The adaptive value of density-dependent habitat specialization and social network centrality.

Density dependence is a fundamental ecological process. In particular, animal habitat selection and social behavior often affect fitness in a density-dependent manner. The Ideal Free Distribution (IFD) and niche variation hypothesis (NVH) present distinct predictions associated with Optimal Foraging Theory about how the effect of habitat selection on fitness varies with population density. Using caribou (Rangifer tarandus) in Canada as a model system, we test competing hypotheses about how habitat specialization, social behavior, and annual reproductive success (co)vary across a population density gradient. Within a behavioral reaction norm framework, we estimate repeatability, behavioral plasticity, and covariance among social behavior and habitat selection to investigate the adaptive value of sociality and habitat selection. In support of NVH, but not the IFD, we find that at high density habitat specialists had higher annual reproductive success than generalists, but were also less social than generalists, suggesting the possibility that specialists were less social to avoid competition. Our study supports niche variation as a mechanism for density-dependent habitat specialization.

Genomic correlates for migratory direction in a free-ranging cervid.

Animal migrations are some of the most ubiquitous and one of the most threatened ecological processes globally. A wide range of migratory behaviours occur in nature, and this behaviour is not uniform among and within species, where even individuals in the same population can exhibit differences. While the environment largely drives migratory behaviour, it is necessary to understand the genetic mechanisms influencing migration to elucidate the potential of migratory species to cope with novel conditions and adapt to environmental change. In this study, we identified genes associated with a migratory trait by undertaking pooled genome-wide scans on a natural population of migrating mule deer. We identified genomic regions associated with variation in migratory direction, including FITM1, a gene linked to the formation of lipids, and DPPA3, a gene linked to epigenetic modifications of the maternal line. Such a genetic basis for a migratory trait contributes to the adaptive potential of the species and might affect the flexibility of individuals to change their behaviour in the face of changes in their environment.

Conceptual and methodological advances in habitat-selection modeling: guidelines for ecology and evolution.

Habitat selection is a fundamental animal behavior that shapes a wide range of ecological processes, including animal movement, nutrient transfer, trophic dynamics and population distribution. Although habitat selection has been a focus of ecological studies for decades, technological, conceptual and methodological advances over the last 20 yr have led to a surge in studies addressing this process. Despite the substantial literature focused on quantifying the habitat-selection patterns of animals, there is a marked lack of guidance on best analytical practices. The conceptual foundations of the most commonly applied modeling frameworks can be confusing even to those well versed in their application. Furthermore, there has yet to be a synthesis of the advances made over the last 20 yr. Therefore, there is a need for both synthesis of the current state of knowledge on habitat selection, and guidance for those seeking to study this process. Here, we provide an approachable overview and synthesis of the literature on habitat-selection analyses (HSAs) conducted using selection functions, which are by far the most applied modeling framework for understanding the habitat-selection process. This review is purposefully non-technical and focused on understanding without heavy mathematical and statistical notation, which can confuse many practitioners. We offer an overview and history of HSAs, describing the tortuous conceptual path to our current understanding. Through this overview, we also aim to address the areas of greatest confusion in the literature. We synthesize the literature outlining the most exciting conceptual advances in the field of habitat-selection modeling, discussing the substantial ecological and evolutionary inference that can be made using contemporary techniques. We aim for this paper to provide clarity for those navigating the complex literature on HSAs while acting as a reference and best practices guide for practitioners.

Tracking snowmelt to jump the green wave: phenological drivers of migration in a northern ungulate.

In northern climates, spring is a time of rapid environmental change: for migrating terrestrial animals, melting snow facilitates foraging and travel, and newly emergent vegetation provides a valuable nutritional resource. These changes result in selection on the timing of important life-history events such as migration and parturition occurring when high-quality resources are most abundant. We examined the timing of female caribou (Rangifer tarandus, n = 94) migration and parturition in five herds across 7 yr in Newfoundland, Canada, as a function of two measures of environmental change-snowmelt and vegetation green-up. We generated resource selection functions to test whether caribou selected for areas associated with snowmelt and green-up during migration and following calving. We found that caribou migrated approximately 1 wk prior to snowmelt, with the flush of emergent vegetation occurring during the weeks following parturition. The results indicate that caribou "jump" the green wave of emergent forage and do so by tracking the receding edge of melting snow, likely reducing movement and foraging costs related to snow cover. Our research further broadens the ecological scope of resource tracking in animals. We demonstrate that resource tracking extends beyond resources directly related to foraging to those related to movement. We also show that snowmelt provides an environmental cue that may provide a buffer against changing environmental conditions.

Geometry of the ideal free distribution: individual behavioural variation and annual reproductive success in aggregations of a social ungulate.

