Evidence for pseudopregnancy and induced ovulation in captive wolverines (Gulo gulo)

Progesterone levels and vaginal smears were monitored to detect estrus and formation of corpora lutea during the first year of a 4-year study of reproduction in captive wolverines. No evidence of spontaneous ovulation was detected during the first year, and most females did not attain complete vaginal cornification. Follicle stimulating hormone was used in subsequent years to induce estrus in several females, and human chorionic gonadotropin (hCG) was used to induce ovulation. Females treated with hCG were artificially inseminated with fresh wolverine semen. Prolonged elevation of serum progesterone above 1 ng/ml was only observed in females that received hCG. The profiles and duration of the progesterone secretory pattern of these females closely resembled that of other mustelids that exhibit a prolonged delay of implantation. Progesterone remained below 1 ng/ml throughout the year in all females that did not receive hCG. No kits were produced. The data suggest that ovulation in this species is normally induced by coitus, and that pseudopregnancy can occur, lasting as long as pregnancy. © 1993 Wiley-Liss, Inc.     

The diet of breeding female wolverines (Gulo gulo) in two areas of Finland

The wolverine (Gulo gulo) hunts and scavenges for food. We examined how the presence of a semi-domesticated reindeer (Rangifer tarandus tarandus) influences the diet of breeding female wolverines in Finland. We compared the dietary composition of wolverines breeding in the reindeer management area (northern Finland) with that of wolverines breeding outside this area (eastern Finland) by analyzing 421 scats collected from six active dens in northern Finland and four dens in eastern Finland during 2004–2006. The occurrence of food sources in the diet was assessed for both areas using logistic mixed-effects models. Diet breadth per study area was also calculated. As expected, semi-domesticated reindeer was the most important food source (66 %) in northern Finland. Mountain hares (Lepus timidus) were the second most important food item (16 %). Wolverines also utilized grouse (7 %), small rodents (6 %), and moose (Alces alces; 4 %) in their diet. In eastern Finland, the most utilized prey species was moose (55 %), while the diet also comprised mountain hares (13 %), domestic pigs (Sus scrofa; 13 %), grouse (11 %), and small rodents (2 %). The diet of wolverines in eastern Finland included more moose and less semi-domesticated reindeers and small rodents than in northern Finland. The niche breadth did not differ between the study areas. In eastern Finland, the diet of wolverines shifted to 55 % moose and 0 % semi-domesticated reindeer; compared to 4 % moose and 66 % semi-domesticated reindeer in northern Finland. Our study highlights the important role of ungulates in the diet of breeding female wolverines. In areas with a low density of medium-sized ungulates, scavenging for wolf- and human-killed moose and carcasses on feeding sites plays an essential role in food acquisition by wolverines.     

Assessing population structure and gene flow in Montana wolverines (Gulo gulo) using assignment-based approaches

In North America, wolverines once occupied a continuous range from Alaska southward to New Mexico. In the lower 48 states, small remnant populations remain only in the northwestern United States. Among these remnant populations, the Montana population has the highest probability of long-term persistence given its size and proximity to healthy populations in Canada. In this study, we evaluate population genetic structure and gene flow among Montana wolverines using 10 polymorphic microsatellite loci. Bayesian and frequency-based assignment tests revealed significant population substructure and provide support for at least three subpopulations in Montana. FST values between subpopulations ranged from 0.08 to 0.10 and provide evidence for male-biased dispersal. The high degree of population substructure and low levels of gene flow contrast results from wolverine population genetic studies in less fragmented landscapes of Alaska and Canada. This study provides additional support for the hypothesis that large carnivore populations of Montana are becoming increasingly fragmented due to human development and disturbance.     

