Capture and medetomidine-ketamine anesthesia of free-ranging wolverines (Gulo gulo)

Capture and anesthesia with medetomidine-ketamine were evaluated in free-ranging wolverines (Gulo gulo) immobilized for marking with radiocollars or intraperitoneal radiotransmitters in Norrbotten, Sweden, during early June 2004 and 2005. Twelve juvenile wolverines were captured by hand and injected with 0.14 +/- 0.03 mg/kg (mean +/- SD) medetomidine and 7.5 +/- 2.0 mg/kg ketamine. Twelve adult wolverines were darted from a helicopter or the ground, or captured by hand. Adults received 0.37 +/- 0.06 mg/kg medetomidine and 9.4 +/- 1.4 mg/kg ketamine. Arterial blood samples were collected between 15 min and 30 min and between 45 min and 60 min after drug administration and immediately analyzed for selected hematologic and plasma variables. Hyperthermia was recorded initially in one juvenile wolverine and 11 adults. Rectal temperature, heart rate, and lactate decreased significantly during anesthesia, whereas hemoglobin oxygen saturation, pH, partial pressure of arterial carbon dioxide, and base excess increased. Adult wolverines darted from a helicopter had a significantly higher rectal temperature, higher glucose and hematocrit values, and a lower heart rate than juveniles captured by hand. Impaired arterial oxygenation was evident in all wolverines. This study provides baseline data on physiologic variables in adult and juvenile wolverines captured with different methods and anesthetized with medetomidine-ketamine.     

Craniometric variation in Norwegian wolverines Gulo gulo L.

Craniometric variation in Norwegian wolverines was analysed using univariate and multivariate techniques. The wolverine skull seems to be full-grown after about 9 months in both sexes although some dimensions continued to grow at a slow rate after the first year. Size allometry was studied in skulls older than 11 months. Allometric coefficients were calculated from different methods. The multivariate allometric coefficients were found to reflect the same allometric patterns as were found from the reduced major axis and from the direction cosines of PCI, whereas the coefficients seem to be underestimated by the least-square regression method. Sexual dimorphism was found to be highly significant, males being the larger sex. The female skulls were found to be relatively broader and to have a relatively higher coronoid process than the male skulls. It seems that the females have a relatively stronger feeding apparatus than the males, which leads to less absolute differences in strength, as reflected in a very high degree of dietary overlap. Two discriminant functions were calculated which both classified the material into their correct sex with nearly 100% accuracy. Geographical variation between samples from Nordland and Finnmark counties were studied. No significant multivariate variation was found between the female skulls. The male variation was highly significant; the Nordland skulls seem to have longer carnassials than those from Finnmark.     

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.     

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.     

Home range and habitat use of wolverines Gulo gulo in Yukon, Canada

Home ranges and habitat use are described for three adult female, one adult male, and one subadult male wolverines in the Kluane Game Sanctuary, Yukon. When long distance excursions are not included, home ranges of wolverines in the Kluane Game Sanctuary were between 76 and 269 km² for females and 209 and 269 km² for males. Habitat use of females was similar to habitat availability. Males used subalpine coniferous habitats more frequently than other habitat types during winter. Although individual variation in the use of forest cover types, aspects, slopes and elevations was apparent, seasonal use did not differ from availability for each sex. Within the 1590 km² study area, three adult males and six adult females were present, corresponding to a density of one resident wolverine 177 km^-2.     

Lack of Spatial Immunogenetic Structure among Wolverine (Gulo gulo) Populations Suggestive of Broad Scale Balancing Selection

