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Many animals use color to signal their quality and/or behavioral motivations. Colorful signals have been well studied in the contexts of competition and mate choice; however, the role of these signals in nonsexual, affiliative relationships is not as well understood. Here, we used wild social groups of the cichlid fish Neolamprologus pulcher to investigate whether the size of a brightly colored facial patch was related to 1) individual quality, 2) social dominance, and/or 3) affiliative relationships. Individuals with larger patches spent more time foraging and tended to perform more aggressive acts against conspecific territory intruders. We did not find any evidence that the size of these yellow patches was related to social rank or body size, but dominant males tended to have larger patches than dominant females. Additionally, patch size had a rank-specific relationship with the number of affiliative interactions that individuals engaged in. Dominant males with large patches received fewer affiliative acts from their groupmates compared to dominant males with small patches. However, subordinates with large patches tended to receive more affiliative acts from their groupmates while performing fewer affiliative acts themselves. Taken together, our results suggest that patch size reflects interindividual variation in foraging effort in this cichlid fish and offer some of the first evidence that colorful signals may shape affiliative relationships within wild social groups., Competing Interests: The authors declare no competing interests., (© The Author(s) 2022. Published by Oxford University Press on behalf of Editorial Office, Current Zoology.)

Size-based dominance hierarchies influence fitness, group size and population dynamics and link dominance structure to evolutionary and ecological outcomes. While larger individuals often gain dominance, social status may influence growth and size in return, resulting in feedbacks among status, growth and size. Here, we present two models evaluating how these feedbacks influence the emergence of size structure in a dominance hierarchy. In the first, size influences competition for food and investment in suppressing growth of groupmates. Stable size differences emerged when suppression was greatest for similarly sized individuals and size had little effect on competition for food. The model predicted size divergence when size strongly affected competition for food. In the second model, we used a dynamic game to solve for optimal investment in growth suppression as a function of size structure. Investment in growth suppression was favoured only when dominants and subordinates were similar in size, generating size ratios different than those expected by chance. Variation in the feedbacks among growth, size and status can explain variation in emergent size structure of dominance hierarchies and its consequences for conflict within groups. This article is part of the theme issue 'The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies'.

Our understanding of how projected climatic warming will influence the world's biota remains largely speculative, owing to the many ways in which it can directly and indirectly affect individual phenotypes. Its impact is expected to be especially severe in the tropics, where organisms have evolved in more physically stable conditions relative to temperate ecosystems. Lake Tanganyika (eastern Africa) is one ecosystem experiencing rapid warming, yet our understanding of how its diverse assemblage of endemic species will respond is incomplete. Herein, we conducted a laboratory experiment to assess how anticipated future warming would affect the mirror-elicited aggressive behaviour of Julidochromis ornatus, a common endemic cichlid in Lake Tanganyika. Given linkages that have been established between temperature and individual behaviour in fish and other animals, we hypothesized that water warming would heighten average individual aggression. Our findings support this hypothesis, suggesting the potential for water warming to mediate behavioural phenotypic expression through negative effects associated with individual health (body condition). We ultimately discuss the implications of our findings for efforts aimed at understanding how continued climate warming will affect the ecology of Lake Tanganyika fishes and other tropical ectotherms.

Conflict is an inherent part of social life in group-living species. Group members may mediate conflict through submissive and affiliative behaviors, which can reduce aggression, stabilize dominance hierarchies, and foster group cohesion. The frequency and resolution of within-group conflict may vary with the presence of neighboring groups. Neighbors can threaten the territory or resources of the whole group, promoting behaviors that foster within-group cohesion. However, neighbors may also foster conflict of interests among group members: opportunities for subordinate dispersal may alter conflict among dominants and subordinates while opportunities for extra-pair reproduction may increase conflict between mates. To understand how neighbors mediate within-group conflict in the cooperatively breeding fish Neolamprologus pulcher , we measured behavioral dynamics and social network structure in isolated groups, groups recently exposed to neighbors, and groups with established neighbors. Aggression and submission between the dominant male and female pair were high in isolated groups, but dominant aggression was directly primarily at subordinates when groups had neighbors. This suggests that neighbors attenuate conflict between mates and foster conflict between dominants and subordinates. Further, aggression and submission between similarly sized group members were most frequent when groups had neighbors, suggesting that neighbors induce rank-related conflict. We found relatively little change in within-group affiliative networks across treatments, suggesting that the presence of neighbors does not alter behaviors associated with promoting group cohesion. Collectively, these results provide some of the first empirical insights into the extent to which intragroup behavioral networks are mediated by intergroup interactions and the broader social context.

