Variation in farming damselfish behaviour creates a competitive landscape of risk on coral reefs.

Interspecific interactions are fundamental drivers of animal space use. Yet while non-consumptive effects of predation risk on prey space use are well-known, the risk of aggressive interactions on space use of competitors is largely unknown. We apply the landscape of risk framework to competition-driven space use for the first time, with the hypothesis that less aggressive competitors may alter their behaviour to avoid areas of high competitor density. Specifically, we test how aggressive risk from territorial algal-farming damselfishes can shape the spatial distribution of herbivore fish competitors. We found that only the most aggressive damselfish had fewer competitors in their surrounding area, demonstrating that individual-level behavioural variation can shape spatial distributions. In contradiction to the landscape of risk framework, abundances of farming damselfish and other fishes were positively associated. Our results suggest that reef fishes do not simply avoid areas of high damselfish abundance, but that spatial variation in aggressive behaviour, rather than of individuals, created a competitive landscape of risk. We emphasize the importance of individual-level behaviour in identifying patterns of space use and propose expanding the landscape of risk framework to non-predatory interactions to explore cascading behavioural responses to aggressive risk.

Movement diversity and partial sympatry of coastal and Northeast Arctic cod ecotypes at high latitudes.

Movement diversity within species represent an important but often neglected, component of biodiversity that affects ecological and genetic interactions, as well as the productivity of exploited systems. By combining individual tracking data from acoustic telemetry with novel genetic analyses, we describe the movement diversity of two Atlantic cod Gadus morhua ecotypes in two high-latitude fjord systems: the highly migratory Northeast Arctic cod (NEA cod) that supports the largest cod fishery in the world, and the more sedentary Norwegian coastal cod, which is currently in a depleted state. As predicted, coastal cod displayed a higher level of fjord residency than NEA cod. Of the cod tagged during the spawning season, NEA cod left the fjords permanently to a greater extent and earlier compared to coastal cod, which to a greater extent remained resident and left the fjords temporarily. Despite this overall pattern, horizontal movements atypical for the ecotypes were common with some NEA cod remaining within the fjords year-round and some coastal cod displaying a low fjord fidelity. Fjord residency and exit timing also differed with spawning status and body size, with spawning cod and large individuals tagged during the feeding season more prone to leave the fjords and earlier than non-spawning and smaller individuals. While our results confirm a lower fjord dependency for NEA cod, they highlight a movement diversity within each ecotype and sympatric residency between ecotypes, previously undetected by population-level monitoring. This new knowledge is relevant for the management, which should base their fisheries advice for these interacting ecotypes on their habitat use and seasonal movements.

Conspecific density and environmental complexity impact behaviour of turquoise killifish (Nothobranchius furzeri).

Fish models are essential for research in many biological and medical disciplines. With a typical lifespan of only 6 months, the Turquoise killifish (Nothobranchius furzeri) was recently established as a time- and cost-efficient model to facilitate whole-life and multigenerational studies in several research fields, including behavioural ecotoxicology. Essential information on the behavioural norm and on how laboratory conditions affect behaviour, however, is deficient. In the current study, we examined the impact of the social and structural environment on a broad spectrum of behavioural endpoints in N. furzeri. While structural enrichment affected only fish boldness and exploratory behaviour, fish rearing density affected the total body length, locomotor activity, boldness, aggressiveness and feeding behaviour of N. furzeri individuals. Overall, these results contribute to compiling a behavioural baseline for N. furzeri that increases the applicability of this new model species. Furthermore, our findings will fuel the development of improved husbandry protocols to maximize the welfare of N. furzeri in a laboratory setting.

Cortisol during adolescence organises personality traits and behavioural syndromes.

Despite the growing evidence for the importance of developmental experiences shaping consistent individual differences in behaviour and physiology, the role of endocrine factors underlying the development and maintenance of such differences across multiple traits, remains poorly understood. Here, we investigated how an experimental manipulation of circulating glucocorticoids during early adolescence affects behavioural and physiological variation and covariation later in life in the precocial cavy (Cavia aperea). Plasma cortisol concentrations were experimentally elevated by administering cortisol via food for 3 weeks. Struggle docility, escape latency, boldness, exploration and social behaviour were then tested three times after individuals attained sexual maturity. In addition, blood samples were taken repeatedly to monitor circulating cortisol concentrations. Exogenous cortisol affected mean trait expression of plasma cortisol levels, struggle docility and escape latency. Repeatability of cortisol and escape latency was increased and repeatability of struggle docility tended to be higher (approaching significance) in treated individuals. Increased repeatability was mainly caused by an increase of among-individual variance. Correlations among docility, escape latency and cortisol were stronger in treated animals compared to control animals. These results suggest that exposure to elevated levels of cortisol during adolescence can alter animal personality traits as well as behavioural syndromes. Social and risk-taking traits showed no correlation with cortisol levels and were unaffected by the experimental manipulation, indicating behavioural modularity. Taken together, our data highlight that cortisol can have organising effects during adolescence on the development of personality traits and behavioural syndromes, adding to the increasing evidence that not only early life but also adolescence is an important sensitive period for behavioural development.

