Ethology

Tortoises do not show parental care and live solitary except for the context of reproduction. Despite their limited need to interact with conspecifics, we previously observed that young tortoises, at their first experiences with conspecifics, can discriminate between familiar and unfamiliar conspecifics after just one encounter with another tortoise. Tortoise hatchlings ignored familiar conspecifics, while they first explored and then actively avoided unfamiliar conspecifics. It remains to be established whether the different reactions to unfamiliar and familiar individuals in tortoise hatchlings are reactions to novelty, or whether they are specific to the interactions with living animals. To test this, we familiarized one-month-old tortoise hatchlings with an object (a brown cone vs. a blue sphere) and then tested them in a novel arena once with the familiar object and once with an unfamiliar one. To measure the reactions toward familiar and unfamiliar objects, we measured the distance between the tortoise and the object throughout the test. Differently from what happened with unfamiliar and familiar conspecifics, we found no difference in behavior toward familiar and unfamiliar objects. This shows that the different reactions toward familiar and unfamiliar conspecifics previously observed are specific for social interactions and are not a mere reaction to the novelty effect. The behavioral responses displayed by young tortoises for unfamiliar conspecifics, but not for unfamiliar objects, show the relevance of social behavior from the beginning of life, even for solitary species.

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.

Time is an important and limited resource that can drive the trade-off between various essential activities in the lives of animals. Group-living animals need to perform different behaviour to meet their individual needs and also participate in group activities. They must, therefore, partition the available time between these activities which may vary considerably with environmental and ecological conditions. We examined time-activity budget of a cooperative passerine, Jungle Babbler (Argya striata) and how their behaviour vary across diel and seasonal scales. A repertoire of 13 behaviour was recorded of which 12 behaviour that occur throughout the year were examined further in detail. This included individual behaviour such as foraging, grooming, rest, shower and group behaviour such as allogrooming, movement, play, sentinel, mobbing and inter-group fight. Our results indicate that most of the time (about 70%) was spent performing individual behaviour and the remaining time (about 30%) was allocated to social behaviour. We also found almost all behaviour varied across diel and seasonal scale with respect to proportion of time spent performing them. This highlights the impact of environmental factors on how animals partition their time to perform various activities. Our study also lays the foundation for future studies examining the role of ecological factors such as habitat type and predation pressure in driving these patterns of behaviour in Jungle Babblers.

Intraspecific agonistic interactions are common in the animal kingdom and have significant consequences for animals. With a growing population and limited resources, these interactions have been observed more frequently among smooth-coated otters in Singapore. However, little is known about the behavioral patterns of these interactions and the factors that may affect their outcomes. To address this, we analyzed intraspecific agonistic interactions through publicly available videos from citizen scientists. Our analysis revealed different behavioral patterns and typical behaviors between winning and losing family groups in different environments. We also found suggestive evidence that larger family groups with more individuals than opponents have an advantage in these interactions. The environment in which an interaction occurs may also affect its intensity.

Mate search tactics and association preferences among organisms in natural habitats can be dynamic and are determined by inherent trait preferences as well as the cost-benefit trade-offs associated with each mating decision. Two of the prime factors regulating mating decisions are the presence of competing conspecifics and predatory threats, both of which have important fitness consequences for the individual. We studied the influence of these two factors separately in mate search tactics and association preferences among zebrafish males. Male zebrafish were presented with a choice of two patches, consisting of different number of females, of which one patch was also associated with a predatory threat. We found that males made a preferential choice for the patch with more number of females only when the numerical difference between choices are starkly different, irrespective of the predatory threat associated with the patch. This points towards the role of numerical cognition in assessing cost-benefit tradeoffs in male zebrafish. We also studied the association preference of males in a multi-choice setup, consisting of four separate mixed-sex groups of zebrafish varying in densities. Our results showed that while test males preferred to visit the male-biased patches more often, they spent more time near female-biased patches or patches with equal sex ratio patches indicating the role of complex interplay of social cues in determining the associative behavior of males to a patch. This study, thus, sheds further light on the interactive roles of social cues and cognitive abilities in mate association patterns in this species.

