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2020 Elsevier Inc. Individuals often respond to social disturbances by increasing prosociality, which can strengthen social bonds, buffer against stress, and promote overall group cohesion. Given their importance in mediating stress responses, glucocorticoids have received considerable attention as potential proximate regulators of prosocial behaviour during disturbances. However, previous investigations have largely focused on mammals and our understanding of the potential prosocial effects of glucocorticoids across vertebrates more broadly is still lacking. Here, we assessed whether experimentally elevated glucocorticoid levels (simulating endogenous cortisol responses mounted following disturbances) promote prosocial behaviours in wild groups of the cichlid fish, Neolamprologus pulcher. Using SCUBA in Lake Tanganyika, we observed how subordinate group members adjusted affiliation, helping, and submission (all forms of prosocial behaviour) following underwater injections of either cortisol or saline. Cortisol treatment reduced affiliative behaviours-but only in females-suggesting that glucocorticoids may reduce overall prosociality. Fish with elevated glucocorticoid levels did not increase performance of submission or helping behaviours. Taken together, our results do not support a role for glucocorticoids in promoting prosocial behaviour in this species and emphasize the complexity of the proximate mechanisms that underlie prosociality.

© 2020 The Fisheries Society of the British Isles The formation of mixed-species social groups, whereby heterospecifics form and maintain either transient or stable groups with each other, can confer substantial fitness benefits to individuals. Such benefits may arise via multiple mechanisms associated with both predation avoidance and foraging efficiency. In fishes, mixed-species shoaling reportedly occurs where displaced tropical species (known as “vagrants”) interact with resident temperate species, although little is known about the nature and frequency of such interactions. To investigate this phenomenon, we used displaced tropical Indo-pacific Sergeant Abudefduf vaigiensis settling in temperate south-eastern Australia as a model system. Underwater visual surveys revealed shoal composition and size differed significantly between open-water and reef habitats, with shoals in open habitats being larger and more speciose. Shoals containing A. vaigiensis were mainly mixed-species, and larger and more speciose in open habitats than nearer to reef. Since both foraging efficiency (via access to plankton) and predation threat likely increase with increasing distance from reef habitat, we suggest that mixed-species shoaling mitigates predation risk whilst allowing increased foraging opportunities for A. vaigiensis in open areas. These findings provide support for the importance of mixed-species shoaling to the persistence of tropical reef fishes in temperate regions.

Biological invasions are a major cause of biodiversity loss and, coupled with climate change, will likely have detrimental impacts for native species and the functioning of ecosystems. To mitigate such impacts, it is important to elucidate the behavioural mechanisms underpinning interactions between invasive and native species. Here we examined how competitive interactions between invasive and native species are modified under conditions of near-future elevated water temperature using freshwater crayfish as a model system. Contest experiments between the native Euastacus spinifer and invasive Cherax destructor revealed that the competitive advantage of E. spinifer at current maximum temperatures (22 °C) was reversed at elevated near-future temperatures (26 °C), after controlling for relative body size. In addition, the native crayfish spent twice as long motionless at 26 °C than C. destructor, consistent with physiological challenges underpinning this competitive reversal. Most alarmingly, E. spinifer experienced significant mortality after fighting C. destructor, particularly at 26 °C. Mortality usually ensued two days post-contest even when E. spinifer had won. Mortality was rare when fighting conspecifics. Together, these results suggest that while E. spinifer is more likely to win contests under current conditions, it could suffer considerable impacts if climate change and the spread of invasive C. destructor continue unabated.

2019, The Author(s). Under projected levels of ocean acidification, shifts in energetic demands and food availability could interact to effect the growth and development of marine organisms. Changes to individual growth rates could then flow on to influence emergent properties of social groups, particularly in species that form size-based hierarchies. To test the potential interactive effects of (1) food availability, (2) elevated CO2 during juvenile development, and (3) parental experience of elevated CO2 on the growth, condition and size-based hierarchy of juvenile fish, we reared orange clownfish (Amphiprion percula) for 50 days post-hatching in a fully orthogonal design. Development in elevated CO2 reduced standard length and weight of juveniles, by 9% and 11% respectively, compared to ambient. Development under low food availability reduced length and weight of juveniles by 7% and 15% respectively, compared to high food. Parental exposure to elevated CO2 restored the length of juveniles to that of controls, but it did not restore weight, resulting in juveniles from elevated CO2 parents exhibiting 33% lower body condition when reared in elevated CO2. The body size ratios (relative size of a fish from the rank above) within juvenile groups were not affected by any treatment, suggesting relative robustness of group-level structure despite alterations in individual size and condition. This study demonstrates that both food availability and elevated CO2 can influence the physical attributes of juvenile reef fish, but these changes may not disrupt the emergent group structure of this social species, at least amongst juveniles.

Population densities of invasive species fluctuate spatially and temporally, suggesting that the intensity of their aggressive interactions with native species is similarly variable. Although inter-specific aggression is often thought to increase with population density, it is often theorized that it should be exceeded by intra-specific aggression since conspecifics share a greater degree of resource overlap. Yet, the magnitude of intra-specific aggression is seldom considered when examining aggressive interactions, particularly those between invasive and native species. Here, we manipulated the density of the invasive eastern mosquitofish, Gambusia holbrooki, and observed its aggressive interactions with juveniles of the native Australian bass, Macquaria novemaculeata in a laboratory setting. For both species, the magnitudes of intra- and inter-specific aggression were recorded. Regardless of density, the native M. novemaculeata was more aggressive towards heterospecifics than G. holbrooki was. In addition to this, M. novemaculeata was more aggressive to G. holbrooki than towards conspecifics, at both low- and high-density conditions. In contrast, G. holbrooki was similarly aggressive towards M. novemaculeata and G. holbrooki at a high density, yet at low density, displayed significantly more aggression towards conspecifics than M. novemaculeata. These findings demonstrate the importance of considering intra-specific aggression when exploring behavioural interactions between native and invasive species.

The ability to identify individuals is important for the success of many behavioral and ecological studies. In fishes, there is a lot of variation in body size, shape, skin thickness, behavior, and ecology, which means that any given marking/tagging method may not work well for all species. For the Humbug Damselfish Dascyllus aruanus, a widely used model species of coral-reef fish, we found that standard tagging methodologies (e.g., phenotypic variation, beads and PVC tags, fluorescent elastomer injections) were ineffective as they could not be applied to the fish or easily detected by observers or from videos. In response, we developed a new method to temporarily tag D. aruanus using colored plastic films and topical surgical glue. These films were easily detectable both directly by human observers and indirectly by video/photo-cameras. We tested the efficacy of this new method by examining the survival time of the tags at various positions on the body. Our results showed that the optimal tag locations were dorsal anterior (with a median attachment time of 53 hours) and dorsal middle (with a median attachment time of 49 hours). Total length of fish was not a significant predictor of tag life. In sum, we demonstrate an effective new method for temporarily tagging a widely studied coral reef fish. This method could be applied to other fishes and aquatic organisms (e.g., amphibians) in both marine and freshwater ecosystems.

