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.

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.

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.

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.

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.

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.

Increasing density leads to generalization in both coarse-grained habitat selection and fine-grained resource selection in a large mammal.

Density is a fundamental driver of many ecological processes including habitat selection. Theory on density-dependent habitat selection predicts that animals should be distributed relative to profitability of habitat, resulting in reduced specialization in selection (i.e. generalization) as density increases and competition intensifies. Despite mounting empirical support for density-dependent habitat selection using isodars to describe coarse-grained (interhabitat) animal movements, we know little of how density affects fine-grained resource selection of animals within habitats [e.g. using resource selection functions (RSFs)]. Using isodars and RSFs, we tested whether density simultaneously modified habitat selection and within-habitat resource selection in a rapidly growing population of feral horses (Equus ferus caballus Linnaeus; Sable Island, Nova Scotia, Canada; 42% increase in population size from 2008 to 2012). Among three heterogeneous habitat zones on Sable Island describing population clusters distributed along a west-east resource gradient (west-central-east), isodars revealed that horses used available habitat in a density-dependent manner. Intercepts and slopes of isodars demonstrated a pattern of habitat selection that first favoured the west, which generalized to include central and east habitats with increasing population size consistent with our understanding of habitat quality on Sable Island. Resource selection functions revealed that horses selected for vegetation associations similarly at two scales of extent (total island and within-habitat zone). When densities were locally low, horses were able to select for sites of the most productive forage (grasslands) relative to those of poorer quality. However, as local carrying capacity was approached, selection for the best of available forage types weakened while selection for lower-quality vegetation increased (and eventually exceeded that of grasslands). Isodars can effectively describe coarse-grained habitat selection in large mammals. Our study also shows that the main predictions of density-dependent habitat selection are highly relevant to our interpretation of RSFs in space and time. At low but not necessarily high population size, density will be a leading indicator of habitat quality. Fitness maximization from specialist vs. generalist strategies of habitat and resource selection may well be apparent at multiple spatial extents and grains of resolution.

Density-dependent habitat selection and partitioning between two sympatric ungulates.

Theory on density-dependent habitat selection predicts that as population density of a species increases, use of higher quality (primary) habitat by individuals declines while use of lower quality (secondary) habitat rises. Habitat partitioning is often considered the primary mechanism for coexistence between similar species, but how this process evolves with changes in population density remains to be empirically tested for free-ranging ungulates. We used resource-selection functions to quantify density effects on landscape-scale habitat selection of two sympatric species of ungulates [moose (Alces alces) and elk (Cervus canadensis manitobensis)] in Riding Mountain National Park, Manitoba, Canada (2000-2011). The density of elk was actively reduced from 1.2 to 0.4 elk km(-2) through increased hunting effort during the period of study, while moose density decreased without additional human influence from 1.6-0.7 moose km(-2). Patterns of habitat selection during winter by both species changed in accordance to expectations from density-dependent habitat-selection theory. At low intraspecific density, moose and elk did not partition habitat, as both species selected strongly for mixed forest (primary habitat providing both food and cover), but did so in different areas segregated across an elevational gradient. As intraspecific density increased, selection for primary habitat by both species decreased, while selection for secondary, lower quality habitat such as agricultural fields (for elk) and built-up areas (for moose) increased. We show that habitat-selection strategies during winter for moose and elk, and subsequent effects on habitat partitioning, depend heavily on the position in state space (density) of both species.

Methanogenic patterns in the gut microbiome are associated with survival in a population of feral horses.

Gut microbiomes are widely hypothesised to influence host fitness and have been experimentally shown to affect host health and phenotypes under laboratory conditions. However, the extent to which they do so in free-living animal populations and the proximate mechanisms involved remain open questions. In this study, using long-term, individual-based life history and shallow shotgun metagenomic sequencing data (2394 fecal samples from 794 individuals collected between 2013-2019), we quantify relationships between gut microbiome variation and survival in a feral population of horses under natural food limitation (Sable Island, Canada), and test metagenome-derived predictions using short-chain fatty acid data. We report detailed evidence that variation in the gut microbiome is associated with a host fitness proxy in nature and outline hypotheses of pathogenesis and methanogenesis as key causal mechanisms which may underlie such patterns in feral horses, and perhaps, wild herbivores more generally.

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.

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.

Considering ecological dynamics in resource selection functions.

1. Describing distribution and abundance is requisite to exploring interactions between organisms and their environment. Recently, the resource selection function (RSF) has emerged to replace many of the statistical procedures used to quantify resource selection by animals. 2. A RSF is defined by characteristics measured on resource units such that its value for a unit is proportional to the probability of that unit being used by an organism. It is solved using a variety of techniques, particularly the binomial generalized linear model. 3. Observing dynamics in a RSF - obtaining substantially different functions at different times or places for the same species - alerts us to the varying ecological processes that underlie resource selection. 4. We believe that there is a need for us to reacquaint ourselves with ecological theory when interpreting RSF models. We outline a suite of factors likely to govern ecologically based variation in a RSF. In particular, we draw attention to competition and density-dependent habitat selection, the role of predation, longitudinal changes in resource availability and functional responses in resource use. 5. How best to incorporate governing factors in a RSF is currently in a state of development; however, we see promise in the inclusion of random as well as fixed effects in resource selection models, and matched case-control logistic regression. 6. Investigating the basis of ecological dynamics in a RSF will allow us to develop more robust models when applied to forecasting the spatial distribution of animals. It may also further our understanding of the relative importance of ecological interactions on the distribution and abundance of species.

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.

Maternal and individual effects in selection of bed sites and their consequences for fawn survival at different spatial scales.

We examined the relationship between survival of roe deer (Capreolus capreolus) fawns at Trois Fontaines, Champagne-Ardennes, France, and factors related to bed-site selection (predator avoidance and thermoregulation) and maternal food resources (forage availability in the maternal home range). Previous studies have demonstrated that at small scales, the young of large herbivores select bed sites independently from their mothers, although this selection takes place within the limits of their mother's home range. Fawn survival was influenced largely by the availability of good bed sites within the maternal home range, not by the fawn's selection of bed sites; however, selection for thermal cover when selecting bed sites positively influenced survival of young fawns. Typical features of a good home range included close proximity to habitat edges, which is related to forage accessibility for roe deer. The availability of bed sites changed as fawns aged, probably due to an increased mobility of the fawn or a different use of the home range by the mother; sites offering high concealment and thermal protection became less available in favor of areas with higher forage accessibility. Despite the minor influence of bed-site selection on survival, roe deer fawns strongly selected their bed sites according to several environmental factors linked to predator avoidance and thermoregulation. Fawns selected for sites providing concealment, light penetration, and avoided signs of wild boar (Sus scrofa) activity. Avoidance of sites with high light penetration by young fawns positively affected their survival, confirming a negative effect on thermoregulation due to reduced thermal cover. Selection for light penetration by older fawns was less clear. We discuss these results in the context of cross-generational effects in habitat selection across multiple scales, and the potential influence of the 'ghost of predation past'.