Behavioral Ecology

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.

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 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.

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.

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.

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.

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.

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 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.

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.

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.

1. Organisms rely on environmental cues, such as moonlight illumination and weather, to modulate nocturnal behaviors that reduce predation risk and conserve energy during demanding periods. Urbanization can alter the reliability of cues, potentially impairing organisms' ability to respond appropriately with consequences in increasingly urbanized landscapes. 2. We leveraged the Motus Wildlife Tracking System to identify roost initiation, roost departure and nocturnal restlessness of dark-eyed juncos (Junco hyemalis) overwintering along an urban to rural gradient to determine if activity patterns are affected by weather, moonlight illumination, and urbanization. 3. Nocturnal behavior of overwintering juncos varied with environmental factors, including moonlight illumination and temperature. However, the strength of these responses differed among sites, suggesting that natural cues of moonlight illumination were masked in urban areas, where individuals were more responsive to temperature. 4. Our results suggest that urbanization weakens birds' responsiveness to natural light cues, likely because artificial lighting reduces the visibility of moonlight; representing a key mechanism by which human-altered landscapes can disrupt nocturnal ecology in birds.

Foraging hematophagous flies such as the stable fly, Stomoxys calcitrans, are shown or expected to sense multiple characteristics of visual host cues. These include relative intensity, spectral composition, and the angle and degree of polarization. Here, we investigated the role of each of these visual cue characteristics on stable fly attraction. In laboratory two-choice bioassays, we offered groups of female or male stable flies a choice between paired photostimuli that differed in their visual characteristics, recording the flies' landing responses on stimuli as the response criterion. Bioassay flies preferentially selected lower-intensity visual stimuli even when paired stimuli markedly differed in spectral composition and polarization characteristics (a high degree of horizontal or vertical polarization). When paired stimuli had identical relative intensity, flies failed to discriminate between the stimulus that presented ultraviolet, blue, green, and red wavebands and a corresponding paired stimulus lacking one of these wavebands. Our results support the hypothesis that foraging stable flies may respond to colour and polarization characteristics of visual stimuli as modulations of perceived intensity. Acquired knowledge on the mechanisms that underlie stable fly visual attraction may inform the design of vision-based traps so that they become optimally attractive to stable flies and other biting flies.

When mating among genetically related individuals results in inbreeding depression, various behaviors and life-history traits evolve to favor outbreeding. Long-range sexual communication following natal dispersal is one such adaptation. In parasitoid wasps with single-locus complementary sex determination (sl-CSD), inbreeding increases the production of unviable or sterile diploid males. Mastrus ridens is such a species, with a genetic load underpinned by sl-CSD, combined with a high potential for sibmating because siblings emerge in close proximity from the same host and become sexually active shortly after emergence. With this context in mind, we investigated the mate-finding strategy of M. ridens using complementary behavioral, chemical, and electrophysiological approaches. Wind-tunnel bioassays and field experiments showed that males orient toward virgin females and, to a lesser extent, mated females. This attraction from a distance appeared to be the sole form of conspecific attraction. Chemical analyses using static and dynamic headspace sampling and solvent extracts, followed by GC-MS, revealed several potential pheromonal candidates, including methyl 6-methylsalicylate. This compound elicited reproducible antennal responses in males in GC-EAD recordings. Altogether, our results suggest that mate finding in M. ridens is guided by a volatile sex pheromone emitted by females, with methyl 6-methylsalicylate as one putative component. Such mate-finding mechanisms promote outbreeding and may explain the low proportion of unfit diploid males observed in natural populations. Surprisingly, chemical analyses suggested that males also emit this compound, whereas neither males nor females showed behavioral responses to males. This apparent discrepancy suggests that attraction may depend on a more complex pheromonal signal and opens further avenues for investigation.

