Animal Behaviour

Approaching conspecifics is a fundamental component of social interactions. Such decisions are typically investigated through the study of social relationships (e.g., kinship, dominance, affiliation) and their impact. In the framework, the role of social affordances, i.e., interaction opportunities arising from the immediate social environment, remains poorly understood. We investigated how context-dependent factors shape approach decisions in wild crested macaques (Macaca nigra) in North Sulawesi, Indonesia. From September 2024 to May 2025, we collected data on 27 adults (20 females, 7 males) using focal observations and an approach-specific protocol recording neighbours identities and behaviours at the initiation of close-proximity approaches (<1 body length). We first compared data collected during approaches with focal data, then conducted within-approach analyses contrasting approached partners and bystanders. We further tested the effects of behavioural state, subgroup size and composition, including infant presence. Bayesian models revealed that affiliative, neutral, and grooming behaviours were more frequent in approach contexts than in general observations. Within approaches, individuals avoided congeners already engaged in grooming and approached individuals in neutral or affiliative states (e.g., lip-smacking, embracing). Macaques globally approached more isolated individuals or dyads. When multiple subgroups were present (i.e., multiple choices), the likelihood of being approached increased with subgroup size and was higher in subgroups containing infants. These findings indicate that macaques use behavioural cues and local social density as affordance signals, adjusting approach decisions to maximize interaction opportunities and navigate complex social environments.

To make adaptive foraging decisions, naive individuals need to gather information about the local prey community. Besides sampling prey personally, the young could gather information about prey profitability by observing the foraging behaviour of other individuals, and parental provisioning provides the first opportunity to acquire this social information. Still, previous research on vertical transmission of prey preferences from parents has provided mixed results that are often confounded with other information sources, such as siblings and peers. It is also not known whether information from parents can change potential innate biases against certain prey types, such as avoidance of warningly coloured insects. Here, we tested whether social information acquired by offspring during parental provisioning influences the development of prey preferences in a generalist predator, the Great Tit (Parus major). We brought 15 great tit broods and their parents into captivity at late nestling stage (14 days old) and divided them into three social information treatments where parents were provided with either brown, red or yellow palatable maggots to feed to their dependent young for 8 days. Once foraging independently from parents, we conducted a preference test where juveniles were offered the full array of coloured maggots. Regardless of palatable exposure to typical warning-coloured maggots (i.e. red and yellow), juveniles consistently preferred yellow over red, and preferred brown maggots the most (i.e. lacking warning coloration). This supports the existence of innate biases against typical warning colours, and that social information from parents is unlikely to override these, at least when alternative prey is easily available.

Risk-taking behaviour permeates daily decision-making across many taxa and has significant impact on fitness outcomes. Previous research finds that individuals display consistent tendencies in their propensity to take risks, often labelled as a "Boldness-Shyness" personality trait. In this experimental study, we investigated repeatability in risk-taking behaviour in 241 wild vervet monkeys by measuring their responses to capture threat (via human-initiated darting procedure) and predation threat (via two predator models). We further examined the influence of socio-demographic, group identity and hormonal factors on risk-taking behaviour. We found that multiple behavioural responses, including the likelihood of approaching a food source under threat, were consistent across contexts. Risk-taking behaviour was influenced by age, sex, dominance rank, and hormonal profiles: juveniles, males, and higher-ranking individuals were more likely to approach food under (perceived) predation risk. Additionally, we observed significant among-group differences, suggesting that individuals within the same social group exhibit similar risk sensitivities, potentially due to social facilitation or shared environmental effects. The findings contribute to our understanding of animal personality and the ecological and social drivers underlying variation in risk-related behaviours.

