Decision making in conservation science relies on the best available information. This may include using models that were not designed for purpose and are not accompanied by an assessment of limitations. To begin addressing these issues, we sought to reproduce and evaluate the spatial transferability of the two best available models for predicting impacts of proposed mining on boreal woodland caribou (Rangifer tarandus caribou) in northern Ontario. We evaluated their suitability for projecting the impacts of development in the Ring of Fire region. To aid in accessibility, we developed an R package for data preparation, modeling of resource selection, and demographic modeling. We found models were either ill-suited or lacking for ongoing regional planning. The specificity of the regional resource selection model limited its usefulness for predicting impacts of development, and the high variability across caribou ranges limited the usefulness of a national aspatial demographic model for predicting range-specific impacts. These existing models are not enough to provide spatially explicit information needed to minimize detrimental effects of anthropogenic development on caribou recovery in northern Ontario. Models designed for forecasting that are regularly updated with range-specific demographic and habitat information are required.

Animals behave ‘optimally’ when they minimize their costs while maximizing their energetic gain. Optimal foraging theory predicts that with decreasing resource abundance, animals will increase 1) niche breadth, 2) territory size and movement distance, and 3) time spent at resource patches. We used optimal foraging as a guiding framework to test what drives differences in behaviour of GPS collared wolves in two predator populations. As expected, niche breadth and territory sizes were larger, movement rates were greater and cluster durations were longer in the study area with relatively lower resource abundance Our comparative test through the lens of optimal foraging theory led to some unexpected discoveries. Specifically, differences in these responses were not explained by the density of the large primary prey (moose) but instead anthropogenic disturbance (linear feature density) was a ubiquitous influence on wolf behaviours. Wolves in higher linear feature density responded by reducing their cluster residency and increased step lengths and territory sizes with increases in linear feature density. Our work provides evidence that large carnivores can mediate changes in resources by adjusting their space use and time budgets in a way that corresponds with optimal behaviour - maximizing their energy gain and minimizing costs.

Anthropogenic linear features often alter wildlife behaviour and movement. Landscape features, such as habitat, can have important mediating effects on wildlife response to disturbance and yet are rarely explicitly considered in how habitat and disturbance interact. We tested the movement and space‐use responses of GPS‐collared grey wolves to linear features with respect to adjacent habitat variation. We simultaneously modelled wolf movement and selection within a conditional logistic regression framework (integrated Step Selection Analysis). We explicitly considered how adjacent habitat alters these responses through putative effects, such as movement friction. Classifying linear features based on the selection and movement response of wolves revealed that pairing transmission lines and primary roads increased the avoidance response to be greater than either feature on its own and provided evidence of a semi‐permeable barrier to movement. In contrast, features with reduced human activity, including secondary and tertiary roads, were highly selected for and may function as movement corridors. Synthesis and applications . Explicitly parameterizing adjacent habitat provides evidence that where a linear feature is routed and which habitats it interacts with will have the greatest implications for wolf behavioural responses. Reduced avoidance behaviour in highly risky environments signifies the importance of habitat for maintaining landscape connectivity, particularly when routing multiple different features parallel and near each other. Increased vegetation density along linear features also reduces movement advantages putatively by increasing friction, indicating that actively decommissioning other features, such as secondary roads, could be an effective mitigation strategy for reducing wolf encounters with prey. Knowing the influence of adjacent habitats on the likelihood of wolves selecting for a given linear feature creates context to minimize the impact of new anthropogenic features on behaviour.

Animals behave ‘optimally’ when they minimize their costs while maximizing their energetic gain. Optimal foraging theory predicts that with decreasing resource abundance, animals will increase 1) niche breadth, 2) territory size and movement distance, and 3) time spent at resource patches. We test these predictions by investigating clusters from GPS collared wolves ( Canis lupus ) in two predator populations with marked differences in their prey composition and abundance. As expected, wolves in a less abundant system increase niche breadth, territory size, step lengths, and time spent at each kill. Our work provides evidence of optimal behavior in an apex predator which can support population resilience across changing landscapes. ### Competing Interest Statement The authors have declared no competing interest.

