Behavioral Ecology and Sociobiology

Female-female competition is increasingly recognized as a key driver of female ornamentation, including birdsong, which often functions in intrasexual competition. However, the specific resources females use elaborate traits to compete for remain unclear. In addition, few studies have simultaneously investigated the use of multiple competitive traits in females, despite growing independent interest in these traits (e.g., female song and aggression). We investigated the competitive contexts of female song, aggression and calling behavior in northern house wrens (Troglodytes aedon) to determine which resources females compete for across the breeding season. We simulated conspecific territorial intrusions using female song at three breeding stages representing different contexts: arrival (mate and territory acquisition), nest building (nest site and breeding status defense), and egg laying (brood defense). We tested whether female song and physical aggression varied as reproductive resources shifted across the breeding cycle. Females were significantly more aggressive during nest building, showing 5.8 times greater odds of a higher-intensity aggressive response during nest building compared to arrival. Female song output was similar across early stages but declined during egg laying, though this was not statistically significant after correction for multiple comparisons and individuals varied substantially in overall singing propensity. Non-song vocalizations varied by call type and breeding stage. Calls associated with aggression occurred most frequently during nest building, consistent with peak physical aggression responses. Together, these results identify nest building as the stage of highest female aggression, consistent with heightened competition over nest cavities and associated breeding status in this cavity-nesting species. In contrast, female song occurred across all stages and appears to function in multiple competitive contexts. This study provides evidence for context and mode-specific female signaling in a temperate songbird and highlights that females strategically use aggression, calls, and song to mediate social conflict across breeding contexts.

Foraging and feeding are often associated with vocalisations across animal species. Food calls may function to claim ownership of resources or recruit others to feeding sites, yet their potential role in facilitating food sharing remains poorly understood. This study investigates the relationship between food calls and food-sharing behaviour in common marmosets (Callithrix jacchus), focusing on adult-infant interactions, where food transfers are most frequent. We analysed over 1000 food interactions across six groups, categorising events as non-sharing (without begging or despite begging) or sharing (proactive, resisted, or facilitated). We examined the spectro-temporal structure of food calls produced immediately before each interaction. Food transfers were more likely to occur when preceded by a food call, suggesting a functional link between food calls and sharing. Random forest models reliably classified food calls according to interaction type, while generalized linear mixed models identified acoustic variation associated with sharing outcomes. Our results suggest that food calls encode information to either encourage or deter conspecific (infant) approach, influencing food transfer outcomes. These findings highlight the role of food calls in modulating social interactions around food in marmosets, advancing our understanding of vocal communication and its integration with cooperative behaviours in primates.

Colony defence is critical for the survival of wild honeybees but presents a challenge for local populations and beekeepers. Apis mellifera scutellata is an important pollinator in its native range, a large area of Southern and Eastern Africa 1, but is reputed for its explosive temperament. Here, we present data demonstrating that this behaviour is only exhibited by a small fraction of colonies, while others are relatively docile. This sheds new light on this subspecies and on its likely evolutionary trajectory. We further characterize the different elements of this defensive response and compare them to that of A. m. carnica and A. m. mellifera. We show that each subspecies invests differently in the two steps of the behavioural sequence (alert and attack). Furthermore, our results suggest that the task of attacking is carried out by specialized workers in carnica, but not in the other subspecies tested, highlighting subtle differences in colony organization across subspecies. Finally, we discovered that defenders from A. m. scutellata and scutellata-hybrid colonies can locate a target that was stung up to 2 h and 3 h earlier, respectively, whereas European bees could only do so for 30 min. This long-term marking could confer an advantage against returning predators with few weak points, such as honey badgers and humans wearing protective clothing. Overall, we offer a fresh perspective on this controversial but important subspecies of honeybees, with implications for both future studies and daily management.

