Coursing the mottled mosaic: Generalist predators track pulses in availability of neonatal ungulates.

The density and distribution of resources shape animal movement and behavior and have direct implications for population dynamics. Resource availability often is "pulsed" in space and time, and individuals should cue in on resource pulses when the energetic gain of doing so exceeds that of stable resources. Birth pulses of prey represent a profitable but ephemeral resource and should thereby result in shifting functional responses by predators. We evaluated movements and resource selection of coyotes (Canis latrans) across a gradient of reproductive stages ranging from late gestation to peak lactation of female mule deer (Odocoileus hemionus) in southwest Wyoming, USA, to test whether coyotes exhibited shifts in selection and movement behavior relative to the availability and vulnerability of neonatal mule deer. We expected coyotes to track pulses in availability of neonatal mule deer, and such behavior would be represented by shifts in resource selection and search behavior of coyotes that would be strongest during peak parturition of mule deer. Coyotes selected areas of high relative probability of use by female mule deer and did so most strongly during peak parturition. Furthermore, searching behavior of coyotes intensified during pulses of availability of deer neonates. Our findings support the notion that coyotes exploit pulses of neonatal deer, presumably as an attempt to capitalize on a vulnerable, energy-rich resource. Our work quantifies the behavioral mechanisms by which coyotes consume ungulate neonates and provides one of the first examples of a mammalian predator-prey system centered on a pulsed resource.

Risky business: How an herbivore navigates spatiotemporal aspects of risk from competitors and predators.

Understanding factors that influence animal behavior is central to ecology. Basic principles of animal ecology imply that individuals should seek to maximize survival and reproduction, which means carefully weighing risk against reward. Decisions become increasingly complex and constrained, however, when risk is spatiotemporally variable. We advance a growing body of work in predator-prey behavior by evaluating novel questions where a prey species is confronted with multiple predators and a potential competitor. We tested how fine-scale behavior of female mule deer (Odocoileus hemionus) during the reproductive season shifted depending upon spatial and temporal variation in risk from predators and a potential competitor. We expected female deer to avoid areas of high risk when movement activity of predators and a competitor were high. We used GPS data collected from 76 adult female mule deer, 35 adult female elk, 33 adult coyotes, and six adult mountain lions. Counter to our expectations, female deer exhibited selection for multiple risk factors, however, selection for risk was dampened by the exposure to risk within home ranges of female deer, producing a functional response in habitat selection. Furthermore, temporal variation in movement activity of predators and elk across the diel cycle did not result in a shift in movement activity by female deer. Instead, the average level of risk within their home range was the predominant factor modulating the response to risk by female deer. Our results counter prevailing hypotheses of how large herbivores navigate risky landscapes and emphasize the importance of accounting for the local environment when identifying effects of risk on animal behavior. Moreover, our findings highlight additional behavioral mechanisms used by large herbivores to mitigate multiple sources of predation and potential competitive interactions.

Migration corridors of adult Golden Eagles originating in northwestern North America.

There has been increasing concern for Golden Eagle (Aquila chrysaetos) populations in North America due to current and future projections of mortality risk and habitat loss from anthropogenic sources. Identification of high-use movement corridors and bottlenecks for the migratory portion of the eagle population in western North America is an important first step to help habitat conservation and management efforts to reduce the risk of eagle mortality. We used dynamic Brownian Bridge movement models to estimate utilization distributions of adult eagles migrating across the western North America and identified high-use areas by calculating the overlap of individuals on population and regional levels. On a population level, the Rocky Mountain Front from east-central British Columbia to central Montana and southwestern Yukon encompassed the most used migration corridors with our study extent for both spring and fall. Regional analysis on a 100 x 200 km scale revealed additional moderate and high-level use corridors in the central British Columbia plateaus. Eagles were more dispersed in the spring until their routes converged in southern Alberta. High-use fall corridors extended farther south into central Wyoming. Knowledge of these high-use areas can aid in conservation and site planning to help maintain and enhance migratory Golden Eagle populations in western North America.

