Asymmetrical predation intensity produces divergent antipredator behaviours in primary and secondary prey.

It is widely recognized that predators can influence prey through both direct consumption and by inducing costly antipredator behaviours, the latter of which can produce nonconsumptive effects that cascade through trophic systems. Yet, determining how particular prey manage risk in natural settings remains challenging as empirical studies disproportionately focus on single predator-prey dyads. Here, we contrast foraging strategies within the context of a primary and secondary prey to explore how antipredator behaviours emerge as a product of predation intensity as well as the setting in which an encounter takes place. We studied the effects of spotted owls (Strix occidentalis) on two species experiencing asymmetrical risk: dusky-footed woodrats (Neotoma fuscipes; primary prey) and deer mice (Peromyscus spp.; alternative prey). Woodrats are most abundant within young forests, but predominantly captured by owls foraging within mature forests; in contrast, deer mice occur in high densities across forest types and seral stages and are consumed at lower per-capita rates overall. We deployed experimental foraging patches within areas of high and low spotted owl activity, created artificial risky and safe refuge treatments, and monitored behaviour throughout the entirety of prey foraging bouts. Woodrats were more vigilant and foraged less within mature forests and at riskier patches, although the effect of refuge treatment was contingent upon forest type. In contrast, deer mice only demonstrated consistent behavioural responses to riskier refuge treatments; forest type had little effect on perceived risk or the relative importance of refuge treatment. Thus, habitat can interact with predator activity to structure antipredator responses differently for primary versus secondary prey. Our findings show that asymmetrical predation can modulate both the magnitude of perceived risk and the strategies used to manage it, thus highlighting an important and understudied contingency in risk effects research. Evaluating the direct and indirect effects of predation through the paradigm of primary and secondary prey may improve our understanding of how nonconsumptive effects can extend to population- and community-level responses.

Landscape heterogeneity provides co-benefits to predator and prey.

Predator populations are imperiled globally, due in part to changing habitat and trophic interactions. Theoretical and laboratory studies suggest that heterogeneous landscapes containing prey refuges acting as source habitats can benefit both predator and prey populations, although the importance of heterogeneity in natural systems is uncertain. Here, we tested the hypothesis that landscape heterogeneity mediates predator-prey interactions between the California spotted owl (Strix occidentalis occidentalis)-a mature forest species-and one of its principal prey, the dusky-footed woodrat (Neotoma fuscipes)-a younger forest species-to the benefit of both. We did so by combining estimates of woodrat density and survival from live trapping and very high frequency tracking with direct observations of prey deliveries to dependent young by owls in both heterogeneous and homogeneous home ranges. Woodrat abundance was ~2.5 times higher in owl home ranges (14.12 km2 ) featuring greater heterogeneity in vegetation types (1805.0 ± 50.2 SE) compared to those dominated by mature forest (727.3 ± 51.9 SE), in large part because of high densities in young forests appearing to act as sources promoting woodrat densities in nearby mature forests. Woodrat mortality rates were low across vegetation types and did not differ between heterogeneous and homogeneous home ranges, yet all observed predation by owls occurred within mature forests, suggesting young forests may act as woodrat refuges. Owls exhibited a type 1 functional response, consuming ~2.5 times more woodrats in heterogeneous (31.1/month ± 5.2 SE) versus homogeneous (12.7/month ± 3.7 SE) home ranges. While consumption of smaller-bodied alternative prey partially compensated for lower woodrat consumption in homogeneous home ranges, owls nevertheless consumed 30% more biomass in heterogeneous home ranges-approximately equivalent to the energetic needs of producing one additional offspring. Thus, a mosaic of vegetation types including young forest patches increased woodrat abundance and availability that, in turn, provided energetic and potentially reproductive benefits to mature forest-associated spotted owls. More broadly, our findings provide strong empirical evidence that heterogeneous landscapes containing prey refuges can benefit both predator and prey populations. As anthropogenic activities continue to homogenize landscapes globally, promoting heterogeneous systems with prey refuges may benefit imperiled predators.

Spatial variation in diet-microbe associations across populations of a generalist North American carnivore.

Generalist species, by definition, exhibit variation in niche attributes that promote survival in changing environments. Increasingly, phenotypes previously associated with a species, particularly those with wide or expanding ranges, are dissolving and compelling greater emphasis on population-level characteristics. In the present study, we assessed spatial variation in diet characteristics, gut microbiome and associations between these two ecological traits across populations of coyotes Canis latrans. We highlight the influence of the carnivore community in shaping these relationships, as the coyote varied from being an apex predator to a subordinate, mesopredator across sampled populations. We implemented a scat survey across three distinct coyote populations in Michigan, USA. We used carbon (δ13 C) and nitrogen (δ15 N) isotopic values to reflect consumption patterns and trophic level, respectively. Corresponding samples were also paired with 16S rRNA sequencing to describe the microbial community and correlate with isotopic values. Although consumption patterns were comparable, we found spatial variation in trophic level among coyote populations. Specifically, δ15 N was highest where coyotes were the apex predator and lowest where coyotes co-occurred with grey wolves Canis lupus. The gut microbial community exhibited marked spatial variation across populations with the lowest operational taxonomic units diversity found where coyotes occurred at their lowest trophic level. Bacteriodes and Fusobacterium dominated the microbiome and were positively correlated across all populations. We found no correlation between δ13 C and microbial community attributes. However, positive associations between δ15 N and specific microbial genera increased as coyotes ascended trophic levels. Coyotes provide a model for exploring implications of niche plasticity because they are a highly adaptable, wide-ranging omnivore. As coyotes continue to vary in trophic position and expand their geographic range, we might expect increased divergence within their microbial community, changes in physiology and alterations in behaviour.