Social Eavesdropping: A Game-Theoretic Analysis

Here we extend the classic Hawk-Dove model of animal conflict to allow for continuous variation in fighting strengths. Whereas the winner of a fight is chosen at random in the discrete game, in our continuous game, the winner of any fight is the stronger individual, and costs are higher for more evenly matched opponents. We identify the evolutionary stable strength threshold beyond which an animal should be prepared to engage in aggressive behaviour and show that this threshold increases with variance in fighting strength when the costs of aggression are insensitive to the level of strength asymmetry, but decreases with variance when the costs are sensitive to the level of asymmetry. In contrast to the classic discrete game, population-wide aggressive behaviour occurs only when the costs of fighting are zero. It is now known that animals can eavesdrop on the outcome of contests between neighbours and modify their behaviour towards observed winners and losers. We therefore further extend our model to allow for social eavesdropping within networks comprising three individuals. Whereas earlier work showed that eavesdropping increases the frequency of mutually aggressive contests in the discrete game by enhancing the value of victory, here we show that aggression thresholds in the continuous game are always higher with eavesdropping than without it: for sufficiently weak animals, avoiding the costs of challenging an observed winner over-rides the potential benefit of winning, so that eavesdropping reduces the frequency of aggressive encounters. Thus, even though strength is not directly observable, information is extracted from the variation in fighting ability that the classic Hawk-Dove game ignores.     

The coevolution of juvenile play–fighting and adult competition

Although play–fighting is widespread among juvenile mammals, its adaptive significance remains unclear. It has been proposed that play is beneficial for developing skills to improve success in adult contests (motor‐training hypothesis), but the links between juvenile play–fighting and adult aggression are complex and not well understood. In this theoretical study, we investigate the coevolution between juvenile play–fighting and adult aggression using evolutionary computer simulations. We consider a simple life history with two sequential stages: a juvenile phase in which individuals play–fight with other juveniles to develop their fighting skills; and an adult phase in which individuals engage in potentially aggressive contests over access to resources and ultimately mating opportunities, leading to reproductive success. The simulations track genetic evolution in key traits affecting adult contests, such as the level of aggression, as well as juvenile investment in play–fighting, capturing the coevolutionary feedbacks between juvenile and adult decisions. We find that coevolution leads to one of two outcomes: a high‐play, high‐aggression situation with highly aggressive adult contests preceded by a prolonged period of juvenile play–fighting to improve fighting ability, or a low‐play, low‐aggression situation in which adult contests are resolved without fighting and there is minimal investment in play–fighting before individuals mature. Which of these outcomes is favoured depends on the mortality costs and on the type of societal structure: societies with strong reproductive skew, favouring monopolization of resources, show high levels of adult aggression and high investment in juvenile play–fighting, whereas societies with low reproductive skew have both low adult aggression and low levels of play–fighting. A review of empirical evidence, particularly in the primate genus Macaca, highlights some limitations of our model and suggests that other, complementary functional explanations are needed to account for the full range of competitive and cooperative forms of play–fighting. Our study illustrates the power of evolutionary simulations to shed light on the long‐standing puzzle of animal play.     

Social evolution: cooperation by conflict

A recent study suggests that aggression between wasps depends upon the costs and benefits of fighting, as determined by the position of individuals in a dominance hierarchy.     

Intrasexual selection favours an immune‐correlated colour ornament in a dragonfly

Sexual signalling is predicted to shape the evolution of sex‐specific ornamentation, and establishing the costs and benefits of ornamentation and the information that ornamentation provides to receivers is necessary to evaluating this adaptive function. Here, we assessed the adaptive function of a common colour ornament in insects, melanin wing ornamentation, using the dragonfly Pachydiplax longipennis. We hypothesized that greater ornamentation would improve territory‐holding success by decreasing aggression that males receive from territorial rivals, but that more ornamented males may have shorter lifespans. Using mark–recapture field observations, we found that more ornamented males had greater territory‐holding success and that viability selection did not act on wing melanization. We then compared the aggression of territorial rivals to decoy males before and after experimentally augmenting wing melanization, finding that males significantly reduced aggression following the manipulation. We next hypothesized that wing melanization would signal fighting ability to territorial rivals by reflecting condition via investment in the costly melanin synthesis pathway. We observed a positive relationship between ornamentation and the likelihood of winning territorial disputes, suggesting that wing melanization provides information about fighting ability to rivals. We also found a positive relationship between melanin‐based immune defence and ornamentation, supporting a link between the signal and condition. We conclude that wing melanization is a condition‐related signal of fighting ability and suggest that this may be a common mechanism promoting the evolution of melanin ornamentation.     