Elucidating the adaptive genetic potential of wildlife populations to environmental selective pressures is fundamental for species conservation. Genes of the major histocompatibility complex (MHC) are highly polymorphic, and play a key role in the adaptive immune response against pathogens. MHC polymorphism has been linked to balancing selection or heterogeneous selection promoting local adaptation. However, spatial patterns of MHC polymorphism are also influenced by gene flow and drift. Wolverines are highly vagile, inhabiting varied ecoregions that include boreal forest, taiga, tundra, and high alpine ecosystems. Here, we investigated the immunogenetic variation of wolverines in Canada as a surrogate for identifying local adaptation by contrasting the genetic structure at MHC relative to the structure at 11 neutral microsatellites to account for gene flow and drift. Evidence of historical positive selection was detected at MHC using maximum likelihood codon-based methods. Bayesian and multivariate cluster analyses revealed weaker population genetic differentiation at MHC relative to the increasing microsatellite genetic structure towards the eastern wolverine distribution. Mantel correlations of MHC against geographical distances showed no pattern of isolation by distance (IBD: r = -0.03, p = 0.9), whereas for microsatellites we found a relatively strong and significant IBD (r = 0.54, p = 0.01). Moreover, we found a significant correlation between microsatellite allelic richness and the mean number of MHC alleles, but we did not observe low MHC diversity in small populations. Overall these results suggest that MHC polymorphism has been influenced primarily by balancing selection and to a lesser extent by neutral processes such as genetic drift, with no clear evidence for local adaptation. This study contributes to our understanding of how vulnerable populations of wolverines may respond to selective pressures across their 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.     

Diseases and mortality in free-ranging brown bear (Ursus arctos), gray wolf (Canis lupus), and wolverine (Gulo gulo) in Sweden

Ninety-eight brown bears (Ursus arctos), 20 gray wolves (Canis lupus), and 27 wolverines (Gulo gulo), all free-ranging, were submitted to the National Veterinary Institute, Uppsala, Sweden, during 1987-2001 for investigation of diseases and causes of mortality. The most common cause of natural death in brown bears was infanticide. Infanticide also was observed in wolverines but not in wolves. Traumatic injuries, originating from road or railway accidents, were the most common cause of death in wolves and occurred occasionally in brown bears. Most wolverines were submitted as forensic cases in which illegal hunting/poaching was suspected. Sarcoptic mange was observed in several wolves but not in brown bears or wolverines. Sarcoptic mange most likely was acquired from infected red foxes (Vulpes vulpes) that were killed by wolves. Other parasites and infectious diseases were only found sporadically.     

Space use and territoriality of wolverines (<Emphasis Type="Italic">Gulo gulo</Emphasis>) in northern Scandinavia

Space use and territoriality influence population structure and dynamics and is therefore an important aspect in understanding the ecology of animals. We investigated spatial and temporal space use of wolverines (Gulo gulo) in northern Scandinavia. We estimated home ranges of 24 radio-marked individuals (17 females and seven males). Male home ranges (mean 669 km²; SE = 211) were significantly larger than female home ranges (mean 170 km²; Wilcoxon–Mann–Whitney; P = 0.001) and encompassed or included parts of up to five different females. Home range sizes of reproducing (170 km²; SE = 51) and barren (171 km²; SE = 63) adult females did not differ. Wolverines in Scandinavia exhibit intrasexual territoriality, with male home ranges totally exclusive and female home ranges either exclusive or with little home range overlap. Overlap between wolverine territories is most likely explained by intrasexual tolerance and kinship.     

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.     

Blue mussels (Mytilus edulis) in the diet of roach (Rutilus rutilus) in outer archipelago areas of the western Gulf of Finland,Baltic Sea

The blue mussel (Mytilus edulis) is one of the key species in the Baltic Sea ecosystem and it is living at the edge of its range in the western Gulf of Finland. Roach (Rutilus rutilus) is a freshwater fish species that has benefited from recent coastal eutrophication and is at present highly abundant in the outer archipelago of the Gulf of Finland. In 2000 and 2001, a total of 516 roach were sampled for diet analysis in three study areas. Shelled molluscs formed over 95% of the diet of roach, blue mussels being the dominant single species. The proportion of this species in the food of roach in the three study areas ranged between 38–61% for smaller roach (<225 mm) and 39–85% for larger (>225 mm) roach, indicating that blue mussel is a highly important food source for roach in outer archipelago areas of the western Gulf of Finland, in contrast to reports from other parts of the northern Baltic Sea. The scarcity of large blue mussels in mussel beds in the easternmost study area was reflected in the lower proportion of blue mussels in the diet of larger roach. However, the growth of roach was not affected by the availability of blue mussels. The twofold differences observed in the annual growth of roach between warm and cold years demonstrated that temperature is an important factor controlling the growth of roach in the western Gulf of Finland.     

