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Sexual selection has a critical role in evolution, and it is fundamental to identify what ecological factors drive its variation. Disentangling the ecological correlates of sexual selection over the long term, however, is challenging and has rarely been done in nature. We sought to assess how demographic changes influenced the intensity, direction and form of sexual selection and whether selective pressures varied with age. We tested whether breeder sex ratio, number of competitors and age structure influenced selection differentials on horn length of wild bighorn rams ( Ovis canadensis) of different age classes on Ram Mountain, Alberta. We used 21 years of data including a detailed pedigree, demographic parameters and repeated morphological measurements. Sexual selection on horn length of males of all ages was directional and positive. Selection intensity increased with the number of competitors, reflecting male-male encounter rate during the rut, but was independent of breeder sex ratio or age structure. This result can also be linked to changes in population size because the number of competitors was highly correlated to total number of sheep. This demographic effect likely arises from age-dependent mating tactics. Males aged 2-4 years are weakly competitive and experienced stronger sexual selection as they accounted for a greater proportion of all males. Selection experienced by mature males appeared independent of demography. Our study provides a rare description of the demographic determinants of sexual selection in nature. [ABSTRACT FROM AUTHOR]

The adaptive nature of sociality has long been a central question in ecology and evolution. However, the relative importance of social behavior for fitness, compared to morphology and environment, remains largely unknown. We assessed the importance of sociality for fitness (lamb production and survival) in a population of marked bighorn sheep (Ovis canadensis) over 16 years (n ¼ 1022 sheep-years). We constructed social networks from observations (n ¼ 38 350) of group membership (n ¼ 3150 groups). We then tested whether consistent individual differences in social behavior (centrality) exist and evaluated their relative importance compared to factors known to affect fitness: mass, age, parental effects, and population density. Sheep exhibited consistent individual differences in social centrality. Controlling for maternal carryover effects and age, the positive effect of centrality in a social network on adult female lamb production and survival was equal or greater than the effect of body mass or population density. Social centrality had less effect on male survival and no effect on adult male lamb production or lamb survival. Through its effect on lamb production and survival, sociality in fission-fusion animal societies may ultimately influence population dynamics equally or more than morphological or environmental effects. [ABSTRACT FROM AUTHOR]

Heterozygosity-fitness correlations (HFCs) are often used to link individual genetic variation to differences in fitness. However, most studies examining HFCs find weak or no correlations. Here, we derive broad theoretical predictions about how many loci are needed to adequately measure genomic heterozygosity assuming different levels of identity disequilibrium (ID), a proxy for inbreeding. We then evaluate the expected ability to detect HFCs using an empirical data set of 200 microsatellites and 412 single nucleotide polymorphisms (SNPs) genotyped in two populations of bighorn sheep (Ovis canadensis), with different demographic histories. In both populations, heterozygosity was significantly correlated across marker types, although the strength of the correlation was weaker in a native population compared with one founded via translocation and later supplemented with additional individuals. Despite being bi-allelic, SNPs had similar correlations to genome-wide heterozygosity as microsatellites in both populations. For both marker types, this association became stronger and less variable as more markers were considered. Both populations had significant levels of ID; however, estimates were an order of magnitude lower in the native population. As with heterozygosity, SNPs performed similarly to microsatellites, and precision and accuracy of the estimates of ID increased as more loci were considered. Although dependent on the demographic history of the population considered, these results illustrate that genome-wide heterozygosity, and therefore HFCs, are best measured by a large number of markers, a feat now more realistically accomplished with SNPs than microsatellites. [ABSTRACT FROM AUTHOR]

