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  1. Abstract

    Conservation of migratory species exhibiting wide‐ranging and multidimensional behaviors is challenged by management efforts that only utilize horizontal movements or produce static spatial–temporal products. For the deep‐diving, critically endangered eastern Pacific leatherback turtle, tools that predict where turtles have high risks of fisheries interactions are urgently needed to prevent further population decline. We incorporated horizontal–vertical movement model results with spatial–temporal kernel density estimates and threat data (gear‐specific fishing) to develop monthly maps of spatial risk. Specifically, we applied multistate hidden Markov models to a biotelemetry data set (n = 28 leatherback tracks, 2004–2007). Tracks with dive information were used to characterize turtle behavior as belonging to 1 of 3 states (transiting, residential with mixed diving, and residential with deep diving). Recent fishing effort data from Global Fishing Watch were integrated with predicted behaviors and monthly space‐use estimates to create maps of relative risk of turtle–fisheries interactions. Drifting (pelagic) longline fishing gear had the highest average monthly fishing effort in the study region, and risk indices showed this gear to also have the greatest potential for high‐risk interactions with turtles in a residential, deep‐diving behavioral state. Monthly relative risk surfaces for all gears and behaviors were added to South Pacific TurtleWatch (SPTW) (https://www.upwell.org/sptw), a dynamic management tool for this leatherback population. These modifications will refine SPTW's capability to provide important predictions of potential high‐risk bycatch areas for turtles undertaking specific behaviors. Our results demonstrate how multidimensional movement data, spatial–temporal density estimates, and threat data can be used to create a unique conservation tool. These methods serve as a framework for incorporating behavior into similar tools for other aquatic, aerial, and terrestrial taxa with multidimensional movement behaviors.

     
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  2. Abstract Background

    Animal movement is a key ecological process that is tightly coupled to local environmental conditions. While agriculture, urbanisation, and transportation infrastructure are critical to human socio-economic improvement, these have spurred substantial changes in animal movement across the globe with potential impacts on fitness and survival. Notably, however, human disturbance can have differential effects across species, and responses to human activities are thus largely taxa and context specific. As human disturbance is only expected to worsen over the next decade it is critical to better understand how species respond to human disturbance in order to develop effective, case-specific conservation strategies.

    Methods

    Here, we use an extensive telemetry dataset collected over 22 years to fill a critical knowledge gap in the movement ecology of lowland tapirs (Tapirus terrestris) across areas of varying human disturbance within three biomes in southern Brazil: the Pantanal, Cerrado, and Atlantic Forest.

    Results

    From these data we found that the mean home range size across all monitored tapirs was 8.31 km2(95% CI 6.53–10.42), with no evidence that home range sizes differed between sexes nor age groups. Interestingly, although the Atlantic Forest, Cerrado, and Pantanal vary substantially in habitat composition, levels of human disturbance, and tapir population densities, we found that lowland tapir movement behaviour and space use were consistent across all three biomes. Human disturbance also had no detectable effect on lowland tapir movement. Lowland tapirs living in the most altered habitats we monitored exhibited movement behaviour that was comparable to that of tapirs living in a near pristine environment.

    Conclusions

    Contrary to our expectations, although we observed individual variability in lowland tapir space use and movement, human impacts on the landscape also had no measurable effect on their movement. Lowland tapir movement behaviour thus appears to exhibit very little phenotypic plasticity in response to human disturbance. Crucially, the lack of any detectable response to anthropogenic disturbance suggests that human modified habitats risk being ecological traps for tapirs and this information should be factored into conservation actions and species management aimed towards protecting lowland tapir populations.

     
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  3. Abstract

    Resource selection functions (RSFs) are among the most commonly used statistical tools in both basic and applied animal ecology. They are typically parameterized using animal tracking data, and advances in animal tracking technology have led to increasing levels of autocorrelation between locations in such data sets. Because RSFs assume that data are independent and identically distributed, such autocorrelation can cause misleadingly narrow confidence intervals and biased parameter estimates.

    Data thinning, generalized estimating equations and step selection functions (SSFs) have been suggested as techniques for mitigating the statistical problems posed by autocorrelation, but these approaches have notable limitations that include statistical inefficiency, unclear or arbitrary targets for adequate levels of statistical independence, constraints in input data and (in the case of SSFs) scale‐dependent inference. To remedy these problems, we introduce a method for likelihood weighting of animal locations to mitigate the negative consequences of autocorrelation on RSFs.

    In this study, we demonstrate that this method weights each observed location in an animal's movement track according to its level of non‐independence, expanding confidence intervals and reducing bias that can arise when there are missing data in the movement track.

    Ecologists and conservation biologists can use this method to improve the quality of inferences derived from RSFs. We also provide a complete, annotated analytical workflow to help new users apply our method to their own animal tracking data using thectmm Rpackage.

     
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  4. Abstract

    Projects focused on movement behaviour and home range are commonplace, but beyond a focus on choosing appropriate research questions, there are no clear guidelines for such studies. Without these guidelines, designing an animal tracking study to produce reliable estimates of space‐use and movement properties (necessary to answer basic movement ecology questions), is often done in an ad hoc manner.

