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Foraging is a key driver of animal movement patterns, with specific challenges for predators which must search for mobile prey. These patterns are increasingly impacted by global changes, principally in land use and climate. Understanding the degree of flexibility in predator foraging and social strategies is pertinent to wildlife conservation under global change, including potential top-down effects on wider ecosystems. Here we propose key future research directions to better understand foraging strategies and social flexibility in predators. In particular, rapid continued advances in biologging technology are helping to record and understand dynamic behavioural and movement responses of animals to environmental changes, and their energetic consequences. Data collection can be optimised by calibrating behavioural interpretation methods in captive settings and strategic tagging decisions within and between social groups. Importantly, many species' social systems are increasingly being found to be more flexible than originally described in the literature, which may be more readily detectable through biologging approaches than behavioural observation. Integrating the effects of the physical landscape and biotic interactions will be key to explaining and predicting animal movements and energetic balance in a changing world. [ABSTRACT FROM AUTHOR]

Uncovering the mechanisms behind animal space use patterns is of vital importance for predictive ecology, thus conservation and management of ecosystems. Movement is a core driver of those patterns so understanding how movement mechanisms give rise to space use patterns has become an increasingly active area of research.This study focuses on a particular strand of research in this area, based around step selection analysis (SSA). SSA is a popular way of inferring drivers of movement decisions, but, perhaps less well appreciated, it also parametrises a model of animal movement. Of key interest is that this model can be propagated forwards in time to predict the space use patterns over broader spatial and temporal scales than those that pertain to the proximate movement decisions of animals.Here, we provide a guide for understanding and using the various existing techniques for scaling up step selection models to predict broad‐scale space use patterns. We give practical guidance on when to use which technique, as well as specific examples together with code in R and Python.By pulling together various disparate techniques into one place, and providing code and instructions in simple examples, we hope to highlight the importance of these techniques and make them accessible to a wider range of ecologists, ultimately helping expand the usefulness of SSA. [ABSTRACT FROM AUTHOR]

Addressing ongoing biodiversity loss requires collaboration between conservation scientists and practitioners. However, such collaboration has proved challenging. Despite the potential importance of tracking animal movements for conservation, reviews of the tracking literature have identified a gap between the academic discipline of movement ecology and its application to biodiversity conservation. Through structured conversations with movement ecologists and conservation practitioners, we aimed to understand whether the identified gap is also perceived in practice, and if so, what factors hamper collaboration and how these factors can be remediated. We found that both groups are motivated and willing to collaborate. However, because their motivations differ, there is potential for misunderstandings and miscommunications. In addition, external factors such as funder requirements, academic metrics, and journal scopes may limit the applicability of scientific results in a conservation setting. Potential solutions we identified included improved communication and better presentation of results, acknowledging each other's motivations and desired outputs, and adjustment of funder priorities. Addressing gaps between science and implementation can enhance collaboration and support conservation action to address the global biodiversity crisis more effectively. [ABSTRACT FROM AUTHOR]

The ability to predict animal space use patterns is a fundamental concern in changing environments. Such predictions require a detailed understanding of the movement mechanisms from which spatial distributions emerge. However, these are typically complex, multifaceted, and therefore difficult to uncover.Here, we provide a methodological framework for uncovering the movement mechanisms necessary for building predictive models of animal space use. Our procedure begins by parametrising a movement model of each individual in a population using step selection analysis, from which we build an individual‐based model (IBM) of interacting individuals, derive predicted broad‐scale space use patterns from the IBM and then compare the predicted and empirical patterns. Importantly, discrepancies between these predicted and empirical patterns are used to formulate new hypotheses about the drivers of animal movement decisions and thus iteratively improve the model's predictive power. We demonstrate our method on a population of feral pigs in Mississippi, USA.Our technique incorporates both social interactions between individuals and environmental drivers of movement. At each iteration of model construction, we were able to identify missing features to improve model prediction by analysing the IBM output. These include overuse‐avoidance effects of self‐attractive mechanisms (i.e. attraction to previously visited sites becomes repulsion if there have been multiple visits in quick succession), which were vital for ensuring predicted occurrence distributions do not become vanishingly small.Overall, we have provided a general method for iteratively improving the predictive power of step selection models. This will enable future researchers to maximise the information obtained from step selection analyses and to highlight potentially missing data for uncovering the drivers of movement decisions and emergent space use patterns. Ultimately, this provides a fundamental step towards the general aim of constructing predictive models of animal space use. [ABSTRACT FROM AUTHOR]

