Density-dependent habitat selection alters drivers of population distribution in northern Yellowstone elk.

Although it is well established that density dependence drives changes in organismal abundance over time, relatively little is known about how density dependence affects variation in abundance over space. We tested the hypothesis that spatial trade-offs between food and safety can change the drivers of population distribution, caused by opposing patterns of density-dependent habitat selection (DDHS) that are predicted by the multidimensional ideal free distribution. We addressed this using winter aerial survey data of northern Yellowstone elk (Cervus canadensis) spanning four decades. Supporting our hypothesis, we found positive DDHS for food (herbaceous biomass) and negative DDHS for safety (openness and roughness), such that the primary driver of habitat selection switched from food to safety as elk density decreased from 9.3 to 2.0 elk/km2 . Our results demonstrate how population density can drive landscape-level shifts in population distribution, confounding habitat selection inference and prediction and potentially affecting community-level interactions.

A 'How to' guide for interpreting parameters in habitat-selection analyses.

Habitat-selection analyses allow researchers to link animals to their environment via habitat-selection or step-selection functions, and are commonly used to address questions related to wildlife management and conservation efforts. Habitat-selection analyses that incorporate movement characteristics, referred to as integrated step-selection analyses, are particularly appealing because they allow modelling of both movement and habitat-selection processes. Despite their popularity, many users struggle with interpreting parameters in habitat-selection and step-selection functions. Integrated step-selection analyses also require several additional steps to translate model parameters into a full-fledged movement model, and the mathematics supporting this approach can be challenging for many to understand. Using simple examples, we demonstrate how weighted distribution theory and the inhomogeneous Poisson point process can facilitate parameter interpretation in habitat-selection analyses. Furthermore, we provide a 'how to' guide illustrating the steps required to implement integrated step-selection analyses using the amt package By providing clear examples with open-source code, we hope to make habitat-selection analyses more understandable and accessible to end users.

Habitat selection patterns are density dependent under the ideal free distribution.

Despite being widely used, habitat selection models are rarely reliable and informative when applied across different ecosystems or over time. One possible explanation is that habitat selection is context-dependent due to variation in consumer density and/or resource availability. The goal of this paper is to provide a general theoretical perspective on the contributory mechanisms of consumer and resource density-dependent habitat selection, as well as on our capacity to account for their effects. Towards this goal we revisit the ideal free distribution (IFD), where consumers are assumed to be omniscient, equally competitive and freely moving, and are hence expected to instantaneously distribute themselves across a heterogeneous landscape such that fitness is equalised across the population. Although these assumptions are clearly unrealistic to some degree, the simplicity of the structure in IFD provides a useful theoretical vantage point to help clarify our understanding of more complex spatial processes. Of equal importance, IFD assumptions are compatible with the assumptions underlying common habitat selection models. Here we show how a fitness-maximising space use model, based on IFD, gives rise to resource and consumer density-dependent shifts in consumer distribution, providing a mechanistic explanation for the context-dependent outcomes often reported in habitat selection analysis. Our model suggests that adaptive shifts in consumer distribution patterns would be expected to lead to nonlinear and often non-monotonic patterns of habitat selection. These results indicate that even under the simplest of assumptions about adaptive organismal behaviour, habitat selection strength should critically depend on system-wide characteristics. Clarifying the impact of adaptive behavioural responses may be pivotal in making meaningful ecological inferences about observed patterns of habitat selection and allow reliable transferability of habitat selection predictions across time and space.

Corridors or risk? Movement along, and use of, linear features varies predictably among large mammal predator and prey species.

Space-use behaviour reflects trade-offs in meeting ecological needs and can have consequences for individual survival and population demographics. The mechanisms underlying space use can be understood by simultaneously evaluating habitat selection and movement patterns, and fine-resolution locational data are increasing our ability to do so. We use high-resolution location data and an integrated step-selection analysis to evaluate caribou, moose, bear, and wolf habitat selection and movement behaviour in response to anthropogenic habitat modification, though caribou data were limited. Space-use response to anthropogenic linear features (LFs) by predators and prey is hypothesized to increase predator hunting efficiency and is thus believed to be a leading factor in woodland caribou declines in western Canada. We found that all species moved faster while on LFs. Wolves and bears were also attracted towards LFs, whereas prey species avoided them. Predators and prey responded less strongly and consistently to natural features such as streams, rivers and lakeshores. These findings are consistent with the hypothesis that LFs facilitate predator movement and increase hunting efficiency, while prey perceive such features as risky. Understanding the behavioural mechanisms underlying space-use patterns is important in understanding how future land-use may impact predator-prey interactions. Explicitly linking behaviour to fitness and demography will be important to fully understand the implications of management strategies.

