Between 1965 and the present, wolves in the study area were neither protected nor bountied, and the influence of trapping and hunting is thought to have been negligible.
THE STUDY AREA
This study was conducted in the Superior National Forest (fig. 1) in northern St. Louis, Lake, and Cook Counties of northeastern Minnesota (92 deg. west longitude, 48 deg. north latitude), an area well described by Stenlund (1955). Most of the data were collected from within and immediately south of the Boundary Waters Canoe Area, a special wilderness region in which travel by motorized vehicles is restricted. The total study area encompasses approximately 1.5 million acres, and numerous lakes and rivers comprise about 15 percent of this area (fig. 2). The topography varies from large stretches of swamps to rocky ridges, with altitudes ranging from 1,000 to 2,300 feet above sea level (fig. 3). Winter temperatures lower than -30 deg. F. are not unusual, and snow depths generally range from 20 to 30 inches on the level. However, an important exception occurred in early 1969 when depths of 45 inches and more accumulated in much of the area. Further details on snow conditions in the study area during the period of this investigation are given by Mech et al. (see page 51). Conifers predominate in the forest overstory, with the following species present: jack pine (Pinus banksiana Lamb.), white pine (P. strobus L.), red pine (P. resinosa Ait.), black spruce (Picea mariana (Mill.) B.S.P.), white spruce (P. glauca (Moench) Voss), balsam fir (Abies balsamea (L.) Mill.), white cedar (Thuja occidentalis L.), and tamarack (Larix laricina (DuRoi) K. Koch). However, as a result of extensive cutting and fires much of the conifer cover is interspersed with large stands of white birch (Betula papyrifera Marsh.) and aspen (Populus tremuloides Michx.). Detailed descriptions of the forest vegetation were presented by Ohmann and Ream (1969).
METHODS
The observations discussed in this paper were all made from aircraft, the method of flying being that reported by Burkholder (1959) and Mech (1966a). The following aircraft were used (in order of size): Aeronca Champ, Supercub, Cessna 172, Cessna 180, and Cessna 206. The smaller aircraft were excellent for holding in tight circles during observations but had the disadvantage of being slow and cold; the larger planes could cover the study area much more quickly and were more comfortable, but were not as maneuverable during observations. For radiotracking, to be discussed below, the best compromise seemed to be a Cessna 172.
To make observations of wolves, we flew over frozen waterways until tracks were found, and then followed the tracks until we lost them or saw the wolves (fig. 4). Several times we located wolves directly just by scanning the lakes. However, because there seemed to be a number of packs in the area, and because most wolves were the same color (with the exception of a few black or white individuals) (see Mech and Frenzel, page 60), it usually was not possible to follow packs from one day to the next and be certain of identification. Moreover, it was impossible to locate any pack at will because most wolves also spent much time inland.
Therefore, to facilitate our observations and to obtain data on wolf movements and extent of range, we began a radiotracking program in 1968-69. A professional trapper, Robert Himes, was employed to capture the wolves. Using Newhouse No. 4 and 14 steel traps at scent-post sets, he caught two wolves, and captured another with a live-snare similar to that used by Nellis (1968); the senior author trapped two additional wolves (fig. 5).
The four wolves held in steel traps were restrained by a choker (fig. 6), and then anesthetized by intramuscular injections (fig. 7A, B) of a combination of 30 mg. of phencyclidine hydrochloride (Sernylan, Parke-Davis Co.) and 25 mg. promazine hydrochloride (Sparine, Wyeth Laboratories) as prescribed by Seal and Erickson (1969); these drugs proved most satisfactory.
The fifth wolf (a female), which was captured around the chest by the live-snare, was handled without drugs. A forked stick was used to hold down her head (Kolenosky and Johnston 1967), and she offered no resistance (fig. 8). Evidently she went into shock or some other psychophysiological state of unconsciousness, for after her release she remained on her side and did not move for 1.5 hours, despite our prodding during the first few minutes (fig. 9). Then suddenly she leaped up and ran off.
Each wolf was examined, outfitted with a radio transmitter collar 15 inches inside circumference (fig. 10) and tagged with identification numbers in both ears (fig. 11). Each transmitter was of a different frequency in the 150 MHz range, emitted a pulsed signal ranging from 75 to 350 pulses per minute, and had a calculated life of at least 300 days (fig. 12). Two types of 12-inch whip antennas were used on the transmitters: one type extended up the side of the collar and then stuck out above for 6 inches; the other was fully attached inside the collar and extended up one side, around the top, and partly down the other side. The transmitter, batteries, and antenna were molded into a collar of acrylic weighing 11 ounces (Mech et al._ 1965). All radio equipment functioned flawlessly for at least 5 months, and one transmitter continued operating for at least 9 months.
