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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'. [ABSTRACT FROM AUTHOR]

The spatial availability of social resources is speculated to structure animal movement decisions, but the effects of social resources on animal movements are difficult to identify because social resources are rarely measured. Here, we assessed whether varying availability of a key social resource—access to receptive mates—produces predictable changes in movement decisions among bighorn sheep in Nevada, the United States. We compared the probability that males made long-distance 'foray' movements, a critical driver of connectivity, across three ecoregions with varying temporal duration of a socially mediated factor, breeding season. We used a hidden Markov model to identify foray events and then quantified the effects of social covariates on the probability of foray using a discrete choice model. We found that males engaged in forays at higher rates when the breeding season was short, suggesting that males were most responsive to the social resource when its existence was short lived. During the breeding season, males altered their response to social covariates, relative to the non-breeding season, though patterns varied, and age was associated with increased foray probability. Our results suggest that animals respond to the temporal availability of social resources when making the long-distance movements that drive connectivity. This article is part of the theme issue 'The spatial–social interface: a theoretical and empirical integration'. [ABSTRACT FROM AUTHOR]

Human disturbance is contributing to widespread, global changes in the distributions and densities of wild animals. These anthropogenic impacts on wildlife arise from multiple bottom-up and top-down pathways, including habitat loss, resource provisioning, climate change, pollution, infrastructure development, hunting and our direct presence. Animal behaviour is an important mechanism linking these disturbances to population outcomes, although these behavioural pathways are often complex and can remain obscured when different aspects of behaviour are studied in isolation from one another. The spatial–social interface provides a lens for understanding how an animal's spatial and social environments interact to determine its spatial and social phenotype (i.e. measurable characteristics of an individual), and how these phenotypes interact and feed back to reshape environments. Here, we review studies of animal behaviour at the spatial–social interface to understand and predict how human disturbance affects animal movement, distribution and intraspecific interactions, with consequences for the conservation of populations and ecosystems. By understanding the spatial–social mechanisms linking human disturbance to conservation outcomes, we can better design management interventions to mitigate undesired consequences of disturbance. This article is part of the theme issue 'The spatial–social interface: a theoretical and empirical integration'. [ABSTRACT FROM AUTHOR]

Managing pathogen spillover at the wildlife–livestock interface is a key step towards improving global animal health, food security and wildlife conservation. However, predicting the effectiveness of management actions across host–pathogen systems with different life histories is an on-going challenge since data on intervention effectiveness are expensive to collect and results are system-specific. We developed a simulation model to explore how the efficacies of different management strategies vary according to host movement patterns and epidemic growth rates. The model suggested that fast-growing, fast-moving epidemics like avian influenza were best-managed with actions like biosecurity or containment, which limited and localized overall spillover risk. For fast-growing, slower-moving diseases like foot-and-mouth disease, depopulation or prophylactic vaccination were competitive management options. Many actions performed competitively when epidemics grew slowly and host movements were limited, and how management efficacy related to epidemic growth rate or host movement propensity depended on what objective was used to evaluate management performance. This framework offers one means of classifying and prioritizing responses to novel pathogen spillover threats, and evaluating current management actions for pathogens emerging at the wildlife–livestock interface. This article is part of the theme issue 'Dynamic and integrative approaches to understanding pathogen spillover'. [ABSTRACT FROM AUTHOR]

