Droughts reshape apex predator space use and intraguild overlap.

Droughts are increasing in frequency and severity globally due to climate change, leading to changes in resource availability that may have cascading effects on animal ecology. Resource availability is a key driver of animal space use, which in turn influences interspecific interactions like intraguild competition. Understanding how climate-induced changes in resource availability influence animal space use, and how species-specific responses scale up to affect intraguild dynamics, is necessary for predicting broader community-level responses to climatic changes. Although several studies have demonstrated the ecological impacts of drought, the behavioural responses of individuals that scale up to these broader-scale effects are not well known, particularly among animals in top trophic levels like large carnivores. Furthermore, we currently lack understanding of how the impacts of climate variability on individual carnivore behaviour are linked to intraguild dynamics, in part because multi-species datasets collected at timescales relevant to climatic changes are rare. Using 11 years of GPS data from four sympatric large carnivore species in southern Africa-lions (Panthera leo), leopards (Panthera pardus), African wild dogs (Lycaon pictus) and cheetahs (Acinonyx jubatus)-spanning 4 severe drought events, we test whether drought conditions impact (1) large carnivore space use, (2) broad-scale intraguild spatial overlap and (3) fine-scale intraguild interactions. Drought conditions expanded space use across species, with carnivores increasing their monthly home range sizes by 35% (wild dogs) to 66% (leopards). Drought conditions increased the amount of spatial overlap between lions and subordinate felids (cheetahs and leopards) by up to 119%, but only lion-cheetah encounter rates were affected by these changes, declining in response to drought. Our findings reveal that drought has a clear signature on the space use of multiple sympatric large carnivore species, which can alter spatiotemporal partitioning between competing species. Our study thereby illuminates the links between environmental change, animal behaviour and intraguild dynamics. While fine-scale avoidance strategies may facilitate intraguild coexistence during periodic droughts, large carnivore conservation may require considerable expansion of protected areas or revised human-carnivore coexistence strategies to accommodate the likely long-term increased space demands of large carnivores under projected increases in drought intensity.

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'.

Anthropogenic impacts at the interface of animal spatial and social behaviour.

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'.

Evidence for seasonal migration by a cryptic top predator of the deep sea.

BACKGROUND: In ecosystems influenced by strong seasonal variation in insolation, the fitness of diverse taxa depends on seasonal movements to track resources along latitudinal or elevational gradients. Deep pelagic ecosystems, where sunlight is extremely limited, represent Earth's largest habitable space and yet ecosystem phenology and effective animal movement strategies in these systems are little understood. Sperm whales (Physeter macrocephalus) provide a valuable acoustic window into this world: the echolocation clicks they produce while foraging in the deep sea are the loudest known biological sounds on Earth and convey detailed information about their behavior. METHODS: We analyze seven years of continuous passive acoustic observations from the Central California Current System, using automated methods to identify both presence and demographic information from sperm whale echolocation clicks. By integrating empirical results with individual-level movement simulations, we test hypotheses about the movement strategies underlying sperm whales' long-distance movements in the Northeast Pacific. RESULTS: We detect foraging sperm whales of all demographic groups year-round in the Central California Current System, but also identify significant seasonality in frequency of presence. Among several previously hypothesized movement strategies for this population, empirical acoustic observations most closely match simulated results from a population undertaking a "seasonal resource-tracking migration", in which individuals move to track moderate seasonal-latitudinal variation in resource availability. DISCUSSION: Our findings provide evidence for seasonal movements in this cryptic top predator of the deep sea. We posit that these seasonal movements are likely driven by tracking of deep-sea resources, based on several lines of evidence: (1) seasonal-latitudinal patterns in foraging sperm whale detection across the Northeast Pacific; (2) lack of demographic variation in seasonality of presence; and (3) the match between simulations of seasonal resource-tracking migration and empirical results. We show that sperm whales likely track oceanographic seasonality in a manner similar to many surface ocean predators, but with dampened seasonal-latitudinal movement patterns. These findings shed light on the drivers of sperm whales' long-distance movements and the shrouded phenology of the deep-sea ecosystems in which they forage.

Increasing environmental variability inhibits evolutionary rescue in a long-lived vertebrate.