Behaviour innovation plays a pivotal role in a species adaptability to dynamic environments. Investigating innovative behaviour and its underlying mechanism is therefore crucial for elucidating the development of cognitive flexibility across animals. Animal personality--which shapes how individuals perceive and engage with their surroundings--could offer insights into individual variation in this process. This study used a three-step foraging puzzle to evaluate the innovation capacity in 28 rosy-faced lovebirds (Agapornis roseicollis), specifically examining their capacity to recombine individually innovated component behaviours into integrated, more sophisticated techniques. We found that nearly half of the individuals spontaneously innovated multiple component behaviours to solve novel puzzles. Crucially, when challenged with a more sophisticated task, they recombined these behaviours into functionally dependent sequences without prior social demonstration. We further identified sex, persistence, and asocial learning capacity as key predictors of innovative problem-solving performance, with females, persistent individuals, and superior asocial learners excelled at problem-solving. Our findings demonstrate that behavioural innovation is not a static event, but a dynamic process--modulated by physical, cognitive, and personality variables--in which behaviours are flexibly transferred and recombined into increasingly complex forms to enable rapid individual adaptation.

Species in stable social groups engage in diverse social interactions, where social competence - the ability to adjust behaviour using social information - can influence fitness. Yet, whether adaptive behavioural flexibility is expressed across contexts within individuals remains relatively untested. To address this, we exposed cooperatively breeding cichlids (Neolamprologus pulcher) to a role-reversal paradigm. In this species, the early social environment shapes social competence, with more competent individuals adjusting behaviour flexibly to social challenges, while individuals also show consistent differences in traits such as aggression. In the present study, individuals were successively assigned to two contrasting roles, smaller territory owners (TOs) and larger intruders (INTs). We predicted role-specific social competence metrics based on behaviours facilitating shelter acquisition. Social competence metrics correlated within, but not across the two roles. Competent TOs showed shorter latencies to submit, more submissive responses to received aggression, and low aggression after initial submission. Competent INTs escalated quickly and relied more on overt aggression rather than displays, allowing faster shelter acquisition. Across roles, individuals competent as TOs were not competent as INTs. In contrast, consistent individual differences in aggression across social roles suggest that stable behavioural tendencies ( animal personalities) may constrain how social competence shapes behavioural strategies.

Social relationships are consequential and affect survival, longevity and reproductive success in many species. Prior studies have successfully identified the fitness consequences of variation in sociality relative to other individuals. However, we have yet to quantify the fitness consequences of social variation within an individual over time. By using a within-individual centering approach, we can identify the consequences of deviations from an individuals average sociality. This approach has not been applied to the study of animal sociality. We used 20 years of data from a population of yellow-bellied marmots (Marmota flaviventer) to understand the fitness consequences of an individuals average affiliative social network position over their lifetime, and of individuals annual deviations from their average social position. We found that an individuals deviation from its average sociality was very strongly associated with both winter and summer survival. Deviations from some average measures of sociality were also associated with the likelihood of reproducing, but not in a consistent direction. Lifetime average social network positions had limited associations with survival and reproduction. Interestingly, litter size results varied between males to females; neither lifetime averages of social network traits nor deviations from these averages were associated with litter size in females, but deviations in sociality were associated with larger numbers of weaned offspring in males. Future studies of the consequences of social relationships may benefit from the use of an individual-centering approach, and further work is required to understand the environmental and social drivers of this intra-individual adaptive plasticity. Graphical abstract O_FIG O_LINKSMALLFIG WIDTH=200 HEIGHT=184 SRC="FIGDIR/small/728517v1_ufig1.gif" ALT="Figure 1"> View larger version (44K): org.highwire.dtl.DTLVardef@380d04org.highwire.dtl.DTLVardef@1deba24org.highwire.dtl.DTLVardef@18ea2dborg.highwire.dtl.DTLVardef@165eb20_HPS_FORMAT_FIGEXP M_FIG C_FIG HighlightsO_LILifetime averages of social network traits are not associated with fitness C_LIO_LIAnnual deviations from lifetime average sociality improves survival and reproduction C_LIO_LIIntra-individual behavioral plasticity may be an understudied fitness driver C_LIO_LIWithin-individual centering is a useful approach for the study of animal behavior C_LI