Animals within social groups respond to costs and benefits of sociality by adjusting the proportion of time they spend in close proximity to other individuals in the group (cohesion). Variation in cohesion between individuals, in turn, shapes important group-level processes such as subgroup formation and fission-fusion dynamics. Although critical to animal sociality, a comprehensive understanding of the factors influencing cohesion remains a gap in our knowledge of cooperative behavior in animals. We tracked 574 individuals from six species within the genus Canis in 15 countries on four continents with GPS telemetry to estimate the time that pairs of individuals within social groups spent in close proximity and test hypotheses regarding drivers of cohesion. Pairs of social canids (Canis spp.) varied widely in the proportion of time they spent together (5%-100%) during seasonal monitoring periods relative to both intrinsic characteristics and environmental conditions. The majority of our data came from three species of wolves (gray wolves, eastern wolves, and red wolves) and coyotes. For these species, cohesion within social groups was greatest between breeding pairs and varied seasonally as the nature of cooperative activities changed relative to annual life history patterns. Across species, wolves were more cohesive than coyotes. For wolves, pairs were less cohesive in larger groups, and when suitable, small prey was present reflecting the constraints of food resources and intragroup competition on social associations. Pair cohesion in wolves declined with increased anthropogenic modification of the landscape and greater climatic variability, underscoring challenges for conserving social top predators in a changing world. We show that pairwise cohesion in social groups varies strongly both within and across Canis species, as individuals respond to changing ecological context defined by resources, competition, and anthropogenic disturbance. Our work highlights that cohesion is a highly plastic component of animal sociality that holds significant promise for elucidating ecological and evolutionary mechanisms underlying cooperative behavior.

BACKGROUND:Movement links the distribution of habitats with the social environment of animals using those habitats. Despite the links between movement, habitat selection, and socioecology, their integration remains a challenge due to lack of shared vocabulary across fields, methodological gaps, and the implicit (rather than explicit) historical development of theory in the fields of social and spatial ecology. Given these challenges can be addressed, opportunity for further study will provide insight about the links between social, spatial, and movement ecology. Here, our objective was to disentangle the roles of habitat selection and social association as drivers of movement in caribou (Rangifer tarandus). METHODS:To accomplish our objective, we modelled the relationship between collective movement and selection of foraging habitats using socially informed integrated step selection function (iSSF). Using iSSF, we modelled the effect of social processes, i.e., nearest neighbour distance and social preference, and movement behaviour on patterns of habitat selection. RESULTS:By unifying social network analysis with iSSF, we identified movement-dependent social association, where individuals took shorter steps in lichen habitat and foraged in close proximity to more familiar individuals. CONCLUSIONS:Our study demonstrates that social preference is context-dependent based on habitat selection and foraging behaviour. We therefore surmise that habitat selection and social association are drivers of collective movement, such that movement is the glue between habitat selection and social association. Here, we put these concepts into practice to demonstrate that movement is the glue connecting individual habitat selection to the social environment.

[This corrects the article DOI: 10.1093/cz/zoaa052.].

final release prior to publication, including data

A bstract Hunger is a frequent state for many predators and increasing hunger is likely to motivate costly behaviour to acquire necessary resources. Generalist predators must balance the costs and gains of hunting different prey, including increasing encounter rates and improving success rates by seeking areas with greater prey catchability. Large carnivores face threats when they interact with humans or conspecifics. We use integrated step selection analysis to describe spatiotemporal factors that influence wolf ( Canis lupus ) hunting behavior in Riding Mountain National Park, a natural area that wolves share with moose ( Alces alces ) and elk ( Cervus canadensis ). If hunger generates more risky behavior by wolves, as time-from-kill increases we expect wolves will: (1) search for and kill a prey that poses higher risk of injury, (2) use the periphery of their range, (3) use areas closer to the park boundary. Hunger alters wolf space use and drives a fine scale change in prey tracking. Movement patterns of hungry wolves are indicative of search behavior, i.e., shorter steps and more turning. Contrary to our predictions, hungry wolves moved further into the park. As wolves become hungrier, they switch their response from a weak selection to avoidance of elk. In contrast, the response to the primary and emergent prey, moose varied between individuals with some pack level similarities. Therefore, the state-based response to a pervasive risk and a historical resource was conserved in a population residing in a prey rich ‘island’ interfacing with human disturbance.

