Summary 1.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. 2. 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. 3. 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. 4. 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.

1. 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.2. 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).3. We highlight that multisensor approaches are a new frontier in biologging, while identifying current limitations and avenues for future development in sensor technology.4. 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.

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Inferring the role of interactions in territorial animals relies upon accurate recordings of the behaviour of neighbouring individuals. Such accurate recordings are rarely available from field studies. As a result, quantification of the interaction mechanisms has often relied upon theoretical approaches, which hitherto have been limited to comparisons of macroscopic population-level predictions from un-tested interaction models. Here we present a quantitative framework that possesses a microscopic testable hypothesis on the mechanism of conspecific avoidance mediated by olfactory signals in the form of scent marks. We find that the key parameters controlling territoriality are two: the average territory size, i.e. the inverse of the population density, and the time span during which animal scent marks remain active. Since permanent monitoring of a territorial border is not possible, scent marks need to function in the temporary absence of the resident. As chemical signals carried by the scent only last a finite amount of time, each animal needs to revisit territorial boundaries frequently and refresh its own scent marks in order to deter possible intruders. The size of the territory an animal can maintain is thus proportional to the time necessary for an animal to move between its own territorial boundaries. By using an agent-based model to take into account the possible spatio-temporal movement trajectories of individual animals, we show that the emerging territories are the result of a form of collective animal movement where, different to shoaling, flocking or herding, interactions are highly heterogeneous in space and time. The applicability of our hypothesis has been tested with a prototypical territorial animal, the red fox (Vulpes vulpes).

SUMMARY BackgroundAlthough short-term outcome in severe alcoholic hepatitis (SAH) is well described, its long-term course remains uncharacterised.

Predicting space use patterns of animals from their interactions with the environment is fundamental for understanding the effect of habitat changes on ecosystem functioning. Recent attempts to address this problem have sought to unify resource selection analysis, where animal space use is derived from available habitat quality, and mechanistic movement models, where detailed movement processes of an animal are used to predict its emergent utilization distribution. Such models bias the animal's movement towards patches that are easily available and resource-rich, and the result is a predicted probability density at a given position being a function of the habitat quality at that position. However, in reality, the probability that an animal will use a patch of the terrain tends to be a function of the resource quality in both that patch and the surrounding habitat.We propose a mechanistic model where this non-local effect of resources naturally emerges from the local movement processes, by taking into account the relative utility of both the habitat where the animal currently resides and that of where it is moving. We give statistical techniques to parametrize the model from location data and demonstrate application of these techniques to GPS location data of caribou (Rangifer tarandus) in Newfoundland.Steady-state animal probability distributions arising from the model have complex patterns that cannot be expressed simply as a function of the local quality of the habitat. In particular, large areas of good habitat are used more intensively than smaller patches of equal quality habitat, whereas isolated patches are used less frequently. Both of these are real aspects of animal space use missing from previous mechanistic resource selection models.Whilst we focus on habitats in this study, our modelling framework can be readily used with any environmental covariates and therefore represents a unification of mechanistic modelling and step selection approaches to understanding animal space use.

Mathematical models of spatial population dynamics typically focus on the interplay between dispersal events and birth/death processes. However, for many animal communities, significant arrangement in space can occur on shorter timescales, where births and deaths are negligible. This phenomenon is particularly prevalent in populations of larger, vertebrate animals who often reproduce only once per year or less. To understand spatial arrangements of animal communities on such timescales, we use a class of diffusion–taxis equations for modelling inter-population movement responses between populations. These systems of equations incorporate the effect on animal movement of both the current presence of other populations and the memory of past presence encoded either in the environment or in the minds of animals. We give general criteria for the spontaneous formation of both stationary and oscillatory patterns, via linear pattern formation analysis. For , we classify completely the pattern formation properties using a combination of linear analysis and nonlinear energy functionals. In this case, the only patterns that can occur asymptotically in time are stationary. However, for , oscillatory patterns can occur asymptotically, giving rise to a sequence of period-doubling bifurcations leading to patterns with no obvious regularity, a hallmark of chaos. Our study highlights the importance of understanding between-population animal movement for understanding spatial species distributions, something that is typically ignored in species distribution modelling, and so develops a new paradigm for spatial population dynamics. Electronic supplementary material The online version of this article (10.1007/s11538-019-00626-9) contains supplementary material, which is available to authorized users.

Collective phenomena, whereby agent-agent interactions determine spatial patterns, are ubiquitous in the animal kingdom. On the other hand, movement and space use are also greatly influenced by the interactions between animals and their environment. Despite both types of interaction fundamentally influencing animal behaviour, there has hitherto been no unifying framework for the models proposed in both areas. Here, we construct a general method for inferring population-level spatial patterns from underlying individual movement and interaction processes, a key ingredient in building a statistical mechanics for ecological systems. We show that resource selection functions, as well as several examples of collective motion models, arise as special cases of our framework, thus bringing together resource selection analysis and collective animal behaviour into a single theory. In particular, we focus on combining the various mechanistic models of territorial interactions in the literature with step selection functions, by incorporating interactions into the step selection framework and demonstrating how to derive territorial patterns from the resulting models. We demonstrate the efficacy of our model by application to a population of insectivore birds in the Amazon rainforest.