Expanding theory, methodology and empirical systems at the spatial-social interface.

All animals exhibit some combination of spatial and social behaviours. A diversity of interactions occurs between such behaviours, producing emergent phenomena at the spatial-social interface. Untangling and interrogating these complex, intertwined processes can be vital for identifying the mechanisms, causes and consequences of behavioural variation in animal ecology. Nevertheless, the integrated study of the interactions between spatial and social phenotypes and environments (at the spatial-social interface) is in its relative infancy. In this theme issue, we present a collection of papers chosen to expand the spatial-social interface along several theoretical, methodological and empirical dimensions. They detail new perspectives, methods, study systems and more, as well as offering roadmaps for applied outputs and detailing exciting new directions for the field to move in the future. In this Introduction, we outline the contents of these papers, placing them in the context of what comes before, and we synthesize a number of takeaways and future directions for the spatial-social interface. This article is part of the theme issue 'The spatial-social interface: a theoretical and empirical integration'.

Scale at the interface of spatial and social ecology.

Animals simultaneously navigate spatial and social environments, and their decision-making with respect to those environments constitutes their spatial (e.g. habitat selection) and social (e.g. conspecific associations) phenotypes. The spatial-social interface is a recently introduced conceptual framework linking these components of spatial and social ecology. The spatial-social interface is inherently scale-dependent, yet it has not been integrated with the rich body of literature on ecological scale. Here, we develop a conceptual connection between the spatial-social interface and ecological scale. We propose three key innovations that incrementally build upon each other. First, the use-availability framework that underpins a large body of literature in behavioural ecology can be used in analogy to the phenotype-environment nomenclature and is transferable across the spatial and social realms. Second, both spatial and social phenotypes are hierarchical, with nested components that are linked via constraints-from the top down-or emergent properties-from the bottom up. Finally, in both the spatial and social realms, the definitions of environment and phenotype depend on the focal scale of inquiry. These conceptual innovations cast our understanding of the relationships between social and spatial dimensions of animal ecology in a new light, allowing a more holistic understanding and clearer hypothesis development for animal behaviour. This article is part of the theme issue 'The spatial-social interface: a theoretical and empirical integration'.

Defining null expectations for animal site fidelity.

Site fidelity-the tendency to return to previously visited locations-is widespread across taxa. Returns may be driven by several mechanisms, including memory, habitat selection, or chance; however, pattern-based definitions group different generating mechanisms under the same label of 'site fidelity', often assuming memory as the main driver. We propose an operational definition of site fidelity as patterns of return that deviate from a null expectation derived from a memory-free movement model. First, using agent-based simulations, we show that without memory, intrinsic movement characteristics and extrinsic landscape characteristics are key determinants of return patterns and that even random movements may generate substantial probabilities of return. Second, we illustrate how to implement our framework empirically to establish ecologically meaningful, system-specific null expectations for site fidelity. Our approach provides a conceptual and operational framework to test hypotheses on site fidelity across systems and scales.

Recent trends in movement ecology of animals and human mobility.

Movement is fundamental to life, shaping population dynamics, biodiversity patterns, and ecosystem structure. In 2008, the movement ecology framework (MEF Nathan et al. in PNAS 105(49):19052-19059, 2008) introduced an integrative theory of organismal movement-linking internal state, motion capacity, and navigation capacity to external factors-which has been recognized as a milestone in the field. Since then, the study of movement experienced a technological boom, which provided massive quantities of tracking data of both animal and human movement globally and at ever finer spatio-temporal resolutions. In this work, we provide a quantitative assessment of the state of research within the MEF, focusing on animal movement, including humans and invertebrates, and excluding movement of plants and microorganisms. Using a text mining approach, we digitally scanned the contents of [Formula: see text] papers from 2009 to 2018 available online, identified tools and methods used, and assessed linkages between all components of the MEF. Over the past decade, the publication rate has increased considerably, along with major technological changes, such as an increased use of GPS devices and accelerometers and a majority of studies now using the R software environment for statistical computing. However, animal movement research still largely focuses on the effect of environmental factors on movement, with motion and navigation continuing to receive little attention. A search of topics based on words featured in abstracts revealed a clustering of papers among marine and terrestrial realms, as well as applications and methods across taxa. We discuss the potential for technological and methodological advances in the field to lead to more integrated and interdisciplinary research and an increased exploration of key movement processes such as navigation, as well as the evolutionary, physiological, and life-history consequences of movement.

