Behavioral Ecology

Non-avian reptiles have been assumed to be non-social for many years, yet recent studies show diverse social behaviours in squamates, crocodilians, and turtles. Here, we investigate social structure within feeding aggregations of three freshwater turtle species caught in baited traps in a coastal marsh over 12 years. In 488 instances in which traps contained turtles, 45% contained multiple individuals, and these aggregations were strongly positively assorted by species. midland painted turtles (Chrysemys picta) and Blandings turtles (Emydoidea blandingii) were captured with conspecifics more often than expected in a non-social null model. Snapping turtles (Chelydra serpentina), the largest species in this study, were caught with conspecifics at rates consistent with the non-social null model, and were avoided by heterospecifics. This suggests that species differences play a role in how feeding aggregations are structured, with painted and Blandings turtles driving positive species assortment while avoiding snapping turtles around food sources. We did not detect significant intraspecific sex assortment in any of the three species, nor did turtle body size strongly affect the probability of aggregating with conspecifics at the perceived food source in the traps. Our study illustrates how long-term monitoring data can be used to investigate social structure in wild populations, an approach that may be valuable for species of conservation concern. Significance StatementReptile sociality has been historically overlooked, but recent work has revealed intriguing social behaviours in non-avian reptiles. We investigated associations among three species of freshwater turtle, captured in baited traps over 12 years of trapping. Turtles in these feeding aggregations grouped with their own species more often than expected by chance. This result was driven by the two smaller-bodied species (midland painted turtle and Blandings turtle), which were more likely to be caught with conspecifics than with other species. In contrast, the largest species (snapping turtle) showed no evidence of attraction to other snapping turtles, and midland painted turtles and Blandings turtles showed avoidance of the larger snapping turtles. Our analyses illustrate how long-term monitoring data can be used to uncover previously unrecognized social behaviour in turtles and other species in which behaviour is difficult to observe directly.

Acoustic signaling is key to individual and species recognition, playing a major role in sexual and social communication. Since reproductive isolation is often maintained through pre-mating mechanisms, song can be an early isolating trait leading to assortative mating, promoting reproductive divergence, and potentially contributing to speciation. However, whether song differences alone are sufficient to prevent interbreeding or if other traits also contribute, remains a matter of debate. Playback experiments provide a more direct way to test the role of song as a reproductive barrier. Here, we use playback experiments to test the hypothesis that song acts as a pre-mating barrier in two recently diverged populations of an island passerine, the Canary Islands Chaffinch (Fringilla canariensis palmae), which inhabit ecologically distinct laurel and pine forests within the island of La Palma. Assuming that male song has diverged in the two habitats, we tested if territorial males from a given habitat responded differently to songs from intruding males from their own habitat or from the other habitat type, using a closely related mainland species as a control. We found that probability of response was weaker to songs of the closely related species and to the different-habitat birds than to songs of the same-habitat birds, but differences for the latter were weak. The intensity of response followed the same pattern. Overall, song divergence between laurel and pine forest chaffinches does not appear strong enough to cause clear behavioural discrimination against individuals from the alternative habitat. Other factors such as morphological and ecological divergence associated with adaptation to local resources might better explain population differentiation. However, testing female responses will be essential to determine whether songs convey lineage-specific information that may elicit assortative mating.

Escape behaviour directly influences survival, yet individuals often vary substantially in escape performance. Laboratory studies have documented trade-offs between anti-predator responses and life-history traits, but it remains unclear whether such trade-offs occur under natural predation risk. We studied a natural population of the field cricket Gryllus campestris. Mortality risk and behavioural performance are known to change with age in this species. We aimed to determine whether individuals expressing a higher escape response pay a cost in terms of a faster increase in mortality risk with age or a shorter lifespan. We quantified escape speed in response to a vibrational predation cue. We found no clear evidence for a trade-off between escape performance and lifespan or age-specific mortality risk. The relationship between escape speed and the among-individual effect of age differed between sexes: older males showed faster escape speeds compared with younger males, whereas younger females were faster than older females. This pattern is consistent with sex-specific selective disappearance. Individual baseline mortality risk varied with sex and escape speed, but age-dependent mortality did not. It suggests that such trade-offs in the wild may be context- or condition-dependent rather than reflecting a universal life-history trade-off.

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.

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.