Self-recognition is the ability to recognise stimuli originating from oneself. Humans and most great apes show evidence of true self-recognition in the mirror test. They use their reflection to remove a mark that is only visible in the mirror. Not all animals, however, rely primarily on vision. In lizards, chemical cues are important in social interactions. A number of lizard species show chemical self-recognition but it has never been investigated in a gecko species. Here, we test the tokay gecko (Gekko gecko) a territorial species with parental care on their ability to discriminate their own skin and faecal chemicals from those of same-sex, unfamiliar conspecifics. Geckos show a higher response rate towards chemicals from unfamiliar individuals compared to self-produced chemicals and a water control. Lizards showed self-directed behaviour, responded stronger to skin chemicals and females responded more than males. Our study provides first evidence towards self-recognition and for a social function of chemical present on faeces in tokay geckos but further tests are needed to confirm true self-recognition. Tokay geckos are an excellent model species to investigate individual recognition to demonstrate more sophisticated social cognitive abilities than have previously been attributed to reptiles.

Individuals of many species live in groups to obtain anti-predatory advantages, foraging benefits and for social reasons. Living in large groups can reduce predation, but as group size increases, competition for resources also increases. The trade-off between the advantages of group living for an individual and competition caused by it can determine group dynamics, and this trade-off can vary with environmental conditions. Shoaling behaviour, the tendency of fish to form groups, is shown to be affected by factors such as resource availability, presence of predators and conspecifics. Although studies indicate that both predation and starvation pressure in an environment can determine whether fish choose to shoal or not, whether prolonged exposure to such conditions influences shoaling behaviour remains little explored. Here, we test how predation pressure and resource availability may interactively shape the shoaling behaviour of zebrafish (Danio rerio) when exposed to combinations of these pressures over a two-week period. We find that shoal size increases with predation and decreases with starvation, and that greater predation pressure increases shoaling tendencies even under reduced food availability. Overall, we show that prolonged exposure to varying predation pressure and resource availability can together influence shoaling tendencies of fish even when such pressures are relaxed. Significant statementIn group living species, group structure and dynamics depend on various intrinsic factors and environmental stressors. Shoaling behaviour in fish, where individuals aggregate to form groups, is shown to be altered with environmental factors such as predation and resource availability. Although studies have examined the effects of these cues on shoaling behaviour, the ecological circumstances experienced by fish could also influence shoaling tendencies. We here show that shoaling behaviour is also shaped by previous experience of fish to predation and food resource availability. We check how shoaling behaviour varies with differences in predation pressure and resource availability after prolonged exposure to these conditions by measuring the shoal size and shoal cohesion in zebrafish. This study illuminates how shoaling tendencies of individuals shaped by the environmental conditions persist even when these environmental pressures are removed.

Asian small-clawed otters (Aonyx cinereus) engage in intricate activities using their paws for purposes such as food extraction, exploration, and tactile communication within their social group. However, some expressions of these behaviours do not appear to be immediately functional in captive populations. Here, we explore the expression of object manipulation behaviours (OMBs), such as rock manipulation (or juggling); the factors that may be driving their expression; and what OMBs may represent. A troop of six otters was observed at Lincolnshire Wildlife Park (United Kingdom) using camera-traps placed at three key locations in their enclosure: outside of their den, at the main feeding area, and at a rock-garden foraging area. A total of 2895 video clips were analysed using a hierarchical ethogram consisting of the behavioural categories of locomotion, stationary, vocalisation, physical affiliative, sustenance, and OMBs. The most commonly observed behaviours were walking (21.5%), lying (20.8%) and feeding (12.2%). OMBs were observed in 3.9% of total observations. Otters performed OMBs significantly more often outside their den than at the other locations. Grooming was also more frequent at this location. There was a negative association between the expression of OMBs and air temperature. Results suggest that in this otter population, OMBs were more likely an expression of play behaviour, contrasting findings reported elsewhere that rock juggling may be driven by hunger.

Conflict is one of the most critical factors affecting the behavior of animals related to their reproduction and survival, with aggressive interactions being central to acquiring resources or mating partners. This phenomenon is more common among males than females, impacting reproduction strategy and beginning of biparental care. Investigating such interactions in species closely related to social species can be illuminating, offering valuable insights into the factors that influence the emergence and maintenance of more complex social behaviors. In this context, we present a field study of male-male agonistic behavior in the wood-feeding cockroach, Panesthia angustipennis. Panesthia is the closest genus to the subsocial genus Salganea, which is known for its biparental care. Our field observations reveal a characteristic behavior where one male pushes a rival away from a female. The victorious male repeatedly returns to a specific site near the female, suggesting a strategy to minimize unnecessary conflict or protect the female. This behavior provides insights into the potential evolutionary strategies that may have evolved in the common ancestor shared by Salganea and Panesthia. Notably, the displaced males persistently reengage, highlighting the high resource value attributed to females and the consequential intensity of male competition. This study not only sheds light on the aggressive and pacifist tendencies in P. angustipennis but also contributes to understanding the evolutionary development of social structures in Salganea. Further experimental investigations into the aggressive behaviors of P. angustipennis will enhance our comprehension of the factors shaping the evolution of sociality in these species.