Current theoretical models of the commons assert that common-pool resources can only be managed sustainably with clearly defined boundaries around both communities and the resources that they use. In these theoretical models, open access inevitably leads to a tragedy of the commons. However, in many open-access systems, use of common-pool resources seems to be sustainable over the long term (i.e., current resource use does not threaten use of common-pool resources for future generations). Here, we outline the conditions that support sustainable resource use in open property regimes. We use the conceptual framework of complex adaptive systems to explain how processes within and couplings between human and natural systems can lead to the emergence of efficient, equitable, and sustainable resource use. We illustrate these dynamics in eight case studies of different social-ecological systems, including mobile pastoralism, marine and freshwater fisheries, swidden agriculture, and desert foraging. Our theoretical framework identifies eight conditions that are critical for the emergence of sustainable use of common-pool resources in open property regimes. In addition, we explain how changes in boundary conditions may push open property regimes to either common property regimes or a tragedy of the commons. Our theoretical model of emergent sustainability helps us to understand the diversity and dynamics of property regimes across a wide range of social-ecological systems and explains the enigma of open access without a tragedy. We recommend that policy interventions in such self-organizing systems should focus on managing the conditions that are critical for the emergence and persistence of sustainability., Competing Interests: The authors declare no conflict of interest., (Copyright © 2018 the Author(s). Published by PNAS.)

Metabolic rate reduction has been considered the mechanism by which sleep conserves energy, similar to torpor or hibernation. This mechanism of energy savings is in conflict with the known upregulation (compared to wake) of diverse functions during sleep and neglects a potential role in energy conservation for partitioning of biological operations by behavioral state. Indeed, energy savings as derived from state-dependent resource allocations have yet to be examined. A mathematical model is presented based on relative rates of energy deployment for biological processes upregulated during either wake or sleep. Using this model, energy savings from sleep-wake cycling over constant wakefulness is computed by comparing stable limit cycles for systems of differential equations. A primary objective is to compare potential energy savings derived from state-dependent metabolic partitioning versus metabolic rate reduction. Additionally, energy conservation from sleep quota and the circadian system are also quantified in relation to a continuous wake condition. As a function of metabolic partitioning, our calculations show that coupling of metabolic operations with behavioral state may provide comparatively greater energy savings than the measured decrease in metabolic rate, suggesting that actual energy savings derived from sleep may be more than 4-fold greater than previous estimates. A combination of state-dependent metabolic partitioning and modest metabolic rate reduction during sleep may enhance energy savings beyond what is achievable through metabolic partitioning alone; however, the relative contribution from metabolic partitioning diminishes as metabolic rate is decreased during the rest phase. Sleep quota and the circadian system further augment energy savings in the model. Finally, we propose that state-dependent resource allocation underpins both sleep homeostasis and the optimization of daily energy conservation across species. This new paradigm identifies an evolutionary selective advantage for the upregulation of central and peripheral biological processes during sleep, presenting a unifying construct to understand sleep function.