Primates maintain cooperative relationships largely through grooming reciprocity, yet studies disagree on the timescales over which grooming is returned. Part of this disagreement stems from statistical issues: inconsistent definitions of a reciprocation window, combined with the exclusion of unreciprocated grooming, leave the debate unresolved. We still do not know if primates exchange grooming within the short windows predicted by biological market theory or over much longer timescales. Whilst survival analysis can identify reciprocation windows objectively by retaining unreciprocated grooming as censored data, previous studies have applied it inconsistently. This inconsistency leaves the true time frames over which primates reciprocate grooming somewhat blurred. Here, we use a statistical approach that addresses these problems to examine grooming reciprocity in crested macaques (Macaca nigra), a socially tolerant macaque species. We applied survival analysis to identify each group's short-term reciprocation window without imposing arbitrary cut offs, and Bayesian survival models to test whether social bond strength and eigenvector centrality predicted reciprocation latency. Short-term reciprocation occurred rarely (29% of events in one group and 5% in the other); most reciprocation occurred over longer timescales than previous research commonly assumes. Stronger bonds predicted faster reciprocation; this finding suggests that crested macaques express social investment in valued partners through how promptly they return grooming, meaning that the timing of exchange carries information about relationship value. Eigenvector centrality showed no credible effect, though our small sample size and dense groups prevent us from interpreting this outcome more robustly. Substantial variation between dyads remained unexplained, pointing to partnerspecific properties we have yet to identify. These findings caution against assuming that reciprocity operates over short windows. They show that social predictors, such as bond strength, remain essential to understanding when primates return grooming. This work provides a baseline for testing whether grooming is exchanged for other social commodities, and for comparing reciprocation timescales across macaque species that differ in social tolerance.

Resource handling time is an important cost to foraging animals and this is especially true for low-ranking individuals who may be displaced before they acquire rewards. Handling may also require visual attention that can lead to a trade-off between scanning the environment and focusing on food acquisition. We ran a foraging experiment on wild vervet monkeys where we simultaneously offered them three puzzle boxes in a straight-line array that increased sequentially in handling difficulty and reward value. We found that differences in competitive ability had a large impact on access to long-handling time resources and willingness to invest time in costly problem-solving when social interference was possible. Resources with lower handling time and smaller rewards were accessed by individuals of all dominance ranks but only dominant monkeys that did not fear displacement attempted the high handling time puzzle box. Handling time improved with experience at each puzzle box and contrary to predictions, when vigilance rates were higher, monkeys were also faster at solving puzzle boxes. Thus, there is not necessarily a trade-off between attention to foraging tasks that require handling and vigilance, if a high scanning frequency to both can be balanced with limited scanning duration. These results have important implications for understanding the decision-making of foraging animals that live in competitive groups and the diets of individuals, which may vary with dominance when important resources require long handling times.

Schooling is a widespread behaviour among pelagic fish that reduces locomotion costs of migration and predation risk. It is characterised by coordinated movements, speed and orientation of group members. Although schooling can vary both between populations and among individuals of the same population, these levels of variation are seldom considered together. Here, we investigated whether schooling tendencies in three-spined sticklebacks (Gasterosteus aculeatus) vary between populations and among individuals within-populations depending on their phenotype. We first compared the schooling behaviour of anadromous (migrants) and recently landlocked (residents) populations and tested whether individual sociability explained within-population variation in schooling. We then studied whether schooling behaviour is modulated when migrants and residents are in mixed groups. Migrant groups displayed better schooling performance than residents with higher alignment. Within migrant groups, less social individuals displayed more leadership, were better aligned, swam closer to others and fatigued less than more social fish. These patterns were absent in residents. When the two populations were mixed, overall schooling performance declined, particularly in migrants, with schools exhibiting reduced compaction and alignment, resembling resident groups. Thus, individuals did not modulate their schooling behaviour to the group social composition and phenotype-dependent schooling patterns were disrupted in migrants when mixed. These findings suggest that migration-related behaviours can disappear quickly post-isolation. They also highlight the importance of individual sociability in self-organisation and shaping collective behaviour, especially in populations that retain migratory behaviour. Considering behavioural variation across multiple hierarchical levels is thus key to understanding complex social systems.

Immune defenses may vary across environments due to differences in life history or abiotic and biotic characteristics, but few studies have compared the strength of immune responses across broad geographical gradients. Here we investigated how bacteria killing ability (BKA), a measure of constitutive innate immune function, varies among populations of tree swallows (Tachycineta bicolor) inhabiting different environments. We hypothesized that populations living in unpredictable environments, which also have shorter breeding seasons and thus fewer opportunities to renest in the event of reproductive failure, would invest in reproduction at the expense of immunity. To test this hypothesis, we measured BKA and monitored reproductive activity in four populations located across most of the breeding range of tree swallows, in Tennessee, New York, Wyoming and Alaska. Consistent with our predictions, BKA was lowest in Alaska and Wyoming and highest in Tennessee. In most populations BKA was also lower during incubation than during nestling provisioning, and lower in males than in females. These results suggest that the high reproductive investment seen in populations experiencing more unpredictable breeding conditions may come at a cost to immune function. While reduced immune investment could increase susceptibility to parasites, this potential cost is likely offset by lower parasite pressure in these environments, limiting the need for maintaining strong constitutive innate defenses like BKA.