The formation of social groups has important impacts on fitness for many animal species, with differences in group compositions resulting in a range of fitness outcomes for individuals. Recent interest in mixed-species grouping, which extends from a large body of literature invested in understanding single-species grouping, highlights novel complexities of group formation which relate to phenotypic, behavioural and physiological differences that naturally exist between species. Among fishes, mixed-species shoaling is a common form of social grouping behaviour displayed across a range of marine and freshwater ecosystems. Research explaining mixed-species shoaling shows some overlap with explanations for single-species shoaling; however, it also demonstrates that distinct differences between species give rise to unique cost-benefit trade-offs which need to be incorporated into conceptual models of mixed-species shoaling behaviour. Unique predation related trade-offs may arise from inefficiency of the confusion effect, variation in vigilance between species and unequal species-preferences shown by predators, whilst unique foraging-related trade-offs may arise from diet partitioning, variations in foraging behaviour and differences in competitive abilities between species. We review the literature on fitness outcomes associated with mixed-species shoaling and present a new theoretical framework to explain the cost-benefit trade-offs for individuals within mixed-species shoals. The framework incorporates both trade-offs arising from differences between species and those arising from group size, the former having been largely ignored due to a focus on single-species shoaling. Our framework is designed to inform future research striving to explain mixed-species shoaling behaviour.

What drives the evolution of sociality in animals? Many robust studies in terrestrial organisms have pointed toward various kinship-based, ecological and life-history traits or phylogenetic constraint which have played a role in the evolution of sociality. These traits are not mutually exclusive and the exact combination of traits is likely taxon-specific. Phylogenetic comparative analyses have been instrumental in identifying social lineages and comparing various traits with non-social lineages to give broad evolutionary perspectives on the development of sociality. Few studies have attempted this approach in marine vertebrate systems. Social marine fishes are particularly interesting because many have a pelagic larval phase and non-conventional life-history strategies (e.g. bi-directional sex-change) not often observed in terrestrial animals. Such strategies provide novel insights into terrestrially-derived theories of social evolution. Here, we assess the strength of the phylogenetic signal of sociality in the Gobiodon genus with Pagel's lambda and Blomberg's K parameters. We found some evidence of a phylogenetic signal of sociality, but factors other than phylogenetic constraint also have a strong influence on the extant social state of each species. We then use phylogenetic generalized least squares analyses to examine several ecological and life-history traits that may have influenced the evolution of sociality in the genus. We found an interaction of habitat size and fish length was the strongest predictor of sociality. Sociality in larger species was more dependent on coral size than in smaller species, but smaller species were more social overall, regardless of coral size. Finally, we comment on findings regarding the validity of the species G. spilophthalmus which arose during the course of our research. These findings in a group of marine fishes add a unique perspective on the evolution of sociality to the excellent terrestrial work conducted in this fiel

Aggressive contests amongst conspecifics are important to understand from an ecological and evolutionary perspective as contest dynamics can directly influence individual fitness. For some species, individual attributes such as relative body size closely predict the outcome, intensity and duration of contests, whereas for others, prior social experiences play a key role. However, disentangling the relative effects of individual attributes and social experiences is challenging from an experimental perspective, and because of this, few studies have investigated relative effects of multiple factors. Rockpool fishes have been well studied in terms of factors governing abundance, distribution and community structure, but much less so in terms of contest behaviour. This is surprising because contest dynamics are likely to directly affect the distribution of fishes along the rocky shore, and hence indirectly govern population and community composition. Here, we explored multiple factors potentially influencing contest behaviour in a numerically dominant, resident intertidal fish species, Bathygobius cocosensis (Gobiidae). Using a series of staged pairwise contest trials, we investigated the effect of size, sex and social experiences (namely prior residency and winner-loser experiences) on contest dynamics. We found no evidence that prior residency influenced contest outcome, suggesting social experiences play a minor role in contest dynamics. Previous winner/loser experience also did not influence contest outcome, although this is likely a product of low sample size. In addition, the likelihood of winning was unrelated to contestant sex, and the combination of sexes in paired contests did not influence contest intensity or duration. Instead, body size was related to contest outcome, intensity and duration in the majority of experimental trials. These results suggest that body size rather than sex or social experiences is the key determinant of contest dynamics in this spec

The present study investigates how the humbug damselfish Dascyllus aruanus, subject of a large number of ecological, evolutionary and behavioural studies, responds to the presence of human observers (effect of scuba diver presence-absence) and how the method of data collection (directly by a scuba diver v. indirectly via video camera) may affect the quality of behavioural data. Scuba diver presence had only subtle effects on fish behaviour. The efficiency of the method of scoring fish behaviour depended on the behaviour under consideration: those behaviours that occur in close proximity to the corals were scored more effectively directly by a scuba diver while those that are performed in a more rapid or repetitive fashion were scored more effectively indirectly via video camera. These results provide a foundation for future behavioural research on D. aruanus and other fishes where scuba divers or video cameras are the prevalent means of data collection.

Rocky intertidal shores are diverse ecosystems that have been extensively studied, yet there is a surprising lack of knowledge on the methods best suited to quantifying species abundance, diversity and assemblages. We compared visual census, unbaited remote underwater video (mini-RUV), baited remote underwater video (mini-BRUV) and draining/collection methods, with the goal of quantifying the effectiveness of non-invasive methods over more typical invasive methods for quantifying rockpool fish assemblages. In addition, we assessed the optimal set time of video deployment for mini-RUV and mini-BRUV. Fieldwork was undertaken from May-June 2016 using 20 rockpools at seven locations on the SE coast of NSW, Australia. Following standardisation of methods, visual census detected the lowest abundance of fishes whereas hand collection detected the highest out of any method. Increasing sampling time of mini-RUV and mini-BRUV from 10 to 25 min improved their effectiveness, with mini-BRUV detecting comparable abundances to invasive hand collection. The methods did not differ with respect to species richness however. Further, 25 min of mini-BRUV yielded greater detection of common species than hand collection. All together, these results suggest that mini-BRUV is the best method if the goal is to provide accurate estimates of overall abundance, richness and assemblages with minimal disturbance to rockpools, with a preferred set time of 25 min (and at least 15 min). The ability to conduct rapid and non-invasive species surveys of natural rockpool fish communities is an important conservation and management tool given that these habitats can be home to a variety of commercially important and rare species.

Social organization is a key factor influencing a species' foraging and reproduction, which may ultimately affect their survival and ability to recover from catastrophic disturbance. Severe weather events such as cyclones can have devastating impacts to the physical structure of coral reefs and on the abundance and distribution of its faunal communities. Despite the importance of social organization to a species' survival, relatively little is known about how major disturbances such as tropical cyclones may affect social structures or how different social strategies affect a species' ability to cope with disturbance. We sampled group sizes and coral sizes of group-forming and pair-forming species of the Gobiid genus Gobiodon at Lizard Island, Great Barrier Reef, Australia, before and after two successive category 4 tropical cyclones. Group sizes of group-forming species decreased after each cyclone, but showed signs of recovery four months after the first cyclone. A similar increase in group sizes was not evident in group-forming species after the second cyclone. There was no change in mean pair-forming group size after either cyclone. Coral sizes inhabited by both group- and pair-forming species decreased throughout the study, meaning that group-forming species were forced to occupy smaller corals on average than before cyclone activity. This may reduce their capacity to maintain larger group sizes through multiple processes. We discuss these patterns in light of two non-exclusive hypotheses regarding the drivers of sociality in Gobiodon, suggesting that benefits of philopatry with regards to habitat quality may underpin the formation of social groups in this genus.