Predicting greater sage-grouse habitat selection at the southern periphery of their range.

Mapping suitable habitat is an important process in wildlife conservation planning. Species distribution reflects habitat selection processes occurring across multiple spatio-temporal scales. Because habitat selection may be driven by different factors at different scales, conservation planners require information at the scale of the intervention to plan effective management actions. Previous research has described habitat selection processes shaping the distribution of greater sage-grouse (Centrocercus urophasianus; sage-grouse) at the range-wide scale. Finer-scale information for applications within jurisdictional units inside the species range is lacking, yet necessary, because state wildlife agencies are the management authority for sage-grouse in the United States. We quantified seasonal second-order habitat selection for sage-grouse across the state of Utah to produce spatio-temporal predictions of their distribution at the southern periphery of the species range. We used location data obtained from sage-grouse marked with very-high-frequency radio-transmitters and lek location data collected between 1998 and 2013 to quantify species habitat selection in relation to a suite of topographic, edaphic, climatic, and anthropogenic variables using random forest algorithms. Sage-grouse selected for greater sagebrush (Artemisia spp.) cover, higher elevations, and gentler slopes and avoided lower precipitations and higher temperatures. The strength of responses to habitat variables varied across seasons. Anthropogenic variables previously reported as affecting their range-wide distribution (i.e., roads, powerlines, communication towers, and agricultural development) were not ranked as top predictors at our focal scale. Other than strong selection for sagebrush cover, the responses we observed differed from what has been reported at the range-wide scale. These differences likely reflect the unique climatic, geographic, and topographic context found in the southern peripheral area of the species distribution compared to range-wide environmental gradients. Our results highlight the importance of considering appropriateness of scale when planning conservation actions for wide-ranging species.

Analysis of movement recursions to detect reproductive events and estimate their fate in central place foragers.

BACKGROUND: Recursive movement patterns have been used to detect behavioral structure within individual movement trajectories in the context of foraging ecology, home-ranging behavior, and predator avoidance. Some animals exhibit movement recursions to locations that are tied to reproductive functions, including nests and dens; while existing literature recognizes that, no method is currently available to explicitly target different types of revisited locations. Moreover, the temporal persistence of recursive movements to a breeding location can carry information regarding the fate of breeding attempts, but it has never been used as a metric to quantify recursive movement patterns. Here, we introduce a method to locate breeding attempts and estimate their fate from GPS-tracking data of central place foragers. We tested the performance of our method in three bird species differing in breeding ecology (wood stork (Mycteria americana), lesser kestrel (Falco naumanni), Mediterranean gull (Ichthyaetus melanocephalus)) and implemented it in the R package 'nestR'. METHODS: We identified breeding sites based on the analysis of recursive movements within individual tracks. Using trajectories with known breeding attempts, we estimated a set of species-specific criteria for the identification of nest sites, which we further validated using non-reproductive individuals as controls. We then estimated individual nest survival as a binary measure of reproductive fate (success, corresponding to fledging of at least one chick, or failure) from nest-site revisitation histories during breeding attempts, using a Bayesian hierarchical modeling approach that accounted for temporally variable revisitation patterns, probability of visit detection, and missing data. RESULTS: Across the three species, positive predictive value of the nest-site detection algorithm varied between 87 and 100% and sensitivity between 88 and 92%, and we correctly estimated the fate of 86-100% breeding attempts. CONCLUSIONS: By providing a method to formally distinguish among revisited locations that serve different ecological functions and introducing a probabilistic framework to quantify temporal persistence of movement recursions, we demonstrated how the analysis of recursive movement patterns can be applied to estimate reproduction in central place foragers. Beyond avian species, the principles of our method can be applied to other central place foraging breeders such as denning mammals. Our method estimates a component of individual fitness from movement data and will help bridge the gap between movement behavior, environmental factors, and their fitness consequences.