Ticks (Acari: Ixodidae) parasitizing migrating and local breeding birds in Finland

Ticks are globally renowned vectors for numerous zoonoses, and birds have been identified as important hosts for several species of hard ticks (Acari: Ixodidae) and tick-borne pathogens. Many European bird species overwinter in Africa and Western Asia, consequently migrating back to breeding grounds in Europe in the spring. During these spring migrations, birds may transport exotic tick species (and associated pathogens) to areas outside their typical distribution ranges. In Finland, very few studies have been conducted regarding ticks parasitizing migrating or local birds, and existing data are outdated, likely not reflecting the current situation. Consequently, in 2018, we asked volunteer bird ringers to collect ticks from migrating and local birds, to update current knowledge on ticks found parasitizing birds in Finland. In total 430 ticks were collected from 193 birds belonging to 32 species, caught for ringing between 2018 and 2020. Furthermore, four Ixodes uriae were collected from two roosting islets of sea birds in 2016 and 2020. Ticks collected on birds consisted of: Ixodes ricinus (n?=?421), Ixodes arboricola (4), Ixodes lividus (2) and Hyalomma marginatum (3). Ixodes ricinus loads (nymphs and larvae) were highest on thrushes (Passeriformes: Turdidae) and European robins (Erithacus rubecula). The only clearly imported exotic tick species was H. marginatum. This study forms the second report of both I. uriae and I. arboricola from Finland, and possibly the northernmost observation of I. arboricola from Europe. The importation of exotic tick species by migrating birds seems a rare occurrence, as over 97% of all ticks collected from birds arriving in Finland during their spring migrations were I. ricinus, a species native to and abundant in Finland.

     

The breeding biology of the starling Sturnus vulgaris in northern Finland

The breeding biology of the starling Sturnus vulgaris was studied in three populations in northern Finland, with additional reference localities in nearby areas. Results from a total of 329 nests are presented from years 1962-1976.
The onset of laying in the Oulu area took place on 5 May, the median date being 10 May, about one week later than in southern Finland and 2–3 weeks later than in central Europe. The clutch-size was 5.29 ± 0.05 in the first nests and 5.06 ± 0.26 in later ones. Incubation time was 12.1 ± 0.6 d. An average of 3.31 ± 0.10 fledglings was produced per pair, 3.69 ± 0.08 in succesful clutches, which is significantly less than usually reported from central Europe. Most of the losses were due to starvation of the young.
No second broods were observed in the Oulu area, and repeated nesting was not common, probably because of the general late start of laying.
The reasons for the recent decline in several Finnish starling populations do not lie in the production capacity but must be sought for elsewhere.     

Host-parasite relationships between two seal populations and two species of Corynosoma (Acanthocephala) in Finland

Ringed seals ( Phoca hispida botnica ), from the Bothnian Bay, north-eastern Baltic Sea were examined for acanthocephalan parasites in 1977–1982, and specimens from Lake Saimaa in south-eastern Finland ( P. h. saimensis ) were examined in 1980–1981. The two seal stocks have been isolated from Arctic stock for 11,000 years and from each other for 8,000 years. Corynosoma strumosum and C. semerme were found in the Baltic seals, but only C. strumosum in those from the lake.
Eighty two of the ringed seals caught in the Bothnian Bay were infected with both Corynosoma species, one with a single infection of C. strumosum and one with C. semerme in autumn. The total number of C. semerme increased in relation to the number of C. strumosum in each set of seals both in autumn and spring. Corynosoma semerme was dominant in most seals in spring, but in only an average of 60% in autumn. The predominance of C. strumosum in 40% of the autumn material (range 33–80%) is of interest because species ratio favouring C. strumosum is not found in any more heavily infected paratenic fish host of Corynosoma in the Bothnian Bay. The change in the Corynosoma species ratio of seals in autumn may be explained by their exceptional feeding behaviour at this time of the year (total or partial fasting) and the suggested longevity of C. strumosum as compared with C. semerme . The stability of the acanthocephalan populations in ringed seals is documented.     

