Otolith-inferred patterns of marine migration frequency in Nunavik Arctic charr.

Alternative migratory tactics in salmonids reflect the large observed interindividual variation in spatial behaviour which may range from strict freshwater residency to uninterrupted anadromy. In Salvelinus, sea migrations are performed during the ice-free period as freshwater overwintering is thought to be obligatory due to physiological constraints. As a result, individuals can either migrate the next spring or remain in freshwater, as anadromy is generally considered facultative. In Arctic charr (Salvelinus alpinus), skipped migrations are known to occur, but limited data are available regarding their frequencies within and among populations. Here, the authors used an otolith microchemistry approach relying on strontium (88 Sr) to infer movements between freshwater and marine habitats, and annual oscillations in zinc (64 Zn) to help with age identification. They determined the age-at-first-migration and the occurrence of subsequent annual migrations in two Nunavik Arctic charr populations sampled in Deception Bay (Salluit) and river systems linked to Hopes Advance Bay (Aupaluk), northern Québec, Canada. The mode for age-at-first-migration was 4+ for both populations, although it exhibited large variation (range: 0+ to 8+). Skipped migrations constituted a rare event, as 97.7% and 95.6% of the examined Arctic charr at Salluit (n = 43, mean age = 10.3 ± 2.0 years) and Aupaluk (n = 45, mean age = 6.0 ± 1.9 years), respectively, were found to have performed uninterrupted annual migrations after initiation of the behaviour. The consistency of the annual migrations suggests that the tactic is sufficiently fitness rewarding to be maintained under current environmental conditions. From a fisheries management perspective, these repeated migrations combined with low site fidelity in this species may lead to large interannual variations in abundance at the local scale, which may represent a challenge for monitoring Arctic charr demographics on a river-by-river basis.

A spatial theory for characterizing predator-multiprey interactions in heterogeneous landscapes.

Trophic interactions in multiprey systems can be largely determined by prey distributions. Yet, classic predator-prey models assume spatially homogeneous interactions between predators and prey. We developed a spatially informed theory that predicts how habitat heterogeneity alters the landscape-scale distribution of mortality risk of prey from predation, and hence the nature of predator interactions in multiprey systems. The theoretical model is a spatially explicit, multiprey functional response in which species-specific advection-diffusion models account for the response of individual prey to habitat edges. The model demonstrates that distinct responses of alternative prey species can alter the consequences of conspecific aggregation, from increasing safety to increasing predation risk. Observations of threatened boreal caribou, moose and grey wolf interacting over 378 181 km(2) of human-managed boreal forest support this principle. This empirically supported theory demonstrates how distinct responses of apparent competitors to landscape heterogeneity, including to human disturbances, can reverse density dependence in fitness correlates.

Evidence for evolutionary convergence at MHC in two broadly distributed mesocarnivores.

Variation within major histocompatibility complex (MHC) genes is important in recognizing pathogens and initiating an immune response. These genes are relevant in enhancing our understanding of how species cope with rapid environmental changes and concomitant fluctuations in selective pressures such as invasive, infectious diseases. Disease-based models suggest that diversity at MHC is maintained through balancing selection arising from the coevolution of hosts and pathogens. Despite intensive balancing selection, sequence motifs or even identical MHC alleles can be shared across multiple species; three potential mechanisms have been put forth to explain this phenomenon: common ancestry, convergent evolution, and random chance. To understand the processes that maintain MHC similarity across divergent species, we examined the variation at two orthologous MHC-DRB genes in widespread North American Musteloid species, striped skunks (Mephitis mephitis), and raccoons (Procyon lotor). These species are often sympatric and exposed to a similar suite of diseases (e.g., rabies, canine distemper, and parvovirus). Given their exposure to similar selective pressures from pathogens, we postulated that similar DRB alleles may be present in both species. Our results indicated that similar motifs are present within both species, at functionally relevant polymorphic sites. However, based on phylogenetic analyses that included previously published MHC sequences of several closely related carnivores, the respective MHC-DRB alleles do not appear to have been maintained through common ancestry and unlikely through random chance. Instead, the similarities observed between the two mesocarnivore species may rather be due to evolutionary convergence.

Temporal dynamics of genetic variability in a mountain goat (Oreamnos americanus) population.