Pathogen spillover between different host species is the trigger for many infectious disease outbreaks and emergence events, and ecosystem boundary areas have been suggested as spatial hotspots of spillover. This hypothesis is largely based on suspected higher rates of zoonotic disease spillover and emergence in fragmented landscapes and other areas where humans live in close vicinity to wildlife. For example, Ebola virus outbreaks have been linked to contacts between humans and infected wildlife at the rural-forest border, and spillover of yellow fever via mosquito vectors happens at the interface between forest and human settlements. Because spillover involves complex interactions between multiple species and is difficult to observe directly, empirical studies are scarce, particularly those that quantify underlying mechanisms. In this review, we identify and explore potential ecological mechanisms affecting spillover of pathogens (and parasites in general) at ecosystem boundaries. We borrow the concept of 'permeability' from animal movement ecology as a measure of the likelihood that hosts and parasites are present in an ecosystem boundary region. We then discuss how different mechanisms operating at the levels of organisms and ecosystems might affect permeability and spillover. This review is a step towards developing a general theory of cross-species parasite spillover across ecosystem boundaries with the eventual aim of improving predictions of spillover risk in heterogeneous landscapes. This article is part of the theme issue 'Dynamic and integrative approaches to understanding pathogen spillover'. [ABSTRACT FROM AUTHOR]

The risk of zoonotic spillover from reservoir hosts, such as wildlife or domestic livestock, to people is shaped by the spatial and temporal distribution of infection in reservoir populations. Quantifying these distributions is a key challenge in epidemiology and disease ecology that requires researchers to make trade-offs between the extent and intensity of spatial versus temporal sampling. We discuss sampling methods that strengthen the reliability and validity of inferences about the dynamics of zoonotic pathogens in wildlife hosts. This article is part of the theme issue 'Dynamic and integrative approaches to understanding pathogen spillover'. [ABSTRACT FROM AUTHOR]

Predicting pathogen spillover requires counting spillover events and aligning such counts with process-related covariates for each spillover event. How can we connect our analysis of spillover counts to simple, mechanistic models of pathogens jumping from reservoir hosts to recipient hosts? We illustrate how the pathways to pathogen spillover can be represented as a directed graph connecting reservoir hosts and recipient hosts and the number of spillover events modelled as a percolation of infectious units along that graph. Percolation models of pathogen spillover formalize popular intuition and management concepts for pathogen spillover, such as the inextricably multilevel nature of cross-species transmission, the impact of covariance between processes such as pathogen shedding and human susceptibility on spillover risk, and the assumptions under which the effect of a management intervention targeting one process, such as persistence of vectors, will translate to an equal effect on the overall spillover risk. Percolation models also link statistical analysis of spillover event datasets with a mechanistic model of spillover. Linear models, one might construct for process-specific parameters, such as the log-rate of shedding from one of several alternative reservoirs, yield a nonlinear model of the log-rate of spillover. The resulting nonlinearity is approximately piecewise linear with major impacts on statistical inferences of the importance of process-specific covariates such as vector density. We recommend that statistical analysis of spillover datasets use piecewise linear models, such as generalized additive models, regression clustering or ensembles of linear models, to capture the piecewise linearity expected from percolation models. We discuss the implications of our findings for predictions of spillover risk beyond the range of observed covariates, a major challenge of forecasting spillover risk in the Anthropocene. This article is part of the theme issue 'Dynamic and integrative approaches to understanding pathogen spillover'. [ABSTRACT FROM AUTHOR]

Group living facilitates pathogen transmission among social hosts, yet temporally stable host social organizations can actually limit transmission of some pathogens. When there are few between-subpopulation contacts for the duration of a disease event, transmission becomes localized to subpopulations. The number of per capita infectious contacts approaches the subpopulation size as pathogen infectiousness increases. Here, we illustrate that this is the case during epidemics of highly infectious pneumonia in bighorn lambs (Ovis canadensis). We classified individually marked bighorn ewes into disjoint seasonal subpopulations, and decomposed the variance in lamb survival to weaning into components associated with individual ewes, subpopulations, populations and years. During epidemics, lamb survival varied substantially more between ewe-subpopulations than across populations or years, suggesting localized pathogen transmission. This pattern of lamb survival was not observed during years when disease was absent. Additionally, group sizes in ewe-subpopulations were independent of population size, but the number of ewe-subpopulations increased with population size. Consequently, although one might reasonably assume that force of infection for this highly communicable disease scales with population size, in fact, host social behaviour modulates transmission such that disease is frequency-dependent within populations, and some groups remain protected during epidemic events. [ABSTRACT FROM AUTHOR]