Prolonged association between mothers and their offspring is common in ungulates, with the level of maternal investment likely to play a central role in shaping this trait. Here we examined patterns of association between mothers and offspring over time, the apparent benefits of association to offspring, and costs to mothers. We analyzed 40 years worth of census data from an individually-monitored, food-limited population of red deer (Cervus elaphus) on the Isle of Rum, Scotland. Starting from birth, female calves associated more frequently with their mothers than male calves in their first year. Calves also associated less with their mothers if the mother did not conceive a new calf. Association frequency decreased with mothers age and population density, and survival over the first year was not related to mother-calf association. Yearlings, now in their second year, were more often associated with their mothers if they were female, if there was no subsequent calf (or the subsequent calf died as a neonate), and if they were still being suckled. Increased association between mothers and yearlings was associated with increased survival to adulthood at 28 months, but suckling a yearling did not improve its probability of survival. For individuals that reached maturity, increased association in the yearling year was associated with slightly shorter adult life spans. The level of association between a calf and mother was not associated with the mothers immediate survival or fecundity. Our findings suggest that juveniles born to poor-condition mothers benefit from prolonged association through improved yearling survival.

Sentinel behaviour occurs when individuals use raised positions to scan for predators while the rest of the group forages. Here, we investigated whether a colonial cooperatively breeding species that forages in large groups, the sociable weaver, Philetairus socius, displays sentinel behaviour. This behaviour has been reported in species with similar ecology, behaviour and foraging habits, (e.g. ground foraging in open habitats where aerial predators are common) and, hence, we expected that it could occur in sociable weavers. On the other hand, sentinel behaviour appears to be less common in species that live in very large groups. We used an experimental set-up consisting of an artificial feeding station and perches to assess occurrence of sentinel related behaviours: (i) perching events > 30s on an elevated position, (ii) head-movements and (iii) alarm calling. Birds were seldom observed perching while others fed, and those that did, perched for periods that were too short to be considered as sentinel behaviour (less than 5s on average). Our results suggest that this behaviour is uncommon or even absent in sociable weavers. We discuss whether other factors such as foraging in very large groups, or interspecific foraging associations might make sentinel behaviour less important in this species.

Protogynous sex change, where individuals first function as females and later as males, is a key life-history strategy among polygynous reef fishes. In haremic systems, sex change is typically socially regulated, with dominants suppressing subordinates sex change through aggression. Females within a harem form a size-based hierarchy that can remain stable in most species through the threat of eviction. We studied a different situation in the cleaner wrasse Labroides dimidiatus, where larger females have incomplete control, as they spend most of their time alone at their own cleaning territory. We tracked over 400 individuals for 12 months, recording growth, behavior, social organization, and sex change. We confirmed earlier reports that both sexes direct aggression primarily at those ranked immediately below them. However, we observed 30 cases where smaller females outgrew larger ones, revealing hierarchy instability. Of 42 sex change events, 43% occurred in presence of the male, and half of these early sex changers were not the largest female, but individuals overlooked by the male. Fast growth relative to harem-mates and harem switching increased the likelihood of sex change. Local population densities also influenced growth and sex change, with individuals in high-density demes growing faster and changing sex at larger sizes. Our findings reveal flexible sex change dynamics in a system with incomplete social dominance. Such incomplete control and observations that becoming male confers both higher reproductive success and survival highlight the need to expand game-theoretical and life-history frameworks to encompass such strategic flexibility. Lay summaryDominant cleaner wrasse cannot fully control subordinates as individuals occupy distinct core areas. Tracking 400 fish for a year, we found that smaller females could outgrow initially larger ones, and early sex change despite a larger male. Fast growth and harem switching increased the chances of becoming male. Population density also shaped these strategies. Our findings reveal flexible sex change dynamics in a system where becoming male confers both higher reproductive success and survival.