Behavioral and endocrine correlates of reproductive failure in social aggregations of captive wolverines (Gulo gulo)

Sociality in mammals is often viewed as a dichotomy, with sociality contrasted against solitariness. However, variation within these broad categories may have strong effects on individual fitness. For example, reproductive suppression of social subordinates is generally associated with group living, but suppression may also occur in solitary species if the behavioral and physiological processes involved can be modulated by the demographic environment. To investigate whether behavioral and physiological traits that normally are associated with group living might be latent even in a solitary species, we explored the level of sociality and investigated causes and mechanisms of reproductive failure in female wolverines Gulo gulo that experienced a highly aggregated social environment in captivity. Behaviorally, females showed low levels of aggression and intermediate levels of social interactions. Reproductive failure seemed to have been related to low social rank and to have occurred between ovulation and implantation in 13 out of 15 breeding attempts. However, three of eight females observed to mate produced offspring, indicating that no individual female fully managed to monopolize breeding. Reproductive failure was not related to elevated levels of glucocorticoid stress hormones. Rather, elevated glucocorticoid levels during the mating season were associated with successful reproduction. We suggest that social tendencies and physiological mechanisms mediating reproductive suppression may be viewed as reaction norms to the social environment. We further suggest that the social flexibility of solitary carnivores might be greater than is commonly observed, due to ecological constraints that may limit aggregation.     

Phylogeography and Post-Glacial Recolonization in Wolverines (Gulo gulo) from across Their Circumpolar Distribution

Interglacial-glacial cycles of the Quaternary are widely recognized in shaping phylogeographic structure. Patterns from cold adapted species can be especially informative - in particular, uncovering additional glacial refugia, identifying likely recolonization patterns, and increasing our understanding of species’ responses to climate change. We investigated phylogenetic structure of the wolverine, a wide-ranging cold adapted carnivore, using a 318 bp of the mitochondrial DNA control region for 983 wolverines (n = 209 this study, n = 774 from GenBank) from across their full Holarctic distribution. Bayesian phylogenetic tree reconstruction and the distribution of observed pairwise haplotype differences (mismatch distribution) provided evidence of a single rapid population expansion across the wolverine’s Holarctic range. Even though molecular evidence corroborated a single refugium, significant subdivisions of population genetic structure (0.01< Φ_ST <0.99, P<0.05) were detected. Pairwise Φ_ST estimates separated Scandinavia from Russia and Mongolia, and identified five main divisions within North America - the Central Arctic, a western region, an eastern region consisting of Ontario and Quebec/Labrador, Manitoba, and California. These data are in contrast to the nearly panmictic structure observed in northwestern North America using nuclear microsatellites, but largely support the nuclear DNA separation of contemporary Manitoba and Ontario wolverines from northern populations. Historic samples (c. 1900) from the functionally extirpated eastern population of Quebec/Labrador displayed genetic similarities to contemporary Ontario wolverines. To understand these divergence patterns, four hypotheses were tested using Approximate Bayesian Computation (ABC). The most supported hypothesis was a single Beringia incursion during the last glacial maximum that established the northwestern population, followed by a west-to-east colonization during the Holocene. This pattern is suggestive of colonization occurring in accordance with glacial retreat, and supports expansion from a single refugium. These data are significant relative to current discussions on the conservation status of this species across its range.     

Two new species of Sarcocystis (Apicomplexa: Sarcocystidae) infecting the wolverine (Gulo gulo) from Nunavut, Canada