Behavioural variation and plasticity along an invasive ant introduction pathway.

Once established in new areas, introduced species may exhibit changes in their biology due to phenotypic plasticity, novel selection pressures and genetic drift. Moreover, the introduction process itself has been hypothesised to act as a selective filter for traits that promote invasiveness. We tested the hypothesis that behaviours thought to promote invasiveness-such as increased foraging activity and aggression-are selected for during invasion by comparing traits among native and introduced populations of the widespread Argentine ant (Linepithema humile). We studied Argentine ant populations in the native range in Argentina and in three invaded regions along an introduction pathway: California, Australia and New Zealand. In each region, we set up 32 experimental colonies to measure foraging activity and interspecific aggression in a subset of the study regions. These colonies were subject to experimental manipulation of carbohydrate availability and octopamine, a biogenic amine known to modulate behaviour in insects, to measure variation in behavioural plasticity. We found variation in foraging activity among populations, but this variation was not consistent with selection on behaviour in relation to the invasion process. We found that colonies with limited access to carbohydrates exhibited unchanged exploratory behaviour, but higher exploitation activity and lower aggression. Colonies given octopamine consistently increased foraging behaviour (both exploration and exploitation), as well as aggression when also sugar-deprived. There was no difference in the degree of behavioural response to our experimental treatments along the introduction pathway. We did not find support for selection of behavioural traits associated with invasiveness along the Argentine ant's introduction pathway or clear evidence for an association between the introduction process and variation in behavioural plasticity. These results indicate that mechanisms promote behavioural variation in a similar fashion both in native and introduced ranges. Our results challenge the assumption that introduced populations always perform better in key behavioural traits hypothesised to be associated with invasion success.

Metabolic rate associates with, but does not generate covariation between, behaviours in western stutter-trilling crickets, Gryllus integer.

The causes and consequences of among-individual variation and covariation in behaviours are of substantial interest to behavioural ecology, but the proximate mechanisms underpinning this (co)variation are still unclear. Previous research suggests metabolic rate as a potential proximate mechanism to explain behavioural covariation. We measured the resting metabolic rate (RMR), boldness and exploration in western stutter-trilling crickets, Gryllus integer, selected differentially for short and fast development over two generations. After applying mixed-effects models to reveal the sign of the covariation, we applied structural equation models to an individual-level covariance matrix to examine whether the RMR generates covariation between the measured behaviours. All traits showed among-individual variation and covariation: RMR and boldness were positively correlated, RMR and exploration were negatively correlated, and boldness and exploration were negatively correlated. However, the RMR was not a causal factor generating covariation between boldness and exploration. Instead, the covariation between all three traits was explained by another, unmeasured mechanism. The selection lines differed from each other in all measured traits and significantly affected the covariance matrix structure between the traits, suggesting that there is a genetic component in the trait integration. Our results emphasize that interpretations made solely from the correlation matrix might be misleading.

Individual versus pseudo-repeatability in behaviour: Lessons from translocation experiments in a wild insect.

