The evolution of white-tailed jackrabbit camouflage in response to past and future seasonal climates.

The genetic basis of adaptive traits has rarely been used to predict future vulnerability of populations to climate change. We show that light versus dark seasonal pelage in white-tailed jackrabbits (Lepus townsendii) tracks snow cover and is primarily determined by genetic variation at endothelin receptor type B (EDNRB), corin serine peptidase (CORIN), and agouti signaling protein (ASIP). Winter color variation was associated with deeply divergent alleles at these genes, reflecting selection on both ancestral and introgressed variation. Forecasted reductions in snow cover are likely to induce widespread camouflage mismatch. However, simulated populations with variation for darker winter pelage are predicted to adapt rapidly, providing a trait-based genetic framework to facilitate evolutionary rescue. These discoveries demonstrate how the genetic basis of climate change adaptation can inform conservation.

Contrasting seasonal effects of climate change influence density in a cold-adapted species.

Many ecological processes are profoundly influenced by abiotic factors, such as temperature and snow. However, despite strong evidence linking shifts in these ecological processes to corresponding shifts in abiotic factors driven by climate change, the mechanisms connecting population size to season-specific climate drivers are little understood. Using a 21-year dataset and a Bayesian state space model, we identified biologically informed seasonal climate covariates that influenced densities of snowshoe hares (Lepus americanus), a cold-adapted boreal herbivore. We found that snow and temperature had strong but conflicting season-dependent effects. Reduced snow duration in spring and fall and warmer summers were associated with lowered hare density, whereas warmer winters were associated with increased density. When modeled simultaneously and under two climate change scenarios, the negative effects of reduced fall and spring snow duration and warmer summers overwhelm the positive effect of warmer winters, producing projected population declines. Ultimately, the contrasting population-level impacts of climate change across seasons emphasize the critical need to examine the entire annual climate cycle to understand potential long-term population consequences of climate change.

Snow-mediated plasticity does not prevent camouflage mismatch.

Global reduction in snow cover duration is one of the most consistent and widespread climate change outcomes. Declining snow duration has severe negative consequences for diverse taxa including seasonally color molting species, which rely on snow for camouflage. However, phenotypic plasticity may facilitate adaptation to reduced snow duration. Plastic responses could occur in the color molt phenology or through behavior that minimizes coat color mismatch or its consequences. We quantified molt phenology of 200 wild snowshoe hares (Lepus americanus), and measured microhabitat choice and local snow cover. Similar to other studies, we found that hares did not show behavioral plasticity to minimize coat color mismatch via background matching; instead they preferred colder, snow free areas regardless of their coat color. Furthermore, hares did not behaviorally mitigate the negative consequences of mismatch by choosing resting sites with denser vegetation cover when mismatched. Importantly, we demonstrated plasticity in the initiation and the rate of the molt and established the direct effect of snow on molt phenology; greater snow cover was associated with whiter hares and this association was not due to whiter hares preferring snowier areas. However, despite the observed snow-mediated plasticity in molt phenology, camouflage mismatch with white hares on brown snowless ground persisted and was more frequent during early snowmelt. Thus, we find no evidence that phenotypic plasticity in snowshoe hares is sufficient to facilitate adaptive rescue to camouflage mismatch under climate change.

Winter color polymorphisms identify global hot spots for evolutionary rescue from climate change.

Maintenance of biodiversity in a rapidly changing climate will depend on the efficacy of evolutionary rescue, whereby population declines due to abrupt environmental change are reversed by shifts in genetically driven adaptive traits. However, a lack of traits known to be under direct selection by anthropogenic climate change has limited the incorporation of evolutionary processes into global conservation efforts. In 21 vertebrate species, some individuals undergo a seasonal color molt from summer brown to winter white as camouflage against snow, whereas other individuals remain brown. Seasonal snow duration is decreasing globally, and fitness is lower for winter white animals on snowless backgrounds. Based on 2713 georeferenced samples of known winter coat color-from eight species across trophic levels-we identify environmentally driven clinal gradients in winter coat color, including polymorphic zones where winter brown and white morphs co-occur. These polymorphic zones, underrepresented by existing global protected area networks, indicate hot spots for evolutionary rescue in a changing climate.

Within-sample variation in snowshoe hare faecal glucocorticoid metabolite measurements.

Faecal glucocorticoid metabolite (FGM) concentrations are used increasingly as a non-invasive measure to index physiological stress experienced by diverse taxa. However, FGM may not be evenly distributed throughout a faecal mass or faecal pellet group. Moreover, within-sample variation in FGM measurements associated with different sampling and/or processing techniques is rarely reported despite potentially having important implications for inferring stress levels in free-ranging wildlife. Using a captive collection of snowshoe hares (Lepus americanus), we (i) assessed repeatability of FGM measurements (i.e. precision) from two processing techniques (measurements derived from dividing whole pellet groups into equal proportions prior to processing [G1], measurements from subsamples derived from thoroughly homogenized whole pellet groups [G2]) and (ii) conducted a power analysis to estimate sample-size requirements for detecting statistically significant differences in FGM concentrations at a population level. Our results indicate that the mean percent coefficient of variation (%CV) for within-sample FGM variation was slightly higher for G1 (%CV = 35, range 13.45-65.37) than for G2 (%CV = 23, range 7.26-47.94), though not statistically significant (two sample t-test, n = 8, t = 1.57, P = 0.16). Thus, FGM is relatively evenly distributed within snowshoe hare faecal pellet groups. However, subsampling from homogenized whole pellet groups may be more appropriate when the sampling time frame is less controlled (e.g. multiple defecation events) because a subsample derived from a homogenized whole pellet group might be more representative of the animal's 'average' physiological state compared to FGM concentrations derived from a few haphazardly selected faecal pellets. Power analysis results demonstrated the importance of a priori consideration of sample sizes. Relatively small effect sizes (e.g. <20%) may require sampling that is logistically and/or cost prohibitive. Yet for many situations of ecological or conservation interest, treatment effects may be substantial (>25%) and thus moderate sample sizes may be sufficient for testing research hypotheses regarding changes FGM concentrations.