Accurate stock identification is important for forest management, yet this can be a challenge for tree species that hybridize naturally. Species discriminating molecular markers provide a means to identify stock with high accuracy. In Canada, lodgepole pine ( Pinus contorta Dougl. ex Loud. var. latifolia) and jack pine ( Pinus banksiana Lamb) form a large hybrid zone in Alberta and Northwest Territories; within this hybrid zone, the identification of parentals and hybrids is difficult due to an overlap in morphological characteristics. Pure and hybrid ancestry can be resolved using microsatellite markers, but these are difficult and costly to type. We have developed a panel of SNP markers using 454 transcriptome sequence data that are more cost-effective, easier to score, and have greater discriminating power for differentiating species than microsatellites. Our SNP panel provides accurate and cost-efficient forest seed stock identification and will thereby facilitate reforestation and our pipeline can be applied to other hybrid systems globally. [ABSTRACT FROM AUTHOR]

Recent advances in technology facilitated development of large sets of genetic markers for many taxa, though most often model or domestic organisms. Cross-species application of genomic technologies may allow for rapid marker discovery in wild relatives of taxa with well-developed resources. We investigated returns from cross-species application of three commercially available SNP chips (the Ovine SNP50, Bovine SNP50 and Equine SNP50 Bead Chips) as a function of divergence time between the domestic source species and wild target species. Across all three chips, we observed a consistent linear decrease in call rate (~1.5% per million years), while retention of polymorphisms showed an exponential decay. These results will allow researchers to predict the expected amplification rate and polymorphism of cross-species application for their taxa of interest, as well as provide a resource for estimating divergence times. [ABSTRACT FROM AUTHOR]

Genetic rescue is a management intervention whereby a small population is supplemented with individuals from other populations in an attempt to reverse the effects of inbreeding and increased genetic load. One such rescue was recently documented in the population of bighorn sheep ( Ovis canadensis) within the National Bison Range wildlife refuge (Montana, USA). Here, we examine the locus-specific effects of rescue in this population using a newly developed genome-wide set of 195 microsatellite loci and first-generation linkage map. We found that the rate of introgression varied among loci and that 111 loci, 57% of those examined, deviated from patterns of neutral inheritance. The most common deviation was an excess of homozygous genotypes relative to neutral expectations, indicative of directional selection. As in previous study of this rescue, individuals with more introduced alleles had higher reproductive success and longevity. In addition, we found 30 loci, distributed throughout the genome, which seem to have individual effects on these life history traits. Although the potential for outbreeding depression is a major concern when translocating individuals between populations, we found no evidence of such effects in this population. [ABSTRACT FROM AUTHOR]

Dissecting the genetic architecture of fitness-related traits in wild populations is key to understanding evolution and the mechanisms maintaining adaptive genetic variation. We took advantage of a recently developed genetic linkage map and phenotypic information from wild pedigreed individuals from Ram Mountain, Alberta, Canada, to study the genetic architecture of ecologically important traits (horn volume, length, base circumference and body mass) in bighorn sheep. In addition to estimating sex-specific and cross-sex quantitative genetic parameters, we tested for the presence of quantitative trait loci (QTLs), colocalization of QTLs between bighorn sheep and domestic sheep, and sex × QTL interactions. All traits showed significant additive genetic variance and genetic correlations tended to be positive. Linkage analysis based on 241 microsatellite loci typed in 310 pedigreed animals resulted in no significant and five suggestive QTLs (four for horn dimension on chromosomes 1, 18 and 23, and one for body mass on chromosome 26) using genome-wide significance thresholds (Logarithm of odds (LOD) >3.31 and >1.88, respectively). We also confirmed the presence of a horn dimension QTL in bighorn sheep at the only position known to contain a similar QTL in domestic sheep (on chromosome 10 near the horns locus; nominal P<0.01) and highlighted a number of regions potentially containing weight-related QTLs in both species. As expected for sexually dimorphic traits involved in male-male combat, loci with sex-specific effects were detected. This study lays the foundation for future work on adaptive genetic variation and the evolutionary dynamics of sexually dimorphic traits in bighorn sheep. [ABSTRACT FROM AUTHOR]