Reproductive Character Displacement (RCD) often occurs when species with mating-related polymorphism come into secondary contact, leading to divergence in reproductive traits. Ischnura elegans and Ischnura graellsii have formed two independent hybrid zones in Spain where reinforcement has strengthened a mechanical barrier, and RCD has shaped mating-related structures, although reinforcement is asymmetric only in gynochrome females. This study examines the link between asymmetric reinforcement and asymmetric RCD. Using geometric morphometrics, we analyze prothorax shape and size in both female morphs and males, and male caudal appendages, to assess morphological divergence, determine whether gynochrome females show stronger divergence, and test for morphological covariation between male traits involved in the tandem position. Our results reveal consistent patterns of size and shape variation across species and zones: in I. elegans, androchromes are larger and resemble males in size, with clear shape differentiation between female morphs that diminishes in hybrid zones. In contrast, I. graellsii shows less consistent size differences between males and morphs, and weaker shape differentiation. Our results confirm RCD in prothorax shape in I. elegans females from both hybrid zones, but reveal that RCD in prothorax size is asymmetric, occurring only in gynochrome females from the NC hybrid zone. We also detected RCD in the prothorax shape of I. elegans males from the NC hybrid zone, extending previous evidence of RCD in male caudal appendages, while morphological covariation between male cerci and the prothorax was limited to size in I. elegans. Together, these findings illustrate how hybridization may generate morph-specific patterns of reproductive divergence.

BackgroundPollen production is a costly investment in angiosperm reproduction, yet only a small fraction of grains reach conspecific stigmas. While pollen loss to floral visitors is well studied, the role of abiotic factors in shaping pollen fate has been largely overlooked. Understanding the relative contributions of biotic and abiotic pollen loss pathways is essential for interpreting floral trait evolution and pollination efficiency. ResultsWe quantified abiotic pollen loss in four animal-pollinated species with contrasting floral longevities and reproductive phase dynamics. Flowers were monitored for five hours after anthesis, with pollen loss compared between unvisited flowers and those receiving single pollinator visits. Across all species, substantial pollen loss occurred in the absence of visitation, ranging from 37-57% of grains. In several cases, losses to legitimate pollinators were indistinguishable from abiotic loss alone, whereas pollen-foraging honeybees removed significantly greater fractions. ConclusionAbiotic factors can account for a large proportion of total pollen loss, sometimes equalling or exceeding pollinator-mediated removal. These findings challenge the assumption that pollen loss is primarily driven by pollinator activity and suggest that floral traits such as closure, gradual pollen release, and pollen packaging may function as adaptations that minimize environmental loss. Incorporating abiotic pollen loss into studies of pollen presentation and pollinator effectiveness provides a more complete understanding of selective pressures shaping floral evolution.

Parental care can shape post-fledging behaviour through provisioning, guidance and social information, yet its absence may alter how young birds establish space use and habitat preferences. We tested the consequences of absent parental care by comparing, hand-reared juvenile herring gulls released without parents with wild, parent-reared conspecifics, focusing on the first two months after fledging. Wild juveniles frequently revisited their natal nest during the first month, whereas hand-reared birds rarely returned to the release site; revisits declined in both groups by the second month but remained more common in wild birds. Wild juveniles used smaller ranges that subsequently expanded, while hand-reared birds began with larger ranges that later contracted, leading to convergence. Contrary to expectation, wild juveniles occurred in areas with higher human population density than hand-reared birds. Habitat use also differed between groups and changed over time. Early on, wild juveniles concentrated activity in anthropogenic and marine habitats, whereas hand-reared birds used rural green habitats more. Later, both groups shifted away from marine areas towards rural green habitats, reducing but not eliminating between-group differences. Short-term survival, did not differ between hand-reared and wild juveniles, indicating that parental care primarily reshaped early space use and habitat choice rather than immediate survival.

Social interactions are ubiquitous in nature and have the potential to affect trait evolution, particularly in group-living animals such as cooperative breeders. Interactions among conspecific individuals can affect the amount of additive genetic variation for a trait when the phenotype of an individual is also affected by the genotype of its social partner(s) via indirect genetic effects. Thus, quantifying both direct and indirect genetic effects of social partners is critical for understanding and predicting evolutionary trajectories. While much is known about maternal indirect genetic effects, empirical estimates of indirect genetic effects from other social partners remain limited, particularly in wild populations. Here, we use animal models to assess the contribution of indirect genetic effects from all social partners in a family group (mothers, fathers, and helpers) on juvenile morphometric traits across ontogeny in the cooperatively-breeding Florida scrub-jay (Aphelocoma coerulescens). We found indirect genetic effects of helpers and fathers on nestling weight, but no indirect genetic effect of mothers. Across ontogeny, we found increasing additive genetic variation in both weight and tarsus length. Our study provides a comprehensive assessment of within-group indirect genetic effects in a cooperative breeder and highlights the importance of considering indirect genetic effects beyond maternal effects.