Infection with Sarcocystis species is common in many species of animals, but it has not yet been reported in wolverines (Gulo gulo). Histological sections of tongues from 41 wolverines in the Kitikmeot Region, Nunavut, Canada, were examined for sarcocysts. Sarcocysts were found in 33 (80.4%) wolverines. Two structurally distinct types of sarcocysts were found. Type A sarcocysts were thin (<1 μm thick) walled. Ultrastructurally, the parasitophorous vacuolar membrane (Pvm) had minute undulations, but it lacked villar protrusions and was not invaginated into the granular layer. The bradyzoites were slender, about 5 × 1 μm in size. Structurally, these sarcocysts were distinct from known species of Sarcocystis and possessed a novel 18S and ITS-1 sequence, sharing 98% and 78% sequence similarity with Sarcocystis canis . A new species name, Sarcocystis kalvikus, is proposed for type A sarcocysts. In contrast, type B sarcocysts had relatively thicker (about 2 μm) cyst walls and larger bradyzoites, each about 10 × 2-3 μm. Ultrastructurally, the Pvm on the sarcocyst wall had villar protrusions that were either mushroom-like or sloping. Molecular analysis identified a unique 18S and ITS-1 sequence that placed them in a clade within the Sarcocystidae. Based on histology, TEM, and genetic data, the new name, Sarcocystis kitikmeotensis, is proposed. Sarcocystis kalvikus was found in 14 (34.1%), S. kitikmeotensis was found in 7 (17%), and both species were found in 12 (29.2%) of 41 wolverines.     

Serologic survey for infectious canine hepatitis virus in grizzly bears (Ursus arctos) from Alaska, 1973 to 1987

Serum antibody prevalence of infectious canine hepatitis virus was 12% (90 of 725) for grizzly bears (Ursus arctos) from Alaska (USA) during the period 1973 to 1987. Prevalence was highest on Kodiak Island at 29% (37 of 127). Prevalence of exposure at individual collection areas did not change significantly over time. There were no significant sex-specific differences in prevalence. Prevalence was directly related to age, but it was 0% for bears less than 2-yr-old. Young bears which are exposed to the virus may develop clinical disease and die as a result of the infection. This disease may be a factor affecting grizzly bear population dynamics.     

Quantification of character-species in vegetation at the tree limit,Brooks Range,Northern Alaska

Quantification of character-species using Dahl's uniformity index is tested on phytosociological data using species composition of tree limit communities on the south slope of the Brooks Range, Northern Alaska as an example. Species in 246 relevés are grouped by minimum spanning tree (MST) analysis. The resulting six species groups are ordinated in a congruent pattern by detrended correspondence analysis along the two main axes of mesotopographic-scale nutrient and moisture gradients. The eutraphent species are restricted to the more fertile sites of the undulating topography in this permafrost area. Uniformity indices are highest for the species in eutrophic sites with average frequencies. In contrast, most oligotraphent species are ubiquitous and occur even within the more eutrophic communities on raised mounds, shallow soils and in the exposed rocky areas. In general, their high frequency and low uniformity indices confirm that they are not useful as characterspecies. Dahl's uniformity index seems to be useful for quantifying a plant's value as a character-species.Abbreviations DCA Detrended correspondence analysis - MST Minimum spanning tree     

Serologic survey for canine coronavirus in wolves from Alaska.

Wolves (Canis lupus) were captured in three areas of Interior Alaska (USA). Four hundred twenty-five sera were tested for evidence of exposure to canine coronavirus by means of an indirect fluorescent antibody procedure. Serum antibody prevalence averaged 70% (167/240) during the spring collection period and 25% (46/185) during the autumn collection period. Prevalence was 0% (0/42) in the autumn pup cohort (age 4-5 mo), and 60% (58/97) in the spring pup cohort (age 9-10 mo). Prevalence was lowest in the Eastern Interior study area. A statistical model indicates that prevalence increased slightly each year in all three study areas. These results indicate that transmission occurs primarily during the winter months, antibody decay is quite rapid, and reexposure during the summer is rare.     

Prevalence of antibodies to Toxoplasma gondii in wolverines from Nunavut, Canada

The prevalence of antibodies to Toxoplasma gondii was determined in blood and tissue exudates recovered from the spleens of 41 wolverines (Gulo gulo) collected in Nunavut, Canada, using a modified agglutination test (MAT). Antibodies to T. gondii were found in 17 (41.5%) of the 41 wolverines with MAT titers of 1:25 in 1, 1:50 in 4, 1:100 in 5, 1:200 in 6, and 1:400 in 1. This is the first report of antibodies to T. gondii in wolverines, and the results indicate that exposure is common.     