    We developed ‘movedesign’, a user‐friendly Shiny application, which can be utilized to investigate the precision of three estimates regularly reported in movement and spatial ecology studies: home range area, speed and distance travelled. Conceptually similar to statistical power analysis, this application enables users to assess the degree of estimate precision that may be achieved with a given sampling design; that is, the choices regarding data resolution (sampling interval) and battery life (sampling duration).

    Leveraging the ‘ctmmRpackage, we utilize two methods proven to handle many common biases in animal movement datasets: autocorrelated kernel density estimators (AKDEs) and continuous‐time speed and distance (CTSD) estimators. Longer sampling durations are required to reliably estimate home range areas via the detection of a sufficient number of home range crossings. In contrast, speed and distance estimation requires a sampling interval short enough to ensure that a statistically significant signature of the animal's velocity remains in the data.

    This application addresses key challenges faced by researchers when designing tracking studies, including the trade‐off between long battery life and high resolution of GPS locations collected by the devices, which may result in a compromise between reliably estimating home range or speed and distance. ‘movedesign’ has broad applications for researchers and decision‐makers, supporting them to focus efforts and resources in achieving the optimal sampling design strategy for their research questions, prioritizing the correct deployment decisions for insightful and reliable outputs, while understanding the trade‐off associated with these choices.

     
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  5. Abstract

    Vertical movements can expose individuals to rapid changes in physical and trophic environments—for aquatic fauna, dive profiles from biotelemetry data can be used to quantify and categorize vertical movements. Inferences on classes of vertical movement profiles typically rely on subjective summaries of parameters or statistical clustering techniques that utilize Euclidean matching of vertical movement profiles with vertical observation points. These approaches are prone to subjectivity, error, and bias. We used machine learning approaches on a large dataset of vertical time series (N = 28,217 dives) for 31 post‐nesting leatherback turtles (Dermochelys coriacea). We applied dynamic time warp (DTW) clustering to group vertical movement (dive) time series by their metrics (depth and duration) into an optimal number of clusters. We then identified environmental covariates associated with each cluster using a generalized additive mixed‐effects model (GAMM). A convolutional neural network (CNN) model, trained on standard dive shape types from the literature, was used to classify dives within each DTW cluster by their shape. Two clusters were identified with the DTW approach—these varied in their spatial and temporal distributions, with dependence on environmental covariates, sea surface temperature, bathymetry, sea surface height anomaly, and time‐lagged surface chlorophyllaconcentrations. CNN classification accuracy of the five standard dive profiles was 95%. Subsequent analyses revealed that the two clusters differed in their composition of standard dive shapes, with each cluster dominated by shapes indicative of distinct behaviors (pelagic foraging and exploration, respectively). The use of these two machine learning approaches allowed for discrete behaviors to be identified from vertical time series data, first by clustering vertical movements by their movement metrics (DTW) and second by classifying dive profiles within each cluster by their shapes (CNN). Statistical inference for the identified clusters found distinct relationships with environmental covariates, supporting hypotheses of vertical niche switching and vertically structured foraging behavior. This approach could be similarly applied to the time series of other animals utilizing the vertical dimension in their movements, including aerial, arboreal, and other aquatic species, to efficiently identify different movement behaviors and inform habitat models.

     
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  6. Abstract

    Warming temperatures and advancing spring are affecting annual snow and ice cycles, as well as plant phenology, across the Arctic and boreal regions. These changes may be linked to observed population declines in wildlife, including barren‐ground caribou (Rangifer tarandus), a key species of Arctic environments. We quantified how barren‐ground caribou, characteristically both gregarious and migratory, synchronize births in time and aggregate births in space and investigated how these tactics are influenced by variable weather conditions. We analyzed movement patterns to infer calving dates for 747 collared female caribou from seven herds across northern North America, totaling 1255 calving events over a 15‐year period. By relating these events to local weather conditions during the 1‐year period preceding calving, we examined how weather influenced calving timing and the ability of caribou to reach their central calving area. We documented continental‐scale synchrony in calving, but synchrony was greatest within an individual herd for a given year. Weather conditions before and during gestation had contrasting effects on the timing and location of calving. Notably, a combination of unfavorable weather conditions during winter and spring, including the pre‐calving migration, resulted in a late arrival on the calving area or a failure to reach the greater calving area in time for calving. Though local weather conditions influenced calving timing differently among herds, warm temperatures and low wind speed, which are associated with soft, deep snow, during the spring and pre‐calving migration, generally affected the ability of female caribou to reach central calving areas in time to give birth. Delayed calving may have potential indirect consequences, including reduced calf survival. Overall, we detected considerable variability across years and across herds, but no significant trend for earlier calving by caribou, even as broad indicators of spring and snow phenology trend earlier. Our results emphasize the importance of monitoring the timing and location of calving, and to examine how weather during summer and winter are affecting calving and subsequent reproductive success.