Social living has evolved numerous times across a diverse array of animal taxa. An open question is how the transition to a social lifestyle has shaped, and been shaped by, the underlying neurohormonal machinery of social behaviour. The nonapeptide neurohormones, implicated in the regulation of social behaviours, are prime candidates for the neuroendocrine substrates of social evolution. Here, we examined the brains of eight cichlid fish species with divergent social systems, comparing the number and size of preoptic neurons that express the nonapeptides isotocin and vasotocin. While controlling for the influence of phylogeny and body size, we found that the highly social cooperatively breeding species ( n  = 4) had fewer parvocellular isotocin neurons than the less social independently breeding species ( n  = 4), suggesting that the evolutionary transition to group living and cooperative breeding was associated with a reduction in the number of these neurons. In a complementary analysis, we found that the size and number of isotocin neurons significantly differentiated the cooperatively breeding from the independently breeding species. Our results suggest that isotocin is related to sociality in cichlids and may provide a mechanistic substrate for the evolution of sociality., Competing Interests: The authors declare no competing interests.

The degree to which group members share reproduction is dictated by both within-group (e.g. group size and composition) and between-group(e.g. density and position of neighbours) characteristics. While many studies have investigated reproductive patterns within social groups, few have simultaneously explored how within-group and between-group social structure influence these patterns. Here, we investigated how group size and composition, along with territory density and location within the colony, influenced parentage in 36 wild groups of a colonial, cooperatively breeding fish Neolamprologus pulcher. Dominant males sired 76% of offspring in their group, whereas dominant females mothered 82% of offspring in their group. Subordinate reproduction was frequent, occurring in 47% of sampled groups. Subordinate males gained more paternity in groups located in high-density areas and in groups with many subordinate males. Dominant males and females in large groups and in groups with many reproductively mature subordinates had higher rates of parentage loss, but only at the colony edge. Our study provides, to our knowledge,the first comprehensive quantification of reproductive sharing among groups of wild N. pulcher, a model species for the study of cooperation and social behaviour. Further, we demonstrate that the frequency of extra-pair parentage differs across small social and spatial scales.

In many non-monogamous systems, males invest less in progeny than do females. This leaves males with higher potential rates of reproduction, and a likelihood of sexual conflict, including, in some systems, coercive matings. If coercive matings are costly, the best female strategy may be to avoid male interaction. We present a model that demonstrates female movement in response to male harassment as a mechanism to lower the costs associated with male coercion, and the effect that female movement has on selection in males for male harassment. We found that, when females can move from a habitat patch to a refuge to which males do not have access, there may be a selection for either high, or low harassment male phenotype, or both, depending on the relationship between the harassment level of male types in the population and a threshold level of male harassment. This threshold harassment level depends on the relative number of males and females in the population, and the relative resource values of the habitat; the threshold increases as the sex ratio favours females, and decreases with the value of the refuge patch or total population. Our model predicts that selection will favour the harassment level that lies closest to this threshold level of harassment, and differing harassment levels will coexist within the population only if they lie on the opposite sides of the threshold harassment. Our model is consistent with empirical results suggesting that an intermediate harassment level provides maximum reproductive fitness to males when females are mobile.

Behaviour is typically regarded as among the most flexible of animal phenotypic traits. In particular, expression of cooperative behaviour is often assumed to be conditional upon the behaviours of others. This flexibility is a key component of many hypothesized mechanisms favouring the evolution of cooperative behaviour. However, evidence shows that cooperative behaviours are often less flexible than expected and that, in many species, individuals show consistent differences in the amount and type of cooperative and non-cooperative behaviours displayed. This phenomenon is known as 'animal personality' or a 'behavioural syndrome'. Animal personality is evolutionarily relevant, as it typically shows heritable variation and can entail fitness consequences, and hence, is subject to evolutionary change. Here, we review the empirical evidence for individual variation in cooperative behaviour across taxa, we examine the evolutionary processes that have been invoked to explain the existence of individual variation in cooperative behaviour and we discuss the consequences of consistent individual differences on the evolutionary stability of cooperation. We highlight that consistent individual variation in cooperativeness can both stabilize or disrupt cooperation in populations. We conclude that recognizing the existence of consistent individual differences in cooperativeness is essential for an understanding of the evolution and prevalence of cooperation.