The energetic gains from foraging and costs of movement are expected to be key drivers of animal decision‐making, as their balance is a large determinant of body condition and survival. This fundamental perspective is often missing from habitat selection studies, which mainly describe correlations between space use and environmental features, rather than the mechanisms behind these correlations.To address this gap, we present a novel parameterisation of step selection functions (SSFs), that we term the energy selection function (ESF). In this model, the likelihood of an animal selecting a movement step depends directly on the corresponding energetic gains and costs, and we can therefore assess how moving animals choose habitat based on energetic considerations.The ESF retains the mathematical convenience and practicality of other SSFs and can be quickly fitted using standard software. In this article, we outline a workflow, from data gathering to statistical analysis, and use a case study of polar bears Ursus maritimus to demonstrate application of the model.We explain how defining gains and costs at the scale of the movement step allows us to include information about resource distribution, landscape resistance and movement patterns. We further demonstrate this process with a case study of polar bears and show how the parameters can be interpreted in terms of selection for energetic gains and against energetic costs.The ESF is a flexible framework that combines the energetic consequences of both movement and resource selection, thus incorporating a key mechanism into habitat selection analysis. Further, because it is based on familiar habitat selection models, the ESF is widely applicable to any study system where energetic gains and costs can be derived, and has immense potential for methodological extensions. [ABSTRACT FROM AUTHOR]

Background: Understanding what animals do in time and space is important for a range of ecological questions, however accurate estimates of how animals use space is challenging. Within the use of animal-attached tags, radio telemetry (including the Global Positioning System, 'GPS') is typically used to verify an animal's location periodically. Straight lines are typically drawn between these 'Verified Positions' ('VPs') so the interpolation of space-use is limited by the temporal and spatial resolution of the system's measurement. As such, parameters such as route-taken and distance travelled can be poorly represented when using VP systems alone. Dead-reckoning has been suggested as a technique to improve the accuracy and resolution of reconstructed movement paths, whilst maximising battery life of VP systems. This typically involves deriving travel vectors from motion sensor systems and periodically correcting path dimensions for drift with simultaneously deployed VP systems. How often paths should be corrected for drift, however, has remained unclear. Methods and results: Here, we review the utility of dead-reckoning across four contrasting model species using different forms of locomotion (the African lion Panthera leo, the red-tailed tropicbird Phaethon rubricauda, the Magellanic penguin Spheniscus magellanicus, and the imperial cormorant Leucocarbo atriceps). Simulations were performed to examine the extent of dead-reckoning error, relative to VPs, as a function of Verified Position correction (VP correction) rate and the effect of this on estimates of distance moved. Dead-reckoning error was greatest for animals travelling within air and water. We demonstrate how sources of measurement error can arise within VP-corrected dead-reckoned tracks and propose advancements to this procedure to maximise dead-reckoning accuracy. Conclusions: We review the utility of VP-corrected dead-reckoning according to movement type and consider a range of ecological questions that would benefit from dead-reckoning, primarily concerning animal–barrier interactions and foraging strategies. [ABSTRACT FROM AUTHOR]

Background: Fine-scale data on animal position are increasingly enabling us to understand the details of animal movement ecology and dead-reckoning, a technique integrating motion sensor-derived information on heading and speed, can be used to reconstruct fine-scale movement paths at sub-second resolution, irrespective of the environment. On its own however, the dead-reckoning process is prone to cumulative errors, so that position estimates quickly become uncoupled from true location. Periodic ground-truthing with aligned location data (e.g., from global positioning technology) can correct for this drift between Verified Positions (VPs). We present step-by-step instructions for implementing Verified Position Correction (VPC) dead-reckoning in R using the tilt-compensated compass method, accompanied by the mathematical protocols underlying the code and improvements and extensions of this technique to reduce the trade-off between VPC rate and dead-reckoning accuracy. These protocols are all built into a user-friendly, fully annotated VPC dead-reckoning R function; Gundog.Tracks, with multi-functionality to reconstruct animal movement paths across terrestrial, aquatic, and aerial systems, provided within the Additional file 4 as well as online (GitHub). Results: The Gundog.Tracks function is demonstrated on three contrasting model species (the African lion Panthera leo, the Magellanic penguin Spheniscus magellanicus, and the Imperial cormorant Leucocarbo atriceps) moving on land, in water and in air. We show the effect of uncorrected errors in speed estimations, heading inaccuracies and infrequent VPC rate and demonstrate how these issues can be addressed. Conclusions: The function provided will allow anyone familiar with R to dead-reckon animal tracks readily and accurately, as the key complex issues are dealt with by Gundog.Tracks. This will help the community to consider and implement a valuable, but often overlooked method of reconstructing high-resolution animal movement paths across diverse species and systems without requiring a bespoke application. [ABSTRACT FROM AUTHOR]

