1. Roads are a prevalent, ever-increasing form of human disturbance on the landscape. In many places in western North America, energy development has brought human and road disturbance into seasonal ...winter range areas for migratory elk. 2. We sought to evaluate the predictions from the risk-disturbance hypothesis when studying elk response to roads during winter. Road proximity and crossing were used to evaluate these behaviours, which offered a rare comparison between two common measures of roads. We used integrated step selection analysis (iSSA) to evaluate four alternative hypotheses regarding the influence of roads on space-use behaviour across 175 elk-years of elk telemetry data, and we quantified both population-level and individual-level variations in responses. 3. We demonstrated, for the first time, how iSSA can be used to combine movement analysis in a refined approach to habitat selection. Elk responded to roads as they would natural predation risk. Elk selected areas farther from roads at all times of day with avoidance being greatest during twilight. In addition, elk sought cover and moved more when in the vicinity of roads. Road crossings were generally avoided, but this avoidance was weakest during daytime when elk were both moving and closer to roads. 4. Synthesis and applications. Energy development is transforming landscapes in western North America with the proliferation of roads, which we show is having substantial and multifaceted negative effects on elk movement and behaviour. These adverse effects can be mitigated by minimizing new road construction and by restricting traffic on roads as well as providing the protection of tree cover on elk winter ranges.
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.
Predators use different spatial tactics to track the prey on the landscape. Three hypotheses describe spatial tactics: prey abundance for prey that are aggregated in space; prey habitat for uniformly ...distributed prey; and prey catchability for prey that are difficult to catch and kill. The gray wolf (
Canis lupus
) is a generalist predator that likely employs more than one spatial hunting tactic to match their diverse prey with distinct distributions and behavior that are available. We conducted a study on 17 GPS collared wolves in 6 packs in Riding Mountain National Park, Manitoba, Canada where wolves prey on moose (
Alces alces
) and elk (
Cervus canadensis
). We evaluated wolf selection for prey density, habitat selection and catchability on the landscape through within-territory habitat selection analysis. We reveal support for both the
prey habitat
and
prey catchability
hypotheses. For moose, their primary prey, wolves employed a mixed habitat and catchability tactic. Wolves used spaces described by the intersection of moose habitat and moose catchability. Wolves selected for the catchability of elk, their secondary prey, but not elk habitat. Counter to our predictions, wolves avoided areas of moose and elk density, likely highlighting the ongoing space race between predator and prey. We illustrate that of the three hypotheses the primary driver was prey catchability, where the interplay of both prey habitat with catchability culminate in predator spatial behaviour in a multiprey system.
Predicting future space use by animals requires models that consider both habitat availability and individual differences in habitat selection. The functional response in habitat selection posits ...animals adjust their habitat selection to availability, but population-level responses to availability may differ from individual responses. Generalized functional response (GFR) models account for functional responses by including fixed effect interactions between habitat availability and selection. Population-level resource selection functions instead account for individual selection responses to availability with random effects. We compared predictive performance of both approaches using a functional response in elk (
Cervus canadensis
) selection for mixed forest in response to road proximity, and avoidance of roads in response to mixed forest availability. We also investigated how performance changed when individuals responded differently to availability from the rest of the population. Individual variation in road avoidance decreased performance of both models (random effects:
β
= 0.69, 95% CI 0.47, 0.91; GFR:
β
= 0.38, 95% CI 0.05, 0.71). Changes in individual road and forest availability affected performance of neither model, suggesting individual responses to availability different from the functional response mediated performance. We also found that overall, both models performed similarly for predicting mixed forest selection (
F
1, 58
= 0.14,
p
= 0.71) and road avoidance (
F
1, 58
= 0.28,
p
= 0.60). GFR estimates were slightly better, but its larger number of covariates produced greater variance than the random effects model. Given this bias-variance trade-off, we conclude that neither model performs better for future space use predictions.
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.
Context
In southwestern Alberta, human development, including roads, is encroaching on the landscape and into the range of a partially migratory population of elk (
Cervus elaphus
).
Objectives
To ...quantify factors influencing among- and within-home-range selection of winter range in this population.
Methods
We studied individual habitat selection and road avoidance at two biologically relevant spatial scales. We outlined availability extents for 107 individual elk-years based on observed fall migration distance, and based on a minimum convex polygon around winter telemetry relocations. To model the response by elk to road disturbance, we fit a habitat-selection model to each elk-year at each of the two availability extents, and examined population-level and individual variation in space-use. We then evaluated the relationship between inferred selection at the two scales and the functional response in selection.
Results
Roads had a ubiquitous influence on elk across scales. Elk, individually and as a population, avoided roads when migrating to their winter range and within this seasonal home range. Individual elk that avoided roads more strongly relative to the population did so at both scales of analysis.Further, the avoidance of low-use roads decreased with increasing road density. These results support bottom-up habitat-selection patterns (i.e., scale-independent) and functional response in habitat selection.