Background: In many cooperatively breeding vertebrates, subordinates assist a dominant pair to raise the dominants' offspring. Previously, it has been suggested that subordinates may help in payment for continued residency on the territory (the 'pay-to-stay hypothesis'), but payment might also be reciprocated or might allow subordinates access to reproductive opportunities., Methodology/principal Findings: We measured dominant and subordinate female alloparental brood care and reproductive success in four separate experiments and show that unrelated female dominant and subordinate cichlid fish care for each other's broods (alloparental brood care), but that there is no evidence for reciprocal 'altruism' (no correlation between alloparental care received and given). Instead, subordinate females appear to pay with alloparental care for own direct reproduction., Conclusions/significance: Our results suggest subordinate females pay with alloparental care to ensure access to the breeding substrate and thereby increase their opportunities to lay their own clutches. Subordinates' eggs are laid, on average, five days after the dominant female has produced her first brood. We suggest that immediate reproductive benefits need to be considered in tests of the pay-to-stay hypothesis.

1. Tiger sharks Galeocerdo cuvier are important predators in a variety of nearshore communities, including the seagrass ecosystem of Shark Bay, Western Australia. Because tiger sharks are known to influence spatial distributions of multiple prey species, it is important to understand how they use habitats at a variety of spatial scales. We used a combination of catch rates and acoustic tracking to determine tiger shark microhabitat use in Shark Bay. 2. Comparing habitat-use data from tracking against the null hypothesis of no habitat preference is hindered in Shark Bay, as elsewhere, by the difficulty of defining expected habitat use given random movement. We used randomization procedures to generate expected habitat use in the absence of habitat preference and expected habitat use differences among groups (e.g. males and females). We tested the performance of these protocols using simulated data sets with known habitat preferences. 3. The technique correctly classified sets of simulated tracks as displaying a preference or not and was a conservative test for differences in habitat preferences between subgroups of tracks (e.g. males vs. females). 4. Sharks preferred shallow habitats over deep ones, and preferred shallow edge microhabitats over shallow interior ones. The use of shallow edges likely increases encounter rates with potential prey and may have profound consequences for the dynamics of Shark Bay's seagrass ecosystem through indirect effects transmitted by grazers that are common prey of tiger sharks. 5. Females showed a greater tendency to use shallow edge microhabitats than did males; this pattern was not detected by traditional analysis techniques. 6. The randomization procedures presented here are applicable to many field studies that use tracking by allowing researchers both to determine overall habitat preferences and to identify differences in habitat use between groups within their sample.

How cooperation and altruism among non-relatives can persist in the face of cheating remains a key puzzle in evolutionary biology. Although mechanisms such as direct and indirect reciprocity and limited movement have been put forward to explain such cooperation, they cannot explain cooperation among unfamiliar, highly mobile individuals. Here we show that cooperation may be evolutionarily stable if decisions taken to cooperate and to change group membership are both dependent on anonymous social experience (generalized reciprocity). We find that a win-stay, lose-shift rule (where shifting is either moving away from the group or changing tactics within the group after receiving defection) evolves in evolutionary simulations when group leaving is moderately costly (i.e. the current payoff to being alone is low, but still higher than that in a mutually defecting group, and new groups are rarely encountered). This leads to the establishment of widespread cooperation in the population. If the costs of group leaving are reduced, a similar group-leaving rule evolves in association with cooperation in pairs and exploitation of larger anonymous groups. We emphasize that mechanisms of assortment within populations are often behavioural decisions and should not be considered independently of the evolution of cooperation.

Unrelated subordinates may invest in costly help to avoid being evicted from groups (the 'pay-to-stay' hypothesis). However, the effectiveness of eviction to enforce help should depend on its being applied accurately and on the costs it imposes on both dominants and subordinates. The relative cost of being evicted is a function of the population frequency of eviction when population growth is limited by density-dependent factors. We describe a stage-structured pay-to-stay model incorporating density-dependent population growth, costly eviction and occasional errors. Breeders demand some amount of help and evict subordinates that do not provide it. Helpers decide on the amount of help they will provide. The threat of eviction alone is sufficient to enforce helping. However, helping will not be favoured if helpers do not impose costs on breeders. The amount of help provided is less than the cost that subordinates impose upon breeders, when any help is provided. Thus, the net fitness effect of a helper under pay-to-stay alone is negative, even if it is investing in cooperative behaviour. Constraints on dispersal have no effect on the amount of help, although they may influence the tolerance threshold of breeders and group stability, depending on the mechanism of density dependence.