Step selection analysis (SSA) is a fundamental technique for uncovering the drivers of animal movement decisions. Its typical use has been to view an animal as 'selecting' each measured location, given its current (and possibly previous) locations. Although an animal is unlikely to make decisions precisely at the times its locations are measured, if data are gathered at a relatively low frequency (every few minutes or hours) this is often the best that can be done. Nowadays, though, tracking data are increasingly gathered at very high frequencies, often ≥1 Hz, so it may be possible to exploit these data to perform more behaviourally‐meaningful step selection analysis.Here, we present a technique to do this. We first use an existing algorithm to determine the turning‐points in an animal's movement path. We define a 'step' to be a straight‐line movement between successive turning‐points. We then construct a generalised version of integrated SSA (iSSA), called time‐varying iSSA (tiSSA), which deals with the fact that turning‐points are usually irregularly spaced in time. We demonstrate the efficacy of tiSSA by application to data on both simulated animals and free‐ranging goats Capra aegagrus hircus, comparing our results to those of regular iSSA with locations that are separated by a constant time‐interval.Using (regular) iSSA with constant time‐steps can give results that are misleading compared to using tiSSA with the actual turns made by the animals. Furthermore, tiSSA can be used to infer covariates that are dependent on the time between turns, which is not possible with regular iSSA. As an example, we show that our study animals tend to spend less time between successive turns when the ground is rockier and/or the temperature is hotter.By constructing a step selection technique that works between observed turning‐points of animals, we enable step selection to be used on high‐frequency movement data, which are becoming increasingly prevalent in modern biologging studies. Furthermore, since turning‐points can be viewed as decisions, our method places step selection analysis on a more behaviourally‐meaningful footing compared to previous techniques. [ABSTRACT FROM AUTHOR]

In environments that vary unpredictably, many animals are nomadic, moving in an irregular pattern that differs from year to year. Exploring the mechanisms of nomadic movement is needed to understand how animals survive in highly variable environments, and to predict behavioural and population responses to environmental change.We developed a network model to identify plausible mechanisms of nomadic animal movement by comparing the performance of multiple movement rules along a continuum from nomadism to residency. Using simulations and analytical results, we explored how different types of habitat modifications (that augment or decrease resource availability) might affect the abundance and movement rates of animals following each of these rules.Movement rules for which departure from patches depended on resource availability and/or competition performed almost equally well and better than residency or uninformed movement under most conditions, even though animals using each rule moved at substantially different rates. Habitat modifications that stabilized resources, either by resource supplementation or degradation, eroded the benefits of informed nomadic movements, particularly for movements based on resource availability alone.These results suggest that simple movement rules can explain nomadic animal movements and determine species' responses to environmental change. In particular, landscape stabilization and supplementation might be useful strategies for promoting populations of resident animals, but would be less beneficial for managing highly mobile species, many of which are threatened by habitat disruption and changes in climate. [ABSTRACT FROM AUTHOR]

Mathematical analysis of partial differential equations (PDEs) has led to many insights regarding the effect of organism movements on spatial population dynamics. However, their use has mainly been confined to the community of mathematical biologists, with less attention from statistical and empirical ecologists. We conjecture that this is principally due to the inherent difficulties in fitting PDEs to data.To help remedy this situation, in the context of movement ecology, we show how the popular technique of step selection analysis (SSA) can be used to parametrize a class of PDEs, called diffusion‐taxis models, from an animal's trajectory. We examine the accuracy of our technique on simulated data, then demonstrate the utility of diffusion‐taxis models in two ways. First, for non‐interacting animals, we derive the steady‐state utilization distribution in a closed analytic form. Second, we give a recipe for deriving spatial pattern formation properties that emerge from interacting animals: specifically, do those interactions cause heterogeneous spatial distributions to emerge and if so, do these distributions oscillate at short times or emerge without oscillations? The second question is applied to data on concurrently tracked bank voles Myodes glareolus.Our results show that SSA can accurately parametrize diffusion‐taxis equations from location data, providing the frequency of the data is not too low. We show that the steady‐state distribution of our diffusion‐taxis model, where it is derived, has an identical functional form to the utilization distribution given by resource selection analysis (RSA), thus formally linking (fine scale) SSA with (broad scale) RSA. For the bank vole data, we show how our SSA‐PDE approach can give predictions regarding the spatial aggregation and segregation of different individuals, which are difficult to predict purely by examining results of SSA.Our methods provide a user‐friendly way into the world of PDEs, via a well‐used statistical technique, which should lead to tighter links between the findings of mathematical ecology and observations from empirical ecology. By providing a non‐speculative link between observed movement behaviours and space use patterns on larger spatio‐temporal scales, our findings will also aid integration of movement ecology into understanding spatial species distributions. [ABSTRACT FROM AUTHOR]