The development of genomic resources for wild species is still in its infancy. However, cross-species utilization of technologies developed for their domestic counterparts has the potential to unlock the genomes of organisms that currently lack genomic resources. Here, we apply the OvineSNP50 BeadChip, developed for domestic sheep, to two related wild ungulate species: the bighorn sheep ( Ovis canadensis) and the thinhorn sheep ( Ovis dalli). Over 95% of the domestic sheep markers were successfully genotyped in a sample of fifty-two bighorn sheep while over 90% were genotyped in two thinhorn sheep. Pooling the results from both species identified 868 single-nucleotide polymorphisms (SNPs), 570 were detected in bighorn sheep, while 330 SNPs were identified in thinhorn sheep. The total panel of SNPs was able to discriminate between the two species, assign population of origin for bighorn sheep and detect known relationship classes within one population of bighorn sheep. Using an informative subset of these SNPs ( n = 308), we examined the extent of genome-wide linkage disequilibrium (LD) within one population of bighorn sheep and found that high levels of LD persist over 4 Mb. [ABSTRACT FROM AUTHOR]

To successfully manage wildlife diseases, it is necessary to understand factors that influence spread. One approach is to analyze host movement and social structure, as these behaviors can be associated with the probability of transmission. Some populations of mule deer (Odocoileus hemionus hemionus (Rafinesque, 1817)) in western Canada are infected with chronic wasting disease (CWD), a transmissible and fatal neurodegenerative disease. We used population analysis of spatial genetic structure of mule deer at broad and local scales to understand factors that influence spread. We genotyped 2535 mule deer sampled from Alberta, Saskatchewan, and portions of British Columbia using 16 microsatellite loci. We found weak genetic structure at broad spatial scales (overall FST = 0.008) that was well defined by geographic distance, indicating the risk of CWD spread from the focus of infection will decline gradually with increasing distance, but there are no barriers to the spread over time. At the local scale of approximately 2 km, elevated relatedness among CWD-infected individuals suggests transmission rates within social groups. Sex-biased spatial autocorrelation in genetic relatedness also indicates that female philopatry underlies the social structure, and therefore transmission among relatives is potentially driving local disease persistence. Afin de gérer avec succès les maladies de la faune sauvage, il est nécessaire de comprendre les facteurs qui influencent leur dissémination. Une méthode est d'analyser les déplacements et la structure sociale des hôtes puisque ces comportements peuvent être associés à la probabilité de transmission. Certaines populations de cerfs-mulets (Odocoileus hemionus hemionus (Rafinesque, 1817)) de l'Ouest canadien sont infectées par la maladie du dépérissement chronique (CWD), une maladie neurodégénérative transmissible et fatale. Une analyse de la structure spatiale génétique de la population de cerfs-mulets à des échelles étendues et locales nous a servi à comprendre les facteurs qui affectent la dissémination. Nous avons déterminé le génotype de 2535 cerfs-mulets échantillonnés en Alberta, en Saskatchewan et dans des régions de Colombie-Britannique à l'analyse de 16 locus microsatellites. Il existe une faible structure génétique aux échelles spatiales étendues (FST global = 0,008) qui est bien définie par la distance géographique, ce qui indique que le risque de transmission de la CWD à partir du foyer d'infection diminue graduellement en fonction de l'accroissement de la distance, mais qu'il n'y a pas de barrière à la transmission dans le temps. À l'échelle locale d'environ 2 km, la parenté accrue entre les individus infectés de la CWD laisse croire à une transmission à l'intérieur des groupes sociaux. L'autocorrélation spatiale de la parenté génétique avec biais sexuel indique aussi que la philopatrie des femelles sous-tend la structure spatiale et qu'ainsi la transmission entre individus apparentés explique potentiellement la persistance locale de la maladie. [ABSTRACT FROM AUTHOR]