Hawk-dove games have been extensively used to predict the conditions under which group-living animals should defend their resources against potential usurpers. Typically, game-theoretic models on aggression consider that resource defense may entail energetic and injury costs. However, intruders may also take advantage of owners who are busy fighting to sneak access to unguarded resources, imposing thereby an additional cost on the use of the escalated hawk strategy. In this article we modify the two-strategy hawk-dove game into a three-strategy hawk-dove-sneaker game that incorporates a distraction-sneaking tactic, allowing us to explore its consequences on the expected level of aggression within groups. Our model predicts a lower proportion of hawks and hence lower frequencies of aggressive interactions within groups than do previous two-strategy hawk-dove games. The extent to which distraction sneakers decrease the frequency of aggression within groups, however, depends on whether they search only for opportunities to join resources uncovered by other group members or for both unchallenged resources and opportunities to usurp.

Abstract: Individuals that produce resources are often exploited by several individuals; such exploitation may or may not be tolerated. We modeled the decision of a resource owner to accept one scrounger (the "satellite") and of both of these to accept being joined by another (the "floater"). In general, satellites tolerated floaters when competition between them was low, while owners tolerated satellites when facilitation between satellite and floater was high. When floaters were likely to find resources without joining, owners were more likely to resist satellites. In some cases, Nash equilibria were also mutually beneficial for two of the three individuals. Our model makes the counterintuitive prediction that mutually beneficial coalitions between satellites and floaters can only arise when the net benefits arising from the other's presence are low. When facilitation between satellites and floaters is high, satellites and owners may form mutually beneficial groups and groups with division of labor, or alternatively, owners may benefit from encouraging floaters to join. Finally, our model suggests there must be differences in competitive ability or some benefit of familiarity for owners to tolerate satellites but not floaters. We discuss empirical evidence for these and other predictions of the model.

Foraging animals must often balance the conflicting demands of finding food and avoiding predators. Temporal variation in predation risk is expected to influence how animals allocate time to these behaviours. Counterintuitively, the proportion of time spent foraging during both high- and low-risk periods should increase with increasing time exposed to high risk. We tested this prediction using intertidal marine snails (Littorina spp.) that were exposed to temporal variation in perceived predation risk from crabs (Cancer productus and Cancer magister). Our results were consistent with those predicted for high-risk, but not low-risk, periods. During high-risk periods, a greater number of snails foraged (versus those that left the water or remained in their shells) as time at high perceived risk increased. For low-risk periods, there was no relationship between the number of snails foraging and time at high risk. This might be due to snails in all treatments foraging maximally in the low-risk periods. As a consequence, the difference in the number of snails foraging between high- and low-risk periods decreased with increasing time subject to high risk. These results indicate that the commonly used protocol of exposing foragers to a single pulse of heightened risk might tend to overestimate their typical investment in anti-predator behaviour.