Animal behavior is elicited, in part, in response to external conditions, but understanding how animals perceive the environment and make the decisions that bring about these behavioral responses is challenging. Animal heads often move during specific behaviors and, additionally, typically have sensory systems (notably vision, smell, and hearing) sampling in defined arcs (normally to the front of their heads). As such, head‐mounted electronic sensors consisting of accelerometers and magnetometers, which can be used to determine the movement and directionality of animal heads (where head "movement" is defined here as changes in heading [azimuth] and/or pitch [elevation angle]), can potentially provide information both on behaviors in general and also clarify which parts of the environment the animals might be prioritizing ("environmental framing"). We propose a new approach to visualize the data of such head‐mounted tags that combines the instantaneous outputs of head heading and pitch in a single intuitive spherical plot. This sphere has magnetic heading denoted by "longitude" position and head pitch by "latitude" on this "orientation sphere" (O‐sphere). We construct the O‐sphere for the head rotations of a number of vertebrates with contrasting body shape and ecology (oryx, sheep, tortoises, and turtles), illustrating various behaviors, including foraging, walking, and environmental scanning. We also propose correcting head orientations for body orientations to highlight specific heading‐independent head rotation, and propose the derivation of O‐sphere‐metrics, such as angular speed across the sphere. This should help identify the functions of various head behaviors. Visualizations of the O‐sphere provide an intuitive representation of animal behavior manifest via head orientation and rotation. This has ramifications for quantifying and understanding behaviors ranging from navigation through vigilance to feeding and, when used in tandem with body movement, should provide an important link between perception of the environment and response to it in free‐ranging animals. [ABSTRACT FROM AUTHOR]

Animal movements towards goals or targets are based upon either maximization of resource acquisition or risk avoidance, and the way animals move can reveal information about their motivation.We use hidden Markov models (HMMs) fitted in a Bayesian framework and hourly Global Positioning System fixes to distinguish animal movements into distinct states and analyse the influence of environmental variables on being in, and switching to, a particular state. Specifically, we apply our models to understand elephant movement decisions around agricultural fields, and crop consumption. As it is unclear what the role of habitat features are on this complex process, we analyse whether elephants target agricultural crops for consumption, or simply pass through them in search of water.Our HMMs separate elephant movements into two states: exploratory movements that are fast and directional, and encamped movements that are slow and meandering. For each elephant, we ran 16 models with each possible combination of selected habitat features (river, elephant corridor, agricultural field, trees), and repeated these analyses including interaction effects with both season and time of day. We used cross‐validation to select the best model.In corridors, exploratory movements are dominant. Elephants mainly showed encamped movements at the river during the dry season, when temporary water sources have dried out and elephants relied on this permanent water source. In fields, males most often exhibited exploratory movements to and from the river, while females showed an increase in the frequency of encamped behaviour during the dry season and at night—the times when most crop consumption and movements through fields occur.Adaptation to risk could explain this behaviour, since foraging in fields is likely less risky under the cover of darkness and during the dry season when farmers are absent. This sex segregation in elephant movement decisions highlights the importance of predation risk in shaping movement patterns, which can result in sex segregation in responses to mitigation methods. The increase in encamped movements in the dry season suggests the importance of agricultural timing, and shows the potential for early ploughing and early‐harvest crop types in order to reduce elephant crop consumption. Taking this into account could increase efficiency of elephant crop consumption mitigation. [ABSTRACT FROM AUTHOR]

