Hair Cortisol and Testosterone Concentrations in Relation to Maturity and Breeding Status of Male Feral Horses.

Steroid-hormone concentrations from non-invasively obtained biomarkers, like hair, can provide a representation of circulating hormones diffused over relatively long time periods (e.g., weeks or months). The hormone cortisol is often associated with physiological or even psychological stress, while testosterone is strongly associated with male development and reproductive success. Increasingly, studies are using hormone levels derived from hair to make inferences among both domestic animals and wildlife. For horses, all previous hair hormone analysis has been done on companion or working animals. We evaluated the levels of hair cortisol (n = 153) and testosterone (n = 48) from 136 feral horses living on Sable Island, Canada that have been part of a long-term individual-based study since 2008. This population has been undisturbed and unmanaged for over 50 years, and exhibits the natural social organization for horses, harem defense polygyny. Hair samples were collected in mid to late summer and the segment analyzed corresponds with hair grown during, and following, the peak of the reproductive season. Social position was determined based on the male's role as either a dominant breeding Stallion (Stallion), a non-breeding subordinate male (tag), adult Bachelor (5 years old or older), or Immature male (2-4 years of age). While there was no difference in hair-cortisol concentration among any class of adult males (i.e., Stallion, tag, or Bachelor), Immature males had significantly lower hair cortisol concentrations than the other groups (p = 0.001). Hair testosterone levels among the four social positions were significantly higher among Stallions (p = 0.04). Hair testosterone concentration was also significantly related to the probability of a male being either a Bachelor or Stallion and was the only variable remaining in AICc model selection (p = 0.016, AICc = 32.3, Null AICc = 38.8). While not a significant relationship, Stallions had a negative correlation between hair cortisol concentrations and testosterone (R2 = -0.20, p = 0.383), and Bachelors, conversely, had a positive association (R2 = 0.43, p = 0.246). Our observations of hormone concentrations in relation to physiological, social, or reproductive parameters in this population suggest trends that are similar to what has been established using blood or other matrices.

Hair Cortisol Concentrations in Feral Horses and the Influence of Physiological and Social Factors.

Cortisol is a glucocorticoid hormone produced during activation of the hypothalamic-pituitary-adrenal axis (HPA) in response to psychological or physiological demands. High amounts of circulating cortisol can be found in individuals experiencing energetically demanding physiological events, such as pregnancy, lactation, injury, or starvation, but, also, in individuals who may have less obvious HPA activation from social situations. The feral horse population on Sable Island (Nova Scotia, Canada) provides an opportunity to look at hair cortisol concentration (HCC) as a proxy for circulating cortisol concentration to better understand physiological correlates. The horse's complex social structure also allows us to look at how the population and group structure may influence HPA activation. Hair samples (n = 282) were analyzed from 113 females and 135 males. Females with dependent offspring (foals) had higher HCC than those females without dependent offspring (p = 0.005). Horses in poor body condition were also more likely to have higher HCC (females: p < 0.001, males: p = 0.028); females had greater variation in the body condition index (BCI), which also correlated with foal production. In general, the top-ranked models describing female cortisol levels included age, BCI, presence of a foal, as well as social measures such as harem size and the number of bachelors in the vicinity. The top model describing male cortisol levels included age, BCI, and year of collection only, and the number of bachelors in the home range appeared in subsequent, though still high-ranked, models. Among the variables not of direct interest, we found some significant results relating to hair color and hair texture. Differences in HCC patterns between feral and domestically kept horses (e.g., age and sex) are likely linked to periods of resource limitations, particularly for individuals experiencing energetically demanding processes such as reproduction, illness/parasitism, or related to experiencing the full range of social and reproductive behaviors.

Shallow shotgun sequencing of the microbiome recapitulates 16S amplicon results and provides functional insights.