Evolutionary rescue, whereby adaptive evolutionary change rescues populations from extinction, is theorized to enable imperiled animal populations to persist under increasing anthropogenic change. Despite a large body of evidence in theoretical and laboratory settings, the potential for evolutionary rescue to be a viable adaptation process for free-ranging animals remains unknown. Here, we leverage a 38-year dataset following the fates of 53,959 Magellanic penguins (Spheniscus magellanicus) to investigate whether a free-ranging vertebrate species can morphologically adapt to long-term environmental change sufficiently to promote population persistence. Despite strong selective pressures, we found that penguins did not adapt morphologically to long-term environmental changes, leading to projected population extirpation. Fluctuating selection benefited larger penguins in some environmental contexts, and smaller penguins in others, ultimately mitigating their ability to adapt under increasing environmental variability. Under future climate projections, we found that the species cannot be rescued by adaptation, suggesting similar constraints for other long-lived species. Such results reveal how fluctuating selection driven by environmental variability can inhibit adaptation under long-term environmental change. Our eco-evolutionary approach helps explain the lack of adaptation and evolutionary rescue in response to environmental change observed in many animal species.

Global expansion of human-wildlife overlap in the 21st century.

Understanding the extent to which people and wildlife overlap in space and time is critical for the conservation of biodiversity and ecological services. Yet, how global change will reshape the future of human-wildlife overlap has not been assessed. We show that the potential spatial overlap of global human populations and 22,374 terrestrial vertebrate species will increase across ~56.6% and decrease across only ~11.8% of the Earth's terrestrial surface by 2070. Increases are driven primarily by intensification of human population densities, not change in wildlife distributions caused by climate change. The strong spatial heterogeneity of future human-wildlife overlap found in our study makes it clear that local context is imperative to consider, and more targeted area-based land-use planning should be integrated into systematic conservation planning.

Spatial match-mismatch between predators and prey under climate change.

Climate change is driving a rapid redistribution of life on Earth. Variability in the rates, magnitudes and directions of species' shifts can alter spatial overlap between predators and prey, with the potential to decouple trophic interactions. Although phenological mismatches between predator requirements and prey availability under climate change are well-established, 'spatial match-mismatch' dynamics remain poorly understood. We synthesize global evidence for climate-driven changes in spatial predator-prey overlap resulting from species redistribution across marine and terrestrial domains. We show that spatial mismatches can have vastly different outcomes for predator populations depending on their diet specialization and role within the wider ecosystem. We illustrate ecosystem-level consequences of climate-driven changes in spatial predator-prey overlap, from restructuring food webs to altering socio-ecological interactions. It remains unclear how predator-prey overlap at the landscape scale relates to prey encounter and consumption rates at local scales, or how the spatial reorganization of food webs affects ecosystem function. We identify key research directions necessary to resolve the scale of ecological impacts caused by species redistribution under climate change.

Long-distance communication can enable collective migration in a dynamic seascape.

Social information is predicted to enhance the quality of animals' migratory decisions in dynamic ecosystems, but the relative benefits of social information in the long-range movements of marine megafauna are unknown. In particular, whether and how migrants use nonlocal information gained through social communication at the large spatial scale of oceanic ecosystems remains unclear. Here we test hypotheses about the cues underlying timing of blue whales' breeding migration in the Northeast Pacific via individual-based models parameterized by empirical behavioral data. Comparing emergent patterns from individual-based models to individual and population-level empirical metrics of migration timing, we find that individual whales likely rely on both personal and social sources of information about forage availability in deciding when to depart from their vast and dynamic foraging habitat and initiate breeding migration. Empirical patterns of migratory phenology can only be reproduced by models in which individuals use long-distance social information about conspecifics' behavioral state, which is known to be encoded in the patterning of their widely propagating songs. Further, social communication improves pre-migration seasonal foraging performance by over 60% relative to asocial movement mechanisms. Our results suggest that long-range communication enhances the perceptual ranges of migrating whales beyond that of any individual, resulting in increased foraging performance and more collective migration timing. These findings indicate the value of nonlocal social information in an oceanic migrant and suggest the importance of long-distance acoustic communication in the collective migration of wide-ranging marine megafauna.

Hunting mode and habitat selection mediate the success of human hunters.

BACKGROUND: As a globally widespread apex predator, humans have unprecedented lethal and non-lethal effects on prey populations and ecosystems. Yet compared to non-human predators, little is known about the movement ecology of human hunters, including how hunting behavior interacts with the environment. METHODS: We characterized the hunting modes, habitat selection, and harvest success of 483 rifle hunters in California using high-resolution GPS data. We used Hidden Markov Models to characterize fine-scale movement behavior, and k-means clustering to group hunters by hunting mode, on the basis of their time spent in each behavioral state. Finally, we used Resource Selection Functions to quantify patterns of habitat selection for successful and unsuccessful hunters of each hunting mode. RESULTS: Hunters exhibited three distinct and successful hunting modes ("coursing", "stalking", and "sit-and-wait"), with coursings as the most successful strategy. Across hunting modes, there was variation in patterns of selection for roads, topography, and habitat cover, with differences in habitat use of successful and unsuccessful hunters across modes. CONCLUSIONS: Our study indicates that hunters can successfully employ a diversity of harvest strategies, and that hunting success is mediated by the interacting effects of hunting mode and landscape features. Such results highlight the breadth of human hunting modes, even within a single hunting technique, and lend insight into the varied ways that humans exert predation pressure on wildlife.