Avoiding death affects biological processes, including behavior. Habitat selection, movement, and sociality are highly flexible behaviors that influence the mortality risks and subsequent fitness of individuals. In the Anthropocene, animals are experiencing increased risks from direct human causes and increased spread of infectious diseases. Using integrated step selection analysis, we tested how the habitat selection, movement, and social behaviors of gray wolves vary in the two months prior to death due to humans (being shot or trapped) or canine distemper virus (CDV). We further tested how those behaviors vary as a prelude to death. We studied populations of wolves that occurred under two different management schemes: a national park managed for conservation and a provincially managed multi-use area. Behaviors that changed prior to death were strongly related to how an animal eventually died. Wolves killed by humans moved slower than wolves that survived and selected to be nearer roads closer in time to their death. Wolves that died due to CDV moved progressively slower as they neared death and reduced their avoidance of wet habitats. All animals, regardless of dying or living, maintained selection to be near packmates across time, which seemingly contributed to disease dynamics in the packs infected with CDV. There were no noticeable differences in behavior between the two management areas. Overall, habitat selection, movement, and sociality interact to put individuals and groups at greater risks, influencing their cause-specific mortality.

Consistent individual differences in behavior, commonly termed animal personality, are a widespread phenomenon across taxa that have important consequences for fitness, natural selection, and trophic interactions. Animal personality research may prove useful in several conservation contexts, but which contexts remains to be determined. We conducted a structured literature review of 654 studies identified by combining search terms for animal personality and various conservation subfields. We scored the relevance of personality and conservation issues for each study to identify which studies meaningfully integrated the 2 fields as opposed to surface-level connections or vague allusions. We found a taxonomic bias toward mammals (29% of all studies). Very few amphibian or reptile studies applied personality research to conservation issues (6% each). Climate change (21%), invasive species (15%), and captive breeding and reintroduction (13%) were the most abundant conservation subfields that occurred in our search, though a substantial proportion of these papers weakly integrated conservation and animal personality (climate change 54%, invasive species 51%, captive breeding and reintroduction 40%). Based on our results, we recommend that researchers strive for consistent and broadly applicable terminology when describing consistent behavioral differences to minimize confusion and improve the searchability of research. We identify several gaps in the literature that appear to be promising and fruitful avenues for future research, such as disease transmission as a function of sociability or exploration as a driver of space use in protected areas. Practitioners can begin informing future conservation efforts with knowledge gained from animal personality research.

Environmental impact assessments often rely on best available information, which may include models that were not designed for purpose and are not accompanied by an assessment of limitations. We reproduced available models of boreal woodland caribou resource selection and demography and evaluated their suitability for projecting impacts of development in the Ring of Fire on boreal caribou in the Missisa range (Ontario, Canada). The specificity of the resource selection model limited usefulness for predicting impacts, and high variability in model coefficients among ranges suggests responses vary with habitat availability. The aspatial demographic model projects decreasing survival and recruitment with increasing disturbance, but high variability among populations implies the importance of these impacts depends on population status, and there is no current status estimate. New models that are designed for forecasting, informed by more current herd status information and information from neighbouring ranges, are required to better inform decisions. To demonstrate how open-source tools and reproducible workflows can improve the transparency and reusability of models we developed an R package for data preparation, resource selection, and demographic calculations. Open-source tools, reproducible workflows, and reuseable forecasting models can improve our collective ability to inform wildlife management decisions in a timely manner.### Competing Interest StatementRob Rempel is principal of FERIT Consulting

Movement links the distribution of habitats with the social environment of animals using those habitats; yet integrating movement, habitat selection, and socioecology remains an opportunity for further study. Here, our objective was to disentangle the roles of habitat selection and social association as drivers of collective movement in a gregarious ungulate. To accomplish this objective, we (1) assessed whether socially familiar individuals form discrete social communities and whether social communities have high spatial, but not necessarily temporal, overlap; and (2) we modelled the relationship between collective movement and selection of foraging habitats using socially informed integrated step selection analysis. Based on assignment of individuals to social communities and home range overlap analyses, individuals assorted into discrete social communities, and these communities had high spatial overlap. By unifying social network analysis with movement ecology, we identified movement-dependent social association, where individuals foraged with more familiar individuals, but moved collectively with any between foraging patches. Our study demonstrates that social behaviour and space use are inter-related based on spatial overlap of social communities and movement-dependent habitat selection. Movement, habitat selection, and social behaviour are linked in theory. Here, we put these concepts into practice to demonstrate that movement is the glue connecting individual habitat selection to the social environment.