Variation in social environment can mitigate risks and rewards associated with occupying a particular patch. We aim to integrate Ideal Free Distribution (IFD) and Geometry of the Selfish Herd (GSH) to address an apparent conflict in their predictions of equal mean fitness between patches (IFD) and declining fitness benefits within a patch (GSH). We tested these hypotheses in a socio-spatial context using individual caribou that were aggregated or disaggregated during calving and varied in their annual reproductive success (ARS). We then tested individual consistency of these spatial tactics. We reveal that two socio-spatial tactics accorded similar mean ARS (IFD); however, ARS for aggregated individuals declined near the periphery (GSH). Individuals near the aggregation periphery exhibited flexibility, whereas others were consistent. The integration of classical theories through a contemporary lens of consistent individual differences provides evidence for an integrated GSH and IFD strategy that may represent an evolutionary stable state.

The Ecology of Individual Differences Empirically Applied to Space-Use and Movement Tactics.

Movement provides a link between individual behavioral ecology and the spatial and temporal variation in an individual's landscape. Individual variation in movement traits is an important axis of animal personality, particularly in the context of foraging ecology. We tested whether individual caribou (Rangifer tarandus) displayed plasticity in movement and space-use behavior across a gradient of resource aggregation. We quantified first-passage time and range-use ratio as proxies for movement-related foraging behavior and examined how these traits varied at the individual level across a foraging resource gradient. Our results suggest that individuals adjusted first-passage time but not range-use ratio to maximize access to high-quality foraging resources. First-passage time was repeatable, and intercepts for first-passage time and range-use ratio were negatively correlated. Individuals matched first-passage time but not range-use ratio to the expectations of our patch-use model that maximized access to foraging resources, a result that suggests that individuals acclimated their movement patterns to accommodate both intra- and interannual variation in foraging resources on the landscape. Collectively, we highlight repeatable movement and space-use tactics and provide insight into how individual plasticity in movement interacts with landscape processes to affect the distribution of behavioral phenotypes and potentially fitness and population dynamics.

Implementing a novel movement-based approach to inferring parturition and neonate caribou calf survival.

In ungulates, parturition is correlated with a reduction in movement rate. With advances in movement-based technologies comes an opportunity to develop new techniques to assess reproduction in wild ungulates that are less invasive and reduce biases. DeMars et al. (2013, Ecology and Evolution 3:4149-4160) proposed two promising new methods (individual- and population-based; the DeMars model) that use GPS inter-fix step length of adult female caribou (Rangifer tarandus caribou) to infer parturition and neonate survival. Our objective was to apply the DeMars model to caribou populations that may violate model assumptions for retrospective analysis of parturition and calf survival. We extended the use of the DeMars model after assigning parturition and calf mortality status by examining herd-wide distributions of parturition date, calf mortality date, and survival. We used the DeMars model to estimate parturition and calf mortality events and compared them with the known parturition and calf mortality events from collared adult females (n = 19). We also used the DeMars model to estimate parturition and calf mortality events for collared female caribou with unknown parturition and calf mortality events (n = 43) and instead derived herd-wide estimates of calf survival as well as distributions of parturition and calf mortality dates and compared them to herd-wide estimates generated from calves fitted with VHF collars (n = 134). For our data, the individual-based method was effective at predicting calf mortality, but was not effective at predicting parturition. The population-based method was more effective at predicting parturition but was not effective at predicting calf mortality. At the herd-level, the predicted distributions of parturition date from both methods differed from each other and from the distribution derived from the parturition dates of VHF-collared calves (log-ranked test: χ2 = 40.5, df = 2, p < 0.01). The predicted distributions of calf mortality dates from both methods were similar to the observed distribution derived from VHF-collared calves. Both methods underestimated herd-wide calf survival based on VHF-collared calves, however, a combination of the individual- and population-based methods produced herd-wide survival estimates similar to estimates generated from collared calves. The limitations we experienced when applying the DeMars model could result from the shortcomings in our data violating model assumptions. However despite the differences in our caribou systems, with proper validation techniques the framework in the DeMars model is sufficient to make inferences on parturition and calf mortality.

The effect of terrain and female density on survival of neonatal white-tailed deer and mule deer fawns.

Juvenile survival is a highly variable life-history trait that is critical to population growth. Antipredator tactics, including an animal's use of its physical and social environment, are critical to juvenile survival. Here, we tested the hypothesis that habitat and social characteristics influence coyote (Canis latrans) predation on white-tailed deer (Odocoileus virginianus) and mule deer (O. hemionus) fawns in similar ways during the neonatal period. This would contrast to winter when the habitat and social characteristics that provide the most safety for each species differ. We monitored seven cohorts of white-tailed deer and mule deer fawns at a grassland study site in Alberta, Canada. We used logistic regression and a model selection procedure to determine how habitat characteristics, climatic conditions, and female density influenced fawn survival during the first 8 weeks of life. Fawn survival improved after springs with productive vegetation (high integrated Normalized Difference Vegetation Index values). Fawns that used steeper terrain were more likely to survive. Fawns of both species had improved survival in years with higher densities of mule deer females, but not with higher densities of white-tailed deer females, as predicted if they benefit from protection by mule deer. Our results suggest that topographical variation is a critical resource for neonates of many ungulate species, even species like white-tailed deer that use more gentle terrain when older. Further, our results raise the possibility that neonatal white-tailed fawns may benefit from associating with mule deer females, which may contribute to the expansion of white-tailed deer into areas occupied by mule deer.