Prevailing 16S rRNA gene-amplicon methods for characterizing the bacterial microbiome of wildlife are economical, but result in coarse taxonomic classifications, are subject to primer and 16S copy number biases, and do not allow for direct estimation of microbiome functional potential. While deep shotgun metagenomic sequencing can overcome many of these limitations, it is prohibitively expensive for large sample sets. Here we evaluated the ability of shallow shotgun metagenomic sequencing to characterize taxonomic and functional patterns in the faecal microbiome of a model population of feral horses (Sable Island, Canada). Since 2007, this unmanaged population has been the subject of an individual-based, long-term ecological study. Using deep shotgun metagenomic sequencing, we determined the sequencing depth required to accurately characterize the horse microbiome. In comparing conventional vs. high-throughput shotgun metagenomic library preparation techniques, we validate the use of more cost-effective laboratory methods. Finally, we characterize similarities between 16S amplicon and shallow shotgun characterization of the microbiome, and demonstrate that the latter recapitulates biological patterns first described in a published amplicon data set. Unlike for amplicon data, we further demonstrate how shallow shotgun metagenomic data provide useful insights regarding microbiome functional potential which support previously hypothesized diet effects in this study system.

Targeted genome-wide SNP genotyping in feral horses using non-invasive fecal swabs.

The development of high-throughput sequencing has prompted a transition in wildlife genetics from using microsatellites toward sets of single nucleotide polymorphisms (SNPs). However, genotyping large numbers of targeted SNPs using non-invasive samples remains challenging due to relatively large DNA input requirements. Recently, target enrichment has emerged as a promising approach requiring little template DNA. We assessed the efficacy of Tecan Genomics' Allegro Targeted Genotyping (ATG) for generating genome-wide SNP data in feral horses using DNA isolated from fecal swabs. Total and host-specific DNA were quantified for 989 samples collected as part of a long-term individual-based study of feral horses on Sable Island, Nova Scotia, Canada, using dsDNA fluorescence and a host-specific qPCR assay, respectively. Forty-eight samples representing 44 individuals containing at least 10 ng of host DNA (ATG's recommended minimum input) were genotyped using a custom multiplex panel targeting 279 SNPs. Genotyping accuracy and consistency were assessed by contrasting ATG genotypes with those obtained from the same individuals with SNP microarrays, and from multiple samples from the same horse, respectively. 62% of swabs yielded the minimum recommended amount of host DNA for ATG. Ignoring samples that failed to amplify, ATG recovered an average of 88.8% targeted sites per sample, while genotype concordance between ATG and SNP microarrays was 98.5%. The repeatability of genotypes from the same individual approached unity with an average of 99.9%. This study demonstrates the suitability of ATG for genome-wide, non-invasive targeted SNP genotyping, and will facilitate further ecological and conservation genetics research in equids and related species. Supplementary Information: The online version contains supplementary material available at 10.1007/s12686-022-01259-2.

Insect-mediated apparent competition between mammals in a boreal food web.

While the important role of animal-mediated interactions in the top-down restructuring of plant communities is well documented, less is known of their ensuing repercussions at higher trophic levels. We demonstrate how typically decoupled ecological interactions may become intertwined such that the impact of an insect pest on forest structure and composition alters predator-prey interactions among large mammals. Specifically, we show how irruptions in a common, cyclic insect pest of the boreal forest, the spruce budworm (Choristoneura fumiferana), modulated an indirect trophic interaction by initiating a flush in deciduous vegetation that benefited moose (Alces alces), in turn strengthening apparent competition between moose and threatened boreal caribou (Rangifer tarandus caribou) via wolf (Canis lupus) predation. Critically, predation on caribou postoutbreak was exacerbated by human activity (salvage logging). We believe our observations of significant, large-scale reverberating consumer-producer-consumer interactions are likely to be common in nature.

Trophic consequences of terrestrial eutrophication for a threatened ungulate.

Changes in primary productivity have the potential to substantially alter food webs, with positive outcomes for some species and negative outcomes for others. Understanding the environmental context and species traits that give rise to these divergent outcomes is a major challenge to the generality of both theoretical and applied ecology. In aquatic systems, nutrient-mediated eutrophication has led to major declines in species diversity, motivating us to seek terrestrial analogues using a large-mammal system across 598 000 km2 of the Canadian boreal forest. These forests are undergoing some of the most rapid rates of land-use change on Earth and are home to declining caribou (Rangifer tarandus caribou) populations. Using satellite-derived estimates of primary productivity, coupled with estimates of moose (Alces alces) and wolf (Canis lupus) abundance, we used path analyses to discriminate among hypotheses explaining how habitat alteration can affect caribou population growth. Hypotheses included food limitation, resource dominance by moose over caribou, and apparent competition with predators shared between moose and caribou. Results support apparent competition and yield estimates of wolf densities (1.8 individuals 1000 km-2) above which caribou populations decline. Our multi-trophic analysis provides insight into the cascading effects of habitat alteration from forest cutting that destabilize terrestrial predator-prey dynamics. Finally, the path analysis highlights why conservation actions directed at the proximate cause of caribou decline have been more successful in the near term than those directed further along the trophic chain.