In social groups, high reproductive skew is predicted to arise when the reproductive output of a group is limited, and dominant individuals can suppress subordinate reproductive efforts. Reproductive suppression is often assumed to occur via overt aggression or the threat of eviction. It is unclear, however, whether the threat of eviction alone is sufficient to induce reproductive restraint by subordinates. Here, we test two assumptions of the restraint model of reproductive skew by investigating whether resource limitation generates reproductive competition and whether the threat of eviction leads to reproductive restraint in the clown anemonefish Amphiprion percula. First, we use a feeding experiment to test whether reproduction is resource limited, which would create an incentive for the dominant pair to suppress subordinate reproduction. We show that the number of eggs laid increased in the population over the study period, but the per cent increase in fed groups was more than twice that in unfed groups (205% and 78%, respectively). Second, we use an eviction experiment to test whether the dominant pair evicts mature subordinates, which would create an incentive for the subordinates to forgo reproduction. We show that mature subordinates are seven times more likely to be evicted than immature subordinates of the same size. In summary, we provide experimental support for the assumptions of the restraint model by showing that resource limitation creates reproductive competition and a credible threat of eviction helps explain why subordinates forego reproduction. Transactional models of reproductive skew may apply well to this and other simple systems.

Invasive species are known to competitively displace native species via direct behavioural interactions, or contests, which can result in limited access to valuable resources such as food or shelter for the native. Here, we aimed to determine the dynamics and outcome of agonistic contests between the critically endangered Fitzroy Falls Spiny Crayfish (Euastacus dharawalus) and the invasive yabby (Cherax destructor) in a controlled laboratory setting. Although C. destructor has been listed as a key threatening process for E. dharawalus, the magnitude and nature of its impact have not been formerly assessed. We recorded the frequency and intensity of aggressive interactions between the species, as well as the winner of the contest and subsequent dominance during staged encounters. In addition to this, we recorded the time individuals spent consuming food and actively moving on the substratum. Contrary to our predictions, the frequency of aggressive behaviours performed by the two species did not significantly differ, but E. dharawalus engaged in more intensely aggressive behaviours and was less submissive than C. destructor. As a result, E. dharawalus won significantly more contests than its invasive rival and subsequently achieved a higher dominance score overall. Despite this result, C. destructor spent significantly more time consuming food than did E. dharawalus. These results suggest that dominance in agonistic contests did not translate into dominance over food resources in our study. Ultimately, reduced access to food, combined with the costs of agonistic contests including injury and the relatively low reproductive rates of E. dharawalus indicate that C. destructor may have a negative effect on the survival and persistence of the critically endangered E. dharawalus.

Competition for limiting resources is a fundamental and well-established driver of niche partitioning, which in turn promotes species coexistence and biodiversity. Although contests are a well-known behavioural mechanism by which organisms compete over limiting resources, there has been surprisingly little application of contest theory to understanding interspecific interactions, niche partitioning, species coexistence and biodiversity. We investigated the link between contest dynamics and community structure using two intertidal rockpool fishes, Bathygobius cocosensis (Gobiidae) and Lepidoblennius haplodactylus (Tripterygiidae), as model species. We assessed the abundance and distribution of the two species on intertidal rocky shores of South-East Australia, and whether distribution patterns were related to abiotic variables. We then conducted interspecific contest experiments between pairs of B. cocosensis and L. haplodactylus of varying size and sex under high and low dissolved oxygen levels to assess the competitive dominance of the species. To relate these results to community structure, we quantified temporal patterns of rockpool fidelity of each species in situ. Field surveys revealed that B. cocosensis was most prevalent in rockpools with higher dissolved oxygen, and L. haplodactylus was most prevalent in rockpools that contained no B. cocosensis. Contest experiments revealed that B. cocosensis displayed higher resource holding power than L. haplodactylus regardless of size asymmetry, sex or dissolved oxygen concentration, although contest intensity was influenced by relative body size of competing individuals. Furthermore, the microhabitat preference of L. haplodactylus was modulated in the presence of B. cocosensis. In the field, B. cocosensis displayed higher rockpool fidelity than L. haplodactylus. In light of these findings, we propose that B. cocosensis is the superior competitor but persistence of both species is likely facilitated by low-cost contest

Interspecific aggression is a critical determinant of the success and competitive superiority of many invasive over native species. While single abiotic stressors can alter aggression levels, the manner in which multiple stressors may alter the strength and outcome of interspecific interactions and hence the invasion potential of a species is still poorly understood, even though multiple stressors are prevalent in many ecosystems. Furthermore, multiple stressors may interact to produce synergistic or antagonistic effects on individual level behaviors, thereby modulating invasive-native species interactions in unexpected ways. Here we examined the effect of two key abiotic stressors in freshwater ecosystems-temperature and salinity-on interspecific aggression between the invasive eastern mosquito fish (Gambusia holbrooki) and juveniles of the native Australian bass (Macquaria novemaculeata). Under controlled laboratory conditions, individuals were exposed to low or high salinity levels (15 and 35‰), and low or high temperatures (21 and 28 °C), and the frequency of interspecific aggressive behaviors was scored. The effect of temperature and salinity on interspecific aggression was antagonistic for both M. novemaculeata and G. holbrooki. While elevated temperature promoted aggression, elevated salinity partially or entirely negated this effect. Moreover, regardless of temperature, M. novemaculeata was more aggressive than G. holbrooki under elevated salinity. In addition to this, the native displayed more aggression to smaller than larger heterospecifics when exposed to elevated salinity alone, while G. holbrooki showed no such preference. These results highlight the importance of considering the interplay between multiple abiotic stressors and behavioral interactions between invasive and native species, combined with the modulating effect of species-specific and size based responses to those stressors.

Determining the extent of repeatable differences in the behavior of animals and the factors that influence behavioral expression is important for understanding individual fitness and population processes, thereby aiding in species conservation. However, little is known about the causes of variation in the repeatability of behavioral differences among species because rarely have comparative studies been undertaken to examine the repeatability of behavioral differences among individuals within their natural ecological settings. Using two species of endemic subtropical anemonefishes, Amphiprion mccullochi and A. latezonatus at Lord Howe and North Solitary Islands, Australia, we conducted an in situ comparative analysis of personality traits, examining the repeatability of boldness, sociability and aggression as well as the potential role of environmental and social factors on behavioral expression. For A. mccullochi, only boldness and aggression were highly repeatable and these behaviors formed a behavioral syndrome. For A. latezonatus, none of the three behaviors were repeatable due to low-inter-individual variation in behavior. We suggest that the harsher and more variable environmental and social conditions experienced by A. latezonatus have resulted in reduced repeatability in behavior, in contrast to A. mccullochi which typically inhabits a more stable lagoonal reef environment. Additionally, group size and size rank, rather than nearest-neighbor distance and anemone size, influenced the expression of these behaviors in both species, suggesting that behavioral variation was more sensitive to social than environmental factors. Overall, differences in repeatability between these closely related species likely reflect adaptations to contrasting environmental and social conditions, although alternative explanations must be considered. The differences in behavioral consistency between these two endemic anemonefishes could lead to disparity in their resilience to enviro