Functional attributes of ungulate migration: landscape features facilitate movement and access to forage.

Long-distance migration by terrestrial mammals is a phenomenon critical to the persistence of populations, but such migrations are declining globally because of over-harvest, habitat loss, and movement barriers. Increasingly, there is a need to improve existing routes, mitigate route segments affected by anthropogenic disturbance, and in some instances, determine whether alternative routes are available. Using a hypothesis-driven approach, we identified landscape features associated with the primary functional attributes, stopovers and movement corridors, of spring migratory routes for mule deer in two study areas using resource selection functions. Patterns of selection for landscape attributes of movement corridors and stopovers mostly were similar; however, landscape features associated with movement corridors aligned better with areas that facilitated movement, whereas selection of stopovers was consistent with sites offering early access to spring forage. For movement corridors, deer selected for dry sites, low elevation, and low anthropogenic disturbance. For stopovers, deer selected for dry sites, with consistently early green-up across years, south-southwesterly aspects, low elevation, and low anthropogenic disturbance. Stopovers and movement corridors of a migratory route presumably promote different functions, but for a terrestrial migrant, patterns of habitat selection indicate that the same general habitat attributes may facilitate both movement and foraging in spring. Our findings emphasize the roles of topographical wetness, vegetation phenology, and anthropogenic disturbance in shaping use of the landscape during migration for this large herbivore. Avoiding human disturbance and tracking ephemeral forage resources appear to be a consistent pattern during migration, which reinforces the notion that movement during migration has a nutritional underpinning and disturbance potentially alters the net benefits of migration.

Large herbivores surf waves of green-up during spring.

The green wave hypothesis (GWH) states that migrating animals should track or 'surf' high-quality forage at the leading edge of spring green-up. To index such high-quality forage, recent work proposed the instantaneous rate of green-up (IRG), i.e. rate of change in the normalized difference vegetation index over time. Despite this important advancement, no study has tested the assumption that herbivores select habitat patches at peak IRG. We evaluated this assumption using step selection functions parametrized with movement data during the green-up period from two populations each of bighorn sheep, mule deer, elk, moose and bison, totalling 463 individuals monitored 1-3 years from 2004 to 2014. Accounting for variables that typically influence habitat selection for each species, we found seven of 10 populations selected patches exhibiting high IRG-supporting the GWH. Nonetheless, large herbivores selected for the leading edge, trailing edge and crest of the IRG wave, indicating that other mechanisms (e.g. ruminant physiology) or measurement error inherent with satellite data affect selection for IRG. Our evaluation indicates that IRG is a useful tool for linking herbivore movement with plant phenology, paving the way for significant advancements in understanding how animals track resource quality that varies both spatially and temporally.

Resource waves: phenological diversity enhances foraging opportunities for mobile consumers.

Time can be a limiting constraint for consumers, particularly when resource phenology mediates foraging opportunity. Though a large body of research has explored how resource phenology influences trophic interactions, this work has focused on the topics of trophic mismatch or predator swamping, which typically occur over short periods, at small spatial extents or coarse resolutions. In contrast many consumers integrate across landscape heterogeneity in resource phenology, moving to track ephemeral food sources that propagate across space as resource waves. Here we provide a conceptual framework to advance the study of phenological diversity and resource waves. We define resource waves, review evidence of their importance in recent case studies, and demonstrate their broader ecological significance with a simulation model. We found that consumers ranging from fig wasps (Chalcidoidea) to grizzly bears (Ursus arctos) exploit resource waves, integrating across phenological diversity to make resource aggregates available for much longer than their component parts. In model simulations, phenological diversity was often more important to consumer energy gain than resource abundance per se. Current ecosystem-based management assumes that species abundance mediates the strength of trophic interactions. Our results challenge this assumption and highlight new opportunities for conservation and management. Resource waves are an emergent property of consumer-resource interactions and are broadly significant in ecology and conservation.