A repeatable and quantitative DNA metabarcoding assay to characterize mixed strongyle infections in horses.

Horses are ubiquitously infected by a diversity of gastro-intestinal parasitic helminths. Of particular importance are nematodes of the family Strongylidae, which can significantly impact horse health and performance. However, knowledge about equine strongyles remains limited due to our inability to identify most species non-invasively using traditional morphological techniques. We developed a new internal transcribed spacer 2 (ITS2) DNA metabarcoding 'nemabiome' assay to characterise mixed strongyle infections in horses and assessed its performance by applying it to pools of infective larvae from fecal samples from an experimental herd in Kentucky, USA and two feral horse populations from Sable Island and Alberta, Canada. In addition to reporting the detection of 33 different species with high confidence, we illustrate the assay's repeatability by comparing results generated from aliquots from the same fecal samples and from individual horses sampled repeatedly over multiple days or months. We also validate the quantitative potential of the assay by demonstrating that the proportion of amplicon reads assigned to different species scales linearly with the number of larvae present. This new tool significantly improves equine strongyle diagnostics, presenting opportunities for research on species-specific anthelmintic resistance and the causes and consequences of variation in mixed infections.

Bacterial dispersal and drift drive microbiome diversity patterns within a population of feral hindgut fermenters.

Studies of microbiome variation in wildlife often emphasize host physiology and diet as proximate selective pressures acting on host-associated microbiota. In contrast, microbial dispersal and ecological drift are more rarely considered. Using amplicon sequencing, we characterized the bacterial microbiome of adult female (n = 86) Sable Island horses (Nova Scotia, Canada) as part of a detailed individual-based study of this feral population. Using data on sampling date, horse location, age, parental status, and local habitat variables, we contrasted the ability of spatiotemporal, life history, and environmental factors to explain microbiome diversity among Sable Island horses. We extended inferences made from these analyses with both phylogeny-informed and phylogeny-independent null modelling approaches to identify deviations from stochastic expectations. Phylogeny-informed diversity measures were correlated with spatial and local habitat variables, but null modelling results suggested that heterogeneity in ecological drift, rather than differential selective pressures acting on the microbiome, was responsible for these correlations. Conversely, phylogeny-independent diversity measures were best explained by host spatial and social structure, suggesting that taxonomic composition of the microbiome was shaped most strongly by bacterial dispersal. Parental status was important but correlated with measures of ß-dispersion rather than ß-diversity (mares without foals had lower alpha diversity and more variable microbiomes than mares with foals). Our results suggest that between host microbiome variation within the Sable Island horse population is driven more strongly by bacterial dispersal and ecological drift than by differential selective pressures. These results emphasize the need to consider alternative ecological processes in the study of microbiomes.

Causes and consequences of an unusually male-biased adult sex ratio in an unmanaged feral horse population.

The adult sex ratio (ASR) is important within ecology due to its predicted effects on behaviour, demography and evolution, but research examining the causes and consequences of ASR bias have lagged behind the studies of sex ratios at earlier life stages. Although ungulate ASR is relatively well-studied, exceptions to the usual female-biased ASR challenge our understanding of the underlying drivers of biased ASR and provide an opportunity to better understand its consequences. Some feral ungulate populations, including multiple horse populations, exhibit unusually male-biased ASR. For example, research suggests that the feral horse Equus ferus caballus population on Sable Island, Nova Scotia, Canada may exhibit a male-biased ASR. Such exceptions to the rule provide a valuable opportunity to reveal the contributions of environmental context and trait differences to ASR bias. We aimed to test for bias in Sable Island horse ASR, identify the demographic drivers of bias, and explore its demographic and social consequences. To do this, we used life history, movement and group membership information for hundreds of horses followed through a long-term individual-based study between 2007 and 2018. Sable Island horse ASR is male biased and this skew has increased over time, reaching 62% male in 2018. Our life table response experiment suggested that ASR skew was driven predominantly by male-biased adult survival. Further analyses pointed to sex-biased survival being driven by reduced female survival post-reproduction. Male-biased ASR was associated with reduced harem sizes, an increase in the number of social groups on the island, and reduced reproduction in young females. Our results support the idea that male-biased ASR in feral ungulate populations may be caused by a combination of high population density and high reproductive output. We suggest that female-biased mortality may be caused by females continuing to reproduce at high density, and thus being more susceptible to resource shortages. Thus, our results highlight the strong context dependence of ASR. Furthermore, our work indicates the potential for ASR to substantially alter a population's social organisation. Such changes in social structure could have knock-on consequences for demography by altering the formation/stability of social relationships, or competition for matings.