Why do we observe so many examples in nature in which individuals routinely delay or completely forgo their own reproductive opportunities in order to join and remain within a group? Cooperative breeding theory provides a rich framework with which to study the factors that may influence the costs and benefits of remaining philopatric as a non-breeder. This is often viewed as an initial step in the development of costly helping behavior provided by non-breeding subordinates. Despite many excellent empirical studies testing key concepts of the theory, there is still debate regarding the relative importance of various evolutionary forces, suggesting that there may not be a general explanation but rather a dynamic and taxonomically varied combination of factors influencing the evolution and maintenance of sociality. Here, we explore two potential improvements in t he study of sociality that could aid in the progress of this field. The first addresses the fact that empirical studies of social evolution are typically conducted using either comparative, observational or manipulative methodologies. Instead, we suggest a holistic approach, whereby observational and experimental studies are designed with the explicit view of advancing comparative analyses of sociality for the taxon, and in tandem, where comparative work informs targeted research effort on specific (usually understudied) species within the lineage. A second improvement relates to the broadening of tests of cooperative breeding theory to include taxa where subordinates do not necessarily provide active cooperation within the group. The original bias toward "helpful subordinates" arose from a focus on terrestrial taxa. However, recent consideration of other taxa, especially marine taxa, is slowly revealing that the theory can and should encompass a continuum of cooperative social systems, including those where subordinates do not actively help. This review summarizes the major hypotheses of cooperative breeding

Competition for limiting resources is a fundamental and well-established driver of niche partitioning, which in turn promotes species coexistence and biodiversity. Although contests are a well-known behavioural mechanism by which organisms compete over limiting resources, there has been surprisingly little application of contest theory to understanding interspecific interactions, niche partitioning, species coexistence and biodiversity. We investigated the link between contest dynamics and community structure using two intertidal rockpool fishes, Bathygobius cocosensis (Gobiidae) and Lepidoblennius haplodactylus (Tripterygiidae), as model species. We assessed the abundance and distribution of the two species on intertidal rocky shores of South-East Australia, and whether distribution patterns were related to abiotic variables. We then conducted interspecific contest experiments between pairs of B. cocosensis and L. haplodactylus of varying size and sex under high and low dissolved oxygen levels to assess the competitive dominance of the species. To relate these results to community structure, we quantified temporal patterns of rockpool fidelity of each species in situ. Field surveys revealed that B. cocosensis was most prevalent in rockpools with higher dissolved oxygen, and L. haplodactylus was most prevalent in rockpools that contained no B. cocosensis. Contest experiments revealed that B. cocosensis displayed higher resource holding power than L. haplodactylus regardless of size asymmetry, sex or dissolved oxygen concentration, although contest intensity was influenced by relative body size of competing individuals. Furthermore, the microhabitat preference of L. haplodactylus was modulated in the presence of B. cocosensis. In the field, B. cocosensis displayed higher rockpool fidelity than L. haplodactylus. In light of these findings, we propose that B. cocosensis is the superior competitor but persistence of both species is likely facilitated by low-cost contest

Many animals form dynamic societies in which the fission and fusion of subgroups occurs on a regular basis. Such societies are intriguing, because it is unclear whether stable dominance relationships that form within subgroups are retained upon fusion with other subgroups, and what the implications of rank instability may be at higher levels of ecological organization. Additionally, little is known about how environmental change affects the fission¿fusion process, even though environments often fluctuate and are predicted to become increasingly variable, in part due to climate change. Here we investigated the social organization, levels of conflict and stability of dominance relationships during group fusion in a globally invasive fish, the eastern mosquitofish, Gambusia holbrooki. To assess the effect of environmental variation, we conducted group fusion experiments at high and low water depths, to simulate normal and drought conditions, and recorded dominance interactions during prefusion, early fusion and late fusion stages. Individuals formed size-based hierarchies within prefusion groups, although the relationship between size and dominance varied with group fusion. Levels of conflict were affected by group fusion and water depth, with higher levels of conflict after fusion and at low depth. Rank relationships in early and late fusion groups were stable and unaffected by water depth. Finally, there was no evidence of coat-tail effects, as familiarity with the alpha dominant in postfusion groups did not lead to a significant increase in subordinate dominance rank. All in all, these results provide key experimental evidence that environmental change in terms of water level is unlikely to impact social organization or rank stability in response to group fusion in this species. More generally, they indicate that sociality in fission¿fusion societies may be relatively robust to changes in both social and environmental contexts.

Even closely related and ecologically similar cichlid species of Lake Tanganyika exhibit an impressive diversity of social systems, and therefore these fishes offer an excellent opportunity to examine the evolution of social behaviour. Sophisticated social relationships are thought to have evolved via a building block design where more fundamental social behaviours and cognitive processes have been combined, incrementally modified, and elaborated over time. Here, we studied two of these putative social building blocks in two closely related species of cichlids: Neolamprologus pulcher, a group-living species, and Telmatochromis temporalis, a non-grouping species. Otherwise well matched in ecology, this pair of species provide an excellent comparison point to understand how behavioural processes may have been modified in relation to the evolution of sociality. Using social assays in both the laboratory and in the field, we explored each species' motivation to interact with conspecifics, and each species' conflict resolution tactics. We found that individuals of the group living species, N. pulcher, displayed higher social motivation and were more likely to produce submission displays than were individuals of the non-grouping species, T. temporalis. We argue that the motivation to interact with conspecifics is a necessary prerequisite for the emergence of group living, and that the use of submission reduces the costs of conflict and facilitates the maintenance of close social proximity. These results suggest that social motivation and conflict resolution tactics are associated with social complexity, and that these behavioural traits may be functionally significant in the evolution and maintenance of sociality.

Estuaries are critical aquatic environments that are used by many fish during their life cycle. However, estuaries often suffer from poor water quality as a result of anthropogenic activities. Fish diversity studies in estuaries are common, although few have examined whether correlations exist between water quality, metal contamination and fish assemblages. In the present study we investigated the effect of abiotic conditions, heavy metals and estuary characteristics on the abundance, diversity and composition of fish in four intermittently open estuaries along the Illawarra coast of south-eastern Australia. The heterogeneity of environmental conditions was reflected in the fish assemblages in each estuary. Environmental variables predicted fish species composition, and estuaries in particularly poor condition contained few species (estuarine residents) in high abundance, indicating their ability to acclimatise and survive in conditions that are hostile to other species. Overall, these findings demonstrate that estuarine fish assemblages may be useful indicators of estuary condition and reveal the importance of managing anthropogenic activities in the surrounding catchment to improve water quality so that biodiversity of fish can be restored in these estuarine environments.