Daphnia inhibits the emergence of spatial pattern in a simple consumer-resource system.

Spatial self-organization can occur in many ecosystems with important effects on food web dynamics and the maintenance of biodiversity. The consumer-resource interaction is known to generate spatial patterning, but only a few empirical studies have investigated the effect of the consumer on resource distribution. Here we report results from a large aquatic mesocosm experiment used to investigate the effect of the consumer Daphnia magna on the distribution of its resource, the green algae Chlorella vulgaris. We maintained large tanks with capacity for 26 ,000 L with either algae or both algae and Daphnia in different temperature conditions. We found that the presence of D. magna inhibited spatial structure in algal distribution that arose as a consequence of increasing temperature. We conjecture that this homogenization effect might be caused by a combination of high mobility combined with high rates of algal consumption by Daphnia. Our study emphasizes the importance of both local constraints on growth and behavioral responses in either promoting or suppressing spatial self-organization in natural populations.

Space-use behaviour of woodland caribou based on a cognitive movement model.

Movement patterns offer a rich source of information on animal behaviour and the ecological significance of landscape attributes. This is especially useful for species occupying remote landscapes where direct behavioural observations are limited. In this study, we fit a mechanistic model of animal cognition and movement to GPS positional data of woodland caribou (Rangifer tarandus caribou; Gmelin 1788) collected over a wide range of ecological conditions. The model explicitly tracks individual animal informational state over space and time, with resulting parameter estimates that have direct cognitive and ecological meaning. Three biotic landscape attributes were hypothesized to motivate caribou movement: forage abundance (dietary digestible biomass), wolf (Canis lupus; Linnaeus, 1758) density and moose (Alces alces; Linnaeus, 1758) habitat. Wolves are the main predator of caribou in this system and moose are their primary prey. Resulting parameter estimates clearly indicated that forage abundance is an important driver of caribou movement patterns, with predator and moose avoidance often having a strong effect, but not for all individuals. From the cognitive perspective, our results support the notion that caribou rely on limited sensory inputs from their surroundings, as well as on long-term spatial memory, to make informed movement decisions. Our study demonstrates how sensory, memory and motion capacities may interact with ecological fitness covariates to influence movement decisions by free-ranging animals.

Wolves adapt territory size, not pack size to local habitat quality.

1. Although local variation in territorial predator density is often correlated with habitat quality, the causal mechanism underlying this frequently observed association is poorly understood and could stem from facultative adjustment in either group size or territory size. 2. To test between these alternative hypotheses, we used a novel statistical framework to construct a winter population-level utilization distribution for wolves (Canis lupus) in northern Ontario, which we then linked to a suite of environmental variables to determine factors influencing wolf space use. Next, we compared habitat quality metrics emerging from this analysis as well as an independent measure of prey abundance, with pack size and territory size to investigate which hypothesis was most supported by the data. 3. We show that wolf space use patterns were concentrated near deciduous, mixed deciduous/coniferous and disturbed forest stands favoured by moose (Alces alces), the predominant prey species in the diet of wolves in northern Ontario, and in proximity to linear corridors, including shorelines and road networks remaining from commercial forestry activities. 4. We then demonstrate that landscape metrics of wolf habitat quality - projected wolf use, probability of moose occupancy and proportion of preferred land cover classes - were inversely related to territory size but unrelated to pack size. 5. These results suggest that wolves in boreal ecosystems alter territory size, but not pack size, in response to local variation in habitat quality. This could be an adaptive strategy to balance trade-offs between territorial defence costs and energetic gains due to resource acquisition. That pack size was not responsive to habitat quality suggests that variation in group size is influenced by other factors such as intraspecific competition between wolf packs.

Towards an energetic landscape: broad-scale accelerometry in woodland caribou.