Large-scale prion protein genotyping in Canadian caribou populations and potential impact on chronic wasting disease susceptibility.

Polymorphisms within the prion protein gene (Prnp) are an intrinsic factor that can modulate chronic wasting disease (CWD) pathogenesis in cervids. Although wild European reindeer (Rangifer tarandus tarandus) were infected with CWD, as yet there have been no reports of the disease in North American caribou (R. tarandus spp.). Previous Prnp genotyping studies on approximately 200 caribou revealed single nucleotide polymorphisms (SNPs) at codons 2 (V/M), 129 (G/S), 138 (S/N), 146 (N/n) and 169 (V/M). The impact of these polymorphisms on CWD transmission is mostly unknown, except for codon 138. Reindeer carrying at least one allele encoding for asparagine (138NN or 138SN) are less susceptible to clinical CWD upon infection by natural routes, with the majority of prions limited to extraneural tissues. We sequenced the Prnp coding region of two caribou subspecies (n = 986) from British Columbia, Saskatchewan, Yukon, Nunavut and the Northwest Territories, to identify SNPs and their frequencies. Genotype frequencies at codon 138 differed significantly between barren-ground (R. t. groenlandicus) and woodland (R. t. caribou) caribou when we excluded the Chinchaga herd (p < .05). We also found new variants at codons 153 (Y/F) and 242 (P/L). Our findings show that the 138N allele is rare among caribou in areas with higher risk of contact with CWD-infected species. As both subspecies are classified as Threatened and play significant roles in North American Indigenous culture, history, food security and the economy, determining frequencies of Prnp genotypes associated with susceptibility to CWD is important for future wildlife management measures.

When the protection of a threatened species depends on the economy of a foreign nation.

A significant challenge of conservation biology is to preserve species in places where their critical habitat also attracts significant economic interest. The problem is compounded when species distributions occur across large spatial extents. Threatened boreal caribou (Rangifer tarandus caribou) epitomize this problem: their critical habitat encompasses a vast expanse of forest that also supplies much of Canada's merchantable timber. Boreal caribou were protected under the Canada Species at Risk Act in 2003. We investigated putative drivers of reduced disturbance for caribou habitat since then. Where the cumulative logging footprint slowed within caribou habitat, this has resulted neither from decreases in annual allowable cut of timber nor the creation or expansion of protected areas. Rather, it has fluctuated with the American economy relative to that of Canada. For each $0.05 US lost over the $CAD, 129 km2 of caribou habitat was not disturbed by logging in a given year. Recent population declines have been occurring even though logging typically remained at <70% of allowed levels. Our study raises concerns about how caribou are functionally being conserved under the current application of existing legislation. In this globalized world, the economy of foreign nations is increasingly likely to govern national conservation objectives.

Functional response of wolves to human development across boreal North America.