In over a thousand avian species, pairs coordinate vocalizations to form duets. Songbirds may use calls (simple, non-learned vocalizations) and songs (complex, learned vocalizations) to answer their partners. In Galapagos Yellow Warblers (Setophaga petechia aureola), song answers in duets appear to have evolved recently in addition to older call answers. Here, we explored the structure and functions of duets in an observational study to understand why song answers in duets may have evolved. We first described the temporal structure of duets initiated by either sex. We then tested whether the distance between pair members, countersinging with neighbours, and timing relative to aggressive encounters predicted answering probability for each sex. Finally, we tested whether the behavioral context of call and song answers differed. We found that most duets were initiated by males, and call answers were almost exclusively given by females. Female answers were also more tightly coordinated than those of males. Female answering probability decreased with increasing distance between pair members, whereas male answering probability did not. In addition, females were less likely to answer with songs when farther from their mate. Females answering probability also increased slightly when their mates initiation was preceded by a neighbors song. Our results show some evidence in support of the joint resource defence hypothesis for female answers. On the other hand, the significance of song in female answers may be related to some other functions that we did not test here, such as signaling quality or commitment to the pair bond. Lay summaryWho turns a song into a duet? In Galapagos Yellow Warblers, it is usually the female: she answers her mates song with a tightly timed call or song. Male replies were rarer and less coordinated. While male replies didnt seem to have a function, female replies were used in territorial disputes with neighbours and may indicate joint ownership of a territory. Female song replies, on the other hand, may signal commitment to the pair bond.

Song evolves rapidly in songbirds, as has been proven for a substantial number of songbird species by demonstrating acoustic differences between current songs and "historical" songs recorded 20 or 30 years previously. In two species of songbirds, white-crowned sparrows and savannah sparrows, it has been further shown that evolutionary changes over such time spans are sufficient to affect the response of receivers, with territorial males responding more aggressively to current songs than to historical ones. These two species, however, have especially low population variability in song, with most males in any population singing the same, single song type; this background of song uniformity makes it especially easy to discern temporal changes. Here we examine response to temporal change in song in a third sparrow species, the song sparrow, in which population variation in song is much higher: males sing 5-13 song types each, with low song-sharing between males, so that hundreds of distinct song types occur in a local population. We find evidence that temporal change has occurred in the songs of our study population, in that 24 song current song types share more introductory phrases with other current song types than do 24 historical song types recorded 27-29 years earlier. Nevertheless, in a song playback experiment, current males showed no difference in response to current and historical songs. The results are in accord with the hypothesis that high levels of population variability in song make temporal changes in song difficult to discern.

Social decisions often require animals to integrate information across multiple attributes of potential partners. Using biological motion stimuli, point-displays generated from tracked live shoals, we tested how adult zebrafish (Danio rerio) weigh shoal size and movement speed during social preference, and whether these preferences are susceptible to contextual manipulation by an asymmetrically placed alternative. In Experiment 1, we established a multi-attribute indifference point by presenting males and females with dichotomous contrasts in which shoal size and movement speed were traded off. Both sexes showed no preference when a larger, slower shoal (4 fish at 0.75x speed) was pitted against a smaller, faster shoal (2 fish at 1.25x speed), but preferred the smaller, faster shoal when the speed difference was greater (4 fish at 0.5x versus 2 fish at 1.25x), indicating that zebrafish are sensitive to graded differences in movement speed relative to numerical cues. In Experiment 2, unidimensional tests confirmed that both sexes preferred larger shoals when speed was held constant but revealed sex-based differences in speed sensitivity: males preferred faster-moving shoals at both shoal sizes tested, whereas females showed no significant speed preference. Male shoal size preferences were stronger at higher movement speeds, suggesting that speed modulates the strength of size preference. In Experiment 3, we tested the asymmetric dominance effect in males, the only sex sensitive to both dimensions, using the indifferent contrast from Experiment 1 as the primary options and four decoy shoals asymmetrically placed along either the size or speed dimension, under counterbalanced presentation orders. No decoy shifted male preference significantly from chance under any condition. These results indicate that zebrafish weigh social cues in a sex-specific manner, and that asymmetric decoy options do not induce preference biases in males when shoals vary along the dimensions of movement speed and size.