Repeatability represents a key parameter in ecological and evolutionary research. Repeatability is underpinned by developmental plasticity and genetic variation but may become biased upwards by repeatable differences in environments to which individuals respond plastically. The extent of upward bias caused by the latter mechanism (causing "pseudo-repeatability") is important yet rarely investigated in ecological research. We repeatedly assayed a key behaviour (flight initiation distance) affecting longevity in a wild cricket population (Gryllus campestris). We used naturally moving, translocated and forced-stationary individuals to study bias in repeatability caused by spatial variability in environmental conditions. Our experiments acknowledged that translocations might themselves bias repeatability estimates if animals respond to handling procedures (a necessary component of translocations). Individuals were, therefore, either (i) repeatedly translocated and assayed or (ii) assayed at multiple burrows as part of natural movements. This enabled estimation of behavioural variance attributable to individual, burrow and residual components within each treatment; comparison across treatments addressed whether translocations caused bias. We also calculated repeatability for individuals that were forced to be stationary to investigate whether this led to upward bias of repeatability. For adult crickets, individual explained 17.8% versus 17.2%, and burrow 8.7% versus 10.3%, of the behavioural variance in translocated versus natural-movement treatments. Repeatability for forced-stationary adults was 31.1%, thereby demonstrating experimentally that certain study designs bias repeatability upwards. For translocated juveniles, individual explained 10.0% and burrow 6.0% of the variance, while in the natural-movement treatment, those components could not be separated as juveniles do not switch burrows. Translocations did not lead to detectable biases in behavioural mean or variance. Repeatability was not biased for adults subjected to the natural-movement treatment because individuals were assayed under many different environments, facilitating the separation of individual from burrow effects. Upward bias would have occurred with less optimal sampling schemes: if individuals had been assayed repeatedly at the same burrow. We, therefore, recommend that translocation experiments are more commonly applied, particularly in stationary species, to ensure the unbiased estimation of repeatability.

Among-individual behavioural variation in the ornamental red cherry shrimp, Neocaridina heteropoda.

Personality variation, defined as among-individual differences in behaviour that are repeatable across time and context, is widely reported across animal taxa. From an evolutionary perspective, characterising the amount and structure of this variation is useful since differences among individuals are the raw material for adaptive behavioural evolution. However, behavioural variation among individuals also has implications for more applied areas of evolution and ecology-from invasion biology to ecotoxicology and selective breeding in captive systems. Here, we investigate the structure of personality variation in the red cherry shrimp, Neocaridina heteropoda, a popular ornamental species that is readily kept and bred under laboratory conditions and is emerging as a decapod crustacean model across these fields, but for which basic biological, ecological and behavioural data are limited. Using two assays and a repeated measures approach, we quantify behaviours putatively indicative of shy-bold variation and test for sexual dimorphism and/or size-dependent behaviours (as predicted by some state-dependent models of personality). We find moderate-to-high behavioural repeatabilities in most traits. Although strong individual-level correlations across behaviours are consistent with a major personality axis underlying these observed traits, the multivariate structure of personality variation does not fully match a priori expectations of a shy-bold axis. This may reflect our ecological naivety with respect to what really constitutes bolder, more risk-prone, behaviour in this species. We find no evidence for sexual dimorphism and only weak support for size-dependent behaviour. Our study contributes to the growing literature describing behavioural variation in aquatic invertebrates. Furthermore, it lays a foundation for further studies harnessing the potential of this emerging model system. In particular, this existing behavioural variation could be functionally linked to life-history traits and invasive success and serve as a target of artificial selection or bioassays. It thus holds significant promise in applied research across ecotoxicology, aquaculture and invasion biology.

The role of social attraction and social avoidance in shaping modular networks.

How interactions between individuals contribute to the emergence of complex societies is a major question in behavioural ecology. Nonetheless, little remains known about the type of immediate social structure (i.e. social network) that emerges from relationships that maximize beneficial interactions (e.g. social attraction towards informed individuals) and minimize costly relationships (e.g. social avoidance of infected group mates). We developed an agent-based model where individuals vary in the degree to which individuals signal benefits versus costs to others and, on this basis, choose with whom to interact depending on simple rules of social attraction (e.g. access to the highest benefits) and social avoidance (e.g. avoiding the highest costs). Our main findings demonstrate that the accumulation of individual decisions to avoid interactions with highly costly individuals, but that are to some extent homogeneously beneficial, leads to more modular networks. On the contrary, individuals favouring interactions with highly beneficial individuals, but that are to some extent homogeneously costly, lead to less modular networks. Interestingly, statistical models also indicate that when individuals have multiple potentially beneficial partners to interact with, and no interaction cost exists, this also leads to more modular networks. Yet, the degree of modularity is contingent upon the variability in benefit levels held by individuals. We discuss the emergence of modularity in the systems and their consequences for understanding social trade-offs.

Short-time development of among-colony behaviour in a high-elevation ant.