The association between population dynamics and genetic variability is of fundamental importance for both evolutionary and conservation biology. We combined long-term population monitoring and molecular genetic data from 123 offspring and their parents at 28 microsatellite loci to investigate changes in genetic diversity over 14 cohorts in a small and relatively isolated population of mountain goats (Oreamnos americanus) during a period of demographic increase. Offspring heterozygosity decreased while parental genetic similarity and inbreeding coefficients (F(IS) ) increased over the study period (1995-2008). Immigrants introduced three novel alleles into the population and matings between residents and immigrants produced more heterozygous offspring than local crosses, suggesting that immigration can increase population genetic variability. The population experienced genetic drift over the study period, reflected by a reduced allelic richness over time and an 'isolation-by-time' pattern of genetic structure. The temporal decline of individual genetic diversity despite increasing population size probably resulted from a combination of genetic drift due to small effective population size, inbreeding and insufficient counterbalancing by immigration. This study highlights the importance of long-term genetic monitoring to understand how demographic processes influence temporal changes of genetic diversity in long-lived organisms.

Genomic data support management of anadromous Arctic Char fisheries in Nunavik by highlighting neutral and putatively adaptive genetic variation.

Distinguishing neutral and adaptive genetic variation is one of the main challenges in investigating processes shaping population structure in the wild, and landscape genomics can help identify signatures of adaptation to contrasting environments. Arctic Char (Salvelinus alpinus) is an anadromous salmonid and the most harvested fish species by Inuit people, including in Nunavik (Québec, Canada), one of the most recently deglaciated regions in the world. Unlike many other anadromous salmonids, Arctic Char occupy coastal habitats near their natal rivers during their short marine phase restricted to the summer ice-free period. Our main objective was to document putatively neutral and adaptive genomic variation in anadromous Arctic Char populations from Nunavik and bordering regions to inform local fisheries management. We used genotyping by sequencing (GBS) to genotype 18,112 filtered single nucleotide polymorphisms (SNP) in 650 individuals from 23 sampling locations along >2000 km of coastline. Our results reveal a hierarchical genetic structure, whereby neighboring hydrographic systems harbor distinct populations grouped by major oceanographic basins: Hudson Bay, Hudson Strait, Ungava Bay, and Labrador Sea. We found genetic diversity and differentiation to be consistent both with the expected postglacial recolonization history and with patterns of isolation-by-distance reflecting contemporary gene flow. Results from three gene-environment association methods supported the hypothesis of local adaptation to both freshwater and marine environments (strongest associations with sea surface and air temperatures during summer and salinity). Our results support a fisheries management strategy at a regional scale, and other implications for hatchery projects and adaptation to climate change are discussed.

Father-offspring phenotypic correlations suggest intralocus sexual conflict for a fitness-linked trait in a wild sexually dimorphic mammal.

In sexually dimorphic and polygynous mammals, sexual selection often favours large males with well-developed weaponry, as these secondary sexual characters confer advantages in intrasexual competition and are often preferred by females. Little is known, however, about the effects of sexually selected paternal traits on offspring phenotype in wild mammals, especially when considering that shared phenotypic traits and selection can also differ greatly between genders. Here, we conducted molecular parentage analyses in a long-term study population of mountain goats (Oreamnos americanus), an ungulate exhibiting high sexual dimorphism in mass, to first assess the determinants of yearly reproductive success (YRS) in males. We then examined the effects of paternal characteristics on offspring mass at 1 year of age. Paternity was highly skewed, with 9 per cent of 57 males siring 51 per cent of 96 offspring assigned over 12 years. Male YRS increased with age until apparent reproductive senescence at 9 years, but mass was a stronger determinant of siring success than age, horn length or social rank. Mass of sons increased with paternal mass, but the mass of daughters was negatively related to that of their father, a finding consistent with recent theory on intralocus sexual conflict. Because early differences in mass persisted to early adulthood, sex-specific effects of paternal mass can have important fitness consequences, as adult mass is positively linked with reproduction in both sexes. Divergent father-offspring phenotypic correlations may partly explain the maintenance of sexual dimorphism in mountain goats and the large variance observed for this homologous trait within each gender in polygynous mammals.

Multilocus heterozygosity, parental relatedness and individual fitness components in a wild mountain goat, Oreamnos americanus population.

Matings between relatives lead to a decrease in offspring genetic diversity which can reduce fitness, a phenomenon known as inbreeding depression. Because alpine ungulates generally live in small structured populations and often exhibit a polygynous mating system, they are susceptible to inbreeding. Here, we used marker-based measures of pairwise genetic relatedness and inbreeding to investigate the fitness consequences of matings between relatives in a long-term study population of mountain goats (Oreamnos americanus) at Caw Ridge, Alberta, Canada. We first assessed whether individuals avoided mating with kin by comparing actual and random mating pairs according to their estimated genetic relatedness, which was derived from 25 unlinked polymorphic microsatellite markers and reflected pedigree relatedness. We then examined whether individual multilocus heterozygosity H, used as a measure of inbreeding, was predicted by parental relatedness and associated with yearling survival and the annual probability of giving birth to a kid in adult females. Breeding pairs identified by genetic parentage analyses of offspring that survived to 1 year of age were less genetically related than expected under random matings. Parental relatedness was negatively correlated with offspring H, and more heterozygous yearlings had higher survival to 2 years of age. The probability of giving birth was not affected by H in adult females. Because kids that survived to yearling age were mainly produced by less genetically related parents, our results suggest that some individuals experienced inbreeding depression in early life. Future research will be required to quantify the levels of gene flow between different herds, and evaluate their effects on population genetic diversity and dynamics.