Living in groups offers individuals a way of reducing their risk of predation. Visual lateralisation, characterised as an asymmetry in eye-use, may offer an additional advantage to group-living animals by enabling them to manage two concurrent visual tasks simultaneously. This could enhance multi-tasking efficiency by facilitating cohesion with group mates while monitoring for threats. In our study, we examined visual lateralisation of Trinidadian guppies (Poecilia reticulata) tested either alone or in groups, in either the presence or absence of a live predator, the blue acara (Andinoacara pulcher). We consistently observed low levels of visual lateralisation across all treatments. Contrary to our expectations, however, guppies exhibited significantly higher absolute lateralisation when tested alone in the absence of the predator compared to the other treatments. Moreover, a significant left-eye bias was observed when the predator was present, and the fish showed a right-eye bias when the predator was absent. Use of a repeated measures design and assessing individual and group ID as random effects provided evidence that both relative and absolute laterality were repeatable at the group level, but there was limited evidence for repeatability at the level of the individuals. This repeatability in lateralisation when tested as a group, but not when individual fish composing these groups were tested alone, suggests social conformity in lateralisation. Our results suggest that social processes may have a significant impact on within-population variation in lateralisation.

When they are confronted to novel environments individuals may express different kinds of behaviors. They can move a little or even stay immobile with few exploration of the new environment or they can increase their activity, move a lot and explore actively the new place. Several studies on neophobia have been conducted on vertebrate species and they distinguish bold and shy patterns in animals reactions. Less studies have been conducted on invertebrates in this area. Here I present the case of the worm Eisenia fetida that I tested in open field tests in order to detect if they present different response profiles when confronted to a novel environment and how these profiles can vary depending on size factor. I was able to distinguish two profiles, a shy/philopatric reaction present in young and adults and a bold/explorer reaction that can be observed in mature worms. From the 311 worms tested one half was bold/explorer and the other half was shy/philopatric. The existence of these two profiles may explain the invasiveness of the species: some indivivuals stay and occupy the known environments and some individuals enhance their activity in order to colonize other environments.

The Great-tailed Grackle (Quiscalus mexicanus) seems to take advantage of inhospitable environments such as cities. However, it is not yet fully understood how these birds exploit hostile environments to their advantage. Casual observation seems to suggest that this species can obtain resources of biological importance such as food or nesting material from the trash. As a first approach to the problem, we located a patch outside a residential building, in a high-density urban area, where the residents left their trash for pick-up, and a group of wild Great-tailed grackles was identified as regular visitors. In total, 25 days were recorded at the site (November 2017 - January 2018). Events such as foraging, number of subjects present at the foraging site, aggressive behaviors between members of the group, and their relation with the presence or absence of the garbage collector truck were registered. The results show a higher number of grackles at the observation site and a higher frequency of foraging behaviors in the presence of garbage collection than in its absence. In its presence, the distribution of foraging during the day follows a normal distribution. In the absence, the distribution shows more variability towards the day. The highest frequency of interactions occurred between two grackles, yet there were records of up to eight subjects. The highest number of aggressions registered took place in the absence of garbage collection than in its presence. Moreover, the focal subject exhibits fewer agonistic behaviors compared to other members of the group, a result expected if the producer-scrounger game literature is considered. The outcome is explained in terms of deprivation and availability of resources. Finally, we conclude that grackles can exploit hazardous environments such as cities due to the highly social behaviors exhibited during foraging.

In some species, such as chameleon and cichlid fish, rapid body colouration changes (within seconds or minutes) serve as visual social signals in male-male competition. This study investigated the relationship between aggressive behaviour and body colouration changes in Oryzias celebensis, an Indonesian medaka fish. We analysed aggressive behaviours and corresponding body colouration changes during attack events in a controlled laboratory setting using groups of 3 adult fish in a small tank. In a triadic relationship consisting of 2 males and a female, males with blackened markings attacked more frequently than males without blackened markings and females. Additionally, we observed that the males with blackened markings were seldom attacked by males without blackened markings and females. These tendencies persisted even in groups consisting of 3 males. Our results suggest that the blackened markings in male O. celebensis not only indicate the level of aggression but also serve as a social signal to suppress attacks by other individuals.

