Brad Shults

SYNTHESIS OF SURVIVAL RATES AND CAUSES OF MORTALITY IN NORTH AMERICAN WOLVERINES

Abstract Understanding population vital rates is fundamental to the evaluation of conservation options for wolverines (Gulo gulo). We estimated survival rates and causes of wolverine mortality in trapped and untrapped populations within montane, boreal, and tundra environments using data from 12 North American radiotelemetry studies conducted between 1972 and 2001. Rates were based on data for 62 mortalities of 239 radiomarked wolverines. Mortalities included 22 wolverines that were trapped or hunted, 3 road or rail killed, 11 that were predated, 18 that starved, and 8 deaths of unknown cause. Annual survivorship rates were estimated for sex and age class using Kaplan-Meier staggered-entry techniques. Survival was substantially lower in trapped (<0.75 for all age–sex classes) than in untrapped (>0.84 for all age–sex classes) populations. Human-caused mortality was mostly additive to natural mortality for wolverines in a management context. Logistic growth rate estimates indicated that trapped populations would decline (λ ≅ 0.88) in the absence of immigration from untrapped populations (λ ≅ 1.06). We recommend a system of spatial harvest controls in northern, continuous populations of wolverines and reduction of harvest along with more spatially explicit conservation measures in southern metapopulations.

Diet of wolverines (Gulo gulo) in the western Brooks Range, Alaska

Migratory caribou herds are an important component of the North American tundra. We investigated the wolverine (Gulo gulo) diet in the migratory range of the Western Arctic Caribou Herd in north-western Alaska. Within this area, caribou are absent or occur at low densities for large parts of the year, and thus show a strong seasonality in abundance. Analyses of stomach and colon contents suggested that wolverines primarily consumed caribou during the winter, and that the dietary dependence was related more to caribou mortality than to caribou abundance in the area. We also found indications that wolverines may switch between moose and caribou during periods of low caribou abundance, but that such a switch did not affect wolverine body condition. Our results thus support previous observations that wolverines primarily consume ungulates. However, a better knowledge of how alternative food sources are utilized will be necessary to predict the dietary and demographic responses of wolverines to variations in caribou abundance. We also suggest that further efforts should be made to investigate the effects of other ungulate-dependent predators on wolverine feeding ecology, because such predators may function both as competitors and as suppliers of carrion for scavenging.

Estimating Sustainable Harvest in Wolverine Populations Using Logistic Regression

Abstract Population viability analysis (PVA) is a common tool to evaluate population vulnerability. However, most techniques require reliable estimates of underlying population parameters, which are often difficult to obtain and PVA are, therefore, best used in a qualitative context. Logistic regression is a powerful alternative to traditional PVA methods but has received surprisingly limited attention. Logistic regression fits regression equations to binary output from PVA models at a specific point in time to predict probability of a binary response over a range of parameter values. We used logistic regression on output from stochastic population models to evaluate the relative importance of demographic parameters for wolverine (Gulo gulo) populations and to estimate sustainable harvest in a wolverine population in Alaska. Our analysis indicated that adult survival is the most important demographic parameter to reliably estimate in wolverine populations because it had a greater effect on population persistence than did both fecundity and subadult survival. In accordance with this, harvest rate had a greater effect on population persistence than did any of the other harvest- and migration-related variables we tested. Furthermore, a high proportion of harvested females strengthened the effect of harvest. Hypothetical wolverine populations suffered high probabilities of both extinction and population decline over a range of realistic population sizes and harvest regimes. We suggest that harvested wolverine populations must be regarded as sink populations and that source populations in combination with sufficient dispersal corridors must be secured for any wolverine harvest to be sustainable.

Evaluation of aerial survey methods for Dall's sheep

Abstract Most Dalls sheep (Ovis dalli dalli) population-monitoring efforts use intensive aerial surveys with no attempt to estimate variance or adjust for potential sightability bias. We used radiocollared sheep to assess factors that could affect sightability of Dalls sheep in standard fixed-wing and helicopter surveys and to evaluate feasibility of methods that might account for sightability bias. Work was conducted in conjunction with annual aerial surveys of Dalls sheep in the western Baird Mountains, Alaska, USA, in 2000–2003. Overall sightability was relatively high compared with other aerial wildlife surveys, with 88% of the available, marked sheep detected in our fixed-wing surveys. Total counts from helicopter surveys were not consistently larger than counts from fixed-wing surveys of the same units, and detection probabilities did not differ for the 2 aircraft types. Our results suggest that total counts from helicopter surveys cannot be used to obtain reliable estimates of detection probabilities for fixed-wing surveys. Groups containing radiocollared sheep often changed in size and composition before they could be observed by a second crew in units that were double-surveyed. Double-observer methods that require determination of which groups were detected by each observer will be infeasible unless survey procedures can be modified so that groups remain more stable between observations. Mean group sizes increased during our study period, and our logistic regression sightability model indicated that detection probabilities increased with group size. Mark–resight estimates of annual population sizes were similar to sightability-model estimates, and confidence intervals overlapped broadly. We recommend the sightability-model approach as the most effective and feasible of the alternatives we considered for monitoring Dalls sheep populations.