Interspecific aggression is a critical determinant of the success and competitive superiority of many invasive over native species. While single abiotic stressors can alter aggression levels, the manner in which multiple stressors may alter the strength and outcome of interspecific interactions and hence the invasion potential of a species is still poorly understood, even though multiple stressors are prevalent in many ecosystems. Furthermore, multiple stressors may interact to produce synergistic or antagonistic effects on individual level behaviors, thereby modulating invasive-native species interactions in unexpected ways. Here we examined the effect of two key abiotic stressors in freshwater ecosystems-temperature and salinity-on interspecific aggression between the invasive eastern mosquito fish (Gambusia holbrooki) and juveniles of the native Australian bass (Macquaria novemaculeata). Under controlled laboratory conditions, individuals were exposed to low or high salinity levels (15 and 35‰), and low or high temperatures (21 and 28 °C), and the frequency of interspecific aggressive behaviors was scored. The effect of temperature and salinity on interspecific aggression was antagonistic for both M. novemaculeata and G. holbrooki. While elevated temperature promoted aggression, elevated salinity partially or entirely negated this effect. Moreover, regardless of temperature, M. novemaculeata was more aggressive than G. holbrooki under elevated salinity. In addition to this, the native displayed more aggression to smaller than larger heterospecifics when exposed to elevated salinity alone, while G. holbrooki showed no such preference. These results highlight the importance of considering the interplay between multiple abiotic stressors and behavioral interactions between invasive and native species, combined with the modulating effect of species-specific and size based responses to those stressors.

Why do we observe so many examples in nature in which individuals routinely delay or completely forgo their own reproductive opportunities in order to join and remain within a group? Cooperative breeding theory provides a rich framework with which to study the factors that may influence the costs and benefits of remaining philopatric as a non-breeder. This is often viewed as an initial step in the development of costly helping behavior provided by non-breeding subordinates. Despite many excellent empirical studies testing key concepts of the theory, there is still debate regarding the relative importance of various evolutionary forces, suggesting that there may not be a general explanation but rather a dynamic and taxonomically varied combination of factors influencing the evolution and maintenance of sociality. Here, we explore two potential improvements in t he study of sociality that could aid in the progress of this field. The first addresses the fact that empirical studies of social evolution are typically conducted using either comparative, observational or manipulative methodologies. Instead, we suggest a holistic approach, whereby observational and experimental studies are designed with the explicit view of advancing comparative analyses of sociality for the taxon, and in tandem, where comparative work informs targeted research effort on specific (usually understudied) species within the lineage. A second improvement relates to the broadening of tests of cooperative breeding theory to include taxa where subordinates do not necessarily provide active cooperation within the group. The original bias toward "helpful subordinates" arose from a focus on terrestrial taxa. However, recent consideration of other taxa, especially marine taxa, is slowly revealing that the theory can and should encompass a continuum of cooperative social systems, including those where subordinates do not actively help. This review summarizes the major hypotheses of cooperative breeding

Even closely related and ecologically similar cichlid species of Lake Tanganyika exhibit an impressive diversity of social systems, and therefore these fishes offer an excellent opportunity to examine the evolution of social behaviour. Sophisticated social relationships are thought to have evolved via a building block design where more fundamental social behaviours and cognitive processes have been combined, incrementally modified, and elaborated over time. Here, we studied two of these putative social building blocks in two closely related species of cichlids: Neolamprologus pulcher, a group-living species, and Telmatochromis temporalis, a non-grouping species. Otherwise well matched in ecology, this pair of species provide an excellent comparison point to understand how behavioural processes may have been modified in relation to the evolution of sociality. Using social assays in both the laboratory and in the field, we explored each species' motivation to interact with conspecifics, and each species' conflict resolution tactics. We found that individuals of the group living species, N. pulcher, displayed higher social motivation and were more likely to produce submission displays than were individuals of the non-grouping species, T. temporalis. We argue that the motivation to interact with conspecifics is a necessary prerequisite for the emergence of group living, and that the use of submission reduces the costs of conflict and facilitates the maintenance of close social proximity. These results suggest that social motivation and conflict resolution tactics are associated with social complexity, and that these behavioural traits may be functionally significant in the evolution and maintenance of sociality.

Estuaries are critical aquatic environments that are used by many fish during their life cycle. However, estuaries often suffer from poor water quality as a result of anthropogenic activities. Fish diversity studies in estuaries are common, although few have examined whether correlations exist between water quality, metal contamination and fish assemblages. In the present study we investigated the effect of abiotic conditions, heavy metals and estuary characteristics on the abundance, diversity and composition of fish in four intermittently open estuaries along the Illawarra coast of south-eastern Australia. The heterogeneity of environmental conditions was reflected in the fish assemblages in each estuary. Environmental variables predicted fish species composition, and estuaries in particularly poor condition contained few species (estuarine residents) in high abundance, indicating their ability to acclimatise and survive in conditions that are hostile to other species. Overall, these findings demonstrate that estuarine fish assemblages may be useful indicators of estuary condition and reveal the importance of managing anthropogenic activities in the surrounding catchment to improve water quality so that biodiversity of fish can be restored in these estuarine environments.

Many animals form dynamic societies in which the fission and fusion of subgroups occurs on a regular basis. Such societies are intriguing, because it is unclear whether stable dominance relationships that form within subgroups are retained upon fusion with other subgroups, and what the implications of rank instability may be at higher levels of ecological organization. Additionally, little is known about how environmental change affects the fission¿fusion process, even though environments often fluctuate and are predicted to become increasingly variable, in part due to climate change. Here we investigated the social organization, levels of conflict and stability of dominance relationships during group fusion in a globally invasive fish, the eastern mosquitofish, Gambusia holbrooki. To assess the effect of environmental variation, we conducted group fusion experiments at high and low water depths, to simulate normal and drought conditions, and recorded dominance interactions during prefusion, early fusion and late fusion stages. Individuals formed size-based hierarchies within prefusion groups, although the relationship between size and dominance varied with group fusion. Levels of conflict were affected by group fusion and water depth, with higher levels of conflict after fusion and at low depth. Rank relationships in early and late fusion groups were stable and unaffected by water depth. Finally, there was no evidence of coat-tail effects, as familiarity with the alpha dominant in postfusion groups did not lead to a significant increase in subordinate dominance rank. All in all, these results provide key experimental evidence that environmental change in terms of water level is unlikely to impact social organization or rank stability in response to group fusion in this species. More generally, they indicate that sociality in fission¿fusion societies may be relatively robust to changes in both social and environmental contexts.

All prey face a fundamental trade-off between avoiding predation and pursuing activities, such as foraging and mating, that enhance fitness. Therefore, the effects of predation can be both consumptive and nonconsumptive and prey need to assess and respond appropriately to predation risk which in turn varies with environmental and social contexts. We tested the effects of predator density and diel cycle on the consumption, interspecific interactions and behavioural responses of a prey species, the native Australian glass shrimp, Paratya australiensis, exposed to a predator, the invasive eastern mosquito fish, Gambusia holbrooki. In the laboratory, P. australiensis were exposed to low or high densities of conspecifics or predators and observed during the day and at night. While P. australiensis experienced more interspecific approaches and nips when exposed to a high density of G. holbrooki and during the day, neither predator density nor diel cycle influenced the actual number of P. australiensis consumed. Similarly, while P. australiensis engaged in significantly more shelter use and swam less, there was no difference in these behaviours in relation to predator density and diel cycle. Foraging by P. australiensis was not related to species composition, but instead depended on the overall number of animals present with more P. australiensis foraging when exposed to a high density of conspecifics and G. holbrooki. These results indicate that the mechanisms by which G. holbrooki exerts negative effects on P. australiensis can be multiple and wide ranging, from direct predation to a reduction in activity and competition for resources. However, as neither predator density nor diurnal variation altered predation rate, P. australiensis did behave in an adaptive manner, by only adjusting its behavioural responses in proportion to the direct risk of predation.