Social animals must identify, evaluate, and appropriately respond to other individuals, often relying on multiple sources of information to guide context-dependent behavior. While individual recognition is thought to be widespread and critical for many social interactions (e.g. territoriality, dominance hierarchies), convincing demonstrations across animals beyond class-level recognition remain rare. Further, the cues animals use to recognize individuals remains poorly understood, especially in natural environments where they are exposed to numerous cues which may require cue integration. Here, we examine how socially dominant males of the African cichlid fish, Astatotilapia burtoni, use individual recognition to modulate aggression and whether this process depends on the spatial context. Using naturalistic experimental setups, we exposed males to familiar and unfamiliar neighbors while systematically manipulating spatial location and presenting a neighbor either singly or simultaneously to simulate competing social cues. Additionally, we measured circulating testosterone to assess endocrine correlates of recognition and aggression. We found that focal males habituated to familiar neighbors only when they remained in a consistent territorial location, demonstrating that spatial context is important for recognition. Further, when familiar and unfamiliar neighbors were presented simultaneously, males preferentially aggressed toward unfamiliar rival and adjust their aggressive response to the perceived threat of their neighbors, indicating that identity-based threat assessment is maintained under competing social demands. Together, our results show that recognition is a flexible, multi-faceted process that depends on the integration of identity and contextual information to guide territorial decision-making under dynamic social conditions.

Vertebrates cope with rapid temperature fluctuations by plastically adjusting their behaviour and physiology. This phenotypic plasticity is partly mediated by plastic changes in glucocorticoid levels; however, whether individual differences in endocrine plasticity are associated with fitness remains unknown. Here, we examined the relationship between individual variation in glucocorticoid plasticity in response to air temperature changes and reproductive success in a wild great tit population (Parus major). Over nine years, we collected repeated measurements of baseline and stress-induced corticosterone during nestling rearing. We found consistent reaction norm patterns in hormone response to temperature variation -- as shown in a previous study conducted over a shorter period. This provides robust evidence of individual differences in average hormone levels (intercepts) and the strength of the plastic response (slopes), indicating potential for evolution. However, neither intercepts nor slopes were associated with reproductive success (fledgling number or brood mass), suggesting that individual variation in endocrine plasticity was not under natural selection, at least within the environmental conditions experienced. This study provides a first empirical test of the evolutionary implications of endocrine plasticity in the wild and highlights the need to assess its fitness relevance under more extreme conditions, such as those expected under future climate scenarios.

Researcher disturbance can alter the behavior, survival, and breeding success of wildlife, with effects that are especially well-documented in waterbirds. Anecdotal reports suggest that disturbance can even cause cavity-nesting species to alter the structure of their nests. We used a long-term study of a burrowing seabird, the Leach's Storm-Petrel (Hydrobates leucorhous), to ask whether storm-petrels renovate existing burrows and whether such renovation is related to a history of researcher disturbance. Our study was conducted at the Bowdoin Scientific Station on Kent Island, New Brunswick, Canada, where in 2024 we measured the burrow architecture of 363 burrows, including length and structure for occupied and unoccupied burrows both inside and outside a long-term study site with consistent research activity. Of these, 150 burrows had been measured in 2006, allowing for direct longitudinal comparison of burrow architecture. Nearly all burrows changed in length over the 18-year study period (mean absolute change = 20.8 cm +- 18.2 SD). A linear model revealed two contrasting processes of burrow length change: more frequently occupied burrows grew longer, whereas initially longer burrows shortened towards mean burrow length (~50 cm) through collapse. Occupied burrows inside the long-term study site were on average 34% longer and significantly more complex than burrows adjacent to the site, suggesting that researcher disturbance directly or indirectly reshapes burrows. Overall, these results show that storm-petrel burrows are shaped by cycles of renovation and subsequent collapse that may be exacerbated by human disturbance. Given ongoing population declines in storm-petrels and other cavity-nesting seabirds, this study emphasizes the need to balance scientific research with minimizing disturbance, particularly in long-term studies.