The advent of miniaturized biologging devices has provided ecologists with unprecedented opportunities to record animal movement across scales, and led to the collection of ever‐increasing quantities of tracking data. In parallel, sophisticated tools have been developed to process, visualize and analyse tracking data; however, many of these tools have proliferated in isolation, making it challenging for users to select the most appropriate method for the question in hand. Indeed, within the r software alone, we listed 58 packages created to deal with tracking data or 'tracking packages'.Here, we reviewed and described each tracking package based on a workflow centred around tracking data (i.e. spatio‐temporal locations (x, y, t)), broken down into three stages: pre‐processing, post‐processing and analysis, the latter consisting of data visualization, track description, path reconstruction, behavioural pattern identification, space use characterization, trajectory simulation and others.Supporting documentation is key to render a package accessible for users. Based on a user survey, we reviewed the quality of packages' documentation and identified 11 packages with good or excellent documentation.Links between packages were assessed through a network graph analysis. Although a large group of packages showed some degree of connectivity (either depending on functions or suggesting the use of another tracking package), one third of the packages worked in isolation, reflecting a fragmentation in the r movement‐ecology programming community.Finally, we provide recommendations for users when choosing packages, and for developers to maximize the usefulness of their contribution and strengthen the links within the programming community. [ABSTRACT FROM AUTHOR]

The paradigm‐changing opportunities of biologging sensors for ecological research, especially movement ecology, are vast, but the crucial questions of how best to match the most appropriate sensors and sensor combinations to specific biological questions and how to analyse complex biologging data, are mostly ignored.Here, we fill this gap by reviewing how to optimize the use of biologging techniques to answer questions in movement ecology and synthesize this into an Integrated Biologging Framework (IBF).We highlight that multisensor approaches are a new frontier in biologging, while identifying current limitations and avenues for future development in sensor technology.We focus on the importance of efficient data exploration, and more advanced multidimensional visualization methods, combined with appropriate archiving and sharing approaches, to tackle the big data issues presented by biologging. We also discuss the challenges and opportunities in matching the peculiarities of specific sensor data to the statistical models used, highlighting at the same time the large advances which will be required in the latter to properly analyse biologging data.Taking advantage of the biologging revolution will require a large improvement in the theoretical and mathematical foundations of movement ecology, to include the rich set of high‐frequency multivariate data, which greatly expand the fundamentally limited and coarse data that could be collected using location‐only technology such as GPS. Equally important will be the establishment of multidisciplinary collaborations to catalyse the opportunities offered by current and future biologging technology. If this is achieved, clear potential exists for developing a vastly improved mechanistic understanding of animal movements and their roles in ecological processes and for building realistic predictive models. [ABSTRACT FROM AUTHOR]

Remote sensing is a valuable tool in movement ecology. However, the way it is combined with animal movement data is far from optimal as most studies overlook differences in the spatial, temporal and thematic resolutions between both data types.rsmove uses a pixel‐based approach to link animal tracking and remote sensing data that bridge the gap between these two disciplines while respecting the limitations of the latest. Additionally, if offers standardize methods to pre‐analyse the connection between animal movement and environmental change.The package guides the choice of study sites, satellite data and environmental predictors though data mining and visualization tools. rsmove offers a simple methodology to analyse animal–environment interactions that helps avoid unnecessary, time‐consuming satellite data processing through an informed decision process.The package offers a basis for a better communication between ecologists and remote sensing experts contributing to the definition of clear remote sensing data requirements. Additionally, it provides future studies on animal movement tools for a more critical approach on the selection of environmental data supporting the reproducibility of animal movement studies. [ABSTRACT FROM AUTHOR]

Automated movement tracking is essential for high‐throughput quantitative analyses of the behaviour and kinematics of organisms. Automated tracking also improves replicability by avoiding observer bias and allowing reproducible workflows. However, few automated tracking programs exist that are open access, open source and capable of tracking unmarked organisms in noisy environments.Tracktor is an image‐based tracking freeware designed to perform single‐object tracking in noisy environments, or multi‐object tracking in uniform environments while maintaining individual identities. Tracktor is code‐based but requires no coding skills other than the user being able to specify tracking parameters in a designated location, much like in a graphical user interface. The installation and use of the software is fully detailed in a user manual.Through four examples of common tracking problems, we show that Tracktor is able to track a variety of animals in diverse conditions. The main strengths of Tracktor lie in its ability to track single individuals under noisy conditions (e.g. when the object shape is distorted), its robustness to perturbations (e.g. changes in lighting conditions during the experiment), and its capacity to track multiple unmarked individuals while maintaining their identities. Additionally, summary statistics and plots allow measuring and visualising common metrics used in the analysis of animal movement (e.g. cumulative distance, speed, acceleration, activity, time spent in specific areas and distance to conspecific, etc.).Tracktor is a versatile, reliable and easy‐to‐use automated tracking software that is compatible with all operating systems and provides many features not available in other existing freeware. Access Tracktor and the complete user manual here: https://github.com/vivekhsridhar/tracktor [ABSTRACT FROM AUTHOR]