AIM: The influence of humans on large carnivores, including wolves, is a worldwide conservation concern. In addition, human-caused changes in carnivore density and distribution might have impacts on prey and, indirectly, on vegetation. We therefore tested wolf responses to infrastructure related to natural resource development (i.e., human footprint). LOCATION: Our study provides one of the most extensive assessments of how predators like wolves select habitat in response to various degrees of footprint across boreal ecosystems encompassing over a million square kilometers of Canada. METHODS: We deployed GPS-collars on 172 wolves, monitored movements and used a generalized functional response (GFR) model of resource selection. A functional response in habitat selection occurs when selection varies as a function of the availability of that habitat. GFRs can clarify how human-induced habitat changes are influencing wildlife across large, diverse landscapes. RESULTS: Wolves displayed a functional response to footprint. Wolves were more likely to select forest harvest cutblocks in regions with higher cutblock density (i.e., a positive functional response to high-quality habitats for ungulate prey) and to select for higher road density in regions where road density was high (i.e., a positive functional response to human-created travel routes). Wolves were more likely to use cutblocks in habitats with low road densities, and more likely to use roads in habitats with low cutblock densities, except in winter when wolves were more likely to use roads regardless of cutblock density. MAIN CONCLUSIONS: These interactions suggest that wolves trade-off among human-impacted habitats, and adaptively switch from using roads to facilitate movement (while also risking encounters with humans), to using cutblocks that may have higher ungulate densities. We recommend that conservation managers consider the contextual and interacting effects of footprints when assessing impacts on carnivores. These effects likely have indirect impacts on ecosystems too, including on prey species.

Environmental and demographic drivers of male mating success vary across sequential reproductive episodes in a polygynous breeder.

Ecological and social factors underpinning the inequality of male mating success in animal societies can be related to sex ratio, sexual conflict between breeders, effects of nonbreeders, resource dispersion, climatic conditions, and the various sequential stages of mating competition that constitute the sexual selection process. Here, we conducted an individual-based study to investigate how local resource availability and demography interact with annual climate conditions to determine the degree of male mating inequality, and thus opportunity for sexual selection across two sequential reproductive episodes (harem and subsequent mate acquisition) in a naturally regulated (feral) horse population in Sable Island National Park Preserve, Canada. Using a 5-year, spatially explicit, mark-resight dataset and hierarchical mixed-effects linear modeling, we evaluated the influence of adult sex ratio (ASR) on mating success and then tested for effects of freshwater availability, density, unpaired male abundance, and precipitation during each breeding season. Unpaired male abundance, freshwater availability, and ASR differed in their effects on male mating success according to year and selection episode. Opportunity for sexual selection in males associated with harem acquisition increased with ASR, and unpaired male abundance further explained weather-related interannual variation after accounting for ASR. In contrast, once a harem was secured, ASR had little effect on male mating inequality in regard to acquiring additional females, while interannual variation in mating inequality increased with decreasing freshwater availability. Our findings show that local demography, resource availability, and weather effect opportunity for sexual selection in males differently depending on selection episode, and can attenuate or accentuate effects of ASR.

Quantitative genetics of gastrointestinal strongyle burden and associated body condition in feral horses.

Variability in host resistance or tolerance to parasites is nearly ubiquitous, and is of key significance in understanding the evolutionary processes shaping host-parasite interactions. While ample research has been conducted on the genetics of parasite burden in livestock, relatively little has been done in free-living populations. Here, we investigate the sources of (co)variation in strongyle nematode faecal egg count (FEC) and body condition in Sable Island horses, a feral population in which parasite burden has previously been shown to negatively correlate with body condition. We used the quantitative genetic "animal model" to understand the sources of (co)variation in these traits, and tested for impacts of an important spatial gradient in habitat quality on the parameter estimates. Although FEC is significantly heritable (h 2 = 0.43 ±â€¯0.11), there was no evidence for significant additive genetic variation in body condition (h 2 = 0.04 ±â€¯0.07), and therefore there was also no significant genetic covariance between the two traits. The negative phenotypic covariance between these traits therefore does not derive principally from additive genetic effects. We also found that both FEC and body condition increase from east to west across the island, which indicates that the longitudinal environmental gradient is not responsible for the negative phenotypic association observed between these traits. There was also little evidence to suggest that quantitative genetic parameters were biased when an individual's location along the island's environmental gradient was not incorporated into the analysis. This research provides new and important insights into the genetic basis and adaptive potential of parasite resistance in free-living animals, and highlights the importance of environmental heterogeneity in modulating host-parasite interactions in wild vertebrate systems.

Partitioning drivers of spatial genetic variation for a continuously distributed population of boreal caribou: Implications for management unit delineation.

