In this article I examine moose—wolf interactions over a broad spectrum of moose densities with the primary objective to test empirically whether wolf predation can regulate moose numbers. I also present four conceptual models of moose population dynamics and outline their specific predictions. Based on 27 studies where moose were the dominant prey species, the functional and numerical responses of wolves to changing moose density were derived using an hyperbolic, Michaelis—Menten function. Per capita killing rate was strongly related (P = .01) to moose density, as was the density of wolves (P < .01). Killing rate plateaued at 3.36 moose°wolf—1°(100 days)—1 when predators were fully satiated. The asymptotic value for wolf density was 58.7 animals/1000 km2. Wolf predation rate, as derived from the total predator response, proved to be density dependent from 0 to 0.65 moose/km2, and inversely density dependent at higher moose densities. Predator: prey ratios reflected wolf predation rate poorly because they did not integrate the functional response. An empirical model based on these results suggests that moose would stabilize at 2.0 moose/km2 in the absence of predators, and at °1.3 moose/km2 in the presence of a single predator, the wolf. Density—dependent food competition creates these two high—density equilibrium conditions. If moose productivity is diminished through either deteriorating habitat quality or bear—induced early calf mortality, then a low—density equilibrium (0.2—0.4 moose/km2) is predicted. The model also suggests that when a low equilibrium develops, a “predator pit” is absent or extremely shallow, thus arguing against the appropriateness of a predation—food two—state model. Further research on the density relationship of bear predation, on the effect of alternate prey on wolf total response, and on the regulatory impact of food competition at high moose densities, is required for a full understanding of moose demography.
Habitat selection is a hierarchical process that may yield various patterns depending on the scales of investigation. We employed satellite radio‐telemetry to examine patterns of habitat selection by female woodland caribou in central Saskatchewan at both coarse (seasonal range) and fine (daily area) scales. At each scale, we converted spatial data describing compositions of available and used habitat to standardised resource selection indices and examined them with multivariate analyses of variance. Seasonal ranges generally showed preferential inclusion of peatlands and black spruce dominated stands relative to recently disturbed stands and early seral stage forests. In all populations, caribou preferred peatlands and black spruce forests to all other habitat types at the daily area scale, in general, these patterns may reveal the effective avoidance of wolves, the primary factor limiting caribou throughout the boreal forest. In three populations where seasonal ranges showed the selective inclusion of either young jack pine stands or clearcuts along with peatlands and black spruce forests, we found a relative avoidance of the clearcuts and young jack pine stands at the daily area scale. As all caribou populations in the area are thought to be relics of a once more continuous distribution, the seasonal range selection by animals in disturbed areas may better describe historic rather than current habitat selection. We found inter‐annual variation in selection at the coarser spatial scale in one population, and inter‐seasonal variation in selection at the finer spatial scale in three populations, indicating that the relative grains of the spatial and temporal scales coincide. We were better able to explain the seasonal variations in finer scale selection by considering available forage, a factor less likely than predation to limit woodland caribou populations. The data agree with the theory that the spatial and temporal hierarchy of habitat selection reflects the hierarchy of factors potentially limiting individual fitness.
We studied genetic structure in polar bear (Ursus maritimus) populations by typing a sample of 473 individuals spanning the species distribution at 16 highly variable microsatellite loci. No genetic discontinuities were found that would be consistent with evolutionarily significant periods of isolation between groups. Direct comparison of movement data and genetic data from the Canadian Arctic revealed a highly significant correlation. Genetic data generally supported existing population (management unit) designations, although there were two cases where genetic data failed to differentiate between pairs of populations previously resolved by movement data. A sharp contrast was found between the minimal genetic structure observed among populations surrounding the polar basin and the presence of several marked genetic discontinuities in the Canadian Arctic. The discontinuities in the Canadian Arctic caused the appearance of four genetic clusters of polar bear populations. These clusters vary in total estimated population size from 100 to over 10 000, and the smallest may merit a relatively conservative management strategy in consideration of its apparent isolation. We suggest that the observed pattern of genetic discontinuities has developed in response to differences in the seasonal distribution and pattern of sea ice habitat and the effects of these differences on the distribution and abundance of seals.
The George River caribou herd in northern QuebeclLabrador increased from about 5000 animals in 1954 to 472 200 (or 1.1 caribou.km.') prior to the 1984 calving season. The range used by the herd expanded from 160 O00 to 442 O00 km2 for the period 1971-84. The exponential rate of increase (r) was estimated at O. 11 in the 1970s. Calkfemale ratio in autumn was relatively constant (x = 0.52) from 1973 to 1983, but decreased to about 0.39 in 1984-86. The harvest rate was relatively low in the 1970s (about 3%.yr"), but seemingly increased in the mid-1980s to 5-7% as a result of more liberal regulations and a greater impetus to exploit caribou for subsistence. The cumulative impact of lower calf recruitment and more intensive hunting may have appreciably depressed the growth rate of the herd in 1984-86. A greater year-round competition for food resources and a greater energy expenditure associated with range expansion are presented as probable regulatory factors for the George River herd. It is argued that the nature of caribou-habitat interactions in continental regions generate long-term fluctuations in caribou numbers if human exploitation remains low. At present, wolf predation does not appear to be an important mortality factor capable of regulating the George River herd.
While some prey species possess an innate recognition of their predators, others require learning to recognize their predators. The specific characteristics of the predators that prey learn and whether prey can generalize this learning to similar predatory threats have been virtually ignored. Here, we investigated whether fathead minnows that learned to chemically recognize a specific predator species as a threat has the ability to generalize their recognition to closely related predators. We found that minnows trained to recognize the odour of a lake trout as a threat (the reference predator) generalized their responses to brook trout (same genus as lake trout) and rainbow trout (same family), but did not generalize to a distantly related predatory pike or non-predatory suckers. We also found that the intensity of antipredator responses to the other species was correlated with the phylogenetic distance to the reference predator; minnows responded with a higher intensity response to brook trout than rainbow trout. This is the first study showing that prey have the ability to exhibit generalization of predator odour recognition. We discuss these results and provide a theoretical framework for future studies of generalization of predator recognition.
Within their circumpolar range, polar bears (Ursus maritimus) are not subject to absolute barriers. However, physiographic features do cause discontinuities in their movements. These discontinuities in distribution can be used to delineate population units. Based on satellite telemetry of the movements of female polar bears carried out in 19891998, we used cluster analysis to identify 6 regions within the Canadian and western Greenland Arctic in which movements appear to be restricted enough to identify distinct populations. These regions generally correspond to management units that have been previously identified as Viscount Melville Sound, Lancaster Sound, Norwegian Bay, Kane Basin, Baffin Bay, and Davis Strait. A northsouth substructure was identified for the Baffin Bay population, but it was weaker than the structure identified for the 6 primary units. The 6 units were consistent with genetic information, except for the Baffin Bay Kane Basin separation, and with markrecapture observations and the traditional knowledge of Inuit hunters. Only 2 of 65 bears that provided telemetry information for more than 1 year were classified in different populations in different years. However, annual rates of exchange, measured as the percentage of locations outside the population boundary, ranged from 0.4 to 8.9%. Analysis of markrecapture movements indicated no difference in large-scale movements between the sexes or long-term movements with age. Although our validation criteria for demographic closure were satisfied, the observed rates of exchange between adjacent populations suggest that population dynamics in adjacent populations may not be completely independent.
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