L. David Mech

Scent-marking in lone wolves and newly formed pairs

Abstract Scent-marking was studied in wolves ( Canis lupus ) along 133 km of tracks in northern Minnesota during winters of 1975 to 1976 and 1976 to 1977 and in two captive packs and four captive pairs for various periods. Lone wolves, which possess neither mates nor territories, rarely marked by raised-leg urination and defaecated and urinated less along roads and trails, where territorial pairs and packs generally marked. Newly formed pairs marked the most, eventually decreasing their rates to those of established packs. Generally, wolves that scent-marked also bred, whereas non-marking wolves usually did not breed. Scent-marking apparently is important to the success of courtship in new pairs and to reproductive synchrony in established pairs, as well as serving a territorial function.

Wolf-Pack Buffer Zones as Prey Reservoirs

In a declining herd, surviving deer inhabited overlapping edges of wolf-pack territories. There, wolves hunted little until desperate, in order to avoid fatal encounters with neighbors. Such encounters reduce wolf numbers and predation pressure and apparently allow surviving deer along territory edges to repopulate the area through dispersal of their prime, less vulnerable offspring into territory cores.

Elk Calf Survival and Mortality Following Wolf Restoration to Yellowstone National Park

Abstract We conducted a 3-year study (May 2003–Apr 2006) of mortality of northern Yellowstone elk (Cervus elaphus) calves to determine the cause for the recruitment decline (i.e., 33 calves to 13 calves/100 adult F) following the restoration of wolves (Canis lupus). We captured, fit with radiotransmitters, and evaluated blood characteristics and disease antibody seroprevalence in 151 calves ≤6 days old (68M:83F). Concentrations (𝑥̄, SE) of potential condition indicators were as follows: thyroxine (T4; 13.8 μg/dL, 0.43), serum urea nitrogen (SUN; 17.4 mg/dL, 0.57), γ-glutamyltransferase (GGT; 66.4 IU/L, 4.36), gamma globulins (GG; 1.5 g/dL, 0.07), and insulin-like growth factor-1 (IGF-1; 253.6 ng/mL, 9.59). Seroprevalences were as follows: brucellosis (Brucella abortus; 3%), bovine-respiratory syncytial virus (3%), bovine-viral-diarrhea virus type 1 (25%), infectious-bovine rhinotracheitis (58%), and bovine parainfluenza-3 (32%). Serum urea nitrogen, GGT, GG, and IGF-1 varied with year; T4, SUN, and GG varied with age (P ≤ 0.01); and SUN varied by capture area (P = 0.02). Annual survival was 0.22 (SE = 0.035, n = 149) and varied by calving area but not year. Neonates captured in the Stephens Creek/Mammoth area of Yellowstone National Park, USA, had annual survival rates >3× higher (0.54) than those captured in the Lamar Valley area (0.17), likely due to the higher predator density in Lamar Valley. Summer survival (20 weeks after radiotagging) was 0.29 (SE = 0.05, n = 116), and calving area, absolute deviation from median birth date, and GG were important predictors of summer survival. Survival during winter (Nov–Apr) was 0.90 (SE = 0.05, n = 42), and it did not vary by calving area or year. Sixty-nine percent (n = 104) of calves died within the first year of life, 24% (n = 36) survived their first year, and 7% (n = 11) had unknown fates. Grizzly bears (Ursus arctos) and black bears (Ursus americanus) accounted for 58–60% (n = 60–62) of deaths, and wolves accounted for 14–17% (n = 15–18). Summer predation (95% of summer deaths) increased, and winter malnutrition (0% of winter deaths) decreased, compared with a similar study during 1987–1990 (72% and 58%, respectively). Physiological factors (e.g., low levels of GG) may predispose calves to predation. Also, the increase in bear numbers since wolf restoration and spatial components finer than the northern range should be considered when trying to determine the causes of the northern Yellowstone elk decline. This is the first study to document the predation impacts from reintroduced wolves on elk calf mortality in an ecosystem already containing established populations of 4 other major predators (i.e., grizzly and black bears, cougars [Puma concolor], and coyotes [Canis latrans]). The results are relevant to resource managers of the Yellowstone ecosystem in understanding the dynamics of the elk population, in providing harvest quota recommendations for local elk hunts to the Montana Department of Fish, Wildlife and Parks, the United States Fish and Wildlife Service regarding wolf and grizzly bear recovery, and to all areas worldwide where predators are increasing, by providing managers with information about potential carnivore impacts on elk populations.

Productivity, Mortality, and Population Trends of Wolves in Northeastern Minnesota

Population parameters, mortality causes, and mechanisms of a population decline were studied in wolves ( Canis lupus lycaon ) from 1968 to 1976 in the Superior National Forest. The main method was aerial radio-tracking of 129 wolves and their packmates. Due to a decline in white-tailed deer ( Odocoileus virginianus ), the wolf population decreased during most of the study. Average annual productivity varied from 1.5 to 3.3 pups per litter, and annual mortality rates from 7 to 65 percent. Malnutrition and intraspecific strife accounted equally for 58 percent of the mortality; human causes accounted for the remainder. As wolf numbers began to decline, pup starvation became apparent, followed by lower pup production, and then by increased intraspecific strife. At higher densities, adult pack wolves were the most secure members of the population, but as the population declined, they became the least secure because of intraspecific strife.