Social aggregation is widespread in vertebrate larvae, yet how genetic relatedness shapes group cohesion remains poorly understood. In particular, the association between genetic similarity and social aggregation may depend on the scale at which genetic relatedness is evaluated. Here, we quantitatively examined aggregation behaviour in tadpoles from three lineages of Japanese toads, Bufo japonicus, B. formosus, and B. gargarizans miyakonis . We experimentally combined tadpole cohorts differing in relatedness and lineage and quantified aggregation using spatial dispersion analyses. We then examined relationships between aggregation behaviour and genetic variation using mitochondrial DNA (mtDNA), major histocompatibility complex (MHC) class II variation, and microsatellite diversity. Overall degree of aggregation did not differ significantly between sibling and non-sibling cohorts, and Morisitas index provided no evidence that tadpoles formed separate clumps according to relatedness. However, the correlation between genetic variation and aggregation differed across scales of genetic relatedness: among non-sibling cohorts, greater mtDNA pairwise distance was associated with weaker aggregation, whereas among sibling cohorts, greater MHC amino-acid difference was associated with weaker aggregation. These results indicate that the degree of aggregation is related to genetic similarity across different scales of relatedness, rather than being explained by a simple sibling/non-sibling distinction. Our findings suggest that tadpole group cohesion is related to genetic similarity in a scale-dependent manner, highlighting the importance of considering the scale of genetic relatedness when investigating social discrimination in vertebrates, while the sensory and molecular mechanisms underlying these relationships remain to be determined.

Foraging decisions often depend on net energy intake, but the need for specific micronutrients means that foods are not always comparable based on energy content. Although sodium is essential for many physiological functions, many organisms face challenges obtaining it, especially those relying on a plant-based diet. Flowering plants often lure pollinators by manipulating nectar carbohydrate content, but nectar sodium might also influence pollinator choice. We experimentally tested how sodium addition and nectar carbohydrate concentration interact to affect bumble bee foraging decisions by allowing bees to forage on artificial flowers containing either low- or high-sucrose nectar, with or without sodium. We also explored whether sugar and sodium content in floral nectar trade-off across plant species. Bees preferred sodium-enriched nectar and preference nearly doubled when sucrose concentration was low. Furthermore, bees switched more often from flowers without sodium to those with sodium when sucrose concentration was low. Given increased fidelity for artificial flowers with nectar sodium, sodium may enhance plant fitness by increasing conspecific pollen receipt. Yet despite behavioral evidence that sodium enhances the perceived quality of low-sucrose nectar for bees, we found no evidence for a nectar sugar-sodium trade-off across plant species. We discuss implications for plant-pollinator interactions and flower evolution.

Ground-nesting wild bees require suitable soil conditions for their below-ground development and overwintering life stages. Understanding how soil translocation and disturbance affect individuals and the following generations of bees is important for identifying conservation measures for ground-nesting wild bees. We investigated the effects of soil translocation and disturbance on the ground-nesting solitary spring mining bee Colletes cunicularius. In a replicated controlled design, we compared the effects of soil translocation with different degrees of disturbance on the number of surviving bees and the emergence time. We found that soil translocation and disturbance reduced bee survival but did not result in complete mortality, with some individuals surviving even under severe soil disturbance. There was no evidence that soil disturbance affected the timing of spring emergence, suggesting that disturbance is unlikely to alter the timing of nest construction and, consequently, the production of the next generation. Our results indicate that translocating soil containing overwintering bees could be a promising conservation measure to compensate for habitat loss caused by land conversion.

Sperm competition can occur when sperm from two or more males compete for fertilization of a batch of eggs. While it has been extensively studied in internal fertilizers, it remains poorly under-stood in external fertilizers. To investigate how sperm competition affects fertilization under sperm limitation, we conducted controlled experiments on two sea urchin species: Paracentrotus lividus and Dendraster excentricus. Eggs from a single female were exposed to sperm from three individ-ual males separately (non-competitive treatment) or to a pooled sample from three males (competi-tive treatment). We measured fertilization success as proportion of eggs fertilized after two hours and assessed sperm motility as percentage of motile sperm and curvilinear velocity. For P. lividus , we estimated the sperm concentrations where fertilization success was 50% of maximum observed fertilization success (F50), and the fertilization efficiency of the sperm (Fe), i.e. the likelihood that a sperm - egg encounter results in a fertilizing contact, using the fertilization kinetics model of Styan (1998). For D. excentricus, fertilization success was measured at a sperm concentration expected to produce 50% fertilization success. Sperm competition reduced fertilization success and increased F50 compared to non-competitive treatments, suggesting antagonistic effects that may compromise fertilization under sperm-limited conditions in external fertilizers.