Decomposing the life track of an animal into behavioral segments is a fundamental challenge for movement ecology. The proliferation of high‐resolution data, often collected many times per second, offers much opportunity for understanding animal movement. However, the sheer size of modern data sets means there is an increasing need for rapid, novel computational techniques to make sense of these data. Most existing methods were designed with smaller data sets in mind and can thus be prohibitively slow. Here, we introduce a method for segmenting high‐resolution movement trajectories into sites of interest and transitions between these sites. This builds on a previous algorithm of Benhamou and Riotte‐Lambert (2012). Adapting it for use with high‐resolution data. The data's resolution removed the need to interpolate between successive locations, allowing us to increase the algorithm's speed by approximately two orders of magnitude with essentially no drop in accuracy. Furthermore, we incorporate a color scheme for testing the level of confidence in the algorithm's inference (high = green, medium = amber, low = red). We demonstrate the speed and accuracy of our algorithm with application to both simulated and real data (Alpine cattle at 1 Hz resolution). On simulated data, our algorithm correctly identified the sites of interest for 99% of "high confidence" paths. For the cattle data, the algorithm identified the two known sites of interest: a watering hole and a milking station. It also identified several other sites which can be related to hypothesized environmental drivers (e.g., food). Our algorithm gives an efficient method for turning a long, high‐resolution movement path into a schematic representation of broadscale decisions, allowing a direct link to existing point‐to‐point analysis techniques such as optimal foraging theory. It is encoded into an R package called SitesInterest, so should serve as a valuable tool for making sense of these increasingly large data streams. We have constructed a new method for identifying sites of interest, tailored specifically for use with high resolution data. This allows for a complicated movement trajectory to be simplified into a "Markov chain‐like" description of behavioural choices, highlighting the animal's broadscale decisions. [ABSTRACT FROM AUTHOR]

Abstract: Recent advances in biologging have resulted in animal location data at unprecedentedly high temporal resolutions, sometimes many times per second. However, many current methods for analysing animal movement (e.g. step selection analysis or state‐space modelling) were developed with lower‐resolution data in mind. To make such methods usable with high‐resolution data, we require techniques to identify features within the trajectory where movement deviates from a straight line. We propose that the intricacies of movement paths, and particularly turns, reflect decisions made by animals so that turn points are particularly relevant to behavioural ecologists. As such, we introduce a fast, accurate algorithm for inferring turning‐points in high‐resolution data. For analysing big data, speed and scalability are vitally important. We test our algorithm on simulated data, where varying amounts of noise were added to paths of straight‐line segments interspersed with turns. We also demonstrate our algorithm on data of free‐ranging oryx Oryx leucoryx. We compare our algorithm to existing statistical techniques for break‐point inference. The algorithm scales linearly and can analyse several hundred‐thousand data points in a few seconds on a mid‐range desktop computer. It identified turnpoints in simulated data with complete accuracy when the noise in the headings had a standard deviation of ±8∘, well within the tolerance of many modern biologgers. It has comparable accuracy to the existing algorithms tested, and is up to three orders of magnitude faster. Our algorithm, freely available in R and Python, serves as an initial step in processing ultra high‐resolution animal movement data, resulting in a rarefied path that can be used as an input into many existing step‐and‐turn methods of analysis. The resulting path consists of points where the animal makes a clear turn, and thereby provides valuable data on decisions underlying movement patterns. As such, it provides an important breakthrough required as a starting point for analysing subsecond resolution data. [ABSTRACT FROM AUTHOR]