Variation in color morph behavior is an important factor in the maintenance of color polymorphism. Alternative anti-predator behaviors are often associated with morphological traits such as coloration, possibly because predator-mediated viability selection favors certain combinations of anti-predator behavior and color. The Aegean wall lizard, Podarcis erhardii, is color polymorphic and populations can have up to three monochromatic morphs: orange, yellow, and white. We investigated whether escape behaviors differ among coexisting color morphs, and if morph behaviors are repeatable across different populations with the same predator species. Specifically, we assessed color morph flight initiation distance (FID), distance to the nearest refuge (DNR), and distance to chosen refuge (DR) in two populations of Aegean wall lizards from Naxos island. We also analyzed the type of refugia color morphs selected and their re-emergence behavior following a standardized intrusion event. We found that orange morphs have different escape behaviors from white and yellow morphs, and these differences are consistent in both of the populations we sampled. Orange morphs have shorter FIDs, DNRs, and DRs, select different refuge types, and re-emerge less often after an intruder event compared to white and yellow morphs. Observed differences in color morph escape behaviors support the idea that morphs have evolved alternative behavioral strategies that may play a role in population-level morph maintenance and loss.

Many rewilding projects emphasize recovering trophic networks through species translocation. Understanding the behaviour and movement patterns of rewilded animals is central to evaluating the success of rewilding efforts. In this study, we compared the behaviour of rewilded Persian leopards (Panthera pardus tulliana) in Russia with their wild counterparts in Iran to assess how they compare in terms of exploratory and cautious movement in new environments. We used Behavioural Change Point Analysis with selection procedures to identify points where individuals switched between movement modes. We analysed intrinsic movement metrics (speed, persistence velocity, turning velocity) and spatiotemporal attributes (duration, net squared displacement, distance travelled). Time spent in each behavioural mode differed between wild and rewilded leopards: on average, rewilded leopards spent more time in the "Ranging" mode and less time "Encamped" compared to wild leopards. Rewilded leopards were also slower than wild leopards, particularly in the "Encamped" state while wild leopards exhibited similar speeds across modes. Importantly, rewilded leopards had larger displacement and spent longer in the "Ranging" mode, a pattern not seen in wild leopards. These findings highlight distinct spatial behaviour patterns between wild and rewilded leopards. Accordingly, rewilded leopards adopted both exploratory and cautious behavioural strategies, reflecting their tendency to range more widely over extended periods, plausibly to gather information from large parts of their environments, albeit at lower speed, arguably to avoid potential risks, suggesting that they are in the process of adapting to new environments. These differences are likely to reflect different ecological circumstances, but even more so the previous experiences and thus different social circumstances of the two samples of Persian leopards. OPEN RESEARCH STATEMENTAll locational data from Iran are publicly available on Movebank: https://www.movebank.org/panel_embedded_movebank_webapp. Project: Persian leopard Tandoureh Iran (accession number 270329098). All the data are available for download as .csv file.

O_LIAnimal personality differences, i.e. consistent among-individual behavioural differences within populations, are prevalent across the animal kingdom. However, we are just beginning to understand the adaptive significance of the observed behavioural variation. We are particularly in need of empirical studies testing hypotheses of proposed theoretical frameworks aiming to understand the existence animal personality differences. In this study, we investigated a hypothesis derived from a framework suggesting that sexual selection may generate and maintain personality variation. The authors of this framework propose parental care as a mechanism linking animal personality and reproductive fitness. C_LIO_LIWe repeatedly measured individual boldness in male and female rainbow kribs, Pelvicachromis pulcher, a West African cichlid known for territorial cave breeding and shared parental care. We then formed 54 breeding pairs with varying behavioural contrasts in boldness. For pairs that produced offspring (N = 20), we repeatedly recorded parental care behaviour (parental boldness and brood guarding) of both parents over one month. Reproductive success was measured as the likelihood to reproduce, the number of offspring, and offspring size at the end of breeding. C_LIO_LIIn both sexes, we found consistent among-individual differences in boldness before breeding and in parental care behaviours. Bolder males were less likely to reproduce but, when breeding was successful, produced bigger broods compared to less bold males. Female parental boldness positively correlated with the number of offspring produced. However, individual boldness before breeding did not predict parental care behaviour in either sex and we found no effects of the pairs contrast in prebreeding boldness on parental care coordination or reproductive success. C_LIO_LIThe outcomes of our study may suggest that among-individual variation in male boldness is maintained by males with different behavioural types playing different reproductive strategies of equal average fitness. Future research should delve into understanding the intricate link between male boldness and reproductive success, exploring for instance underlying genetic mechanisms and interactions with environmental conditions. C_LI