Energetic balance is a central driver of individual survival and population change, yet estimating energetic costs in free- and wide-ranging animals presents a significant challenge. Animal-borne activity monitors (using accelerometer technology) present a promising method of meeting this challenge and open new avenues for exploring energetics in natural settings. To determine the behaviours and estimated energetic costs associated with a given activity level, three captive reindeer (Rangifer tarandus tarandus) at the Toronto Zoo were fitted with collars and observed for 53 h. Activity patterns were then measured over 13 months for 131 free-ranging woodland caribou (R. t. caribou) spanning 450,000 km(2) in northern Ontario. The captive study revealed a positive but decelerating relationship between activity level and energetic costs inferred from previous behavioural studies. Field-based measures of activity were modelled against individual displacement, vegetation abundance (Normalized Difference Vegetation Index), snow depth and temperature, and the best fit model included all parameters and explained over half of the variation in the data. Individual displacement was positively related to activity levels, suggesting that broad differences in energetic demands are influenced by variation in movement rates. After accounting for displacement, activity was highest at intermediate levels of vegetation abundance, presumably due to foraging behaviour. Snow depth, probably associated with digging for winter forage, moderately increased activity. Activity levels increased significantly at the coldest winter temperatures, suggesting the use of behavioural thermoregulation by caribou. These interpretations of proximate causal factors should be regarded as hypotheses subject to validation under normal field conditions. These results illustrate the landscape characteristics that increase energetic demands for caribou and confirm the great potential for the use of accelerometry in studies of animal energetics.

Pursuit and escape drive fine-scale movement variation during migration in a temperate alpine ungulate.

Climate change reduces snowpack, advances snowmelt phenology, drives summer warming, alters growing season precipitation regimes, and consequently modifies vegetation phenology in mountain systems. Elevational migrants track spatial variation in seasonal plant growth by moving between ranges at different elevations during spring, so climate-driven vegetation change may disrupt historic benefits of migration. Elevational migrants can furthermore cope with short-term environmental variability by undertaking brief vertical movements to refugia when sudden adverse conditions arise. We uncover drivers of fine-scale vertical movement variation during upland migration in an endangered alpine specialist, Sierra Nevada bighorn sheep (Ovis canadensis sierrae) using a 20-year study of GPS collar data collected from 311 unique individuals. We used integrated step-selection analysis to determine factors that promote vertical movements and drive selection of destinations following vertical movements. Our results reveal that relatively high temperatures consistently drive uphill movements, while precipitation likely drives downhill movements. Furthermore, bighorn select destinations at their peak annual biomass and maximal time since snowmelt. These results indicate that although Sierra Nevada bighorn sheep seek out foraging opportunities related to landscape phenology, they compensate for short-term environmental stressors by undertaking brief up- and downslope vertical movements. Migrants may therefore be impacted by future warming and increased storm frequency or intensity, with shifts in annual migration timing, and fine-scale vertical movement responses to environmental variability.

Spatial memory and animal movement.

Memory is critical to understanding animal movement but has proven challenging to study. Advances in animal tracking technology, theoretical movement models and cognitive sciences have facilitated research in each of these fields, but also created a need for synthetic examination of the linkages between memory and animal movement. Here, we draw together research from several disciplines to understand the relationship between animal memory and movement processes. First, we frame the problem in terms of the characteristics, costs and benefits of memory as outlined in psychology and neuroscience. Next, we provide an overview of the theories and conceptual frameworks that have emerged from behavioural ecology and animal cognition. Third, we turn to movement ecology and summarise recent, rapid developments in the types and quantities of available movement data, and in the statistical measures applicable to such data. Fourth, we discuss the advantages and interrelationships of diverse modelling approaches that have been used to explore the memory-movement interface. Finally, we outline key research challenges for the memory and movement communities, focusing on data needs and mathematical and computational challenges. We conclude with a roadmap for future work in this area, outlining axes along which focused research should yield rapid progress.

Environmental and individual drivers of animal movement patterns across a wide geographical gradient.

Within the rapidly developing field of movement ecology, much attention has been given to studying the movement of individuals within a subset of their population's occupied range. Our understanding of the effects of landscape heterogeneity on animal movement is still fairly limited as it requires studying the movement of multiple individuals across a variety of environmental conditions. Gaining deeper understanding of the environmental drivers of movement is a crucial component of predictive models of population spread and habitat selection and may help inform management and conservation. In Ontario, woodland caribou (Rangifer tarandus caribou) occur along a wide geographical gradient ranging from the boreal forest to the Hudson Bay floodplains. We used high-resolution GPS data, collected from 114 individuals across a 450000 km(2) area in northern Ontario, to link movement behaviour to underlying local environmental variables associated with habitat permeability, predation risk and forage availability. We show that a great deal of observed variability in movement patterns across space and time can be attributed to local environmental conditions, with residual individual differences that may reflect spatial population structure. We discuss our results in the context of current knowledge of movement and caribou ecology and highlight potential applications of our approach to the study of wide-ranging animals.

Optimal prey switching: Predator foraging costs provide a mechanism for functional responses in multi-prey systems.