Increasing ambient temperatures trigger shifts in activity patterns and temporal partitioning in a large carnivore guild.

Shifts in species' interactions are implicated as an important proximate cause underpinning climate-change-related extinction. However, there is little empirical evidence on the pathways through which climate conditions, such as ambient temperature, impact community dynamics. The timing of activities is a widespread behavioural adaptation to environmental variability, and temporal partitioning is a key mechanism that facilitates coexistence, especially within large carnivore communities. We investigated temperature impacts on community dynamics through its influence on the diel activity of, and temporal partitioning amongst, four sympatric species of African large carnivores: lions (Panthera leo), leopards (Panthera pardus), cheetahs (Acinonyx jubatus) and African wild dogs (Lycaon pictus). Activity of all species was shaped by a combination of light availability and temperature, with most species becoming more nocturnal and decreasing activity levels with increasing temperatures. A nocturnal shift was most pronounced in cheetahs, the most diurnal species during median temperatures. This shift increased temporal overlap between cheetahs and other carnivore species by up to 15.92%, highlighting the importance of considering the responses of interacting sympatric species when inferring climate impacts on ecosystems. Our study provides evidence that temperature can significantly affect temporal partitioning within a carnivore guild by generating asymmetrical behavioural responses amongst functionally similar species.

Selection of planning unit size in dynamic management strategies to reduce human-wildlife conflict.

Conservation planning traditionally relies upon static reserves; however, there is increasing emphasis on dynamic management (DM) strategies that are flexible in space and time. Due to its novelty, DM lacks best practices to guide design and implementation. We assessed the effect of planning unit size in a DM tool designed to reduce entanglement of protected whales in vertical ropes of surface buoys attached to crab traps in the lucrative U.S. Dungeness crab (Metacarcinus magister) fishery. We conducted a retrospective analysis from 2009 to 2019 with modeled distributions of blue (Balaenoptera musculus) and humpback (Megaptera novaeangliae) whales and observed fisheries effort and revenue to evaluate the effect of 7 planning unit sizes on DM tool performance. We measured performance as avoided whale entanglement risk and protected fisheries revenue. Small planning units avoided up to $47 million of revenue loss and reduced entanglement risk by up to 25% compared to the large planning units currently in use by avoiding the incidental closure of areas with low biodiversity value and high fisheries revenue. However, large planning units were less affected by an unprecedented marine heat wave in 2014-2016 and by delays in information on the distributions of whales and the fishery. Our findings suggest that the choice of planning unit size will require decision-makers to navigate multiple socioecological considerations-rather than a one-size-fits-all approach-to separate wildlife from threats under a changing climate.

Selección del tamaño de la unidad de planeación en las estrategias dinámicas de manejo para reducir el conflicto humano-fauna Resumen La planeación de la conservación depende por tradición de las reservas estáticas; sin embargo, cada vez hay más énfasis en estrategias de manejo dinámico (MD) que son flexibles con el tiempo y el espacio. Ya que es novedoso, el MD carece de buenas prácticas que guíen el diseño y la implementación. Analizamos el efecto del tamaño de la unidad de planeación en una herramienta de MD diseñada para reducir el número de ballenas que se enredan en las cuerdas verticales de las boyas amarradas a las trampas para cangrejos de la pesquería lucrativa del cangrejo Dungeness (Metacarcinus magister) en los Estados Unidos. Realizamos un análisis retrospectivo de 2009 a 2019 con modelos de distribución de la ballena azul (Balaenoptera musculus) y la ballena jorobada (Megaptera novaeangliae) y observamos los esfuerzos y ganancias de la pesquería para evaluar el efecto del tamaño de siete unidades de planeación sobre el desempeño de una herramienta de MD. Medimos el desempeño como el riesgo de enredamiento evitado y los ingresos protegidos de la pesquería. Las unidades pequeñas de planeación evitaron hasta $47 millones de ingresos perdidos y redujeron el riesgo de enredamiento hasta en 25% en comparación con las unidades grandes que se usan actualmente al evitar el cierre indirecto de áreas con un valor bajo de biodiversidad e ingresos elevados para la pesquería. Sin embargo, las unidades grandes de planeación estuvieron menos afectadas por una ola de calor marino sin precedentes entre 2014 y 2016 y por los retrasos en la información sobre la distribución de las ballenas y la pesquería. Nuestros hallazgos sugieren que la selección del tamaño de la unidad de planeación requerirá que el órgano decisorio navegue múltiples consideraciones socio-ecológicas-en lugar de un enfoque de un-tamaño-para-todos-para separar a la fauna de las amenazas bajo el clima cambiante.