Detailed observational research on free-ranging species of marine megafauna is required to understand their behavioural ecology, including how groups respond to environmental and anthropogenic pressures. New technologies are opening up potential for research on these species in the wild, especially on group-based and collective behaviours. Reef manta rays (Mobula alfredi) are socially interactive elasmobranchs that form groups in coastal reef habitats. Collective behaviours are likely important to their fitness, but may be disturbed by humans. Using small, remotely-piloted drones, we performed aerial observations of manta ray groups in Raja Ampat, West Papua. We empirically quantified patterns of collective movement including relative spatial positions, alignment, speed and leadership positions of conspecifics. We found unique patterns of spatial positioning, alignment and leadership, including differences between sexes, and high levels of local attraction, which were suggestive of distinct collective behaviour states. We suggest that rules of interaction in manta rays vary at the individual level and can shift depending on local environmental and social conditions. Leader-follower behaviour likely has broad importance to cohesive movement and social behaviour in M. alfredi. We suggest that further studies on manta ray movement should consider utilising models of collective motion that capture group-level behavioural processes.

We investigated the impact of captive life on behaviour and learning abilities in an enigmatic, aquifer-dwelling blind eel, Rakthamichthys digressus. Of eight major behavioural traits related to exploration and activity in a novel arena, four were significantly altered by life in captivity. While the startle response upon introduction into the arena and overall swimming away from the walls increased after captivity, inactivity exhibited immediately after the startle and the reaction to an external disturbance decreased. We also observed behavioural syndromes between startle responses and horizontal wall following, and between overall activity and vertical wall following; however, these behavioural syndromes were not altered by maintenance in captivity. Interestingly, this blind-eel failed to learn a simple spatial task in a Y-maze apparatus. Captive-associated behavioural changes in R. digressus may influence their survival after reintroduction into natural habitats, and such changes must be taken into account while developing protocols for ex-situ conservation and subsequent release.

Across many taxa, individuals learn how to detect, recognise and respond to predators via social learning. Learning to recognise and interpret predator cues is essential in the accurate assessment of risk. Cues can come directly from a predators presence (visual, acoustic) or from secondary predator cues (SPCs, such as hair/feathers, urine or faeces) left in the environment. Animals show various responses to encountering SPCs, which are thought to act in reducing risk to the individual. Meerkats, Suricata suricatta, show a response to SPCs not described in any other species: they display a mobbing-like behaviour. The function of this behaviour is unclear as unlike mobbing, the response it so closely resembles, it cannot serve to drive predators away. We used experiments to investigate whether adults may use this mobbing-like response to teach naive young how to recognise and respond to predators. Meerkats are known to teach pups hunting skills, but there is as yet no evidence that any species other than humans teaches across multiple contexts. We used experimental presentations of SPCs to test whether wild adult meerkats respond more intensely to SPCs in the presence of naive pups, as would be expected if the behaviour serves to promote learning. Contrary to this prediction, response intensity was lower when pups were present than when they were absent, and declined as the number of pups in the group increased, possibly due to costs associated with foraging with dependent young. Response intensity instead increased with increasing group size, number of group members interacting with the cue, and varied with predator cue type. These results suggest that the mobbing-like response to SPCs is not a form of teaching in meerkats. Instead, this behaviour may function to increase the recruitment of others to investigate the SPC. Exposing group members to SPCs may better inform them of the nature of the threat, facilitating more effective defensive group responses.

Behavior is often linked to gonadal sex; however, ecological or social environments can induce plasticity in sex-biased behaviors. In biparental species, pairs may divide offspring care into two parental roles, in which one parent specializes in territory defense and the other in nest care. The African cichlid fish Julidochromis marlieri displays plasticity in sex-biased behaviors. In Lake Tanganyika, J. marlieri form female-larger pairs in which the female is more aggressive than the male who performs more nest care, but under laboratory conditions, male-larger pairs can be formed in which these sex-biased behaviors are reversed. We investigated the influence of social environment on behavior by observing how individuals in both pair-types respond to conspecific intruders of either sex. We examined behavioral responses to three factors: sex of the subject, relative size of the subject, and the sex of the intruder. We confirm that relative size is a factor in behavior. The larger fish in the pair is more aggressive than the smaller fish is towards an intruder. While neither fish in the female-larger pairs varied their behaviors in response to the sex of the intruder, both members of the male-larger pairs were sensitive to intruder sex. Both individuals in the male-larger pairs engaged in more biting behaviors towards the intruder. Intruder biting behaviors strongly correlated with the biting behavior of the larger individual in the pair and occurred more frequently when encountering pairs with same sex as the larger fish when compared to pairs with the same sex as the smaller fish. Our results support the role of the social environment as a contributor in the expression of sex-biased behavior.