Intra-sexual female agonistic behaviour of the South American sea lion (Otaria flavescens) in two colonies with different breeding substrates

Female agonistic behaviour during pregnancy and lactation is common in several pinnipeds and has been associated to pup or territory defence. Previous studies on female agonistic behaviour in pinniped breeding groups have not usually considered the number of females per breeding group, which could influence the degree of aggressiveness. We compared female agonistic behaviour (open-mouth displays and bites) within two colonies with different breeding substrates (homogeneous, Punta Norte; heterogeneous, Pirámide) of South American sea lions using two methods: weighted means and general linear models. We found that both aggressive interaction rates were significantly affected by female density, which accounted for a greater proportion on the variability in bite rates than in open-mouth rates. Controlling for the number of females, we found with both methods that open-mouth rates were higher than bite rates within the Pirámide colony; however, no differences were found within the Punta Norte colony. In Pirámide, open-mouth rates could be used more often as threats to minimise the chances of more severe aggressions. The conclusion is that females vary their use of agonistic interactions in relation to their density in the colony with heterogeneous substrate, which may be related to the presence of limited numbers of tide pools that heightens competition for thermoregulatory resources.     

Mate preferences in female canaries (Serinus canaria) within a breeding season

Divorce and remating in birds can be described as strategies used to enhance reproductive success. Mate switching often occurs because pairs failed to brood at least one chick during the previous breeding season. In the present study, we evaluated the influence of reproductive success on female preferences in domesticated canaries (Serinus canaria). For that purpose, females previously paired and having reared young were placed in a choice test situation: They were allowed to choose between their previous mate and a familiar male (a male neighbor during the breeding period). During these choice tests, females tended to stay near their previous mate longer than near a male neighbor when their reproductive success was “good” (at least two chicks). On the other hand, females with “poor” reproductive success (one chick) did not show a preference for their previous mate. Furthermore, in the present study, we observed that during choice tests males reacted to the presence of their previous mate in a particular way, by gathering nest material. This behavior was more scarcely observed in neighbor males which, on the contrary, sang significantly more than previous mates did.     

The causation of two scent-marking behaviour patterns in female hamsters (Mesocricetus auratus)

Social, olfactory, and oestrous cycle influences on the frequency of flank-marking and vaginal marking were studied in female hamsters. Vaginal marking was more frequent in the presence of males or their odours than females or their odours. Vaginal marking frequency was greatest the day before oestrus (day 4), intermediate on days 2 and 3, and zero on the oestrous day. The probable sexual advertisement functions of vaginal marking were discussed. In contrast, flank-marking by females was stimulated more by other female odours than by male odours. The frequency of flank-marking was elevated by agonistic encounters and was reduced by sexual encounters. Flank-marking seems to have functions related to functions of aggression among hamsters.     

Effects of habitat fragmentation on provisioning rates, diet and breeding success in two species of tit (great tit and blue tit)

The aim of this study was to examine the effects of forest fragmentation on the ability of parent birds to provide their young with an adequate food supply. To examine whether prey population densities of the great tit (Parus major L.) and the blue tit (P. caeruleus L.) vary between study areas in different forest size classes we compared provisioning rates and chick diet and related these parameters to breeding success. We filmed 217 nests over two breeding seasons and collected data on frass fall as a general estimate of caterpillar availability. Nests which were attended by none or one parent only during filming (n = 46) were excluded from the analyses. In both years and for both species feeding rates were highest in the smallest fragments and lowest in the large forest. There was also a suggestion that differences in feeding rates between areas vary between years. We found no consistent tendency for prey size to change with forest size, although both species brought slightly smaller prey items to the nest in the smallest forest fragments and feeding rates correlated negatively with prey size. Caterpillars were the main item fed to nestlings, in both species. We found no evidence to suggest that either frass fall or the proportion of caterpillars in the diet varied with forest size. There was also no correlation between mean frass fall and the total number of caterpillars brought to the nests, in either species. Breeding success, as measured by clutch size, brood size, fledging weight and fledging success, did not differ between the small fragments and the large forest, in either species. There was also no relationship between provisioning rate (as concerns volume of prey fed to nestlings and the quality of chick diet) and breeding success parameters. In conclusion, this study does not suggest suboptimal foraging or breeding conditions in small fragments compared to a nearby large forest, for either species. Received: 23 June 1997 / Accepted: 29 December 1997