Rayleigh step-selection functions and connections to continuous-time mechanistic movement models.

BACKGROUND: The process known as ecological diffusion emerges from a first principles view of animal movement, but ecological diffusion and other partial differential equation models can be difficult to fit to data. Step-selection functions (SSFs), on the other hand, have emerged as powerful practical tools for ecologists studying the movement and habitat selection of animals. METHODS: SSFs typically involve comparing resources between a set of used and available points at each step in a sequence of observed positions. We use change of variables to show that ecological diffusion implies certain distributions for available steps that are more flexible than others commonly used. We then demonstrate advantages of these distributions with SSF models fit to data collected for a mountain lion in Colorado, USA. RESULTS: We show that connections between ecological diffusion and SSFs imply a Rayleigh step-length distribution and uniform turning angle distribution, which can accommodate data collected at irregular time intervals. The results of fitting an SSF model with these distributions compared to a set of commonly used distributions revealed how precision and inference can vary between the two approaches. CONCLUSIONS: Our new continuous-time step-length distribution can be integrated into various forms of SSFs, making them applicable to data sets with irregular time intervals between successive animal locations.

Demographic partitioning of dynamic energy subsidies revealed with an Ornstein-Uhlenbeck space use model.

In populations across many taxa, a large fraction of sexually mature individuals do not breed but are attempting to enter the breeding population. Such individuals, often referred to as "floaters," can play critical roles in the dynamics and stability of these populations and buffer them through periods of high adult mortality. Floaters are difficult to study, however, so we lack data needed to understand their roles in the population ecology and conservation status of many species. Here, we analyzed satellite telemetry data with a newly developed mechanistic space use model based on an Ornstein-Uhlenbeck process to help overcome the paucity of data in studying the differential habitat selection and space use of floater and territorial golden eagles Aquila chrysaetos. Our sample consisted of 49 individuals tracked over complete breeding seasons across 4 years, totaling 104 eagle breeding seasons. Modeling these data mechanistically was required to disentangle key differences in movement and particularly to separate aspects of movement driven by resource selection from those driven by use of a central place. We found that floaters generally had more expansive space use patterns and larger home ranges, as well as evidence that they partition space with territorial individuals seemingly on fine scales through differential habitat and resource selection. Floater and territorial eagle home ranges overlapped markedly, suggesting that floaters use the interstices between territories. Furthermore, floater and territorial eagles differed in how they selected for uplift variables, key components of soaring birds' energy landscape, with territorial eagles apparently better able to find and use thermal uplift. We also found relatively low individual heterogeneity in resource selection, especially among territorial individuals, suggesting a narrow realized niche for breeding individuals, which varied from the level of among-individual variation present during migration. This work furthers our understanding of floaters' potential roles in the population ecology of territorial species and suggests that conserving landscapes occupied by territorial eagles also protects floaters.

Recursive Bayesian computation facilitates adaptive optimal design in ecological studies.

Optimal design procedures provide a framework to leverage the learning generated by ecological models to flexibly and efficiently deploy future monitoring efforts. At the same time, Bayesian hierarchical models have become widespread in ecology and offer a rich set of tools for ecological learning and inference. However, coupling these methods with an optimal design framework can become computationally intractable. Recursive Bayesian computation offers a way to substantially reduce this computational burden, making optimal design accessible for modern Bayesian ecological models. We demonstrate the application of so-called prior-proposal recursive Bayes to optimal design using a simulated data binary regression and the real-world example of monitoring and modeling sea otters in Glacier Bay, Alaska. These examples highlight the computational gains offered by recursive Bayesian methods and the tighter fusion of monitoring and science that those computational gains enable.

Mechanistic movement models identify continuously updated autumn migration cues in Arctic caribou.

BACKGROUND: Migrations in temperate systems typically have two migratory phases, spring and autumn, and many migratory ungulates track the pulse of spring vegetation growth during a synchronized spring migration. In contrast, autumn migrations are generally less synchronous and the cues driving them remain understudied. Our goal was to identify the cues that migrants use in deciding when to initiate migration and how this is updated while en route. METHODS: We analyzed autumn migrations of Arctic barren-ground caribou (Rangifer tarandus) as a series of persistent and directional movements and assessed the influence of a suite of environmental factors. We fitted a dynamic-parameter movement model at the individual-level and estimated annual population-level parameters for weather covariates on 389 individual-seasons across 9 years. RESULTS: Our results revealed strong, consistent effects of decreasing temperature and increasing snow depth on migratory movements, indicating that caribou continuously update their migratory decision based on dynamic environmental conditions. This suggests that individuals pace migration along gradients of these environmental variables. Whereas temperature and snow appeared to be the most consistent cues for migration, we also found interannual variability in the effect of wind, NDVI, and barometric pressure. The dispersed distribution of individuals in autumn resulted in diverse environmental conditions experienced by individual caribou and thus pronounced variability in migratory patterns. CONCLUSIONS: By analyzing autumn migration as a continuous process across the entire migration period, we found that caribou migration was largely related to temperature and snow conditions experienced throughout the journey. This mechanism of pacing autumn migration based on indicators of the approaching winter is analogous to the more widely researched mechanism of spring migration, when many migrants pace migration with a resource wave. Such a similarity in mechanisms highlights the different environmental stimuli to which migrants have adapted their movements throughout their annual cycle. These insights have implications for how long-distance migratory patterns may change as the Arctic climate continues to warm.

Diffusion modeling reveals effects of multiple release sites and human activity on a recolonizing apex predator.