Conclusions
Overall, using a multi-scale habitat selection analysis, we show that road avoidance is a major determinant of elk space-use behaviour across multiple scales. Consequently, any new road construction or increases in road-use intensity could have detrimental effects on migratory elk populations by restricting space-use.
The COVID-19 pandemic stressed public health systems worldwide due to emergence of several highly transmissible variants of concern. Diverse and complex intervention policies deployed over the last ...years have shown varied effectiveness in controlling the pandemic. However, a systematic analysis and modelling of the combined effects of different viral lineages and complex intervention policies remains a challenge due to the lack of suitable measures of pandemic inequality and nonlinear effects. Using large-scale agent-based modelling and a high-resolution computational simulation matching census-based demographics of Australia, we carried out a systematic comparative analysis of several COVID-19 pandemic scenarios. The scenarios covered two most recent Australian census years (2016 and 2021), three variants of concern (ancestral, Delta and Omicron), and five representative intervention policies. We introduced pandemic Lorenz curves measuring an unequal distribution of the pandemic severity across local areas. We also quantified pandemic biomodality, distinguishing between urban and regional waves, and measured bifurcations in the effectiveness of interventions. We quantified nonlinear effects of population heterogeneity on the pandemic severity, highlighting that (i) the population growth amplifies pandemic peaks, (ii) the changes in population size amplify the peak incidence more than the changes in density, and (iii) the pandemic severity is distributed unequally across local areas. We also examined and delineated the effects of urbanisation on the incidence bimodality, distinguishing between urban and regional pandemic waves. Finally, we quantified and examined the impact of school closures, complemented by partial interventions, and identified the conditions when inclusion of school closures may decisively control the transmission. Public health response to long-lasting pandemics must be frequently reviewed and adapted to demographic changes. To control recurrent waves, mass-vaccination rollouts need to be complemented by partial NPIs. Healthcare and vaccination resources need to be prioritised towards the localities and regions with high population growth and/or high density.
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.
There is strenuous debate among ecologists regarding the inclusion of predator density into the originally prey-dependent functional response. We provided comprehensive empirical comparisons of ...alternative functional response models for the predatory ostracod Heterocypris incongruens (Ramdohr, 1808) and the rotifer Brachionus calyciflorus (Pallas, 1766) as its prey in small freshwater microcosms. Prey killed was measured at factorial combinations of four predator densities and five prey densities, and was recorded at 3 min intervals over 60 min experiments. To support the potential effect of predator interference on per capita kill rate, we recorded ostracod activity and aggression. Kill rate increased following a saturating function with increasing prey density and decreased with increasing predator density. Model evaluation using an information–theoretic approach indicated that the Arditi–Ginzburg type II ratio-dependent model performed best, followed by the Arditi–Akcakaya and Beddington–DeAngelis type II predator-dependent models, suggesting that predator interference was important in predicting kill rates. Interference among predators increased and their activity decreased with increasing predator density, providing confirmation that interference was responsible for the predator-dependent effect. By combining a microcosm experiment and behavioral observations, our results suggest that predator interference at realistic population densities influences ostracod kill rates and this form of interference was best accommodated by predator-dependent models.
Genome-wide association studies (GWAS) of Alzheimer's disease (AD) have identified several risk loci, but many remain unknown. Cerebrospinal fluid (CSF) biomarkers may aid in gene discovery and we ...previously demonstrated that six CSF biomarkers (β-amyloid, total/phosphorylated tau, NfL, YKL-40, and neurogranin) cluster into five principal components (PC), each representing statistically independent biological processes. Here, we aimed to (1) identify common genetic variants associated with these CSF profiles, (2) assess the role of associated variants in AD pathophysiology, and (3) explore potential sex differences.
We performed GWAS for each of the five biomarker PCs in two multi-center studies (EMIF-AD and ADNI). In total, 973 participants (n = 205 controls, n = 546 mild cognitive impairment, n = 222 AD) were analyzed for 7,433,949 common SNPs and 19,511 protein-coding genes. Structural equation models tested whether biomarker PCs mediate genetic risk effects on AD, and stratified and interaction models probed for sex-specific effects.
Five loci showed genome-wide significant association with CSF profiles, two were novel (rs145791381 inflammation and GRIN2D synaptic functioning) and three were previously described (APOE, TMEM106B, and CHI3L1). Follow-up analyses of the two novel signals in independent datasets only supported the GRIN2D locus, which contains several functionally interesting candidate genes. Mediation tests indicated that variants in APOE are associated with AD status via processes related to amyloid and tau pathology, while markers in TMEM106B and CHI3L1 are associated with AD only via neuronal injury/inflammation. Additionally, seven loci showed sex-specific associations with AD biomarkers.
These results suggest that pathway and sex-specific analyses can improve our understanding of AD genetics and may contribute to precision medicine.
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