Many animals undertake movements that are longer scaled and more directed than their typical home ranging behaviour. These movements include seasonal migrations (e.g. between breeding and feeding grounds), natal dispersal, nomadic range shifts and responses to local environmental disruptions. While various heuristic tools exist for identifying range shifts and migrations, none explicitly model the movement of the animals within a statistical framework that facilitates quantitative comparisons., We present the mechanistic range shift analysis (MRSA), a method to estimate a suite of range shift parameters: times of initiation, duration of transitions, centroids and areas of respective ranges. The method can take the autocorrelation and irregular sampling that is characteristic of much movement data into account. The mechanistic parameters suggest an intuitive measure, the range shift index, for the extent of a range shift. The likelihood based estimation further allows for statistical tests of several relevant hypotheses, including a range shift test, a stopover test and a site fidelity test. The analysis tools are provided in an R package ( marcher)., We applied the MRSA to a population of GPS tracked roe deer ( Capreolus capreolus) in the Italian Alps between 2005 and 2008. With respect to seasonal migration, this population is extremely variable and difficult to classify. Using the MRSA, we were able to quantify the behaviours across the population and among individuals across years, identifying extents, durations and locations of seasonal range shifts, including cases that would have been ambiguous to detect using existing tools., The strongest patterns were differences across years: many animals simply did not perform a seasonal migration to wintering grounds during the mild winter of 2006-2007, even though some of these same animals did move extensively in other, harsher winters. For seasonal migrants, however, site fidelity across years was extremely high, even after skipping an entire seasonal migration. These results suggest that for roe deer behavioural plasticity and tactical responses to immediate environmental cues are reflected in the decision of whether rather than where to migrate. The MRSA also revealed a trade-off between the probability of migrating and the size of a home range. [ABSTRACT FROM AUTHOR]

Home range sizes of territorial animals are often observed to vary periodically in response to seasonal changes in foraging opportunities. Here we develop the first mechanistic model focused on the temporal dynamics of home range expansion and contraction in territorial animals. We demonstrate how simple movement principles can lead to a rich suite of range size dynamics, by balancing foraging activity with defensive requirements and incorporating optimal behavioral rules into mechanistic home range analysis. Our heuristic model predicts three general temporal patterns that have been observed in empirical studies across multiple taxa. First, a positive correlation between age and territory quality promotes shrinking home ranges over an individual’s lifetime, with maximal range size variability shortly before the adult stage. Second, poor sensory information, low population density, and large resource heterogeneity may all independently facilitate range size instability. Finally, aggregation behavior toward forage-rich areas helps produce divergent home range responses between individuals from different age classes. This model has broad applications for addressing important unknowns in animal space use, with potential applications also in conservation and health management strategies. [ABSTRACT FROM AUTHOR]

Understanding how individual movement scales with body size is of fundamental importance in predicting ecological relationships for diverse species. One-dimensional movement metrics scale consistently with body size yet vary over different temporal scales. Knowing how temporal scale influences the relationship between animal body size and movement would better inform hypotheses about the efficiency of foraging behaviour, the ontogeny of energy budgets, and numerous life-history trade-offs., We investigated how the temporal scaling of allometric patterns in movement varies over the course of a year, specifically during periods of motivated (directional and fast movement) and unmotivated (stationary and tortuous movement) behaviour. We focused on a recently diverged group of species that displays wide variation in movement behaviour - giant Galapagos tortoises ( Chelonoidis spp.) - to test how movement metrics estimated on a monthly basis scaled with body size., We used state-space modelling to estimate seven different movement metrics of Galapagos tortoises. We used log-log regression of the power law to evaluate allometric scaling for these movement metrics and contrasted relationships by species and sex., Allometric scaling of movement was more apparent during motivated periods of movement. During this period, allometry was revealed at multiple temporal intervals (hourly, daily and monthly), with values observed at daily and monthly intervals corresponding most closely to the expected one-fourth scaling coefficient, albeit with wide credible intervals. We further detected differences in the magnitude of scaling among taxa uncoupled from observed differences in the temporal structuring of their movement rates., Our results indicate that the definition of temporal scales is fundamental to the detection of allometry of movement and should be given more attention in movement studies. Our approach not only provides new conceptual insights into temporal attributes in one-dimensional scaling of movement, but also generates valuable insights into the movement ecology of iconic yet poorly understood Galapagos giant tortoises. [ABSTRACT FROM AUTHOR]

A resource selection function is a model of the likelihood that an available spatial unit will be used by an animal, given its resource value. But how do we appropriately define availability? Step selection analysis deals with this problem at the scale of the observed positional data, by matching each 'used step' (connecting two consecutive observed positions of the animal) with a set of 'available steps' randomly sampled from a distribution of observed steps or their characteristics., Here we present a simple extension to this approach, termed integrated step selection analysis ( iSSA), which relaxes the implicit assumption that observed movement attributes (i.e. velocities and their temporal autocorrelations) are independent of resource selection. Instead, iSSA relies on simultaneously estimating movement and resource selection parameters, thus allowing simple likelihood-based inference of resource selection within a mechanistic movement model., We provide theoretical underpinning of iSSA, as well as practical guidelines to its implementation. Using computer simulations, we evaluate the inferential and predictive capacity of iSSA compared to currently used methods., Our work demonstrates the utility of iSSA as a general, flexible and user-friendly approach for both evaluating a variety of ecological hypotheses, and predicting future ecological patterns. [ABSTRACT FROM AUTHOR]