Foragers must balance the costs and gains inherent in the pursuit of their next meal. Classical functional response formulations describe consumption rates driven by prey density and are naive to predator foraging costs. Here, we integrated foraging costs into functional responses to add mechanism and precision to foundational ideas. Specifically, using a model system with a single predator and two prey, we express a functional response emerging from variable energy and time costs of each predation phase: searching, attacking, or consuming prey. The utility of our model is explored through a focused example where prey can exert variable influence on predator foraging costs through antipredator traits. Dissimilarity between prey in their foraging costs influence the energy gain rate of the predator through optimal prey switching. We found that a small subset of prey antipredator traits and density conditions generated a stabilizing Type III (sigmoidal) functional response-the pattern often thought to typify a generalist predator switching between prey species. The sigmoid functional response occurred for highly profitable prey only when the costly prey (1) were at a high density and (2) their antipredator traits increased energy or time costs following an encounter. We outline testable predictions regarding foraging costs from our model. We provide guidance on how to apply optimal foraging theory to empirical scenarios where predator foraging costs vary due to prey type, predator type, or environmental conditions. Our framework represents a synergy of foundational and contemporary theory across disciplines, facilitating the discovery of shared principles and context-dependent variation across varied predator-prey systems.

Behavioral responses of terrestrial mammals to COVID-19 lockdowns.

COVID-19 lockdowns in early 2020 reduced human mobility, providing an opportunity to disentangle its effects on animals from those of landscape modifications. Using GPS data, we compared movements and road avoidance of 2300 terrestrial mammals (43 species) during the lockdowns to the same period in 2019. Individual responses were variable with no change in average movements or road avoidance behavior, likely due to variable lockdown conditions. However, under strict lockdowns 10-day 95th percentile displacements increased by 73%, suggesting increased landscape permeability. Animals' 1-hour 95th percentile displacements declined by 12% and animals were 36% closer to roads in areas of high human footprint, indicating reduced avoidance during lockdowns. Overall, lockdowns rapidly altered some spatial behaviors, highlighting variable but substantial impacts of human mobility on wildlife worldwide.

Rotifer population spread in relation to food, density and predation risk in an experimental system.

1. Despite the popular use of diffusion models to predict the spatial spread of populations over time, we currently know little about how diffusion rates change with the state of the environment or the internal condition of individuals. To address this gap in our understanding, we measured rates of spread for many populations of the rotifer Brachionus calyciflorus in a suite of well-replicated experiments. 2. In one set of experiments, we manipulated food availability and population density along a continuous range of densities. In a second set, we manipulated the internal state of entire populations via food deprivation and exposure to predator kairomones. 3. Across replicate populations, diffusion rates were positively correlated with conspecific density. Diffusion rates were negatively correlated with food availability, especially when conspecific density was high. Diffusion rates of food-deprived populations or those exposed to predation risk were lower than controls. 4. Our results provide direct experimental evidence that rates of population spread are conditional on population density, food availability, body condition and predation risk.

Resource exploitation efficiency collapses the home range of an apex predator.

Optimizing energy acquisition and expenditure is a fundamental trade-off for consumers, strikingly reflected in how mobile organisms use space. Several studies have established that home range size decreases as resource density increases, but the balance of costs and benefits associated with exploiting a given resource density is unclear. We evaluate how the ability of consumers to exploit their resources through movement (termed "resource exploitation") interacts with resource density to influence home range size. We then contrast two hypotheses to evaluate how resource exploitation influences home range size across a vast gradient of productivity and density of human-created linear features (roads and seismic lines) that are known to facilitate animal movements. Under the Diffusion Facilitation Hypothesis, linear features are predicted to lead to more diffuse space use and larger home ranges. Under the Exploitation Efficiency Hypothesis, linear features are predicted to increase foraging efficiency, resulting in less space being required to meet energetic demands and therefore smaller home ranges. Using GPS telemetry data from 142 wolves (Canis lupus) distributed over more than 500,000 km2 , we found that wolf home range size was influenced by the interaction between resource density and exploitation efficiency. Home range size decreased as linear feature density increased, supporting the Exploitation Efficiency Hypothesis. However, the effect of linear features on home range size diminished in more productive areas, suggesting that exploitation efficiency is of greater importance when resource density is low. These results suggest that smaller home ranges will occur where both linear feature density and primary productivity are higher, thereby increasing regional wolf density.

Linking rates of diffusion and consumption in relation to resources.