Although it is generally expected that inbreeding would lower fitness, few studies have directly quantified the effects of inbreeding in wild mammals. We investigated the effects of inbreeding using long-term data from bighorn sheep on Ram Mountain, Alberta, Canada, over 20 years. This population underwent a drastic decline from 1992 to 2002 and has since failed to recover. We used a pedigree to calculate inbreeding coefficients and examined their impact on lamb growth, birth date and survival. Inbreeding had a substantial effect on female survival: for a given mass in September, the probability of overwinter survival for inbred female lambs was about 40% lower than that of noninbred ones. Contrary to our expectations, inbred female lambs were born earlier than noninbred ones. Earlier birth led to inbred female lambs being heavier by mid-September than noninbred ones. There was a nonsignificant trend for inbred female yearlings to weigh more than noninbred ones. A stronger mass-dependent viability selection for inbred compared to noninbred female lambs may explain why surviving inbred females were heavier than noninbred ones. Survival of male lambs was not affected by inbreeding. Sex-differential effects of inbreeding may be a general pattern in sexually dimorphic mammals, because of sex-biased maternal care or sexual differences in early development strategies. [ABSTRACT FROM AUTHOR]

We tested for cross-species amplification of microsatellite loci located throughout the domestic sheep ( Ovis aries) genome in two north American mountain ungulates (bighorn sheep, Ovis canadensis, and mountain goats, Oreamnos americanus). We identified 247 new polymorphic markers in bighorn sheep (≥ 3 alleles in one of two study populations) and 149 in mountain goats (≥ 2 alleles in a single study population) using 648 and 576 primer pairs, respectively. Our efforts increased the number of available polymorphic microsatellite markers to 327 for bighorn sheep and 180 for mountain goats. The average distance between successive polymorphic bighorn sheep and mountain goat markers inferred from the Australian domestic sheep genome linkage map (mean ± 1 SD) was 11.9 ± 9.2 and 15.8 ± 13.8 centimorgans, respectively. The development of genomic resources in these wildlife species enables future studies of the genetic architecture of trait variation. [ABSTRACT FROM AUTHOR]

The ease of obtaining genotypic data from wild populations has renewed interest in the relationship between individual genetic diversity and fitness-related traits (heterozygosity–fitness correlations, or HFC). Here we present a comprehensive meta-analysis of HFC studies using powerful multivariate techniques which account for nonindependence of data. We compare these findings with those from univariate techniques, and test the influence of a range of factors hypothesized to influence the strength of HFCs. We found small but significantly positive effect sizes for life-history, morphological, and physiological traits; while theory predicts higher mean effect sizes for life-history traits, effect size did not differ consistently with trait type. Newly proposed measures of variation were no more powerful at detecting relationships than multilocus heterozygosity, and populations predicted to have elevated inbreeding variance did not exhibit higher mean effect sizes. Finally, we found evidence for publication bias, with studies reporting weak, nonsignificant effects being under-represented in the literature. In general, our review shows that HFC studies do not generally reveal patterns predicted by population genetic theory, and are of small effect (less than 1% of the variance in phenotypic characters explained). Future studies should use more genetic marker data and utilize sampling designs that shed more light on the biological mechanisms that may modulate the strength of association, for example by contrasting the strength of HFCs in mainland and island populations of the same species, investigating the role of environmental stress, or by considering how selection has shaped the traits under investigation. [ABSTRACT FROM AUTHOR]

We describe the isolation and characterization of 12 highly polymorphic microsatellite loci for the muskrat, Ondatra zibethicus. Microsatellite markers from three other rodent species were cross-amplified in muskrat and one of them was polymorphic. We observed moderate to high levels of genetic variability in these 13 polymorphic loci (five to 22 alleles per locus) with observed heterozygosity ranging from 0.48 to 0.96. These markers will be useful for further studies on population genetic structure in muskrat and potentially in other rodent species. [ABSTRACT FROM AUTHOR]

STRUCTURE is the most widely used clustering software to detect population genetic structure. The last version of this software (STRUCTURE 2.1) has been enhanced recently to take into account the occurrence of linkage disequilibrium (LD) caused by admixture between populations. This last version, however, still does not consider the effects of strong background LD caused by genetic drift, and which may cause spurious results. STRUCTURE authors have, therefore, suggested a rough threshold value of the distance (1.0 cM) between two loci below which the pair of loci should not be used. Because of the sensitiveness of LD to demographic events, the distance between loci is not always a good indicator of the strength of LD. In this study, we examine the link between genomic distance and the strength of the correlation between loci (rLD) in a free-ranging population of mouflon (Ovis aries), and we present an empirical test of effect of rLD on the clustering results provided by the linkage model in STRUCTURE. We showed that a high rLD value increases the probability of detecting spurious clustering. We propose to use rLD as an index to base a decision on whether or not to use a pair of loci in a clustering analysis.Heredity (2007) 99, 374–380; doi:10.1038/sj.hdy.6801010; published online 11 July 2007 [ABSTRACT FROM AUTHOR]