The ability of animals to innovate - solve novel problems - can shape their ecology and evolution. Here we investigate how individual traits and environmental complexity relate to successful solving of a novel problem. We presented foraging bumble bees (Bombus impatiens) with artificial flowers of not-previously-encountered shapes and recorded the bees latency to access nectar. We measured individual foraging traits across multiple trips with simple flowers that did not require innovation, and bees were foraging either in a simple or complex environment (cluttered flight arena). Bees in complex environments took longer to find and were less likely to land on novel flowers, indicating that environmental complexity may take up cognitive resources and make search more difficult. However, we did not find an effect of environmental treatment on the ability or time to access reward in novel flowers once bees had landed on them. In contrast, behavioral traits significantly predicted how quickly bees solved novel flowers. In particular, overall foraging tempo as well as routine formation, i.e. how much bees followed a fixed route on known flowers, predicted innovation - faster bees innovated faster, and bees with more repetitive foraging sequences were slower to solve the novel tasks. Overall, while the degree of evolutionary novelty in tasks or solutions is always hard to evaluate, our findings demonstrate that environment and individual traits may affect innovation in different ways. Individuals in simple environments may be more likely to detect, and individuals that are generally faster and have a lower tendency to develop fixed routines may be more likely to solve, novel tasks.

Seasonal rainfall shapes biological responses in tropical ecosystems, yet how tropical organisms integrate behavioral and physiological responses to cope with seasonality remains poorly understood. We assessed how four poison frog species with contrasting reproductive strategies respond to dry and wet season environmental conditions. We quantified spatial behavior, microhabitat use, hormone concentrations, and chemical defenses in two seasonal breeders (Allobates femoralis and Ameerega trivittata) and two year-round breeders (Ameerega macero and Ameerega shihuemoy). Seasonal breeders exhibited pronounced sex-specific shifts in space use, where males expanded their space use during the wet season, likely to track reproductive opportunities, while A. femoralis females increased their spatial use during the dry season, likely responding to foraging demands when prey resources are sparse. Year-round breeders maintained similar space use across seasons, likely reflecting their ability to access key resources within the same space to reproduce year-round. Microhabitat use was flexible, as seasonal breeders shifted toward humid refugia during the dry season and reproduction-associated microhabitats during the wet season, whereas year-round breeders selected microhabitats that facilitate continuous reproduction across seasons. Despite these behavioral responses, corticosterone, testosterone, and chemical defenses showed no consistent seasonal variation, suggesting that behavioral flexibility is decoupled from seasonal variation in these measured physiological responses. Our study suggests that poison frogs are able to buffer environmental fluctuations through behavioral flexibility. However, given the increasing unpredictability in rainfall timing and intensity as a result of climate change, how these coping strategies will function in the long term is uncertain.

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.

The ectoparasitic mite Varroa destructor is the most significant parasite of the Western honey bee (Apis mellifera) and a major driver of colony losses worldwide. Although extensive research has focused on behavioral and physiological mechanisms of host resistance, comparatively little attention has been paid to potential phenotypic responses of the parasite itself. Here we investigated body size variation in Varroa destructor associated with varroa-resistant and non-resistant honey bee colonies across four European countries. We quantified the dorsal shield area of adult female mites from multiple colonies differing in the honey bee colonies resistance status, using standardized digital image analysis. Across geographically distant non-resistant populations, mite body size was remarkably consistent, with a median dorsal shield area of 1.47 mm2. In contrast, mites originating from varroa-resistant colonies were consistently smaller, with a median dorsal shield area of 1.37 mm2, representing an approximately 6.8% reduction in body size. This pattern was reproducible across different geographical areas, honey bee genetic backgrounds and beekeeping practices. The striking stability of mite body size in non-resistant populations contrasted with the consistent reduction observed in mites associated with resistant hosts, suggesting a host associated shift in parasite phenotype. Because body size in arthropods integrates developmental conditions, nutritional availability and resource allocation, the observed pattern may reflect altered developmental environments and selective pressures imposed by resistant hosts. Our results show a consistent morphological shift in this globally important parasite associated with resistant hosts and suggest that dorsal shield size in Varroa could serve as a new selection marker for varroa-resistant honey bee colonies.