A study of the genetic relationships within and among wolf packs using DNA fingerprinting and mitochondrial DNA

SummaryDNA fingerprinting and mitochondrial DNA analyses have not been used in combination to study relatedness in natural populations. We present an approach that involves defining the mean fingerprint similarities among individuals thought to be unrelated because they have different mtDNA genotypes. Two classes of related individuals are identified by their distance in standard errors above this mean value. The number of standard errors is determined by analysis of the association between fingerprint similarity and relatedness in a population with a known genealogy. We apply this approach to gray wolf packs from Minnesota, Alaska, and the Northwest Territories. Our results show that: (1) wolf packs consist primarily of individuals that are closely related genetically, but some packs contain unrelated, non-reproducing individuals; (2) dispersal among packs within the same area is common; and (3) short-range dispersal appears more common for female than male wolves. The first two of these genetically-based observations are consistent with behavioral data on pack structure and dispersal in wolves, while the apparent sex bias in dispersal was not expected.

Winter severity and wolf predation on a formerly wolf-free elk herd

We studied wolf (Canis lupus) predation on elk (Cervus elaphus) in Yellowstone National Park from 17 March to 15 April 1997 (severe winter conditions) and from 2 to 31 March 1998 (mild winter conditions) 2-3 years after wolves were reintroduced to the park. Elk composed 91% of 117 kills. Data comparisons for 1997 versus 1998 were: hunting success rate, 26% versus 15%; kill rate, 17.1 kg/wolf/day versus 6.1; percent of kill consumed in first day 7 versus 86; percent femur marrow fat of adult kills, 27 versus 70; calf:adult ratios of kills, 2:33 versus 17:23; sex ratio of kills, 14M:19F versus 17M:6F; mean age of elk killed, males 6.1 years, females 15.2 versus males, 4.8, females 13.0. Winter severity influenced the wolf-elk relationship more than the naivete of the elk herd to predation by wolves.

Pack size and wolf pup survival: their relationship under varying ecological conditions

SummaryThe relationship between pack size and two parameters of reproductive success (litter size at 7–8 months and pup weights at 5–6 months) were determined for two wolf (Canis lupus) populations in northern Minnesota. Pup weights were not correlated with pack size for either population. Litter size, however, was correlated with pack size, but the direction of the relationship varied between the two study populations.In the superior National Forest, where prey were scarce and the wolf population was declining from high densities, litter size and pack size were inversely related. Pairs produced more surviving pups than did larger packs with one or more potential helpers. In the Beltrami Island State Forest, where prey were relatively abundant and the wolf population was increasing, pack size and litter size were positively correlated. The results suggest that ecological factors, such as prey availability, affected the ability or willingness of various pack members to provide food or other care for the pups.The lack of correlation between number of auxiliaries and number of pups in canid populations with low and declining prey densities may be explained on the basis of heterogeneous prey density resulting in drastic annual variation in litter production. No study to date has measured the actual benefit that pups derive from helping by auxiliaries, and the costs and benefits of it. The relationships discussed herein can be considered valid only after such research is completed.

Wolf population survival in an area of high road density

-Wolf mortality in a high-road-density area of Minnesota exceeds that in an adjacent wilderness, and is primarily human-caused. The wolf population there is maintained primarily by ingress from the adjacent wilderness areas. A road density of 0.58 km/km2 can be exceeded and the area still support wolves if it is adjacent to extensive roadless areas.

Simulation analyses of space use: Home range estimates, variability, and sample size

Simulations of space use by animals were run to determine the relationship among home range area estimates, variability, and sample size (number of locations). As sample size increased, home range size increased asymptotically, whereas variability decreased among mean home range area estimates generated by multiple simulations for the same sample size. Our results suggest that field workers should ascertain between 100 and 200 locations in order to estimate reliably home range area. In some cases, this suggested guideline is higher than values found in the few published studies in which the relationship between home range area and number of locations is addressed. Sampling differences for small species occupying relatively small home ranges indicate that fewer locations may be sufficient to allow for a reliable estimate of home range. Intraspecific variability in social status (group member, loner, resident, transient), age, sex, reproductive condition, and food resources also have to be considered, as do season, habitat, and differences in sampling and analytical methods. Comparative data still are needed.

RELATIONSHIP BETWEEN SNOW DEPTH AND GRAY WOLF PREDATION ON WHITE-TAILED DEER

Survival of 203 yearling and adult white-tailed deer (Odocoileus virginianus) was monitored for 23,441 deer days from January through April 1975-85 in northeastern Minnesota. Gray wolf (Canis lupus) predation was the primary mortality cause, and from year to year during this period, the mean predation rate ranged from 0.00 to 0.29. The sum of weekly snow depths/month explained 51% of the variation in annual wolf predation rate, with the highest predation during the deepest snow. J. WILDL. MANAGE. 50(3):471-474 Wolves capture more prey during severe winters with deep snow (Pimlott et al. 1969, Mech and Frenzel 1971, Haber 1977, Mech and Karns 1977, Peterson 1977, Eide and Ballard 1982). However, except for extreme snow conditions, the relationship between degree of winter severity and wolf predation rate has only begun to be studied. Increased wolf kill of moose (Alces alces) has been related to increasing snow depths (Peterson and Allen 1974, Haber 1977, Gasaway et al. 1983), but no quantitative measure of this relationship between wolves and white-tailed deer has been made. This paper is the 1st attempt at analyzing such a relationship. This study was supported by the U.S. Fish and Wildl. Serv., U.S. For. Serv. North Central For. Exp. Stn., Mardag Found., Weyerhauser Found., and W. Dayton. Statistical help was provided by G. L. Hensler and R. E. McRoberts. We also thank the Minn. Dep. Nat. Resour. and several field technicians.