Conflicts of interest are part and parcel of living in a social group, yet actual conflict can be rare in established groups. Within limits, individuals can maximize the benefits of group living by resolving conflict with other group members. Thus, understanding what causes conflict, what determines its outcome, and how it is resolved holds the key to understanding the evolution and maintenance of sociality. Here, we investigate these questions using the clown anemonefish Amphiprion percula. Clownfish live in groups composed of a breeding pair and zero to four non-breeders that queue for breeding positions. Within groups, there is potential conflict over rank yet actual conflict is very rare. We staged contests in aquaria between pairs of non-breeding individuals over access to a key resource (an anemone), analogous to contests that would occur at the onset of group formation in the wild. The initial size ratio between individuals tended to predict the intensity, and predicted the outcome and resolution of conflict: conflict intensity was greater when individuals were more similar in size; the probability of the smaller individual winning was greater when individuals were more similar in size; and the loser of the contest grew less than the winner when individuals were more similar in size. These results provide a critical test of foundational assumptions upon which our understanding of clownfish and other fish societies has been built. More generally, the results show that one of the simplest and most effective ways for animals to resolve conflict is to modify the phenotype that triggers conflict.

Predicting the impacts of climate change to biological systems requires an understanding of the ability for species to acclimate to the projected environmental change through phenotypic plasticity. Determining the effects of higher temperatures on individual performance is made more complex by the potential for environmental conditions experienced in previous and current generations to independently affect phenotypic responses to high temperatures. We used a model coral reef fish (Acanthochromis polyacanthus) to investigate the influence of thermal conditions experienced by two generations on reproductive output and the quality of offspring produced by adults. We found that more gradual warming over two generations, +1.5°C in the first generation and then +3.0°C in the second generation, resulted in greater plasticity of reproductive attributes, compared to fish that experienced the same increase in one generation. Reproduction ceased at the projected future summer temperature (31.5°C) when fish experienced +3.0°C for two generations. Additionally, we found that transgenerational plasticity to +1.5°C induced full restoration of thermally affected reproductive and offspring attributes, which was not possible with developmental plasticity alone. Our results suggest that transgenerational effects differ depending on the absolute thermal change and in which life stage the thermal change is experienced.

Predicting the impacts of climate change to biological systems requires an understanding of the ability for species to acclimate to the projected environmental change through phenotypic plasticity. Determining the effects of higher temperatures on individual performance is made more complex by the potential for environmental conditions experienced in previous and current generations to independently affect phenotypic responses to high temperatures. We used a model coral reef fish (Acanthochromis polyacanthus) to investigate the influence of thermal conditions experienced by two generations on reproductive output and the quality of offspring produced by adults. We found that more gradual warming over two generations, +1.5°C in the first generation and then +3.0°C in the second generation, resulted in greater plasticity of reproductive attributes, compared to fish that experienced the same increase in one generation. Reproduction ceased at the projected future summer temperature (31.5°C) when fish experienced +3.0°C for two generations. Additionally, we found that transgenerational plasticity to +1.5°C induced full restoration of thermally affected reproductive and offspring attributes, which was not possible with developmental plasticity alone. Our results suggest that transgenerational effects differ depending on the absolute thermal change and in which life stage the thermal change is experienced.

All prey face a fundamental trade-off between avoiding predation and pursuing activities, such as foraging and mating, that enhance fitness. Therefore, the effects of predation can be both consumptive and nonconsumptive and prey need to assess and respond appropriately to predation risk which in turn varies with environmental and social contexts. We tested the effects of predator density and diel cycle on the consumption, interspecific interactions and behavioural responses of a prey species, the native Australian glass shrimp, Paratya australiensis, exposed to a predator, the invasive eastern mosquito fish, Gambusia holbrooki. In the laboratory, P. australiensis were exposed to low or high densities of conspecifics or predators and observed during the day and at night. While P. australiensis experienced more interspecific approaches and nips when exposed to a high density of G. holbrooki and during the day, neither predator density nor diel cycle influenced the actual number of P. australiensis consumed. Similarly, while P. australiensis engaged in significantly more shelter use and swam less, there was no difference in these behaviours in relation to predator density and diel cycle. Foraging by P. australiensis was not related to species composition, but instead depended on the overall number of animals present with more P. australiensis foraging when exposed to a high density of conspecifics and G. holbrooki. These results indicate that the mechanisms by which G. holbrooki exerts negative effects on P. australiensis can be multiple and wide ranging, from direct predation to a reduction in activity and competition for resources. However, as neither predator density nor diurnal variation altered predation rate, P. australiensis did behave in an adaptive manner, by only adjusting its behavioural responses in proportion to the direct risk of predation.

Conflicts of interest are part and parcel of living in a social group, yet actual conflict can be rare in established groups. Within limits, individuals can maximize the benefits of group living by resolving conflict with other group members. Thus, understanding what causes conflict, what determines its outcome, and how it is resolved holds the key to understanding the evolution and maintenance of sociality. Here, we investigate these questions using the clown anemonefish Amphiprion percula. Clownfish live in groups composed of a breeding pair and zero to four non-breeders that queue for breeding positions. Within groups, there is potential conflict over rank yet actual conflict is very rare. We staged contests in aquaria between pairs of non-breeding individuals over access to a key resource (an anemone), analogous to contests that would occur at the onset of group formation in the wild. The initial size ratio between individuals tended to predict the intensity, and predicted the outcome and resolution of conflict: conflict intensity was greater when individuals were more similar in size; the probability of the smaller individual winning was greater when individuals were more similar in size; and the loser of the contest grew less than the winner when individuals were more similar in size. These results provide a critical test of foundational assumptions upon which our understanding of clownfish and other fish societies has been built. More generally, the results show that one of the simplest and most effective ways for animals to resolve conflict is to modify the phenotype that triggers conflict.

Anthropogenic climate change is expected to result in dramatic shifts in the abiotic conditions within estuaries, including an elevation of temperature and salinity levels. Even so, few studies have addressed the impacts of multiple abiotic stressors on the behaviour and life history of key estuarine species, such as those of biological and commercial importance. Here we used a cross-factored experimental design to tease apart the effects of temperature and salinity on intraspecific aggression and growth rates of a native estuarine fish species, the Australian bass (Macquaria novemaculeata). Juvenile bass were exposed to one of four treatment conditions: 1) baseline temperature and salinity, 2) elevated temperature, 3) elevated salinity, and 4) elevated temperature and salinity. Elevated salinity increased rates of aggression, and elevated temperature decreased rates of growth, although the effects of both factors were mediated by the body size of individual bass. These results therefore highlight complex and variable effects of abiotic stressors and body size, emphasising the importance of considering individual-level attributes when evaluating the impacts of climate change on estuarine fishes.