Environmental effects on CO2 efflux from riparian tundra in the northern foothills of the Brooks Range,Alaska, USA

Summary Carbon dioxide efflux and soil microenvironmental factors were measured diurnally in Carex aquatilus-and Eriophorum angustifolium-dominated riparian tundra communities to determine the relative importance of soil environmental factors controlling ecosystem carbon dioxide exchange with the atmosphere. Measurements were made weekly between 18 June and 24 July 1990. Diurnal patterns in carbon dioxide efflux were best explained by changes in soil temperature, while seasonal changes in efflux were correlated with changes in depth to water table, depth to frozen soil and soil moisture. Carbon dioxide efflux rates were lowest early in the growing season when high water tables and low soil temperatures limited microbial and root activity. Individual rainfall events that raised the water table were found to strongly reduce carbon dioxide efflux. As the growing season progressed, rainfall was low and depth to water table and soil temperatures increased. In response, carbon dioxide efflux increased strongly, attaining rates late in the season of approximately 10 g CO₂ m^–2 day^–1. These rates are as high as maxima recorded for other arctic sites. A mathematical model is developed which demonstrates that soil temperature and depth to water table may be used as efficient predictors of ecosystem CO₂ efflux in this habitat. In parallel with the field measurements of CO₂ efflux, microbial respiration was studied in the laboratory as a function of temperature and water content. Estimates of microbial respiration per square meter under field conditions were made by adjusting for potential respiring soil volume as water table changed and using measured soil temperatures. The results indicate that the effect of these factors on microbial respiration may explain a large part of the diurnal and seasonal variation observed in CO₂ efflux. As in coastal tundra sites, environmental changes that alter water table depth in riparian tundra communities will have large effects on ecosystem CO₂ efflux and carbon balance.     

Serologic survey for antibodies to canine distemper virus in collared peccary (Tayassu tajacu) populations in Arizona

In 1989, a disease outbreak was observed among collared peccaries (javelina, Tayassu tajacu) in southern Arizona (USA) and canine distemper virus (CDV) was isolated from affected animals. Subsequently, 364 sera were collected from hunter-harvested javelina over a 4 yr period (1993-96) and were tested for antibody to CDV. Neutralizing antibody to CDV was detected in 58% of the serum samples suggesting that CDV infection is probably enzootic in the collared peccary populations of southern Arizona.     

Genetic diversity and population structure of wolverine (Gulo gulo) populations at the southern edge of their current distribution in North America with implications for genetic viability

The current range of wolverines (Gulo gulo) within the lower 48 states includes small, remnant populations in Idaho, Washington, Wyoming and Montana. The size and trend of each of these populations and connectivity to adjacent populations in the contiguous United States and Canada are poorly understood. In this study, levels of genetic diversity and population genetic structure were examined in three states (Idaho, Wyoming, and Montana) and two Canadian provinces (Alberta and British Columbia) using both mitochondrial (mtDNA) and nuclear microsatellite DNA. Restricted levels of gene flow were detected among these populations with mitochondrial and nuclear DNA and our observations suggest a pattern of male-mediated gene flow. Populations in the United States appear to be receiving migrants from Canada, however, substantial genetic differentiation suggests that gene flow may not be high enough to prevent genetic drift. Our analyses suggest that at least 400 breeding pairs or 1–2 effective migrants per generation would be needed to ensure genetic viability in the long-term for each of the populations in the United States. Significant matrilineal structuring and restricted female gene flow indicates that demographic viability will depend upon the movement of female wolverines into new territories. Results from this study provide guidelines for conservation and management and indicate the need for more ecological data.     

