Growth Rates and Variances of Unexploited Wolf Populations in Dynamic Equilibria

Several states have begun harvesting gray wolves (Canis lupus), and these states and various European countries are closely monitoring their wolf populations. To provide appropriate perspective for determining unusual or extreme fluctuations in their managed wolf populations, we analyzed natural, longterm, wolf-population-density trajectories totaling 130 years of data from 3 areas: Isle Royale National Park in Lake Superior, Michigan, USA; the east-central Superior National Forest in northeastern Minnesota, USA; and Denali National Park, Alaska, USA. Ratios between minimum and maximum annual sizes for 2 mainland populations (n = 28 and 46 yr) varied from 2.5–2.8, whereas for Isle Royale (n = 56 yr), the ratio was 6.3. The interquartile range (25th percentile, 75th percentile) for annual growth rates, Nt+1/Nt, was (0.88, 1.14), (0.92, 1.11), and (0.86, 1.12) for Denali, Superior National Forest, and Isle Royale respectively. We fit a density-independent model and a Ricker model to each time series, and in both cases we considered the potential for observation error. Mean growth rates from the density-independent model were close to 0 for all 3 populations, with 95% credible intervals including 0. We view the estimated model parameters, including those describing annual variability or process variance, as providing useful summaries of the trajectories of these populations. The estimates of these natural wolf population parameters can serve as benchmarks for comparison with those of recovering wolf populations. Because our study populations were all from circumscribed areas, fluctuations in them represent fluctuations in densities (i.e., changes in numbers are not confounded by changes in occupied area as would be the case with populations expanding their range, as are wolf populations in many states)