Many social animals exhibit fission-fusion dynamics, where group members split into subgroups and come back together. Studying these dynamics can reveal how individuals weigh the costs and benefits of sociality. Most studies infer the drivers of fission-fusion dynamics by examining subgroup association patterns, yet different underlying processes can produce similar patterns. Here we take a fundamentally different approach by focusing on subgrouping events themselves, where decision-making plays out in real time. Using multi-sensor tracking collars, we collected simultaneous movement and vocalisation data from all group members within two wild white-nosed coati groups. We developed analytical tools to extract and characterise fission and fusion events, thus establishing a framework for quantifying the spatiotemporal dynamics of these events and the use of vocalisations before, during, and after them. We found that fissions typically occurred when groups were initially stationary, indicating that splitting does not result from loss of coordination while moving.Subgrouping was associated with reduced aggression, yet aggressive vocalisations did not precede splits, supporting the hypothesis that subgrouping is a pre-emptive mechanism to manage within-group conflict. Contact calls in moving subgroups increased before fissions and fusions, indicating that vocal communication is key to the coordination of these collective movement decisions.

Collective movement requires coordination between individuals, yet how this emerges during early interactions remains poorly understood. We investigated how partner familiarity influences coordination, leader-follower dynamics, and learning in homing pigeon pairs navigating from novel sites. Birds were released repeatedly with either familiar or unfamiliar partners, followed by solo releases to assess learning. By quantifying bidirectional information flow, we found familiarity influenced information-transfer dynamics during the first release: familiar pairs exhibited more asymmetric information transfer, likely reflecting established leader-follower relationships, whereas unfamiliar pairs showed more symmetric exchange. These differences disappeared after one release. Conversely, familiarity had little effect on cohesion or navigational performance. There was some evidence for an influence on learning: birds from familiar pairings had higher homing efficiency on a subsequent solo release. Finally, across partnerships, followership was more predictable than leadership with respect to individual identity and flight speed, indicating stable variation in individuals' tendency to follow rather than lead. This suggests that a shift in emphasis from leadership to followership might enhance our understanding of collective decision-making dynamics. Our results demonstrate how flight partners rapidly coordinate, producing limited downstream effects on navigation and learning, with implications for many animals that travel in fission-fusion transitory collectives.

Interactions between cleaner fish Labroides dimidiatus and client fish, from which cleaners remove ectoparasites and mucus, represent a textbook example of mutualism involving sophisticated strategic decision-making. However, cleaners must also face intraspecific social challenges within a size-based hierarchy, where the largest females may eventually change sex and become males with higher reproductive rates. Following 540 individuals over 11 months, we found that, contrary to expectations, slow-growing females spent more time cleaning and cheated more frequently, without causing more negative client responses than fast-growing females did. Instead, variation in growth was best explained by social factors: fast-growing individuals experienced reduced social control, while slow growers spent more time in proximity to dominant individuals. As there was no evidence that spawning activity affected growth patterns, it appears that fast growth as a viable strategy for becoming a male largely depends on the lack of control by dominants.

For many group living species, vocal signals are a vital form of communication needed to coordinate social behaviours. Investigating these processes first requires a detailed description of the species vocal repertoire. White-nosed coatis (Nasua narica), which forage and move together in forest habitats, are thought to rely on vocalisations to coordinate movements and maintain group cohesion. However, quantitative studies of their vocal repertoire are lacking. We examined the vocal repertoire of white-nosed coatis from wild populations in Panama and Arizona, USA, to gain a more comprehensive view of their calling behaviours. By combining traditional acoustic analyses with an unsupervised approach based on spectrogram structure, we characterised the diversity of calls in this species and described the temporal and structural features of their vocalisations. We identified 19 call types, with some of these calls emitted in multi-syllable call sequences or in fast succession. We found variability in call rates among group members, which may be driven by differences in social status within the group. In addition, our results indicate that white-nosed coatis likely have individually-recognisable vocalisations. This study provides a foundational description of the white-nosed coati vocal repertoire, laying the groundwork for future research on vocal communication in this species.