Modelling the dispersal of the two main hosts of the raccoon rabies variant in heterogeneous environments with landscape genetics.

Predicting the geographic spread of wildlife epidemics requires knowledge about the movement patterns of disease hosts or vectors. The field of landscape genetics provides valuable approaches to study dispersal indirectly, which in turn may be used to understand patterns of disease spread. Here, we applied landscape genetic analyses and spatially explicit models to identify the potential path of raccoon rabies spread in a mesocarnivore community. We used relatedness estimates derived from microsatellite genotypes of raccoons and striped skunks to investigate their dispersal patterns in a heterogeneous landscape composed predominantly of agricultural, forested and residential areas. Samples were collected in an area covering 22 000 km(2) in southern Québec, where the raccoon rabies variant (RRV) was first detected in 2006. Multiple regressions on distance matrices revealed that genetic distance among male raccoons was strictly a function of geographic distance, while dispersal in female raccoons was significantly reduced by the presence of agricultural fields. In skunks, our results suggested that dispersal is increased in edge habitats between fields and forest fragments in both males and females. Resistance modelling allowed us to identify likely dispersal corridors used by these two rabies hosts, which may prove especially helpful for surveillance and control (e.g. oral vaccination) activities.

Oral rabies vaccination in raccoons: comparison of ONRAB® and RABORAL V-RG® vaccine-bait field performance in Québec, Canada and Vermont, USA.

The control of rabies in raccoons (Procyon lotor) and striped skunks (Mephitis mephitis) in North America has been conducted mainly through aerial distribution of oral vaccine-baits. The effectiveness of the vaccine-bait used is therefore of prime importance for disease eradication. In a previous field comparison between the ONRAB(®) bait in the province of New Brunswick, Canada, and RABORAL V-RG(®) bait in the state of Maine, USA, the ONRAB bait produced a higher percentage of antibody-positive raccoons under nearly identical bait distribution for the two vaccines. The main objective of the present study was to conduct a similar cross-border comparison of these two vaccine-baits using raccoon sera collected during post-oral rabies vaccination monitoring in Québec, Canada, and Vermont, USA, where ONRAB and V-RG, respectively, were distributed aerially at a targeted density of 150 baits/km(2). A comparison of the equivalency of two serologic tests used in Canada and the USA was also conducted using sera from raccoons and striped skunks. Rabies virus neutralization assay (USA) yielded similar results to the competitive enzyme-linked immunosorbent assay (Canada), with agreement between the two tests of 92% for raccoon sera and 96% for skunk sera. With both assays, the percentage of antibody-positive raccoons was greater with ONRAB (51%, n=265) than with V-RG (38%, n=66). These new results support the conclusion from the previous study, that ONRAB vaccine-baits may be more effective for the control of rabies in raccoons.

Genetic structure and rabies spread potential in raccoons: the role of landscape barriers and sex-biased dispersal.

Identifying natural barriers to movements of hosts associated with infectious diseases is essential for developing effective control strategies. Raccoon rabies variant (RRV) is a zoonosis of concern for humans because its main vector, the raccoon (Procyon lotor), is found near residential areas. In Québec, Canada, all cases of RRV found in raccoons since 2006 were detected on the eastern side of the Richelieu River, suggesting that this river acts as a barrier to gene flow and thus the potential for RRV to spread. The objectives of this study were to characterize the genetic structure of raccoon populations and assess the effect of the Richelieu River on the population structure in southern Québec, Canada. We also evaluated whether RRV spread potential differed between sex and at a larger spatial scale. Our analyses revealed a weak signal of genetic differentiation among individuals located on each side of the Richelieu River. At a larger spatial scale, genetic structuring was weak. Our results suggest that rivers might not always efficiently restrain raccoon movements and spread of RRV. We suggest that the difference in genetic structure found between sexes can be partly explained by male movements during the breeding season in winter, when ice bridges allow passage over most rivers in Québec.

Lack of genetic structure and female-specific effect of dispersal barriers in a rabies vector, the striped skunk (Mephitis mephitis).