Judgment bias tasks are increasingly used to assess affective states in animals, yet the extent to which they might reflect transient states or stable traits remains unclear. Here, we tested bumblebees (Bombus terrestris) in an active choice task across three repeated sessions to assess individual consistency in the absence of any manipulation. Bees were trained to associate each of two colours with either a high or a low reward, presented in separate chambers. During testing, they were presented with ambiguous colours. Bees were more likely to choose the high reward chamber and to choose more quickly in response to colours closer to the positive colour. The latency to choose the cues showed significant and moderate repeatability across sessions, suggesting a stable, trait-like underlying component. In contrast, the repeatability of the chamber choices was negligible, indicating that such responses might be largely state-dependent and influenced by situational factors. These findings suggest that judgment biases, particularly as assessed through an active choice task reflect states affected by external factors. Active choice tasks may help disentangle stable behavioural traits from transient affective states in invertebrates.

According to sexual selection theory, males should benefit more from mating with multiple partners than females do, as male investment into offspring production is typically lower. For females, empirical evidence indeed often shows diminishing returns or even costs of mating multiply. For males, the assumption often seems to be "the more, the better" - i.e., a steady increase of male reproductive success with mate number - but experimental tests of it are rare. Here we used a laboratory experiment with Trinidadian guppies (Poecilia reticulata), known for being promiscuous, to assess how pairing males weekly with 4 vs. 7 females affects both sexes reproductive performance (n = 32 polygynous males and 170 monogamous females). Increased polygyny delayed females reproductive onset by 9% and tripled their risk of reproductive failure. High-polygyny males fathered offspring with 49% more females and had 73% higher daily reproductive output. Yet, they needed 19% longer to initiate pregnancy, and only accumulated more offspring than low-polygyny males after two months. This study suggests that male mating performance is not unlimited. Especially when high extrinsic mortality selects for fast reproduction, less polygyny might be advantageous, and the strength of sexual selection perhaps more similar between the sexes than often assumed.

Understanding relatedness in sharks is challenging due to uncertainty in distributions, low population densities and difficulties in sampling across life stages. In Fiji, bull sharks (Carcharhinus leucas), with an effective population size estimate of [~]258, aggregate at the Shark Reef Marine Reserve (SRMR), but gravid females disperse at the end of the year to give birth in adjacent rivers. Questions remain regarding reproductive connectivity, female returns across years, and kinship structure. Using population genomics on 296 bull sharks across age classes (neonates, young-of-the-year, juveniles, and adults) collected over a decade at the SRMR and in three adjacent rivers, we assessed familial connections. Direct genetic links, including first- and second-degree relationships, connected SRMR adults with young age classes in the Navua and Rewa rivers, providing evidence of reproductive connectivity. Within rivers, genetic similarities across cohorts revealed reproductive philopatry. Remarkably, several individuals sampled years apart were assigned to the same sire-dam pairs, indicating repeated pairings across breeding seasons. However, the few related links detected between the SRMR and the rivers may reflect incomplete sampling. Altogether, bull shark reproduction in Fiji seems influenced by reproductive philopatry and repeated pairings, suggesting added complexity in their reproductive behaviour.

Group-living animals often coordinate their behavior using "contact calls". Identifying the function of these calls requires testing whether they are intended for any group member or targeted to specific preferred associates. If contact calling is used to coordinate with preferred associates, then higher rates of contact calling are expected between group members with a history of more frequent affiliation and cooperation. To test this hypothesis, we constructed a contact-calling network using synchronized recordings of vocal interactions between all 28 possible pairs of 8 female common vampire bats with well-sampled histories of social grooming and regurgitated food sharing. Bayesian multilevel models show that pairwise rates of contact calling were clearly predicted by social grooming and cooperative allofeeding rates in ways not explained by kinship. These findings show that common vampire bats use contact calls to coordinate with specific same-sex associates, unlike other studied bat species where individuals produce contact calls at similar rates towards different group members. We also found that, compared to white-winged vampire bats, common vampire bats are ten times less likely to rapidly respond to a contact call; this suggests yet-to-be-discovered differences in social behavior between vampire bat species. Finally, we discuss implications for the vocal grooming hypothesis.