Climate presses and pulses mediate the decline of a migratory predator.

Long-term climate changes and extreme climate events differentially impact animal populations, yet whether and why these processes may act synergistically or antagonistically remains unknown. Disentangling these potentially interactive effects is critical for predicting population outcomes as the climate changes. Here, we leverage the "press-pulse" framework, which is used to describe ecological disturbances, to disentangle population responses in migratory Magellanic penguins to long-term changes in climate means and variability (presses) and extreme events (pulses) across multiple climate variables and life history stages. Using an unprecedented 38-y dataset monitoring 53,959 penguins, we show for the first time that the presses and pulses of climate change mediate the rate of population decline by differentially impacting different life stages. Moreover, we find that climate presses and pulses can work both synergistically and antagonistically to affect animal population persistence, necessitating the need to examine both processes in concert. Negative effects of terrestrial heat waves (pulses) on adult survival, for example, were countered by positive effects of long-term changes in oceanographic conditions in migratory grounds (presses) on juvenile and adult survival. Taken together, these effects led to predicted population extirpation under all future climate scenarios. This work underscores the importance of a holistic approach integrating multiple climate variables, life stages, and presses and pulses for predicting the persistence of animals under accelerating climate change.

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.

The influence of social cues on timing of animal migrations.

Animal migration plays a central role in many ecological and evolutionary processes, yet migratory populations worldwide are increasingly threatened. Adjusting migration timing to match ecosystem phenology is key to survival in dynamic and changing ecosystems, especially in an era of human-induced rapid environmental change. Social cues are increasingly recognized as major components of migratory behaviour, yet a comprehensive understanding of how social cues influence the timing of animal migrations remains elusive. Here, we introduce a framework for assessing the role that social cues, ranging from explicit (for example, active cueing) to implicit (for example, competition), play in animals' temporal migration decisions across a range of scales. By applying this theoretical lens to a systematic review of published literature, we show that a broad range of social cues frequently mediate migration timing at a range of temporal scales and across highly diverse migratory taxa. We further highlight that while rarely documented, several social cue mechanisms (for example, social learning and density dependency) play important adaptive roles in matching migration timing with ecosystem dynamics. Thus, social cues play a fundamental role in migration timing, with potentially widespread ecological consequences and implications for the conservation of migratory species. Furthermore, our analysis establishes a theoretical basis on which to evaluate future findings on the role of both conspecific and interspecific social cues in this intersection of behavioural ecology and global change biology.

Blue whales increase feeding rates at fine-scale ocean features.

Marine predators face the challenge of reliably finding prey that is patchily distributed in space and time. Predators make movement decisions at multiple spatial and temporal scales, yet we have a limited understanding of how habitat selection at multiple scales translates into foraging performance. In the ocean, there is mounting evidence that submesoscale (i.e. less than 100 km) processes drive the formation of dense prey patches that should hypothetically provide feeding hot spots and increase predator foraging success. Here, we integrated environmental remote-sensing with high-resolution animal-borne biologging data to evaluate submesoscale surface current features in relation to the habitat selection and foraging performance of blue whales in the California Current System. Our study revealed a consistent functional relationship in which blue whales disproportionately foraged within dynamic aggregative submesoscale features at both the regional and feeding site scales across seasons, regions and years. Moreover, we found that blue whale feeding rates increased in areas with stronger aggregative features, suggesting that these features indicate areas of higher prey density. The use of fine-scale, dynamic features by foraging blue whales underscores the need to take these features into account when designating critical habitat and may help inform strategies to mitigate the impacts of human activities for the species.

Long-term, climate-driven phenological shift in a tropical large carnivore.