Two putative populations of hooded seals ( Cystophora cristata) occur in the North Atlantic. The Greenland Sea population pup and breed on the pack ice near Jan Mayen (‘West Ice’) while the Northwest Atlantic population is thought to pup in the Davis Strait, in the Gulf of St. Lawrence (the ‘Gulf’), and off southern Labrador or northeast Newfoundland (the ‘Front’). We used microsatellite profiling of 300 individuals at 13 loci and mitochondrial DNA sequencing of the control region of 123 individuals to test for genetic differentiation between these four breeding herds. We found no significant genetic differences between breeding areas, nor evidence for cryptic nor higher level genetic structure in this species. The Greenland Sea breeding herd was genetically most distant from the Northwest Atlantic breeding areas; however, the differences were statistically nonsignificant. Our data therefore suggest that the world's hooded seals comprise a single panmictic genetic population. [ABSTRACT FROM AUTHOR]

There has recently been great interest in applying theoretical quantitative genetic models to empirical studies of evolution in wild populations. However, while classical models assume environmental constancy, most natural populations exist in variable environments. Here, we applied a novel analytical technique to a long-term study of birthweight in wild sheep and examined, for the first time, how variation in environmental quality simultaneously influences the strength of natural selection and the genetic basis of trait variability. In addition to demonstrating that selection and genetic variance vary dramatically across environments, our results show that environmental heterogeneity induces a negative correlation between these two parameters. Harsh environmental conditions were associated with strong selection for increased birthweight but low genetic variance, and vice versa. Consequently, the potential for microevolution in this population is constrained by either a lack of heritable variation (in poor environments) or by a reduced strength of selection (in good environments). More generally, environmental dependence of this nature may act to limit rates of evolution, maintain genetic variance, and favour phenotypic stasis in many natural systems. Assumptions of environmental constancy are likely to be violated in natural systems, and failure to acknowledge this may generate highly misleading expectations for phenotypic microevolution. [ABSTRACT FROM AUTHOR]

The confounding effects of population structure complicate efforts to identify regions of the genome under the influence of selection in natural populations. Here we test for evidence of selection in three genes involved in vertebrate immune function — the major histocompatibility complex (MHC), interferon gamma (IFNG) and natural resistance associated macrophage polymorphism (NRAMP) — in highly structured populations of wild thinhorn sheep ( Ovis dalli). We examined patterns of variation at microsatellite loci linked to these gene regions and at the DNA sequence level. Simple Watterson's tests indicated balancing selection at all three gene regions. However, evidence for selection was confounded by population structure, as the Watterson's test statistics from linked markers were not outside of the range of values from unlinked and presumably neutral microsatellites. The translated coding sequences of thinhorn IFNG and NRAMP are fixed and identical to those of domestic sheep ( Ovis aries). In contrast, the thinhorn MHC DRB locus shows significant evidence of overdominance through both an excess of nonsynonymous substitution and trans-species polymorphism. The failure to detect balancing selection at microsatellite loci linked to the MHC is likely the result of recombination between the markers and expressed gene regions. [ABSTRACT FROM AUTHOR]