Animals of many species demonstrate movement behaviour in which decisions are based on a variety of information. Effects of resources have been studied widely, often under the assumption that the environment is constant over the course of the study. Much less understood is the role of dynamic information about continuously changing resource availability and past experiences. Such information can be acquired during movement bouts and used for future decisions via memory., We present a new class of animal movement models, which incorporates a dynamic interplay of movement and information gain processes. Information is contained in a dynamic cognitive map. As an example, we consider time since last visit to locations and how this interacts with environmental information to shape movement patterns. Our models can be fitted to empirical movement trajectories and are therefore amenable to statistical inference (parameter estimation and model selection)., We tested the functionality of our method using simulated data. Parameter estimates were in accordance with true values used in the simulations, and model selection via Bayesian information criterion (BIC) was able to identify true underlying mechanisms of simulated trajectories. Thus, if time since last visit to locations influences movement decisions, our method is able to detect this mechanism., The use of dynamic information such as the one demonstrated in our example models likely requires cognitive abilities such as spatial memory. Therefore, our method can be used to reveal evidence of spatial memory in empirical movement data. Understanding the components of individual movement decisions and their interactions ultimately helps us to predict how population distribution patterns respond to environmental changes, such as landscape fragmentation and changing climate. [ABSTRACT FROM AUTHOR]

The article presents a study which examined the relationship between animal movement and spatially explicit population dynamics across terrestrial and marine environments. It reviews modelling approaches that were used at different levels of organization. It also identifies key approaches for connecting levels as well as integrating empirical and theoretical approaches. Lastly, distilling directions for research are discussed.

Despite its central place in animal ecology no general mechanistic movement model with an emergent home-range pattern has yet been proposed. Random walk models, which are commonly used to model animal movement, show diffusion instead of a bounded home range and therefore require special modifications. Current approaches for mechanistic modeling of home ranges apply only to a limited set of taxa, namely territorial animals and/or central place foragers. In this paper we present a more general mechanistic movement model based on a biased correlated random walk, which shows the potential for home-range behavior. The model is based on an animal tracking a dynamic resource landscape, using a biologically plausible two-part memory system, i.e. a reference- and a working-memory. Our results show that by adding these memory processes the random walker produces home-range behavior as it gains experience, which also leads to more efficient resource use. Interestingly, home-range patterns, which we assessed based on home-range overlap and increase in area covered with time, require the combined action of both memory components to emerge. Our model has the potential to predict home-range size and can be used for comparative analysis of the mechanisms shaping home-range patterns. [ABSTRACT FROM AUTHOR]

1. Although the home range is a fundamental ecological concept, there is considerable debate over how it is best measured. There is a substantial literature concerning the precision and accuracy of all commonly used home range estimation methods; however, there has been considerably less work concerning how estimates vary with sampling regime, and how this affects statistical inferences. 2. We propose a new procedure, based on a variance components analysis using generalized mixed effects models to examine how estimates vary with sampling regime. 3. To demonstrate the method we analyse data from one study of 32 individually marked roe deer and another study of 21 individually marked kestrels. We subsampled these data to simulate increasingly less intense sampling regimes, and compared the performance of two kernel density estimation (KDE) methods, of the minimum convex polygon (MCP) and of the bivariate ellipse methods. 4. Variation between individuals and study areas contributed most to the total variance in home range size. Contrary to recent concerns over reliability, both KDE methods were remarkably efficient, robust and unbiased: 10 fixes per month, if collected over a standardized number of days, were sufficient for accurate estimates of home range size. However, the commonly used 95% isopleth should be avoided; we recommend using isopleths between 90 and 50%. 5. Using the same number of fixes does not guarantee unbiased home range estimates: statistical inferences differ with the number of days sampled, even if using KDE methods. 6. The MCP method was highly inefficient and results were subject to considerable and unpredictable biases. The bivariate ellipse was not the most reliable method at low sample sizes. 7. We conclude that effort should be directed at marking more individuals monitored over long periods at the expense of the sampling rate per individual. Statistical results are reliable only if the whole sampling regime is standardized. We derive practical guidelines for field studies and data analysis. [ABSTRACT FROM AUTHOR]