Infectious disease and the decline of Steller sea lions (Eumetopias jubatus) in Alaska, USA: insights from serologic data

Serologic data were examined to determine whether infectious disease may have played a role in the decline of Steller sea lions (Eumetopias jubatus) in the Gulf of Alaska and Aleutian Islands, USA. Available published data, unpublished data, and recent collections (1997-2000) were compared and reviewed. Data were stratified by geography to compare the declining western Alaskan population in the Aleutian Islands through eastern Prince William Sound to the increasing population in southeastern Alaska. Prevalences of antibodies from the 1970s to the early 1990s were noted for Leptospira interrogans, Chlamydophila psittaci, Brucella spp., phocid herpesvirus-1, and calciviruses. Serum samples collected from 1997-2000 were tested for antibodies to these agents as well as to marine mammal morbilliviruses, canine parvovirus, and canine adenovirus-1 and -2. Conclusions could not be drawn about changes in antibody prevalence to these agents during the decline of Steller sea lions, however, because data were incomplete or not comparable as a result of inconsistencies in testing techniques. Despite these shortcomings, results provided no convincing evidence of significant exposure of Steller sea lions to morbilliviruses, Brucella spp., canine parvovirus, or L. interrogans. Steller sea lions have been exposed to phocid herpesviruses, caliciviruses, canine adenovirus, and C. psittaci or to cross-reactive organisms in regions of both increasing and decreasing sea lion abundance. Based on similar antibody prevalence estimates from the increasing and decreasing populations, these agents are unlikely to have been the primary cause of the population decline. They may have contributed to the decline or impeded population recovery, however, because of undetected mortality and morbidity or reductions of fecundity and body condition in animals under other stresses. Systematic monitoring for disease agents and their effects is needed to determine whether infectious disease currently plays a role in the decline and lack of recovery of Steller sea lions.     

A serologic survey of the island fox (Urocyon littoralis) on the Channel Islands, California.

The island fox is listed as a threatened species in California. A serologic survey of 194 island foxes (Urocyon littoralis) was conducted over the entire range of the species on the Channel Islands (California, USA). Antibody prevalence against canine adenovirus and canine parvovirus reached 97% and 59%, respectively, in some populations sampled. Antibody prevalence of canine herpesvirus, canine coronavirus, leptospirosis and toxoplasmosis were low. Antibodies against canine distemper virus were not detected.     

Serologic survey for Actinobacillus capsulatus in free-ranging snowshoe hares (Lepus americanus) from Alaska and Alberta

A plate agglutination method was developed to test sera from free-ranging snowshoe hares (Lepus americanus) captured in Alaska (USA) or Alberta (Canada) for antibody against Actinobacillus capsulatus. Antiserum against A. capsulatus was prepared in a domestic rabbit. A concentrated suspension of formalin-killed A. capsulatus was prepared for use as an antigen. Serum antibody prevalence for hares was 98 of 239 (41%) in Alaska and 51 of 111 (46%) in Alberta. Prevalence in Alaska peaked in 1981 corresponding to a peak in hare population density. Seasonal prevalence peaked in May in Alaska. Prevalence at one capture site in Alaska was significantly higher than at four other sites. There was no difference in sex-specific prevalence for either Alaska or Alberta.     

Serologic survey for selected disease agents in wolves (Canis lupus) from Alaska and the Yukon Territory, 1984-2000

Wolves (Canis lupus) were captured in several geographic areas of Alaska (USA) and the Yukon Territory (Canada) during 1984-2000. Blood was collected from 1,122 animals. Sera were tested for antibodies against infectious canine hepatitis virus (ICH), canine distemper virus (CDV), canine parvovirus (CPV), Francisella tularensis, and serovars of Leptospira interrogans. Antibody prevalence for ICH was >84% for all areas. Area-specific prevalences of antibodies ranged from 12% to 70% for CPV, from 0% to 41% for CDV, and from 4% to 21% for F. tularensis. There was no evidence of CDV exposure at the two southernmost locations in Alaska. Prevalence of antibodies for ICH increased slightly during the 16-yr course of the survey. There was essentially no evidence of exposure to L. interrogans. Prevalences of antibodies for both CPV and CDV were age-specific, with higher values in the adult cohort compared with the pup cohort. There were no sex-specific differences in prevalence of antibodies for any of the five disease agents.