Courtship is an essential component of reproductive success for many species and many courtship behaviors, such as vocalization, are energetically demanding. In order to successfully balance investment in courtship with feeding and foraging, individuals must be aware of their current energy balance, and hormonal signals of energetic state have been found to influence social signal effort across a variety of species. Ghrelin is one hormone that plays an important role in regulating energy balance by signaling hunger and promoting feeding. Previous studies have investigated the role of ghrelin in courtship, aggression, and mating behavior. In this study, we build on prior work by investigating the effects of ghrelin on ultrasonic vocalizations (USV) and other courtship behaviors of male mice. We expand on previous research by observing ghrelin's effects over longer interactions, allowing us to understand how ghrelin impacts the dynamics of USV production over time. Our results show that while males initially vocally court females at a high rate regardless of treatment, ghrelin reduces overall USV production by increasing the rate of decline of vocalization. Surprisingly, despite its effects on vocal courtship, we found no significant differences in the production or timing of other, non-vocal, courtship behaviors. This study deepens our understanding of the role of hormones traditionally associated with controlling hunger and feeding in regulating social behaviors.

1. Adaptive behavioural strategies require animals to simultaneously navigate social and ecological domains across multiple spatial and temporal scales. Although drones and computer vision have recently transformed the study of wild animal societies, many nocturnal species and those occupying structurally complex habitats remain inaccessible to these approaches, limiting our understanding of behaviour in natural settings. 2. We aimed to determine how behavioural strategies are organised across nested spatial and temporal scales within a wild communication network. 3. We used three-dimensional acoustic localisation and source separation to track individual male Hyperolius sp. A, a nocturnal African reed frog, within a natural rainforest chorus and quantify patterns of site fidelity, movement, spatial organisation, and call-timing interactions. 4. Males exhibited significant site fidelity across nights, while chorus spatial structure varied with local caller density. Within nights, individuals followed a stereotyped behavioural sequence, descending from elevated arboreal refugia before settling into lower calling positions near breeding sites. At finer temporal scales, call-timing interactions varied according to both local competitor density and the proximity of neighbouring rivals. 5. These findings demonstrate that behavioural strategies emerge across nested spatial and temporal scales and that long-term spatial positioning, short-term movement decisions, and moment-to-moment signalling interactions are tightly linked within natural communication networks. More broadly, acoustic localisation provides a powerful framework for studying behaviour in species and habitats that remain difficult to observe using conventional approaches.

Social relationships are strongly linked to health and biological fitness in humans and other animals. For example, social support may buffer individuals against harsh ecological conditions. Reciprocally, better health may allow for greater investment in social relationships. We examined links between health and social behaviour in an endangered population of northern bottlenose whales (Hyperoodon ampullatus). Using photo-identifications spanning 37 years, we visually classified whether individuals had poor body condition on each day they were observed (N = 3,086 whale-days). We then used generalized linear mixed models to explore how body condition varied with sex, age, and within-individual deviations in group size or other social metrics derived from Bayesian social networks. Approximately 11% of individuals were classified as being in poor condition at any time, though the prevalence of poor condition varied across raters and years. Visually assessed condition was successfully validated with morphometric measures, though there was only weak evidence that poor condition predicted apparent survival. Contrary to expectations, whales in poor condition were found in larger groups and had more social partners than their average across the study period. These findings run counter to the expectation that health and social connectedness are positively correlated in social mammals.