Avoiding death infects biological processes, including behavior. Habitat selection, movement, and sociality are highly flexible behaviors that influence the mortality risks and subsequent fitness of individuals. In the Anthropocene, animals are experiencing increased risks from direct human causes and increased spread of infectious diseases. Using integrated step selection analysis, we tested how the habitat selection, movement, and social behaviors of grey wolves vary as an individual dies due to humans or canine distemper virus (CDV) and how those behaviors may vary in the lead up to death. Behaviors that changed prior to death were strongly related to how an animal eventually died. Wolves killed by humans moved slower than wolves that survived and selected to be nearer roads closer in time to their death. Wolves that died due to CDV moved progressively slower as they neared death and reduced their avoidance of wet habitats. All animals, regardless of dying or not maintained strong selection to be near packmates across time, which seemingly contributed to disease dynamics in the packs that became infected with CDV. Habitat selection, movement, and sociality interact to put individuals and groups at greater risks, influencing their cause-specific mortality. Lay Summary Not much is known about behaviors prior to death in wild animals. Grey wolves killed by humans selected to be in riskier areas increasingly prior to their deaths. Wolves that died due to disease moved slower and changed their habitat selection to be in areas with more water as they became sicker. Sick wolves also continued to select for packmates, increasing the chances that the whole pack would succumb to the disease. ### Competing Interest Statement The authors have declared no competing interest.

With global biodiversity declines, government regulations protecting wildlife serve a key role in species persistence. Despite its importance, planning for protection can be a slow process, taking up to several decades. Such delays have led to species declines and extinction. Here, we investigate the factors driving time between conservation listing and the creation of a plan to support species recovery. Using Canada's Species at Risk Act as a case study, we used Cox proportional-hazards models to test how quickly a species designatable unit (DU) would receive a recovery plan given the reasons for its designation, conservation status, taxon, sociopolitical climate, the extent of scientific research, and public awareness. AIC model selection revealed that sociopolitical factors best explained how quickly a DU would receive a plan. We found that the time for a DU to receive a plan decreased the more often a majority government was in power, the fewer environmental bills passed, and the lower average GDP growth rate during planning. Our results highlight the need for greater consistency in species recovery planning based on conservation needs and status, regardless of sociopolitical climate. We recommend further examination of the relationship between time for recovery planning and plan effectiveness to elucidate how planning delays impact species recovery.

Movement links the distribution of habitats with the social environment of animals using those habitats; yet integrating movement, habitat selection, and socioecology remains an opportunity for further study. Here, our objective was to disentangle the roles of habitat selection and social association as drivers of collective movement in a gregarious ungulate. To accomplish this objective, we (1) assessed whether socially familiar individuals form discrete social communities and whether social communities have high spatial, but not necessarily temporal, overlap; and (2) we modelled the relationship between collective movement and selection of foraging habitats using socially informed integrated step selection analysis. We used social network analysis to assign individuals to social communities and determine short and long-term social preference among individuals. Using integrated step selection functions (iSSF), we then modelled the effect of social processes, i.e., nearest neighbour distance and social preference, and movement behaviour on patterns of habitat selection. Based on assignment of individuals to social communities and home range overlap analyses, individuals assorted into discrete social communities, and these communities had high spatial overlap. By unifying social network analysis with iSSF, we identified movement-dependent social association, where individuals foraged with more familiar individuals, but moved collectively with any between foraging patches. Our study demonstrates that social behaviour and space use are inter-related based on spatial overlap of social communities and movement-dependent habitat selection. Movement, habitat selection, and social behaviour are linked in theory. Here, we put these concepts into practice to demonstrate that movement is the glue connecting individual habitat selection to the social environment.