In social groups, hierarchies are the fundamental organizational unit and integral to the structure of social groups. For many social fishes, rank is determined by body size and conflict over rank is resolved via aggressive threats from dominants and growth restraint by subordinates. However, this balance may be offset by an alteration of abiotic factors, such as elevated temperature expected from climate change, which could thereby disrupt the usual mechanisms of conflict resolution. Here, we determined the effect of elevated temperature on hierarchy structure, stability, and conflict resolution in the Eastern mosquitofish, Gambusia holbrooki. Body size was significantly related to dominance rank, and aggression was more commonly directed toward subordinates and was heightened between individuals of adjacent rank, demonstrating that conflict over rank occurs in size-based hierarchies. Temperature did not affect overall levels or directionality/adjacency of aggression but substantially altered subordinate growth patterns. In only the high-temperature groups, growth rates of subordinates decreased as the size ratio between themselves and their immediate dominant approached 1.0, whereas growth rates of dominants were unaffected. This unique finding suggests that only under high temperatures, subordinates may adopt growth regulation to resolve conflict, when the costs of conflict with dominants are greater. This provides the first causal link between abiotic stressors and changes to hierarchical structure and functioning, providing a springboard for further research into implications of temperature-dependent subordinate growth alteration at higher levels of ecological organization.

Anthropogenic climate change is expected to result in dramatic shifts in the abiotic conditions within estuaries, including an elevation of temperature and salinity levels. Even so, few studies have addressed the impacts of multiple abiotic stressors on the behaviour and life history of key estuarine species, such as those of biological and commercial importance. Here we used a cross-factored experimental design to tease apart the effects of temperature and salinity on intraspecific aggression and growth rates of a native estuarine fish species, the Australian bass (Macquaria novemaculeata). Juvenile bass were exposed to one of four treatment conditions: 1) baseline temperature and salinity, 2) elevated temperature, 3) elevated salinity, and 4) elevated temperature and salinity. Elevated salinity increased rates of aggression, and elevated temperature decreased rates of growth, although the effects of both factors were mediated by the body size of individual bass. These results therefore highlight complex and variable effects of abiotic stressors and body size, emphasising the importance of considering individual-level attributes when evaluating the impacts of climate change on estuarine fishes.

In social groups, hierarchies are the fundamental organizational unit and integral to the structure of social groups. For many social fishes, rank is determined by body size and conflict over rank is resolved via aggressive threats from dominants and growth restraint by subordinates. However, this balance may be offset by an alteration of abiotic factors, such as elevated temperature expected from climate change, which could thereby disrupt the usual mechanisms of conflict resolution. Here, we determined the effect of elevated temperature on hierarchy structure, stability, and conflict resolution in the Eastern mosquitofish, Gambusia holbrooki. Body size was significantly related to dominance rank, and aggression was more commonly directed toward subordinates and was heightened between individuals of adjacent rank, demonstrating that conflict over rank occurs in size-based hierarchies. Temperature did not affect overall levels or directionality/adjacency of aggression but substantially altered subordinate growth patterns. In only the high-temperature groups, growth rates of subordinates decreased as the size ratio between themselves and their immediate dominant approached 1.0, whereas growth rates of dominants were unaffected. This unique finding suggests that only under high temperatures, subordinates may adopt growth regulation to resolve conflict, when the costs of conflict with dominants are greater. This provides the first causal link between abiotic stressors and changes to hierarchical structure and functioning, providing a springboard for further research into implications of temperature-dependent subordinate growth alteration at higher levels of ecological organization.

In complex societies, some animals forgo breeding and help others raise young. Although this behavior at first seems maladaptive, evolutionary biologists have developed theory that explains when and how it can evolve. For example, if it helps kin raise more young or if dispersing to breed elsewhere is risky, an individual may cooperate with nearby breeders. After two decades of work on the social behavior of coral reef fish, Peter Buston of Boston University and Marian Wong of University of Wollongong, Australia, have shown that this behavior can unfold under a third scenario: If conflict with dominant breeders is especially risky, then subordinates will politely wait their turn to breed, rather than risk a fight. Buston and Song explain their experimental and observational work in this feature, and then outline how their work influences current theory.

of a presentation that was presented at the Society for Integrative and Comparative Biology 2014 Annual Meeting, 3-7 January, Austin, TX, USA.

In complex societies, some animals forgo breeding and help others raise young. Although this behavior at first seems maladaptive, evolutionary biologists have developed theory that explains when and how it can evolve. For example, if it helps kin raise more young or if dispersing to breed elsewhere is risky, an individual may cooperate with nearby breeders. After two decades of work on the social behavior of coral reef fish, Peter Buston of Boston University and Marian Wong of University of Wollongong, Australia, have shown that this behavior can unfold under a third scenario: If conflict with dominant breeders is especially risky, then subordinates will politely wait their turn to breed, rather than risk a fight. Buston and Song explain their experimental and observational work in this feature, and then outline how their work influences current theory.

of a presentation that was presented at the Society for Integrative and Comparative Biology 2014 Annual Meeting, 3-7 January, Austin, TX, USA.

A major focus in evolutionary ecology lies in explaining the evolution and maintenance of social systems. Although most theoretical formulations of social system evolution were initially inspired by studies of birds, mammals, and insects, incorporating a wider taxonomic perspective is important for testing deeply entrenched theory. Here, we review the contribution of studies of habitat-specialist coral reef fishes to our understanding of the evolutionary ecology of animal social systems. These fishes are ecologically similar but display remarkable variation in mating systems, social organization, and sex allocation strategies. By reviewing recent research, we demonstrate their amenability for experimental testing of key concepts in social evolution and for generating novel insights, including the ultimate reasons for female reproductive suppression, group living, and bidirectional sex change. Habitat-specialist reef fishes are a tried and tested group of model organisms for advancing our understanding of the evolution and ecology of social systems in animals.

A major focus in evolutionary ecology lies in explaining the evolution and maintenance of social systems. Although most theoretical formulations of social system evolution were initially inspired by studies of birds, mammals, and insects, incorporating a wider taxonomic perspective is important for testing deeply entrenched theory. Here, we review the contribution of studies of habitat-specialist coral reef fishes to our understanding of the evolutionary ecology of animal social systems. These fishes are ecologically similar but display remarkable variation in mating systems, social organization, and sex allocation strategies. By reviewing recent research, we demonstrate their amenability for experimental testing of key concepts in social evolution and for generating novel insights, including the ultimate reasons for female reproductive suppression, group living, and bidirectional sex change. Habitat-specialist reef fishes are a tried and tested group of model organisms for advancing our understanding of the evolution and ecology of social systems in animals.