BACKGROUND: Reintroducing predators is a promising conservation tool to help remedy human-caused ecosystem changes. However, the growth and spread of a reintroduced population is a spatiotemporal process that is driven by a suite of factors, such as habitat change, human activity, and prey availability. Sea otters (Enhydra lutris) are apex predators of nearshore marine ecosystems that had declined nearly to extinction across much of their range by the early 20th century. In Southeast Alaska, which is comprised of a diverse matrix of nearshore habitat and managed areas, reintroduction of 413 individuals in the late 1960s initiated the growth and spread of a population that now exceeds 25,000. METHODS: Periodic aerial surveys in the region provide a time series of spatially-explicit data to investigate factors influencing this successful and ongoing recovery. We integrated an ecological diffusion model that accounted for spatially-variable motility and density-dependent population growth, as well as multiple population epicenters, into a Bayesian hierarchical framework to help understand the factors influencing the success of this recovery. RESULTS: Our results indicated that sea otters exhibited higher residence time as well as greater equilibrium abundance in Glacier Bay, a protected area, and in areas where there is limited or no commercial fishing. Asymptotic spread rates suggested sea otters colonized Southeast Alaska at rates of 1-8 km/yr with lower rates occurring in areas correlated with higher residence time, which primarily included areas near shore and closed to commercial fishing. Further, we found that the intrinsic growth rate of sea otters may be higher than previous estimates suggested. CONCLUSIONS: This study shows how predator recolonization can occur from multiple population epicenters. Additionally, our results suggest spatial heterogeneity in the physical environment as well as human activity and management can influence recolonization processes, both in terms of movement (or motility) and density dependence.

Novel step selection analyses on energy landscapes reveal how linear features alter migrations of soaring birds.

Human modification of landscapes includes extensive addition of linear features, such as roads and transmission lines. These can alter animal movement and space use and affect the intensity of interactions among species, including predation and competition. Effects of linear features on animal movement have seen relatively little research in avian systems, despite ample evidence of their effects in mammalian systems and that some types of linear features, including both roads and transmission lines, are substantial sources of mortality. Here, we used satellite telemetry combined with step selection functions designed to explicitly incorporate the energy landscape (el-SSFs) to investigate the effects of linear features and habitat on movements and space use of a large soaring bird, the golden eagle Aquila chrysaetos, during migration. Our sample consisted of 32 adult eagles tracked for 45 spring and 39 fall migrations from 2014 to 2017. Fitted el-SSFs indicated eagles had a strong general preference for south-facing slopes, where thermal uplift develops predictably, and that these areas are likely important aspects of migratory pathways. el-SSFs also provided evidence that roads and railroads affected movement during both spring and fall migrations, but eagles selected areas near roads to a greater degree in spring compared to fall and at higher latitudes compared to lower latitudes. During spring, time spent near linear features often occurred during slower-paced or stopover movements, perhaps in part to access carrion produced by vehicle collisions. Regardless of the behavioural mechanism of selection, use of these features could expose eagles and other soaring species to elevated risk via collision with vehicles and/or transmission lines. Linear features have previously been documented to affect the ecology of terrestrial species (e.g. large mammals) by modifying individuals' movement patterns; our work shows that these effects on movement extend to avian taxa.

Trophic redundancy and predator size class structure drive differences in kelp forest ecosystem dynamics.

Ecosystems are changing at alarming rates because of climate change and a wide variety of other anthropogenic stressors. These stressors have the potential to cause phase shifts to less productive ecosystems. A major challenge for ecologists is to identify ecosystem attributes that enhance resilience and can buffer systems from shifts to less desirable alternative states. In this study, we used the Northern Channel Islands, California, as a model kelp forest ecosystem that had been perturbed from the loss of an important sea star predator due to a sea star wasting disease. To determine the mechanisms that prevent phase shifts from productive kelp forests to less productive urchin barrens, we compared pre- and postdisease predator assemblages as predictors of purple urchin densities. We found that prior to the onset of the disease outbreak, the sunflower sea star exerted strong predation pressures and was able to suppress purple urchin populations effectively. After the disease outbreak, which functionally extirpated the sunflower star, we found that the ecosystem response-urchin and algal abundances-depended on the abundance and/or size of remaining predator species. Inside Marine Protected Areas (MPAs), the large numbers and sizes of other urchin predators suppressed purple urchin populations resulting in kelp and understory algal growth. Outside of the MPAs, where these alternative urchin predators are fished, less abundant, and smaller, urchin populations grew dramatically in the absence of sunflower stars resulting in less kelp at these locations. Our results demonstrate that protected trophic redundancy inside MPAs creates a net of stability that could limit kelp forest ecosystem phase shifts to less desirable, alternative states when perturbed. This highlights the importance of harboring diversity and managing predator guilds.

Local meteorological conditions reroute a migration.

For migrating animals, realized migration routes and timing emerge from hundreds or thousands of movement decisions made along migration routes. Local weather conditions along migration routes continually influence these decisions, and even relatively small changes in en route weather may cumulatively result in major shifts in migration patterns. Here, we analysed satellite tracking data to score a discrete navigation decision by a large migratory bird as it navigated a high-latitude, 5000 m elevation mountain range to understand how those navigational decisions changed under different weather conditions. We showed that wind conditions in particular areas along the migration pathway drove a navigational decision to reroute a migration; conditions encountered predictably resulted in migrants routing either north or south of the mountain range. With abiotic conditions continuing to change globally, simple decisions, such as the one described here, might additively emerge into new, very different migration routes.