Feathers incorporated into bird nests are thought to provide multiple benefits, including thermal insulation and protection against nest-dwelling ectoparasites. For instance, in Swallows, it has been suggested that feathers influence infestations by Protocalliphora blow flies either by acting as a physical barrier between larvae and nestlings or through chemical compounds associated with preen oil. However, empirical evidence for an antiparasitic effect of feathers remains equivocal. Using 1,100 naturally infested Tree Swallow (Tachycineta bicolor) nests collected from a nest-box study system monitored in southern Quebec, Canada, between 2004 and 2018, we tested whether nest mass and feather abundance influenced hematophagous ectoparasites (Protocalliphora spp.) loads. We further assessed whether feather abundance directly affected body size using measurements of 1,513 P. sialia puparia, and emergence success using data from 961 Protocalliphora spp. puparia. Nest mass was positively associated with feather abundance, and both variables showed very weak positive relationships with parasite abundance. Feather abundance was unrelated to puparium size nor parasite emergence success. Our results provide no support for an anti-parasitic effect of feather lining within Tree Swallow nests. Using one of the largest datasets assembled to date on feather-lined nests and avian blow flies, we found no evidence that feather abundance impairs Protocalliphora density, development or emergence success. However, our findings do not preclude potential effects of feathers on other ectoparasites or microorganisms with different life histories. Future studies simultaneously monitoring multiple parasite taxa (such as fleas, mites, or bacteria) may therefore be needed to fully evaluate the ecological consequences of feather lining within avian nests.

1. Group foraging reduces the risks associated with foraging: larger groups dilute individual predation risk, allow group members to share the costs of vigilance, and provide a social structure for exchanging information about predators. The benefits of shared social information over simple safety in numbers should depend on the relationships among group members. Consequently, group composition should affect the selection of risky and safe habitats in social species. 2. We asked whether relationships among elk (Cervus canadensis) group members moving together influence their selection of risky habitats more than grouping alone. We tested three hypotheses. The familiarity hypothesis predicts that elk with close social ties are most likely to select risky habitats together. The inclusive fitness hypothesis predicts that the closest genetic relatives are most likely to select risky habitats together. The dilution hypothesis predicts that familiarity and genetic relationships do not influence habitat selection, with all group foraging benefits derived from proximity to others. 3. We used integrated Step Selection Analysis to test whether the identity of the closest group members influenced habitat selection. We tested whether the relationship of the nearest elk--either genetic or social--incentivized selection of risky habitat more than proximity to the nearest elk. 4. Group foraging clearly reduced risk, with elk more likely to select risky habitat when near others. Elk were also more likely to select risky habitats when near closer relatives, albeit with considerable variation among individuals. However, familiarity had a large and consistent effect: elk were far more likely to select risky habitats when near socially familiar group members. 5. Our results suggest that information shared within foraging groups, especially between socially familiar individuals, may lower the risks associated with habitat selection. Similarly, the benefits of reliable social information for reducing risk likely incentivizes the formation of stronger social ties within foraging groups. This positive feedback between social familiarity and habitat selection could thereby increase access to profitable habitats that are otherwise too risky, cementing what we term the social information landscape as a key driver of movement and habitat selection.

Social relationships can buffer individuals against environmental challenges, reducing the physiological costs of disturbance. Assessing how social and environmental conditions jointly shape physiological stress is critical for predicting individual responses to disturbance, particularly in managed populations. We evaluated whether social and environmental stressors act additively or whether social context modifies physiological responses to environmental challenges. We combined behavioural data and faecal glucocorticoid metabolite samples from a small, intensively monitored population of African elephants (Loxodonta africana) in a fenced reserve. Using a Bayesian framework, we tested how social factors interacted with variation in the biophysical environment, including natural environmental and anthropogenic disturbances, to predict glucocorticoid concentrations. Natural environmental variables, including weather, water availability, and season, did not predict glucocorticoid concentrations. In contrast, helicopter use, a common management intervention, was associated with higher glucocorticoid concentrations. Social effects were context dependent. Following disturbance, low-ranking individuals showed higher glucocorticoid concentrations than high-ranking individuals, whereas this pattern reversed under low-disturbance conditions. Individuals with more social connections had lower glucocorticoid concentrations, but this effect was evident only under non-disturbance conditions. Measures of betweenness centrality did not predict glucocorticoid levels. These results demonstrate that social factors mediate physiological responses to anthropogenic disturbance and that management actions can impose uneven physiological costs across individuals. Quantifying interactions between social factors and the biophysical environment may improve conservation and management strategies for social species. Integrating physiological responses to interactions between social factors and the biophysical environment can guide conservation strategies and improve management interventions in social species.