Standardised assays are often used to characterise aggression in animals. In ants, such assays can be applied at several organisational levels (e.g., colony, population) and at specific times during the season. However, whether the behaviour differs at these levels and changes over a few weeks remains largely unexplored. Here, six colonies from the high-elevation ant Tetramorium alpestre were collected weekly for five weeks from two behaviourally-different populations (aggressive and peaceful in intraspecific encounters). We conducted one-on-one worker encounters at the colony and population levels. When analysing the colony combinations separately, the behaviour was peaceful and remained so within the peaceful population; initial aggression became partially peaceful within the aggressive population; and initial aggression decreased occasionally and increased in one combination but remained constant for most across-population combinations. When analysing all colony combinations together, within-population behaviour remained similar, but across-population behaviour became peaceful. The observed behavioural differences among organisational levels emphasise the relevance of assessing both levels. Moreover, the effect of decreasing aggression is discernible already over a few weeks. Compression of the vegetation period at high elevations may compress such behavioural changes. Addressing both organisational levels and seasonality is important, particularly in studies of behavioural complexity such as in this ant.

Harvester ant colonies differ in collective behavioural plasticity to regulate water loss.

Collective behavioural plasticity allows ant colonies to adjust to changing conditions. The red harvester ant (Pogonomyrmex barbatus), a desert seed-eating species, regulates foraging activity in response to water stress. Foraging ants lose water to evaporation. Reducing foraging activity in dry conditions sacrifices food intake but conserves water. Within a year, some colonies tend to reduce foraging on dry days while others do not. We examined whether these differences among colonies in collective behavioural plasticity persist from year to year. Colonies live 20-30 years with a single queen who produces successive cohorts of workers which live only a year. The humidity level at which all colonies tend to reduce foraging varies from year to year. Longitudinal observations of 95 colonies over 5 years between 2016 and 2021 showed that differences among colonies, in how they regulate foraging activity in response to day-to-day changes in humidity, persist across years. Approximately 40% of colonies consistently reduced foraging activity, year after year, on days with low daily maximum relative humidity; approximately 20% of colonies never did, foraging as much or more on dry days as on humid days. This variation among colonies may allow evolutionary rescue from drought due to climate change.

Inter-community behavioural variation confirmed through indirect methods in four neighbouring chimpanzee communities in Cantanhez NP, Guinea-Bissau.

Culture, while long viewed as exclusively human, has now been demonstrated across diverse taxa and contexts. However, most animal culture data are constrained to well-studied, habituated groups. This is the case for chimpanzees, arguably the most 'cultural' non-human species. While much progress has been made charting wild chimpanzees' cultural repertoire, large gaps remain in our knowledge of the majority of the continent's chimpanzees. Furthermore, few studies have compared neighbouring communities, despite such comparisons providing the strongest evidence for culture, and few have studied communities living in anthropogenic habitats although their culture is in imminent danger of disappearing. Here we combine direct, indirect and remote methods, including camera traps, to study, over 2 years, four unhabituated neighbouring chimpanzee communities inhabiting human-impacted habitats in Cantanhez NP, Guinea-Bissau. From traces collected during 1089 km of reconnaissance walks and 4197 videos from 56 camera trap locations, we identified 18 putative cultural traits. These included some noteworthy novel behaviours for these communities, and behaviours possibly new to the species. We created preliminary behavioural profiles for each community, and found inter-community differences spanning tool use, communication, and social behaviour, demonstrating the importance of comparing neighbouring communities and of studying previously neglected communities including those inhabiting anthropogenic landscapes.

Behavioural variation and learning across the lifespan in wild white-faced capuchin monkeys.

Natural selection has evidently mediated many species characteristics relevant to the evolution of learning, including longevity, length of the juvenile period, social organization, timing of cognitive and motor development, and age-related shifts in behavioural propensities such as activity level, flexibility in problem-solving and motivation to seek new information. Longitudinal studies of wild populations can document such changes in behavioural propensities, providing critical information about the contexts in which learning strategies develop, in environments similar to those in which learning strategies evolved. The Lomas Barbudal Monkey Project provides developmental data for the white-faced capuchin, Cebus capucinus, a species that has converged with humans regarding many life-history and behavioural characteristics. In this dataset, focused primarily on learned aspects of foraging behaviour, younger capuchins are more active overall, more curious and opportunistic, and more prone to inventing new investigative and foraging-related behaviours. Younger individuals more often seek social information by watching other foragers (especially older foragers). Younger individuals are more creative, playful and inventive, and less neophobic, exhibiting a wider range of behaviours when engaged in extractive foraging. Whereas adults more often stick with old solutions, younger individuals often incorporate recently acquired experience (both social and asocial) when foraging. This article is part of the theme issue 'Life history and learning: how childhood, caregiving and old age shape cognition and culture in humans and other animals'.