The functional response is a fundamental model of the relationship between consumer intake rate and resource abundance. The random walk is a fundamental model of animal movement and is well approximated by simple diffusion. Both models are central to our understanding of numerous ecological processes but are rarely linked in ecological theory. To derive a synthetic model, we draw on the common logical premise underlying these models and show how the diffusion and consumption rates of consumers depend on elementary attributes of naturally occurring consumer-resource interactions: the abundance, spatial aggregation, and traveling speed of resources as well as consumer handling time and directional persistence. We show that resource aggregation may lead to increased consumer diffusion and, in the case of mobile resources, reduced consumption rate. Resource-dependent movement patterns have traditionally been attributed to area-restricted search, reflecting adaptive decision making by the consumer. Our synthesis provides a simple alternative hypothesis that such patterns could also arise as a by-product of statistical movement mechanics.

Food availability modulates temperature-dependent effects on growth, reproduction, and survival in Daphnia magna.

Reduced body size and accelerated life cycle due to warming are considered major ecological responses to climate change with fitness costs at the individual level. Surprisingly, we know little about how relevant ecological factors can alter these life history trade-offs and their consequences for individual fitness. Here, we show that food modulates temperature-dependent effects on body size in the water flea Daphnia magna and interacts with temperature to affect life history parameters. We exposed 412 individuals to a factorial manipulation of food abundance and temperature, tracked each reproductive event, and took daily measurements of body size from each individual. High temperature caused a reduction in maximum body size in both food treatments, but this effect was mediated by food abundance, such that low food conditions resulted in a reduction of 20% in maximum body size, compared with a reduction of 4% under high food conditions. High temperature resulted in an accelerated life cycle, with pronounced fitness cost at low levels of food where only a few individuals produced a clutch. These results suggest that the mechanisms affecting the trade-off between fast growth and final body size are food-dependent, and that the combination of low levels of food and high temperature could potentially threaten viability of ectotherms.

Animal movement tools (amt): R package for managing tracking data and conducting habitat selection analyses.

Advances in tracking technology have led to an exponential increase in animal location data, greatly enhancing our ability to address interesting questions in movement ecology, but also presenting new challenges related to data management and analysis. Step-selection functions (SSFs) are commonly used to link environmental covariates to animal location data collected at fine temporal resolution. SSFs are estimated by comparing observed steps connecting successive animal locations to random steps, using a likelihood equivalent of a Cox proportional hazards model. By using common statistical distributions to model step length and turn angle distributions, and including habitat- and movement-related covariates (functions of distances between points, angular deviations), it is possible to make inference regarding habitat selection and movement processes or to control one process while investigating the other. The fitted model can also be used to estimate utilization distributions and mechanistic home ranges. Here, we present the R package amt (animal movement tools) that allows users to fit SSFs to data and to simulate space use of animals from fitted models. The amt package also provides tools for managing telemetry data. Using fisher (Pekania pennanti) data as a case study, we illustrate a four-step approach to the analysis of animal movement data, consisting of data management, exploratory data analysis, fitting of models, and simulating from fitted models.

Density-dependent space use affects interpretation of camera trap detection rates.

Camera traps (CTs) are an increasingly popular tool for wildlife survey and monitoring. Estimating relative abundance in unmarked species is often done using detection rate as an index of relative abundance, which assumes that detection rate has a positive linear relationship with true abundance. This assumption may be violated if movement behavior varies with density, but the degree to which movement behavior is density-dependent across taxa is unclear. The potential confounding of population-level relative abundance indices by movement would depend on how regularly, and by what magnitude, movement rate and home-range size vary with density. We conducted a systematic review and meta-analysis to quantify relationships between movement rate, home-range size, and density, across terrestrial mammalian taxa. We then simulated animal movements and CT sampling to test the effect of contrasting movement scenarios on CT detection rate indices. Overall, movement rate and home-range size were negatively correlated with density and positively correlated with one another. The strength of the relationships varied significantly between taxa and populations. In simulations, detection rates were related to true abundance but underestimated change, particularly for slower moving species with small home ranges. In situations where animal space use changes markedly with density, we estimate that up to thirty percent of a true change in relative abundance may be missed due to the confounding effect of movement, making trend estimation more difficult. The common assumption that movement remains constant across densities is therefore violated across a wide range of mammal species. When studying unmarked species using CT detection rates, researchers and managers should explicitly consider that such indices of relative abundance reflect both density and movement. Practitioners interpreting changes in camera detection rates should be aware that observed differences may be biased low relative to true changes in abundance. Further information on animal movement, or methods that do not depend on assumptions of density-independent movement, may be required to make robust inferences on population trends.