Evaluating the permeability of potential barriers to movement, dispersal and gene exchanges can help describe spreading patterns of wildlife diseases. Here, we used landscape genetics methods to assess the genetic structure of the striped skunk (Mephitis mephitis), which is a frequent vector of rabies, a lethal zoonosis of great concern for public health. Our main objective was to identify landscape elements shaping the genetic structure of this species in Southern Québec, Canada, in an area where the raccoon rabies variant has been detected. We hypothesised that geographic distance and landscape barriers, such as highways and major rivers, would modulate genetic structure. We genotyped a total of 289 individuals sampled across a large area (22,000 km²) at nice microsatellite loci. Genetic structure analyses identified a single genetic cluster in the study area. Major rivers and highways, however, influenced the genetic relatedness among sampled individuals. Sex-specific analyses revealed that rivers significantly limited dispersal only for females while highways only had marginal effects. Rivers and highways did not significantly affect male dispersal. These results support the contention that female skunks are more philopatric than males. Overall, our results suggest that the effects of major rivers and highways on dispersal are sex-specific and rather weak and are thus unlikely to prevent the spread of rabies within and among striped skunk populations.

Oral rabies vaccination of raccoons and striped skunks with ONRAB® baits: multiple factors influence field immunogenicity.

Multiple control methods have been used in North America to manage the spread of rabies caused by the raccoon (Procyon lotor) rabies virus variant (RRVV). Recently, oral vaccination with ONRAB(®) vaccine baits, which contain an adenovirus rabies glycoprotein recombinant, has been made available as an additional tool for rabies control. Our objectives were to estimate rabies antibody prevalence in wild-caught raccoons and striped skunks (Mephitis mephitis), and identify factors influencing the probability of being antibody positive at the individual level in these species, following oral rabies vaccination (ORV) campaigns in which ONRAB was distributed aerially in 2007-2009 in southern Québec, Canada. Following the aerial distribution of 43-155 ONRAB baits/km(2), the annual percentages of antibody-positive raccoons and skunks varied between 35% and 56% and 11% and 17%, respectively. In raccoons, the probability of being antibody positive was positively associated with age and density of ONRAB distributed, and influenced by the number of previous ORV campaigns conducted. Conversely, this probability was negatively associated with estimated abundance of raccoons in the trapping cell and proportion of residential areas near the raccoon capture location. None of the variables examined explained variation in the probability of being antibody positive in skunks. Our results indicate that the ONRAB density applied during ORV campaigns should be adjusted to account for variations in raccoon population density and presence of residential areas to increase the likelihood of creating an effective immunological barrier against RRVV. The high percentage of juvenile raccoons (annual mean =45 ± 3 [SE]%) and skunks (66 ± 2%) captured during post-ORV monitoring suggests that ORV campaigns should be conducted at least annually to account for the recruitment of naïve individuals into the populations. In Québec, the increasing use of ONRAB coincided with the elimination of rabies caused by RRVV. Nonetheless, our results indicate that improvements to this vaccine bait and/or the distribution techniques are required to increase its efficacy, especially in striped skunks.

Oral vaccination against raccoon rabies: landscape heterogeneity and timing of distribution influence wildlife contact rates with the ONRAB vaccine bait.

Aerial distribution of oral vaccine baits is one of the available strategies for controlling the spread of infectious wildlife diseases. This technique has commonly been used to control rabies in wild carnivores and, together with other techniques, was used to immunize wild populations of raccoons (Procyon lotor) and striped skunks (Mephitis mephitis) after the detection of the first rabid raccoon in the province of Quebec, Canada, in 2006. Vaccine bait distribution was conducted over large areas where agricultural land is dominant but interspersed with residual forest patches. Our objective was to evaluate the effect of habitat (forest vs. agricultural crops) in space and time on the contact rate between wildlife and the ONRAB(®) vaccine bait, a recent alternative to the V-RG(®). Four transects of eight vaccine baits each were installed parallel to, and at different distances from, the forest's edge (under forest cover, at field-forest edge, and at 50 and 200 m from forest edge in agricultural crops) at three sites composed of various crop types interspersed with forest patches. This experiment was conducted during three periods (late spring, 1-7 June; summer, 27 July-2 August; and fall, 24-30 October) in 2009. Contact rates with vaccine baits were monitored for 7 days in each period to evaluate the potential temporal variations generated within the habitat types. Contact rates with ONRAB vaccine baits were highest under forest cover and in the fall. Of 13 species observed in proximity to the vaccine baits, raccoons were the most frequent (49.5%, n=55 visits). Our study underlines the importance of taking into account landscape heterogeneity and timing of distribution when planning the distribution of vaccine baits to control rabies in raccoons.