Toxoplasma gondii is hypothesized to manipulate the behavior of warm-blooded hosts to promote trophic transmission into the parasite's definitive feline hosts. A key prediction of this hypothesis is that T. gondii infections of non-feline hosts are associated with costly behavior toward T. gondii's definitive hosts; however, this effect has not been documented in any of the parasite's diverse wild hosts during naturally occurring interactions with felines. Here, three decades of field observations reveal that T. gondii-infected hyena cubs approach lions more closely than uninfected peers and have higher rates of lion mortality. We discuss these results in light of 1) the possibility that hyena boldness represents an extended phenotype of the parasite, and 2) alternative scenarios in which T. gondii has not undergone selection to manipulate behavior in host hyenas. Both cases remain plausible and have important ramifications for T. gondii's impacts on host behavior and fitness in the wild. The parasite causing toxoplasmosis can manipulate prey to behave in ways that promote transmission to the parasite's definitive feline hosts. The first study consistent with this extended phenotype in the wild finds that infected hyena cubs approach lions more closely than uninfected peers and have higher rates of lion mortality.

The Anthropocene marks great changes to environments and the animals that inhabit them. Changes, such as disturbance, can affect the manner in which animals interact with their environments, such as moving and selecting habitats. To test how animals might respond to changing disturbance regimes, we employ an experimental approach to movement ecology. We used integrated step selection analysis (iSSA) to test the behavioural responses of individually-marked grove snails ( Cepaea nemoralis ) exposed to a gradient of physical disturbance in their habitat. We used a before-after control-impact (BACI) experimental design within semi-controlled mesocosms to manipulate edge and disturbance variables by altering the area of the mesocosm covered by bricks. We showed that grove snails perceive edges of enclosures and edges of bricks as risks, and responded to such risks by altering their movement. Grove snails displayed a bimodal response to risk by taking shelter in place or moving faster to be farther from the disturbance. Furthermore, individuals tended to modulate their behavioural response to the degree of risk. Our study highlights the usefulness of experimental mesocosms in movement ecology and in determining the effects of habitat alteration and human-imposed risk on movement behaviour.

Studies in rodents and captive primates suggest that the early-life social environment affects future phenotype, potentially through alterations to DNA methylation. Little is known of these associations in wild animals. In a wild population of spotted hyenas, we test the hypothesis that maternal care during the first year of life and social connectedness during two periods of early development leads to differences in DNA methylation and fecal glucocorticoid metabolites (fGCMs) later in life. Here we report that although maternal care and social connectedness during the den-dependent life stage are not associated with fGCMs, greater social connectedness during the subadult den-independent life stage is associated with lower adult fGCMs. Additionally, more maternal care and social connectedness after den independence correspond with higher global (%CCGG) DNA methylation. We also note differential DNA methylation near 5 genes involved in inflammation, immune response, and aging that may link maternal care with stress phenotype.

There is growing interest in the alteration of host behaviors by parasites, yet crucial gaps remain in our understanding of its ecological and evolutionary significance. Here, we present the first evidence that the enhanced boldness of infected intermediate hosts of Toxoplasma gondii can increase their risk of mortality by the parasite’s definitive feline hosts. In a long-term study of hyenas in Kenya’s Masai Mara region, we found that 65% of hyenas were seropositive for T. gondii in ELISA IgG assays. Seropositive hyenas approached lions more closely than uninfected counterparts, and also showed longer latencies to approach a simulated conspecific territorial intruder. Lastly, although not significant, the ratio of mortalities caused by lions (vs. other sources) was higher for hyenas that were infected by T. gondii. These results accord with a long-standing hypothesis that the manipulation of host boldness and/or ailurophilia evolved to enhance disease transmission. Since hyenas are rarely consumed by lions, however, elevating their boldness toward lions may not be adaptive for T. gondii. Instead, it may reflect “collateral manipulation” that evolved to influence homologous mechanisms underlying behaviors of alternative hosts (e.g. rodents). This model is often invoked to explain T. gondii’s many effects in humans, but is virtually unexplored in natural settings. For T. gondii, these effects could feasibly impact both behavior and fitness in a vast array, and significant proportion, of earth’s mammals and birds. In addition to characterizing behavioral covariates of infection, we examined spatial and temporal patterns of T. gondii prevalence within the Mara landscape. Contrary to our predictions, disease prevalence did not differ 1) at a protected vs. disturbed locality, or 2) over three decades of increasing human activity within the disturbed locality.