     
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  7. Abstract

    Migratory birds have the capacity to shift their migration phenology in response to climatic change. Yet the mechanistic underpinning of changes in migratory timing remain poorly understood. We employed newly developed global positioning system (GPS) tracking devices and long-term dataset of migration passage timing to investigate how behavioral responses to environmental conditions relate to phenological shifts in American robins (Turdus migratorius) during spring migration to Arctic-boreal breeding grounds. We found that over the past quarter-century (1994–2018), robins have migrated ca. 5 d/decade earlier. Based on GPS data collected for 55 robins over three springs (2016–2018), we found the arrival timing and likelihood of stopovers, and timing of arrival to breeding grounds, were strongly influenced by dynamics in snow conditions along migratory paths. These findings suggest plasticity in migratory behavior may be an important mechanism for how long-distance migrants adjust their breeding phenology to keep pace with advancement of spring on breeding grounds.

     
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  8. Abstract

    Ecologists have long been interested in linking individual behaviour with higher level processes. For motile species, this ‘upscaling’ is governed by how well any given movement strategy maximizes encounters with positive factors and minimizes encounters with negative factors. Despite the importance of encounter events for a broad range of ecological processes, encounter theory has not kept pace with developments in animal tracking or movement modelling. Furthermore, existing work has focused primarily on the relationship between animal movement and encounterrateswhile the relationship between individual movement and the spatiallocationsof encounter events in the environment has remained conspicuously understudied.

    Here, we bridge this gap by introducing a method for describing the long‐term encounter location probabilities for movement within home ranges, termed the conditional distribution of encounters (CDE). We then derive this distribution, as well as confidence intervals, implement its statistical estimator into open‐source software and demonstrate the broad ecological relevance of this distribution.

    We first use simulated data to show how our estimator provides asymptotically consistent estimates. We then demonstrate the general utility of this method for three simulation‐based scenarios that occur routinely in biological systems: (a) a population of individuals with home ranges that overlap with neighbours; (b) a pair of individuals with a hard territorial border between their home ranges; and (c) a predator with a large home range that encompassed the home ranges of multiple prey individuals. Using GPS data from white‐faced capuchinsCebus capucinus, tracked on Barro Colorado Island, Panama, and sleepy lizardsTiliqua rugosa,tracked in Bundey, South Australia, we then show how the CDE can be used to estimate the locations of territorial borders, identify key resources, quantify the potential for competitive or predatory interactions and/or identify any changes in behaviour that directly result from location‐specific encounter probability.

    The CDE enables researchers to better understand the dynamics of populations of interacting individuals. Notably, the general estimation framework developed in this work builds straightforwardly off of home range estimation and requires no specialized data collection protocols. This method is now openly available via thectmm Rpackage.

     
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  9. Abstract

    Many important demographic processes are seasonal, including survival. For many species, mortality risk is significantly higher at certain times of the year than at others, whether because resources are scarce, susceptibility to predators or disease is high, or both. Despite the importance of survival modelling in wildlife sciences, no tools are available to estimate the peak, duration and relative importance of these ‘seasons of mortality’.

    We presentcyclomort, anrpackage that estimates the timing, duration and intensity of any number of mortality seasons with reliable confidence intervals. The package includes a model selection approach to determine the number of mortality seasons and to test whether seasons of mortality vary across discrete grouping factors.

    We illustrate the periodic hazard function model and workflow of cyclomort with simulated data. We then estimate mortality seasons of two caribouRangifer taranduspopulations that have strikingly different mortality patterns, including different numbers and timing of mortality peaks, and a marked change in one population over time.

    Thecyclomortpackage was developed to estimate mortality seasons for wildlife, but the package can model any time‐to‐event processes with a periodic component.

     
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  10. Abstract

    Alpine treelines are expected to shift upward due to recent climate change. However, interpretation of changes in montane systems has been problematic because effects of climate change are frequently confounded with those of land use changes. The eastern Himalaya, particularly Langtang National Park, Central Nepal, has been relatively undisturbed for centuries and thus presents an opportunity for studying climate change impacts on alpine treeline uncontaminated by potential confounding factors.

    We studied two dominant species,Abies spectabilis (AS)andRhododendron campanulatum (RC), above and below the treeline on two mountains. We constructed 13 transects, each spanning up to 400 m in elevation, in which we recorded height and state (dead or alive) of all trees, as well as slope, aspect, canopy density, and measures of anthropogenic and animal disturbance.

    All size classes ofRCplants had lower mortality above treeline than below it, and youngRCplants (<2 m tall) were at higher density above treeline than below.ASshows little evidence of a position change from the historic treeline, with a sudden extreme drop in density above treeline compared to below. Recruitment, as measured by size–class distribution, was greater above treeline than below for both species butASis confined to ~25 m above treeline whereasRCis luxuriantly growing up to 200 m above treeline.

    Synthesis. Evidence suggests that the elevational limits ofRChave shifted upward both because (a) young plants above treeline benefited from facilitation of recruitment by surrounding vegetation, allowing upward expansion of recruitment, and (b) temperature amelioration to mature plants increased adult survival. We predict that the current pure stand ofRCgrowing above treeline will be colonized byASthat will, in turn, outshade and eventually relegateRCto be a minor component of the community, as is the current situation below the treeline.

     
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