37]These animals were near the shore of the lake, so others may have been inland where they could not be seen.[38]This group might well have been the same as that seen on Dec. 18, 1968.
[37]These animals were near the shore of the lake, so others may have been inland where they could not be seen.
[37]These animals were near the shore of the lake, so others may have been inland where they could not be seen.
[38]This group might well have been the same as that seen on Dec. 18, 1968.
[38]This group might well have been the same as that seen on Dec. 18, 1968.
Figure 2.—A pack of four blacks with two grays (first and third). (Photo courtesy of John Winship.)Figure 2.—A pack of four blacks with two grays (first and third). (Photo courtesy of John Winship.)
Because black and white color phases have rarely if ever been reported forlycaon, yet were well known fornubilus, it is not unreasonable to conclude that the race of wolves now occupying northeastern Minnesota does show strongnubilusinfluence. Goldman examined the skulls only of 10 Minnesota specimens assignable tolycaonand only one referable tonubilus. Because wolves in the known range ofnubilusare thought to be extinct, and because the animals in northeastern Minnesota are legally unprotected and subject to a control program, it seems highly desirable that the question of their taxonomy be studied intensively while specimens are still available.
This study was supported by Macalester College, the New York Zoological Society, the Minnesota Department of Conservation, the U.S. Bureau of Sport Fisheries and Wildlife, and the USDA Forest Service. Mr. Wallace C. Dayton and Miss Elizabeth Dayton, and the Quetico-Superior Foundation, all of Minneapolis, financed Mech during the preparation of this paper. We would also like to thank Dr. J. L. Paradiso, Dr. H. L. Gunderson, and Mr. M. H. Stenlund for reviewing this manuscript.
Goldman, E. A. 1944. The wolves of North America, Part II. Classification of Wolves. p. 389-636. Washington, D.C.: The Amer. Wildl. Inst.
Pimlott, D. H., Shannon, J. A., and Kolenosky, G. B. 1969. The ecology of the timber wolf in Algonquin Provincial Park. Ont. Dep. Lands and Forests Res. Pap. (Wildl.) 87, 94 p.
Tree Improvement Opportunities in the North-Central States Related to Economic Trends, A Problem Analysis, by David H. Dawson and John A. Pitcher. USDA Forest Serv. Res. Pap. NC-40, 30 p., illus. 1970.
Relation Between the National Fire Danger Spread Component and Fire Activity in the Lake States, by Donald A. Haines, William A. Main, and Von J. Johnson. USDA Forest Serv. Res. Pap. NC-41, 8 p., illus. 1970.
Thinning and Fertilizing Red Pine to Increase Growth and Cone Production, by John H. Cooley. USDA Forest Serv. Res. Pap. NC-42, 8 p., illus. 1970.
The Impact of Estimation Errors on Evaluations of Timber Production Opportunities, by Dennis L. Schweitzer. USDA Forest Serv. Res. Pap. NC-43, 18 p., illus. 1970.
User Evaluation of Campgrounds on Two Michigan National Forests, by Robert C. Lucas. USDA Forest Serv. Res. Pap. NC-44, 15 p., illus. 1970.
System Identification Principles in Studies of Forest Dynamics, by Rolfe A. Leary. USDA Forest Serv. Res. Pap. NC-45, 38 p., illus. 1970.
Skiing in the Great Lakes State: the Industry and the Skier, by William A. Leuschner. USDA Forest Serv. Res. Pap. NC-46, 42 p., illus. 1970.
Proceedings of the Ninth Lake States Forest Tree Improvement Conference, August 22-23, 1969. USDA Forest Serv. Res. Pap. NC-47, 34 p. 1970.
A Water Curtain for Controlling Experimental Forest Fires, by Von J. Johnson. USDA Forest Serv. Res. Pap. NC-48, 7 p., illus. 1970.
Wildness Ecology: A Method of Sampling and Summarizing Data for Plant Community Classification, by Lewis F. Ohmann and Robert R. Ream. USDA Forest Serv. Res. Pap. NC-49, 14 p., illus. 1970.
ABOUT THE FOREST SERVICE....As our Nation grows, people expect and need more from their forests—more wood; more water, fish, and wildlife; more recreation and natural beauty; more special forest products and forage. The Forest Service of the U.S. Department of Agriculture helps to fulfill these expectations and needs through three major activities:Conducting forest and range research at over 75 locations ranging from Puerto Rico to Alaska to Hawaii.Participating with all State forestry agencies in cooperative programs to protect, improve, and wisely use our Country's 395 million acres of State, local, and private forest lands.Managing and protecting the 187-million acre National Forest System.The Forest Service does this by encouraging use of the new knowledge that research scientists develop; by setting an example in managing, under sustained yield, the National Forests and Grasslands for multiple use purposes; and by cooperating with all States and with private citizens in their efforts to achieve better management, protection, and use of forest resources.Traditionally, Forest Service people have been active members of the communities and towns in which they live and work. They strive to secure for all, continuous benefits from the Country's forest resources.For more than 60 years, the Forest Service has been serving the Nation as a leading natural resource conservation agency.
ABOUT THE FOREST SERVICE....
As our Nation grows, people expect and need more from their forests—more wood; more water, fish, and wildlife; more recreation and natural beauty; more special forest products and forage. The Forest Service of the U.S. Department of Agriculture helps to fulfill these expectations and needs through three major activities:
The Forest Service does this by encouraging use of the new knowledge that research scientists develop; by setting an example in managing, under sustained yield, the National Forests and Grasslands for multiple use purposes; and by cooperating with all States and with private citizens in their efforts to achieve better management, protection, and use of forest resources.
Traditionally, Forest Service people have been active members of the communities and towns in which they live and work. They strive to secure for all, continuous benefits from the Country's forest resources.
For more than 60 years, the Forest Service has been serving the Nation as a leading natural resource conservation agency.
This is a compilation of four separate reports, each having their own table and figure numbers. I have retained the original table and figure numbers due to all the references made to them within the text. However I did reindex the footnotes for the complete compilation. I made minor punctuation corrections, modified the table formats, moved some illustrations, and made the following typo corrections:Table ofContents: Changed "Occurence" to "Occurrence".Originally: The Possible Occurence of the Great Plains Wolf in Northeastern MinnesotaPage5: Added missing parenthesis after "individuals".Originally: the same color (with the exception of a few black or white individuals (see Mech and Frenzel, page60)Page27, Deleted repeated word "the".Originally: When still on the the ice about 15 feet from shore,Page34, Literature Cited: Changed "Vegetatation" to "Vegetation".Originally: Ohmann, L. F., and Ream, R. R. 1969 Vegetatation studies in the BWCAPage37: Changed "repreductive" to "reproductive".Originally: lungs, liver, kidneys, repreductive tractsPage40: Changed "wildnerness" to "wilderness".Originally: while in the wildnerness more males were takenPage41: Changed "decidous" to "deciduous".Originally: The deciduous first incisors of fawns and the decidousPage42: Changed "end" to "and".Originally: from wolf-killed deer end examined grossly in the fieldPage42,Figure 9: Changed "discoverd" to "discovered".Originally: A permanent first premolar (arrow) was discoverd in M-8.Page47: Changed "wildnerness" to "wilderness".Originally: not surprising that in the wildnerness areaPage57, Footnote36: Deleted duplicate "to".Originally: Personal correspondence to to L. D. Mech, Oct. 10, 1969.Page58: Changed "diffference" to "difference".Originally: Nevertheless, with extremely deep snow, the diffferencePage59, Literature Cited: Changed "roll" to "role".Originally: Nasimovich, A. A. 1955. The roll of the regime of snow