Isolation by distance (IBD) is a natural pattern not readily incorporated into theoretical models nor traditional metrics for differentiating populations, although clinal genetic differentiation can be characteristic of many wildlife species. Landscape features can also drive population structure additive to baseline IBD resulting in differentiation through isolation-by-resistance (IBR). We assessed the population genetic structure of boreal caribou across western Canada using nonspatial (STRUCTURE) and spatial (MEMGENE) clustering methods and investigated the relative contribution of IBD and IBR on genetic variation of 1,221 boreal caribou multilocus genotypes across western Canada. We further introduced a novel approach to compare the partitioning of individuals into management units (MU) and assessed levels of genetic connectivity under different MU scenarios. STRUCTURE delineated five genetic clusters while MEMGENE identified finer-scale differentiation across the study area. IBD was significant and did not differ for males and females both across and among detected genetic clusters. MEMGENE landscape analysis further quantified the proportion of genetic variation contributed by IBD and IBR patterns, allowing for the relative importance of spatial drivers, including roads, water bodies, and wildfires, to be assessed and incorporated into the characterization of population structure for the delineation of MUs. Local population units, as currently delineated in the boreal caribou recovery strategy, do not capture the genetic variation and connectivity of the ecotype across the study area. Here, we provide the tools to assess fine-scale spatial patterns of genetic variation, partition drivers of genetic variation, and evaluate the best management options for maintaining genetic connectivity. Our approach is highly relevant to vagile wildlife species that are of management and conservation concern and demonstrate varying degrees of IBD and IBR with clinal spatial genetic structure that challenges the delineation of discrete population boundaries.

Examining forest resilience to changing fire frequency in a fire-prone region of boreal forest.

Future changes in climate are widely anticipated to increase fire frequency, particularly in boreal forests where extreme warming is expected to occur. Feedbacks between vegetation and fire may modify the direct effects of warming on fire activity and shape ecological responses to changing fire frequency. We investigate these interactions using extensive field data from the Boreal Shield of Saskatchewan, Canada, a region where >40% of the forest has burned in the past 30 years. We use geospatial and field data to assess the resistance and resilience of eight common vegetation states to frequent fire by quantifying the occurrence of short-interval fires and their effect on recovery to a similar vegetation state. These empirical relationships are combined with data from published literature to parameterize a spatially explicit, state-and-transition simulation model of fire and forest succession. We use this model to ask if and how: (a) feedbacks between vegetation and wildfire may modify fire activity on the landscape, and (b) more frequent fire may affect landscape forest composition and age structure. Both field and GIS data suggest the probability of fire is low in the initial decades after fire, supporting the hypothesis that fuel accumulation may exert a negative feedback on fire frequency. Field observations of pre- and postfire composition indicate that switches in forest state are more likely in conifer stands that burn at a young age, supporting the hypothesis that resilience is lower in immature stands. Stands dominated by deciduous trees or jack pine were generally resilient to fire, while mixed conifer and well-drained spruce forests were less resilient. However, simulation modeling suggests increased fire activity may result in large changes in forest age structure and composition, despite the feedbacks between vegetation-fire likely to occur with increased fire activity.

Individual quality and age but not environmental or social conditions modulate costs of reproduction in a capital breeder.

Costs associated with reproduction are widely known to play a role in the evolution of reproductive tactics with consequences to population and eco-evolutionary dynamics. Evaluating these costs as they pertain to species in the wild remains an important goal of evolutionary ecology. Individual heterogeneity, including differences in individual quality (i.e., among-individual differences in traits associated with survival and reproduction) or state, and variation in environmental and social conditions can modulate the costs of reproduction; however, few studies have considered effects of these factors simultaneously. Taking advantage of a detailed, long-term dataset for a population of feral horses (Sable Island, Nova Scotia, Canada), we address the question of how intrinsic (quality, age), environmental (winter severity, location), and social conditions (group size, composition, sex ratio, density) influence the costs of reproduction on subsequent reproduction. Individual quality was measured using a multivariate analysis on a combination of four static and dynamic traits expected to depict heterogeneity in individual performance. Female quality and age interacted with reproductive status of the previous year to determine current reproductive effort, while no effect of social or environmental covariates was found. High-quality females showed higher probabilities of giving birth and weaning their foal regardless of their reproductive status the previous year, while those of lower quality showed lower probabilities of producing foals in successive years. Middle-aged (prime) females had the highest probability of giving birth when they had not reproduced the year before, but no such relationship with age was found among females that had reproduced the previous year, indicating that prime-aged females bear higher costs of reproduction. We show that individual quality and age were key factors modulating the costs of reproduction in a capital breeder but that environmental or social conditions were not, highlighting the importance of considering multiple factors when studying costs of reproduction.

