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.
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.
Understanding how animals use information about their environment to make movement decisions underpins our ability to explain drivers of and predict animal movement. Memory is the cognitive process ...that allows species to store information about experienced landscapes, however, remains an understudied topic in movement ecology. By studying how species select for familiar locations, visited recently and in the past, we can gain insight to how they store and use local information in multiple memory types. In this study, we analyzed the movements of a migratory mule deer (
Odocoileus hemionus
) population in the Piceance Basin of Colorado, United States to investigate the influence of spatial experience over different time scales on seasonal range habitat selection. We inferred the influence of short and long-term memory from the contribution to habitat selection of previous space use within the same season and during the prior year, respectively. We fit step-selection functions to GPS collar data from 32 female deer and tested the predictive ability of covariates representing current environmental conditions and both metrics of previous space use on habitat selection, inferring the latter as the influence of memory within and between seasons (summer vs. winter). Across individuals, models incorporating covariates representing both recent and past experience and environmental covariates performed best. In the top model, locations that had been previously visited within the same season and locations from previous seasons were more strongly selected relative to environmental covariates, which we interpret as evidence for the strong influence of both short- and long-term memory in driving seasonal range habitat selection. Further, the influence of previous space uses was stronger in the summer relative to winter, which is when deer in this population demonstrated strongest philopatry to their range. Our results suggest that mule deer update their seasonal range cognitive map in real time and retain long-term information about seasonal ranges, which supports the existing theory that memory is a mechanism leading to emergent space-use patterns such as site fidelity. Lastly, these findings provide novel insight into how species store and use information over different time scales.
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.
Animal migrations are ubiquitous and one of the most threatened ecological processes globally. Because of the multifaceted nature of migration – seasonal movements between home ranges – it can be ...difficult to tease apart the underlying mechanisms influencing this behaviour. It is necessary to understand these mechanisms, not only to deepen our fundamental understanding of migration in animals, but also because migrations in many species are vulnerable to environmental change. In Chapter 2, I first systematically identify the broad proximate drivers of migration and offer generalities across vertebrate taxa. I quantitatively reviewed 45 studies and extracted 132 observations of effect sizes for internal and external proximate drivers that influenced migration propensity. Through this meta-analysis, I found that internal and external drivers had a medium and large effect, respectively, on migration propensity. Predator abundance and predation risk had a large effect on migration propensity, as did individual behaviour. Of the studies that examined genetic divergence between migrant and resident populations, 64% found some genetic divergence between groups. In Chapter 3, I explore the genetic basis for migration and identified genes associated with migration direction from pooled genome-wide scans on a population of 233 migrating female mule deer (Odocoileus hemionus) where I identified genomic regions including FITM1, a gene linked to the formation of lipids, and DPPA3, a gene linked to epigenetic modifications of the maternal line. These results are consistent with the underlying genetic basis for a migratory trait which contributes to the additive genetic variance influencing migratory behaviours and can affect the adaptive potential of a species. Finally, in Chapter 4 I used a pedigree-free quantitative genetic approach to estimate heritability and sources of environmental variation in migration distance, timing, and movement rate of the same population of mule deer. I found low heritability for broad patterns of migration timing, and greater variation in heritability for behaviours during migration, with low heritability for distance and duration and high heritability for movement rate along the route. Insights into the genetic and environmental sources of variation for migration are critical both for the eco-evolutionary dynamics of migration behaviour, and for the conservation of species whose migrations may be vulnerable to environmental change. My thesis reveals that broad patterns of migration are driven largely by environmental effects while within these broad patterns, migration behaviour is driven to a measurable degree by genetic variation.
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.
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.
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.
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.
Variation in social environment can mitigate risks and rewards associated with occupying a particular patch. We 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).
Coyote Huang, Richard K.K; Webber, Quinn M.R; Laforge, Michel P ...
Canadian journal of zoology,
05/2021, Volume:
99, Issue:
5
Journal Article
Peer reviewed
The interplay of predator encounters and antipredator responses is an integral part of understanding predatorprey interactions and spatial co-occurrence and avoidance can elucidate these ...interactions. We conducted hard-part dietary analysis of coyotes (Canis latrans Say, 1823) and space use of coyotes and woodland caribou (Rangifer tarandus caribou (Gmelin, 1788)) to test two competing hypotheses about coyote and caribou predator-prey spatial dynamics using resource selection functions. The high encounter hypothesis predicts that coyotes would maximize encounters with caribou via high spatial cooccurrence, whereas the predator stealth hypothesis predicts that through low spatial co-occurrence with caribou, coyotes act as stealth predators by avoiding habitats that caribou typically select. Our dietary analysis revealed that ~46% of sampled coyote diet is composed of caribou. We found that coyote share space with caribou in lichen-barren habitat in both summer and winter and that coyotes co-occur with caribou in forested habitat during summer, but not during winter. Our findings support predictions associated with the high encounter predator hypothesis whereby coyotes and caribou have high spatial co-occurrence promoting caribou in coyote diet.
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