Understanding the degree to which animals are shifting their phenology to track optimal conditions as the climate changes is essential to predicting ecological responses to global change. Species at low latitudes or high trophic levels are theoretically expected to exhibit weaker phenological responses than other species, but limited research on tropical systems or on top predators impedes insight into the contexts in which these predictions are upheld. Moreover, a lack of phenological studies on top predators limits understanding of how climate change impacts propagate through entire ecosystems. Using a 30-y dataset on endangered African wild dogs (Lycaon pictus), we examined changes in reproductive phenology and temperatures during birthing and denning over time, as well as potential fitness consequences of these changes. We hypothesized that their phenology would shift to track a stable thermal range over time. Data from 60 packs and 141 unique pack-years revealed that wild dogs have delayed parturition by 7 days per decade on average in response to long-term warming. This shift has led to temperatures on birthing dates remaining relatively stable but, contrary to expectation, has led to increased temperatures during denning periods. Increased denning temperatures were associated with reduced reproductive success, suggesting that a continued phenological shift in the species may become maladaptive. Such results indicate that climate-driven shifts could be more widespread in upper trophic levels than previously appreciated, and they extend theoretical understanding of the species traits and environmental contexts in which large phenological shifts can be expected to occur as the climate changes.

A spatial capture-recapture model for group-living species.

Group living in species can have complex consequences for individuals, populations, and ecosystems. Therefore, estimating group density and size is often essential for understanding population dynamics, interspecific interactions, and conservation needs of group-living species. Spatial capture-recapture (SCR) has been used to model both individual and group density in group-living species, but modeling either individual-level or group-level detection results in different biases due to common characteristics of group-living species, such as highly cohesive movement or variation in group size. Furthermore, no SCR method currently estimates group density, individual density, and group size jointly. Using clustered point processes, we developed a cluster SCR model to estimate group density, individual density, and group size. We compared the model to standard SCR models using both a simulation study and a data set of detections of African wild dogs (Lycaon pictus), a group-living carnivore, on camera traps in northern Botswana. We then tested the model's performance under various scenarios of group movement in a separate simulation study. We found that the cluster SCR model outperformed a standard group-level SCR model when fitted to data generated with varying group sizes, and mostly recovered previous estimates of wild dog group density, individual density, and group size. We also found that the cluster SCR model performs better as individuals' movements become more correlated with their groups' movements. The cluster SCR model offers opportunities to investigate ecological hypotheses relating group size to population dynamics while accounting for cohesive movement behaviors in group-living species.

An integrated path for spatial capture-recapture and animal movement modeling.

Ecologists and conservation biologists increasingly rely on spatial capture-recapture (SCR) and movement modeling to study animal populations. Historically, SCR has focused on population-level processes (e.g., vital rates, abundance, density, and distribution), whereas animal movement modeling has focused on the behavior of individuals (e.g., activity budgets, resource selection, migration). Even though animal movement is clearly a driver of population-level patterns and dynamics, technical and conceptual developments to date have not forged a firm link between the two fields. Instead, movement modeling has typically focused on the individual level without providing a coherent scaling from individual- to population-level processes, whereas SCR has typically focused on the population level while greatly simplifying the movement processes that give rise to the observations underlying these models. In our view, the integration of SCR and animal movement modeling has tremendous potential for allowing ecologists to scale up from individuals to populations and advancing the types of inferences that can be made at the intersection of population, movement, and landscape ecology. Properly accounting for complex animal movement processes can also potentially reduce bias in estimators of population-level parameters, thereby improving inferences that are critical for species conservation and management. This introductory article to the Special Feature reviews recent advances in SCR and animal movement modeling, establishes a common notation, highlights potential advantages of linking individual-level (Lagrangian) movements to population-level (Eulerian) processes, and outlines a general conceptual framework for the integration of movement and SCR models. We then identify important avenues for future research, including key challenges and potential pitfalls in the developments and applications that lie ahead.

Ontogenetic shifts from social to experiential learning drive avian migration timing.

Migrating animals may benefit from social or experiential learning, yet whether and how these learning processes interact or change over time to produce observed migration patterns remains unexplored. Using 16 years of satellite-tracking data from 105 reintroduced whooping cranes, we reveal an interplay between social and experiential learning in migration timing. Both processes dramatically improved individuals' abilities to dynamically adjust their timing to track environmental conditions along the migration path. However, results revealed an ontogenetic shift in the dominant learning process, whereby subadult birds relied on social information, while mature birds primarily relied on experiential information. These results indicate that the adjustment of migration phenology in response to the environment is a learned skill that depends on both social context and individual age. Assessing how animals successfully learn to time migrations as environmental conditions change is critical for understanding intraspecific differences in migration patterns and for anticipating responses to global change.