Climatic conditions cause spatially dynamic polygyny thresholds in a large mammal.

The polygyny threshold (PT) is a critical transition point in the sexual selection process for many organisms in natural populations, characterizing when females choose to mate with an already mated male over an unmated one to improve fitness. Understanding its causes and consequences is therefore of high interest. While both theoretical and empirical work suggest that the degree of polygyny within a species is plastic and a function of male inequality, the functional relationship between underlying availability of resources occupied by breeding males under variable climatic conditions and the dynamics of PTs across space and time has received less attention. Here, we use a standardized measure of male mating inequality as the culmination of female mate choices to analyse how spatially dynamic PTs in a naturally regulated feral horse (Equus ferus caballus) population emerge along a geographic gradient in a known, limiting resource (freshwater) each year from variable climatic conditions. Polygyny threshold distance from permanent freshwater increased with increasing precipitation during the breeding season of each year, suggesting a relationship between annual resource availability and female mate choice. The mechanism by which climatic conditions underpin the spatial dynamics of PTs was likely through precipitation providing ephemeral freshwater sources across the study area that effectively weakened the gradient in availability of permanent freshwater, thereby providing mating males that occupied home ranges far from permanent water with access to this limiting resource and enabling them to attract and retain females. Increased precipitation also coincided with a decreased proportion of males in the population that experienced sexual selection pressure attributed to female mate choice in relation to the acquisition and/or defence of freshwater sources. Climatic conditions caused spatial shifts in PTs annually along the geographic gradient in resource availability. Our findings reveal that such environmental gradients may either buffer or amplify impacts of climatic variation on selection pressure operating in natural populations, and emphasize the importance of integrating spatially explicit PTs with atmospheric fluctuations when predicting the effect of climatic change on selection processes within populations that occupy environmental gradients.

Density-dependent resource selection by a terrestrial herbivore in response to sea-to-land nutrient transfer by seals.

Sea-to-land nutrient transfers can connect marine food webs to those on land, creating a dependence on marine webs by opportunistic species. We show how nitrogen, imported by gray seals, Halichoerus grypus, and traced through stable isotope (δ15 N) measurements in marram grass, Ammophila breviligulata, significantly alters foraging behavior of a free-roaming megaherbivore (feral horses, Equus ferus caballus) on Sable Island, Canada. Values of δ15 N correlated with protein content of marram and strongly related to pupping-seal densities, and positively influenced selective foraging by horses. The latter was density dependent, consistent with optimal foraging theory. We present the first demonstration of how sea-to-land nutrient transfers can affect the behavioral process of resource selection (resource use relative to availability) of terrestrial consumers. We hypothesize that persistence of horses on Sable Island is being facilitated by N subsidies. Our results have relevance to advancing theory on trophic dynamics in island biogeography and metaecosystem ecology.

Negative covariance between parasite load and body condition in a population of feral horses.

In wild and domestic animals, gastrointestinal parasites can have significant impacts on host development, condition, health, reproduction and longevity. Improving our understanding of the causes and consequences of individual-level variation in parasite load is therefore of prime interest. Here we investigated the relationship between strongyle fecal egg count (FEC) and body condition in a unique, naturalized population of horses that has never been exposed to anthelmintic drugs (Sable Island, Nova Scotia, Canada). We first quantified variation in FEC and condition for 447 individuals according to intrinsic (sex, age, reproductive status, social status) and extrinsic (group size, location, local density) variables. We then quantified the repeatability of measurements obtained over a field season and tested for covariance between FEC and condition. FECs were high relative to other horse populations (mean eggs per gram ± SD = 1543·28 ± 209·94). FECs generally decreased with age, were higher in lactating vs non-lactating females, and unexpectedly lower in males in some part of the island. FECs and condition were both spatially structured, with patterns depending on age, sex and reproductive status. FECs and condition were both repeatable. Most notably, FECs and condition were negatively correlated, especially in adult females.