One challenge of reintroduction programs is ensuring that captive individuals are prepared for life in the wild, including having the ability to forage on wild resources. Foraging proficiency can be especially challenging to foster in species with a wide dietary breadth or reliance on ephemerally available resources. A relatively unexplored obstacle to release readiness may be an animals hesitancy to approach or consume unfamiliar foods, i.e. their neophobia and dietary conservatism. In this study, we examined the responses of a conservation breeding population of the extinct-in-the-wild alal[a] (Corvus hawaiiensis), a dietary generalist, to novel and familiar native and non-native fruits. Birds were similarly willing to contact both novel and familiar fruits, suggesting little aversion to the presence and appearance of novel food items. However, birds showed a clear preference for contacting non-native fruits over native fruits, regardless of familiarity. Additionally, while birds were willing to contact novel items, they were less likely to consume novel native fruits, suggesting that dietary conservatism may limit incorporation of these resources in the wild. Finally, juvenile birds were less likely to approach the testing setup and contact items, suggesting that neophobia may constrain their exploration of novel environments and resources. We discuss the implications of alal[a] preferences for non-native foods and reluctance to consume novel native items for pre-release preparations and the goal of wild species recovery.

Animal vocalisations can convey information about external events, but whether this goes beyond reflecting the emotional state elicited by these events is debated. To explore this, we studied the acoustic structure of common marmoset (Callithrix jacchus) phee (long-distance contact) and ek (alert/mobbing) calls produced in five treatments varying in the emotional valence and arousal they elicit (internal state), as well as food and social context (external events). We measured changes in arousal via nasal temperature and analysed both basic acoustic parameters and Mel-frequency cepstral coefficients (MFCCs) of the calls. Support Vector Machines combined with Linear Mixed effect models revealed that phee calls encode both external events and internal states, while eks reflected predominantly arousal. Notably, an acoustic signature related to food context was present in phees both when provided (positive valence) and teased with highly preferred food items (negative valence), and even when food was not physically present (food call playback treatment). This suggests marmoset long-distant phee calls encode external information beyond emotional arousal and valence, and independently of the presence of an immediately triggering stimulus.

Mother-daughter relationships are central to social organization in rhesus macaques, providing the foundation for kin familiarity, affiliative bias, and intergenerational rank inheritance. Understanding how these bonds are structured is therefore important for identifying the processes that contribute to matrilineal cohesion and for anticipating risks of social instability and matriline fragmentation in captive groups. We studied 141 dam-daughter dyads in large, captive groups of rhesus macaques, using generalized linear models to examine associations between demographic variables and affiliative behaviors, and a multivariate clustering approach to characterize how these behaviors combine into distinct relationship types. Our analyses identified eight distinct dam-daughter relationship types defined by combinations of three behavioral domains: grooming, spatial closeness, and agonistic support. Generalized linear models showed that demographic variables had domain-specific effects, with family size, age gaps, and dam age differentially associated with grooming, proximity, and support. These patterns were reflected in the clustering analysis, in which dyads differed not only in overall levels of behavior but also in how behaviors combined within dyads. For example, some dyads exhibited high grooming and support despite low spatial proximity, whereas others showed consistently high expression across all domains. Together, these results demonstrate that dam-daughter bonds are multidimensional and structured through the combination of multiple behavioral domains influenced by demographic and life-history factors. Because these dyads scaffold interactions among close kin, variation in dam-daughter bond structure may influence the development and stability of broader matrilineal relationships. From an applied perspective, considering multiple behavioral domains when assessing social relationships can help identify dyads that may warrant closer monitoring and inform management strategies to support matrilineal cohesion and reduce the risk of social instability in captive groups.

Many primates exhibit female philopatry and live in stable, female-bonded social groups. Permanent group fusions are rarely documented in these populations. We present a case study on a fusion of two social groups from a hybrid population of baboons (Papio cynocephalus x P. anubis) living in the Amboseli basin of Kenya. The fusion occurred following a period of increased human-induced mortality in one of the two social groups. After the fusion, females from the smaller group became the lowest ranking. We compared female behavior in the months following the fusion to the behavior of females in groups that had not fused and also compared pre- and post-fusion fitness outcomes. Following the fusion, the groups activity budget and patterns of agonistic interactions were typical for the study population. Females preferred familiar grooming partners for a short period following the fusion; however, after three months, patterns in female grooming were comparable to other groups, indicating rapid social integration. With the caveat that our sample size was limited, we observed no detectable fitness-related costs of group fusion in terms of birth rates or offspring survival, and adult female mortality was low following the fusion. These results demonstrate the flexibility of female baboons in navigating exposure to novel same-sex conspecifics despite a species-typic pattern of female philopatry. Based on this and previous examples of group fusions, we propose that group fusions may be most likely to occur when groups are too small to retain adult males, defend against predators, or compete with other groups.