Fluctuating asymmetry (FA) is widely used as a proxy for developmental instability, although reduced rather than increased FA, has occasionally been reported under particularly challenging conditions, possibly because environmental filtering removes less developmentally stable individuals. Here, we investigated temporal variation in wing shape asymmetry in males of Drosophila sturtevanti collected in 2022, 2024, and 2026 at the same sampling site, an urban Atlantic Forest fragment. Sampling in 2022 and 2026 were completed during single field expeditions in early April, whereas in 2024 repeated collections were required until mid-May because of markedly low capture success. We therefore examined whether this difference was associated with unusual environmental conditions and whether FA was higher in 2024, as expected under greater developmental perturbation. Daily temperature and relative humidity were examined from early March onward, encompassing approximately one month before adult emergence. The 2024 sampling period was warmer and drier than the corresponding periods in 2022 and 2026 and coincided with the final phase of the strong 2023-2024 El Ni&ntildeo event. Contrary to our prediction, FA was significantly lower in 2024. The magnitude of directional asymmetry remained similar among years, although the spatial pattern of the DA vector differed between 2024 and 2026. The coincidence of reduced capture success and reduced FA is consistent with the hypothesis that adverse conditions may impose demographic filtering, such that individuals expressing greater developmental instability are less likely to contribute to the sampled adult population.

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.

Whether sexual selection facilitates or hinders adaptation to novel environments is a question of fundamental importance due to current rapid environmental change. Using phylogenetic comparative analysis in 42 Drosophila species, we found that sperm length - a sexually selected ornament - positively associates with the ability of males to survive high temperatures. However, the ability to remain fertile does not increase with sperm length to the same extent, meaning that longer sperm species experience a larger gap between temperatures that kill and temperatures that sterilize males. The findings are consistent with the idea that strong sexual selection on condition-dependent exaggerated traits may facilitate somatic resilience to environmental stressors, but that susceptibility to heat-induced infertility may be a cost of extreme sperm ornamentation. Overall, the data suggests multifaceted links between postcopulatory sexual selection and climate change.

Precocious reproduction, prior to the acquisition of the final adult morphology, occurs in a variety of animals and represents a distinct life-history strategy with important consequences for growth and survival. Larval reproduction is far rarer and usually restricted to groups that go through metamorphosis. In ctenophores, Carl Chun described dissogeny more than 130 years ago, a distinctive life-history pattern involving larval and adult reproductive periods separated by gonadal regression. Yet, the potential plasticity of larval reproduction in ctenophores remains unresolved. Here, we developed a method to follow and quantify reproductive output in Mnemiopsis leidyi larvae and tested environmental factors that may regulate precocious reproduction. We found that conspecific density strongly influences its onset: low-density, well-fed animals can bypass reproduction at the cydippid stage and develop rapidly into the lobed form, whereas higher density promotes precocious reproduction. Food availability and associated growth rate also contribute but cannot alone explain the observed differences. Once reproduction is established, however, high conspecific density suppresses reproductive output, an effect reproduced by exposure to seawater incubated with high M. leidyi densities, consistent with water-borne chemical communication. Salinity and light conditions further modulate spawning. Together, our results reveal substantial environmental plasticity in precocious reproduction in M. leidyi, reconcile long-standing contradictory observations of ctenophore development, and highlight how environmental cues can reshape reproductive life-history transitions.

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.

Heat-induced sterility is widespread across ectotherm species, yet its underlying mechanisms remain poorly understood. In mated females, thermal stress can disrupt female reproductive physiology while also affecting sperm stored within the reproductive tract, making the mechanisms of fertility loss difficult to distinguish. Here, we investigated the extent and recovery dynamics of female heat-induced fertility loss across seven Drosophila species. We tracked fertility after acute heat exposure, tested whether remating restored fertility, quantified stored sperm in two closely related species, and examined whether reproductive thermal sensitivity covaried with adult heat tolerance. We found that heat exposure reduced egg-hatching success in all seven species, but recovery differed among species. Fertility recovered within several days in most species, whereas D. simulans showed persistent declines and D. melanogaster showed limited recovery. Remating restored or maintained fertility in both species. Stored sperm declined through time in both D. simulans and D. mauritiana, but D. mauritiana retained more sperm at later sampling points, consistent with their contrasting patterns of fertility decline and maintenance. Across species, greater adult heat tolerance was associated with smaller immediate reductions in fertility, but not with cumulative reproductive loss across subsequent days. Together, these findings show that similar initial fertility losses can lead to markedly different recovery trajectories and highlight the importance of comparative, multi-species approaches that integrate female physiology, sperm-storage dynamics, mating behavior, and whole-organism thermal tolerance when assessing reproductive resilience to climate warming.