Sex-Specific Dispersal Patterns of Wolverines: Insights from Microsatellite Markers

Abstract Dispersal of individuals can be defined as movement and settling outside the natal home range. Such dispersal is often sex-biased among vertebrates, and is generally expected to be male-biased in polygynous mammals. We used microsatellite markers scored on harvested wolverines (Gulo gulo) to test the prediction of male-biased dispersal in a population in the western Brooks Range, Alaska. Our analyses suggested a high rate of dispersal within the population, but provided no support for sex differences in dispersal tendencies across the sampled spatial scale. Previous studies have implied male-biased dispersal among wolverine populations on an interpopulation scale. We suggest 3, not exclusive, explanations to reconcile these differences: low power to detect sex biases in dispersal tendencies in this panmictic population; a scale-dependent component in dispersal tendencies, where males are overrepresented among interpopulation migrants; and lower reproductive success for dispersing females compared to more philopatric ones.

Diversity of nitrogen isotopes and protein status in caribou: implications for monitoring northern ungulates

Abstract Nutritional condition is an important determinant of productivity and survival in caribou (Rangifer tarandus). We used samples of excreta (n  =  1,150) to estimate diet composition from microhistology and 2 isotopic proxies of protein status for 2 ecotypes of caribou in 4 herds in late winter (2006–2008). Isotopes of nitrogen (&dgr;15N in parts per thousand [‰]) from excreta samples (urea, diet, and body N) were used to estimate indexes of protein status: the proportion of urea N derived from body N (p-UN) and the difference between the &dgr;15N of the body and urinary urea (&Dgr;body-urea). We examined dietary and terrain characteristics, &dgr;15N, p-UN, and &Dgr;body-urea by ecotype, herd, year, and foraging site. Multiple regression and an information-theoretic approach were used to evaluate correlates of protein status at each foraging site. The dietary and terrain characteristics of foraging sites did not vary by ecotype or herd (P > 0.108); diets were dominated by lichens (68% ± 14.1% SD). The &dgr;15N of urea, diet, body N, p-UN, and &Dgr;body-urea varied among foraging sites within each herd (P < 0.001). Although highly variable, the &dgr;15N of urinary urea was typically low (−4.68‰ ± 2.67‰ SD). Dietary N also had low &dgr;15N (−4.18‰ ± 0.92‰ SD), whereas body N was generally heavier in 15N (2.20‰ ± 1.56‰ SD) than urinary urea or the diet. Both measures of protein status were similarly diverse between ecotypes and among herds, which limited their applicability to monitor protein status at the population level. Although we observed limitations to interpreting estimates of p-UN from highly vagile ungulates, the &Dgr;body-urea may prove to be a useful index of protein status at smaller spatial and temporal scales. Indeed, a portion of the observed variance (r2  =  0.26) in &Dgr;body-urea at each foraging site was explained by the proportion of shrubs in the winter diet. There remains potential in using &dgr;15N in excreta as a noninvasive tool for evaluating protein status in northern ungulates; however, considerable analytical and sampling challenges remain for applying these isotopic approaches at large scales.

A serologic survey for antibodies to three canine viruses in wolverines (Gulo gulo) from the brooks range, alaska

Canine distemper virus (CDV), canine parvovirus type 2 (CPV-2), and canine adenovirus type 1 (CAV-1) are pathogens that are typically associated with canids but may cause serious disease in a wide range of other carnivores. From 1998 to 2002, serum samples from 64 wolverines (Gulo gulo) from the Brooks Range, Alaska, were tested for antibodies to CDV, CPV-2, and canine adenovirus (CAV). Four animals tested positive for antibodies to CDV (7%), one for antibodies to CPV-2 (2%), and none for antibodies to CAV. These are similar to antibody prevalence estimates for other large and medium carnivores in North America.