Despite convincing in vitro evidence, a vitamin C-E interaction has not been confirmed in vivo. This study was designed to examine the effects of supplementation with either vitamin C or E on their respective plasma concentrations, other antioxidants, lipids and some haemostatic variables. Fasting blood was collected before and after intervention from thirty healthy adults in a double-blinded crossover study. Baselines for measured variables were established after 2 weeks of placebo supplementation, followed by daily supplementation with 73.5 mg RRR-alpha-tocopherol acetate or 500 mg ascorbic acid, and placebo, for 6 weeks. A 2 month washout preceded supplement crossover. Mean values showed that plasma lipid standardised alpha-tocopherol increased with ascorbic acid supplementation: from 4.09 (sem 0.51) to 4.53 (sem 0.66) micromol/mmol total cholesterol plus triacylglycerol (P < 0.05), and plasma ascorbic acid increased from 62.8 (sem 14.9) to 101.3 (sem 22. 2) micromol/l (P < 0.005). Supplementation with (RRR)-alpha-tocopherol acetate increased plasma alpha-tocopherol from 26.8 (sem 3.9) to 32.2 (sem 3.8) micromol/l (P < 0.05), and lipid-standardised alpha-tocopherol from 4.12 (sem 0.48) to 5.38 (sem 0.52) micromol/mmol (P < 0.001). Mean plasma ascorbic acid also increased with vitamin E supplementation, from 64.4 (sem 13.3) to 76. 4 (sem 18.4) micromol/l (P < 0.05). Plasma ferric reducing (antioxidant) power and glutathione peroxidase (U/g haemoglobin) increased in both groups, while urate, total cholesterol and triacylglycerol levels decreased (P < 0.05 throughout). Results are supportive of an in vivo interaction between vitamins C and E.

I apply skew theory to the division of resources within multimember social foraging and antipredator groups. Resource division is modeled as a game between an individual controlling resources (recruiter) and a potential joiner to the group. If a recruiter benefits from the presence of a joiner, it will allocate sufficient resources to the joiner so that the joiner gains as much from group foraging as it would from foraging alone. Joiners should receive a greater proportion of resources controlled by the recruiter when benefits to grouping are low. If group success is a concave-down function of group size, this framework can be used to predict the stable group size. The stable group size is larger than the optimal group size, given equal division of resources, and smaller than the stable group size, given equal division of resources. Furthermore, both current group members and potential joiners agree on the stable group size, so long as the recruiter is able to control resource division. If the recruiter cannot control resource division, there may be conflict over group size and the opportunity for group members to contribute less to group success than they are able.

Juvenile flying animals, including bats, often cease increasing, or even decrease in body mass after the onset of flight. The low body mass of juvenile bats may be a result of stresses associated with the initiation of flight, or may be an adaptation to reduce flight costs. To test these hypotheses, the body masses of adults and unknown aged juvenile big brown bats (Eptesicus fuscus) were separately compared in August 1990 through to 1995. It was hypothesized that there should be little variation in the body mass of juveniles from year to year if low body mass is an adaption to reduce flight costs. The foraging behaviour and body masses of bats of known or estimated age were also compared in 1994 and 1995. Ambient temperature at sunset was greater in 1994 than in 1995. It was assumed that prey density and predictability increased with temperature, and hypothesized that if low body mass results from energetic stress, then this should be more apparent in 1995. Using radiotelemetry, the foraging behaviour of juvenile, yearling and adult E. fuscus was compared in 1994 and 1995 to determine whether young bats attempt to compensate for energetic shortfalls by foraging for longer, or emerging earlier, than adults. Unknown-aged juveniles did not differ significantly in body mass in August from 1990 through to 1995. Adults were significantly heavier in August in 1990 and 1994 than in 1991, 1993 and 1995. Known-aged juveniles had lower body mass than adults in 1994, and lower wing loading than adults in 1995. Known-aged juveniles did not differ in mass between 1994 and 1995, while adults were heavier in 1994. The foraging times for juvenile and adult bats were both significantly related to ambient temperature. However, the slope of this relationship was steeper for weaned juveniles than for adults. The foraging times of yearling and adults bats were not significantly different. The emergence times of juvenile and adult bats did not differ, nor did the emergence times of yearlings and adults. Emergence time did not differ significantly between 1994 and 1995. The results of the study suggest that juvenile bats maintain a low body mass even under conditions that permit adults to accumulate greater fat deposits. Juveniles do not forage earlier or for longer to compensate for poor foraging ability and increased energy expenditures resulting from the onset of flight. This suggests that, by maintaining a low body mass, juveniles reduce flight costs and the risk of predation at a time when flight and foraging are still developing.