Human activities have profoundly changed the abiotic environment, which in many cases has resulted in the deterioration of natural ecosystems. This global problem requires collaborative efforts between scientific disciplines and between academics and wildlife managers if we are to preserve the remaining biodiversity. Here I discuss a potential application of behavioral ecology to the conservation of social species in the face of abiotic disturbances. I argue that a key step towards an effective application of behavioral ecology lies in uniting perspectives on the effects of individual-level social behaviors at multiple levels of ecological organization, with perspectives on the effects of human-induced abiotic perturbations on individual-level social responses. I illustrate these linkages by describing how dominance interactions between individuals can affect the structure and stability of groups, populations and communities. Then, I discuss how abiotic perturbations brought about by two key anthropogenic impacts can disrupt dominance relationships and hierarchy stability in both terrestrial and aquatic ecosystems. Finally, I describe management implications to highlight the applicability of the framework to predict consequences, prioritize species for conservation effort and suggest management actions. By integrating these perspectives, researchers studying animal societies can articulate a clear conservation message - that is, the impact of human activities on populations and communities can be rapidly and accurately predicted based on the modulation of individual-level social traits caused by abiotic stressors.

Human activities have profoundly changed the abiotic environment, which in many cases has resulted in the deterioration of natural ecosystems. This global problem requires collaborative efforts between scientific disciplines and between academics and wildlife managers if we are to preserve the remaining biodiversity. Here I discuss a potential application of behavioral ecology to the conservation of social species in the face of abiotic disturbances. I argue that a key step towards an effective application of behavioral ecology lies in uniting perspectives on the effects of individual-level social behaviors at multiple levels of ecological organization, with perspectives on the effects of human-induced abiotic perturbations on individual-level social responses. I illustrate these linkages by describing how dominance interactions between individuals can affect the structure and stability of groups, populations and communities. Then, I discuss how abiotic perturbations brought about by two key anthropogenic impacts can disrupt dominance relationships and hierarchy stability in both terrestrial and aquatic ecosystems. Finally, I describe management implications to highlight the applicability of the framework to predict consequences, prioritize species for conservation effort and suggest management actions. By integrating these perspectives, researchers studying animal societies can articulate a clear conservation message - that is, the impact of human activities on populations and communities can be rapidly and accurately predicted based on the modulation of individual-level social traits caused by abiotic stressors.

Social aggression is one of the most conspicuous features of animal societies, yet little is known about the causes of individual variation in aggression within social hierarchies. Recent theory suggests that when individuals form queues for breeding, variation in social aggression by non-breeding group members is related to their probability of inheriting breeding status. However, levels of aggression could also vary as a temporary response to changes in the hierarchy, with individuals becoming more aggressive as they ascend in rank, in order to re-establish dominance relationships. Using the group-living fish, Neolamprologus pulcher, we show that subordinates became more aggressive after they ascended in rank. Female ascenders exhibited more rapid increases in aggression than males, and the increased aggression was primarily directed towards group members of adjacent rather than non-adjacent rank, suggesting that social aggression was related to conflict over rank. Elevated aggression by ascenders was not sustained over time, there was no relationship between rank and aggression in stable groups, and aggression given by ascenders was not sex-biased. Together, these results suggest that the need to re-establish dominance relationships following rank ascension is an important determinant of variation in aggression in animal societies.

A rich theoretical framework exists for understanding animal conflict. When two opponents fight over a resource, the duration, intensity and outcome of the fight ought to be determined in large part by the relative difference in resource-holding power between contestants. While our understanding of one-time conflict resolution is excellent, our knowledge is still limited of how these rules scale up when contests occur in a social context where individuals have long-term interactions. Here, we use a convenient model system, Neolamprologus pulcher, a small cooperatively breeding cichlid fish, to explore decisions in pairwise contests over resources in a species where two individual contestants are likely to remain in the same social group, and regularly and repeatedly interact. Contests began after approximately 1 min, with a short display phase, and continued in an aphasic manner for an average of 10 min before a clear winner emerged. Information about opponents' body size was important when deciding on the giving-up point, but contestants' own body size was not, suggesting that assessment of opponent size is paramount in contest decision making. No sex differences were detected in contest structure, duration or intensity, and contests between males or between females were indistinguishable. These results offer an important window on conflict in a cooperative breeder and shed insight on rules of engagement within hierarchical social groups.

A rich theoretical framework exists for understanding animal conflict. When two opponents fight over a resource, the duration, intensity and outcome of the fight ought to be determined in large part by the relative difference in resource-holding power between contestants. While our understanding of one-time conflict resolution is excellent, our knowledge is still limited of how these rules scale up when contests occur in a social context where individuals have long-term interactions. Here, we use a convenient model system, Neolamprologus pulcher, a small cooperatively breeding cichlid fish, to explore decisions in pairwise contests over resources in a species where two individual contestants are likely to remain in the same social group, and regularly and repeatedly interact. Contests began after approximately 1 min, with a short display phase, and continued in an aphasic manner for an average of 10 min before a clear winner emerged. Information about opponents' body size was important when deciding on the giving-up point, but contestants' own body size was not, suggesting that assessment of opponent size is paramount in contest decision making. No sex differences were detected in contest structure, duration or intensity, and contests between males or between females were indistinguishable. These results offer an important window on conflict in a cooperative breeder and shed insight on rules of engagement within hierarchical social groups.

The conundrum of why subordinate individuals assist dominants at the expense of their own direct reproduction has received much theoretical and empirical attention over the last 50 years. During this time, birds and mammals have taken centre stage as model vertebrate systems for exploring why helpers help. However, fish have great potential for enhancing our understanding of the generality and adaptiveness of helping behaviour because of the ease with which they can be experimentally manipulated under controlled laboratory and field conditions. In particular, the freshwater African cichlid, Neolamprologus pulcher, has emerged as a promising model species for investigating the evolution of cooperative breeding, with 64 papers published on this species over the past 27 years. Here we clarify current knowledge pertaining to the costs and benefits of helping in N. pulcher by critically assessing the existing empirical evidence. We then provide a comprehensive examination of the evidence pertaining to four key hypotheses for why helpers might help: (1) kin selection; (2) pay-to-stay; (3) signals of prestige; and (4) group augmentation. For each hypothesis, we outline the underlying theory, address the appropriateness of N. pulcher as a model species and describe the key predictions and associated empirical tests. For N. pulcher, we demonstrate that the kin selection and group augmentation hypotheses have received partial support. One of the key predictions of the pay-to-stay hypothesis has failed to receive any support despite numerous laboratory and field studies; thus as it stands, the evidence for this hypothesis is weak. There have been no empirical investigations addressing the key predictions of the signals of prestige hypothesis. By outlining the key predictions of the various hypotheses, and highlighting how many of these remain to be tested explicitly, our review can be regarded as a roadmap in which potential paths for future empirical research into the evoluti

The number of group members in an animal society can have a major influence on group members' life history, survival, and reproductive success. Identifying the factors that limit group size is therefore fundamental for a complete understanding of social behavior. Here, I examined the relationships between resource availability, social conflict, and group size in the coral-dwelling fish, Paragobiogon xanthosomus (Gobiidae). The size of the largest (breeding) female and the minimum size difference required for hierarchy stability strongly but not perfectly predicted maximum group size, suggesting that social conflicts and hierarchy structure set the upper limit on group size. Deviations in group size around the predicted maximum were explained by variation in average body size ratios but not by variation in coral size, suggesting that coral size does not directly influence group size. In contrast, coral size was a significant predictor of body size ratios, and possible explanations for this relationship are discussed. Group size may be limited by the social conflicts that characterize size-based dominance hierarchies. Ecological factors, namely coral size, may in turn play an indirect role via an effect on body size ratios.