Multimodal communication plays a central role in animal behavior, particularly when individuals must integrate information from different sensory channels to make rapid decisions. In aquatic environments, chemical and visual cues differ markedly in their spatial and temporal properties, such that chemical signals may be constrained by limited spatial resolution and temporal instability, potentially requiring visual information to reliably guide social decisions. In decapod crustaceans, both cue types are known to mediate reproduction, yet their relative contribution to mate-location behavior remains unclear. Here, we tested how visual and chemical cues from males influence mate-location behavior in females of the prawn Macrobrachium rosenbergii. Females were placed in a central arena and exposed to four stimulus configurations combining visual cues (a life-size photograph of a male or a control background) and chemical cues (water from an aquarium with or without a male). Attraction was quantified as the time spent in each half of the arena. Females showed no directional preference when exposed to chemical cues alone or when visual and chemical cues were spatially incongruent. In contrast, females spent significantly more time near male-associated stimuli only when visual and chemical cues were spatially congruent. These results indicate that mate-location behavior in this species depends on multimodal integration with a strong contextual dependence on visual information, which appears to gate the effectiveness of chemical cues. Spatially congruent multimodal signals are therefore necessary to guide orientation during mate search, suggesting that disruption of visual or chemical information in aquaculture systems may impair mating efficiency.

Acoustic communication is fundamental to social interactions in many animal species, allowing individuals to transmit information about identity, reproductive status, and competitive ability. Because call production incurs inherent costs, individuals are expected to modify their vocalizations depending on the social context. However, while context-dependent call variation has been documented in several taxa, including anurans (frogs and toads), glass frogs (Centrolenidae) remain among those for which the acoustic repertoire across social contexts is poorly characterized. Here, we investigated context-dependent call modification in males of the Emerald glass frog, Espadarana prosoblepon, comparing calls produced across four social contexts: advertisement in isolation, advertisement in a group, courtship interactions with females, and agonistic interactions with other males. By integrating detailed behavioral field observations with a robust analytical framework, we present evidence that males modify multiple acoustic properties in response to the social context. Specifically, males produced longer, louder advertisement calls when calling in a group than when calling in isolation. Courtship calls contained more notes and were louder than other call types, whereas aggressive interactions were characterized by pulseless, low-frequency, soft calls. Our findings demonstrate that the distinct call types of E. prosoblepon are consistently associated with specific social contexts and can be reliably distinguished based on their acoustic structure, providing a framework for future studies investigating the functional significance of context-dependent acoustic signals in anurans.

Colour and contrast are commonly deployed in anti-predator signals like aposematism or deimatism. In oceans, colour information diminishes with depth, leaving blue bioluminescence the most common visual signal, regardless of function. Bioluminescence can deter predators, but without contrasting colours, how so is largely uncharacterized. Here, we test this by observing fish predators responding to prey that use defensive bioluminescence (Ostracods, Cypridinidae). By manipulating potential chemical defences of prey, and by comparing feeding responses to both luminescent and nonluminescent prey, we show that luminescent prey are unpalatable and use facultative bioluminescence as an aposematic signal. We observed active, luminescent prey secrete bioluminescence only after being attacked. Predatory fishes rarely consumed luminescent prey, especially compared to nonluminescent alternatives. Food treatments revealed that luminescent species may possess some unidentified defence over nonluminescent relatives because fishes also readily ate luminescent prey that had been treated (frozen or boiled), which removed such defences. Over the course of four experimental trials, predators were less likely to consume luminescent prey as their cumulative exposure to anti-predator light displays increased, indicative of learning. Despite their intermittency, temporally dynamic signals like aposematic bioluminescence may be as common and effective as better-studied static coloration, especially in marine ecosystems.