For tracking radio-equipped wolves, a directional yagi antenna (fig. 13) was attached to each of the wing struts of an aircraft and connected inside to a portable receiver. The usual tracking technique was to fly at 1,500 to 3,000 feet elevation to the last known location of the wolf being sought (fig. 14). If a signal was not obtained at that point, the aircraft spiraled upward until the signal was found or until 10,000 feet altitude had been reached. If the signal still was not heard, a search pattern was flown at 10,000 feet. The range of the signal from this altitude was 15 to 35 miles; at 3,000 feet it was 10 to 15 miles. Collars with antennas molded fully inside gave only about two-thirds the range of those protruding partly, but could be expected to last longer because the antennas could not break off. It is unknown whether any protruding antennas did break during the study, but on January 5, 1970, one wolf was recaptured, and its antenna had broken.
When a signal was received, the aircraft was headed in the approximate direction of the source until the signal strength reached a peak; a 90 deg. turn was then made in the direction the signal seemed the strongest. A series of these maneuvers soon narrowed the area to the point where visual search was possible. After practice and experience with this technique, we could locate the approximate source of the signal within 10 to 30 minutes after first receiving it.
Even though the radiotagged wolves spent most of their time inland, often in stands of conifers, they were frequently observed from the aircraft. The technique was to circle at 300 to 800 feet altitude around a radius of a quarter mile from the point where the strongest signal emanated. From December through April, 65 percent of the wolves located by radio were sighted; the rate was much higher for more experienced personnel. A pack of five wolves that was tracked was seen 31 times out of 33 attempts during February and March.
Whenever wolves were located, radiotagged or not, observations were made from an altitude that did not disturb them. Packs varied in the concern shown the aircraft, but only one or two ran from it. The radiotagged wolves, and a pack of 10 to 13 animals, were habituated to the aircraft and usually could be observed from altitudes of 500 feet and less without disturbance (fig. 15).
Almost all the radiotracking was done from aircraft, but when inclement weather prevented flying, some attempts from the ground succeeded when wolves were close enough to roads. The usual range on the ground was 0.75 to 1.50 miles. One wolf was approached to within 35 feet through radiotracking.
FOOTNOTES:
Mention of trade names does not constitute endorsement by the USDA Forest Service.
The acrylic collar was fashioned by the Davidson Co., Minneapolis, Minnesota, which also produced some of the transmitters. Other transmitters and two radio receivers were manufactured by the AVM Instrument Co., Champaign, Illinois.
RESULTS AND OBSERVATIONS
Aerial observations made during this study involved 490 hours distributed as follows: January, February, March 1967--124 hours; February 1968--10 hours; December 1968 through August 1969--356 hours. Seventy-seven observations involving a total of 323 wolves were made (table 1), excluding animals located through radiotracking.
One male and four female wolves were radiotagged, and they and their associates were followed intermittently for periods of 5 to 8 months (table 2). All except one initially suffered some injury to a foot. Three of these animals were seen limping, but only in one case was the limp judged extreme enough to have significantly affected the movements or behavior of the animal. In that one case, the wolf (No. 1057) was caught in a steel trap on an extremely cold night, and her foot froze. After that she was often seen hopping on three legs. She was not able to keep up with her pack, which consisted of 10 to 13 members, and her movements were much restricted compared with those of other wolves. However, she was frequently observed feeding on fresh kills, and may even have made them herself.
Table 1.--Sizes of wolf population units observed in northeastern Minnesota
#: Number %: Percent
+------------------+-------------------------------------------+ | | Wolf observations | |Population unit+----------+----------+----------+----------+ |(number of wolves)| Winter | Winter | Total |Winters| | | 1966-67 | 1968-69 | | 1948-53 | +------------------+----------+----------+----------+----------+ | # % # % # % # % | | 1 8 31 17 33 25 32 48 43 | | 2 3 12 6 12 9 12 24 22 | | 3 3 12 2 4 5 6 7 6 | | 4 -- -- 7 14 7 9 7 6 | | 5 2 7 4 8 6 8 8 7 | | 6 3 12 4 8 7 9 7 6 | | 7 2 7 1 2 3 4 4 4 | | 8 3 12 3 6 6 8 3 3 | | 9 -- -- 2 4 2 3 3 3 | | 10 2 7 2 4 4 5 -- -- | | 11 -- -- -- -- -- -- -- -- | | 12 -- -- 1 2 1 1 1 1 | | 13 -- -- 2 4 2 3 -- -- | | --------------------------------------------+ |Total number | | of wolves 109 -- 214 -- 323 -- 318 -- | |Total number | | of observations 26 -- 51 -- 77 -- 112 -- | |Mean population | | unit size 4.2 -- 4.2 -- 4.2 -- 2.8 -- | +--------------------------------------------------------------+
FOOTNOTES:
Because wolf packs sometimes split temporarily, these figures may not strictly represent actual pack sizes; nevertheless they should provide reasonably accurate approximations.
From Stenlund (1955).
Table 2.--Background information on five radiotagged wolves studied in northeastern Minnesota
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