A growing literature now documents the presence of fine-scale genetic structure in wild vertebrate populations. Breeding population size, levels of dispersal and polygyny — all hypothesized to affect population genetic structure — are known to be influenced by ecological conditions experienced by populations. However the possibility of temporal or spatial variation in fine-scale genetic structure as a result of ecological change is rarely considered or explored. Here we investigate temporal variation in fine-scale genetic structure in a red deer population on the Isle or Rum, Scotland. We document extremely fine-scale spatial genetic structure (< 100 m) amongst females but not males across a 24-year study period during which resource competition has intensified and the population has reached habitat carrying capacity. Based on census data, adult deer were allocated to one of three subpopulations in each year of the study. Global FST estimates for females generated using these subpopulations decreased over the study period, indicating a rapid decline in fine-scale genetic structure of the population. Global FST estimates for males were not different from zero across the study period. Using census and genetic data, we illustrate that, as a consequence of a release from culling early in the study period, the number of breeding females has increased while levels of polygyny have decreased in this population. We found little evidence for increasing dispersal between subpopulations over time in either sex. We argue that both increasing female population size and decreasing polygyny could explain the decline in female population genetic structure. [ABSTRACT FROM AUTHOR]

The thinhorn sheep ( Ovis dalli ssp.) provides a rare example of a North American large mammal that occupies most of its native range and maintains close to ancestral population size. There are currently two recognized subspecies, Dall's sheep ( O. d. dalli) and Stone's sheep ( O. d. stonei), the validity of which remains uncertain. We investigated the spatial genetic structure of thinhorn sheep populations representing both subspecies by genotyping individuals ( n = 919) from across the species range at 12 variable microsatellite loci. We found high levels of genetic diversity within ( HE = 0.722) and significant genetic structure among the 24 sampled areas ( FST = 0.160). Genetic distance measures and Bayesian clustering analyses revealed the presence of at least eight subpopulations that are delineated by mountain range topology. A strong overall pattern of isolation-by-distance is evident across the sampling range ( r = 0.75, P<0.001) suggesting limited dispersal and extensive philopatry. Partial Mantel tests of this relationship showed mountain range distinctions represent significant barriers to gene flow ( P = 0.0001), supporting the Bayesian analyses. Genetic structure was more strongly pronounced in southern Yukon and Alaska than elsewhere. We also show evidence for genetic differences between the two currently recognized thinhorn subspecies. [ABSTRACT FROM AUTHOR]

Nearly 70% of the world's mountain ungulate taxa are endangered. The availability of a standard set of DNA markers for forensic and molecular ecology studies would help to establish conservation programs and detect poaching activities of these endangered taxa. We tested 60 published microsatellite primer pairs from bovids (cattle, sheep and goat) on 49 individuals from 11 taxa including six wild goat-like species ( Capra spp.), three divergent wild sheep ( Ovis spp.), and two chamois ( Rupicapra spp.) species. Approximately 30 microsatellites amplified a microsatellite-like PCR product in all three genera, and with the exception of ILST097, nearly all the loci were polymorphic within most of the 11 species. [ABSTRACT FROM AUTHOR]

Abstract The fine-scale genetic structure of wild animal populations has rarely been analysed, yet is potentially important as a confounding factor in quantitative genetic and allelic association studies, as well as having implications for population dynamics, inbreeding and kin selection. In this study, we examined the extent to which the three spatial subunits, or hefts, of the Village Bay population of Soay sheep (Ovis aries ) on St Kilda, Scotland, are genetically structured using data from 20 microsatellite and protein loci. Allele frequencies differed significantly among three hefts in all the study years we considered (1987–2000 inclusive). Small but significantly positive F ST and negative F IS values were observed in most years, indicating that the hefts are genetically differentiated, and that within each heft there is more observed heterozygosity than would be expected if each were an isolated breeding population. Males showed less fidelity to their natal heft, and as a consequence higher levels of relatedness within hefts were observed among females than among males. There was a significant negative relationship between geographical proximity and relatedness in pairwise comparisons involving females, and on average pairs of females located within 50 m of each other were related at the equivalent level of second cousins. Structure is therefore largely driven by incomplete postnatal dispersal by females. Mating appears to be random with respect to the spatial-genetic substructure of the hefts, and therefore genetic structure does not contribute to the overall rate of inbreeding in the population. However, genetic substructure can lead to allelic associations and generate environmental effects within lineages that have the potential to confound heritability analyses and allelic association studies. [ABSTRACT FROM AUTHOR]