The energetic cost of reproductive conflicts in the ant Pachycondyla obscuricornis

In a variety of social animals, individuals can secure reproductive rights through aggressive dominance. Direct individual benefits of aggression are widely recognized, but underlying costs affecting group productivity, and thus indirect benefits, are less clear. Costs of aggressive regulation of reproduction are especially important in small social insect colonies, where individual workers could potentially dominate male production. We estimated the energetic costs associated with the regulation of worker reproduction in the ponerine ant Pachycondyla obscuricornis, using the total CO2 emission of a colony as a measure. The level of CO2 emission of 12 experimental colonies varied significantly during five periods with varying levels of aggression and egg-laying. Overall, CO2 emission increased with the degree of fighting in a colony, but was not associated with differences in egg-laying. Aggressive regulation of reproduction and the formation of a dominance hierarchy thus pose an energetic cost to the colony. Furthermore, workers reduce their work-activities immediately after experimental orphaning, giving a further cost to the colony. These costs might influence the outcome of conflicts over male production in ants. This paper presents the first quantification of energetic costs of aggressive behavior regulating reproduction in ants.     

The Evolution of Sex: a Perspective from the Fungal Kingdom

Summary: Sex is shrouded in mystery. Not only does it preferentially occur in the dark for both fungi and many animals, but evolutionary biologists continue to debate its benefits given costs in light of its pervasive nature. Experimental studies of the benefits and costs of sexual reproduction with fungi as model systems have begun to provide evidence that the balance between sexual and asexual reproduction shifts in response to selective pressures. Given their unique evolutionary history as opisthokonts, along with metazoans, fungi serve as exceptional models for the evolution of sex and sex-determining regions of the genome (the mating type locus) and for transitions that commonly occur between outcrossing/self-sterile and inbreeding/self-fertile modes of reproduction. We review here the state of the understanding of sex and its evolution in the fungal kingdom and also areas where the field has contributed and will continue to contribute to illuminating general principles and paradigms of sexual reproduction.     

Simulating the Evolution of the Human Family: Cooperative Breeding Increases in Harsh Environments

Verbal and mathematical models that consider the costs and benefits of behavioral strategies have been useful in explaining animal behavior and are often used as the basis of evolutionary explanations of human behavior. In most cases, however, these models do not account for the effects that group structure and cultural traditions within a human population have on the costs and benefits of its members'' decisions. Nor do they consider the likelihood that cultural as well as genetic traits will be subject to natural selection. In this paper, we present an agent-based model that incorporates some key aspects of human social structure and life history. We investigate the evolution of a population under conditions of different environmental harshness and in which selection can occur at the level of the group as well as the level of the individual. We focus on the evolution of a socially learned characteristic related to individuals'' willingness to contribute to raising the offspring of others within their family group. We find that environmental harshness increases the frequency of individuals who make such contributions. However, under the conditions we stipulate, we also find that environmental variability can allow groups to survive with lower frequencies of helpers. The model presented here is inevitably a simplified representation of a human population, but it provides a basis for future modeling work toward evolutionary explanations of human behavior that consider the influence of both genetic and cultural transmission of behavior.     

The evolution of intergroup tolerance in nonhuman primates and humans

Primate individuals use a variety of strategies in intergroup encounters, from aggression to tolerance; however, recent focus on the evolution of either warfare or peace has come at the cost of characterizing this variability. We identify evolutionary advantages that may incentivize tolerance toward extra‐group individuals in humans and nonhuman primates, including enhanced benefits in the domains of transfer, mating, and food acquisition. We highlight the role these factors play in the flexibility of gorilla, chimpanzee, bonobo, and human behavior. Given humans have an especially broad range of intergroup behavior, we explore how the human foraging ecology, especially large spatial and temporal fluctuations in resource availability, may have selected for a greater reliance on tolerant between‐community relationships—relationships reinforced by status acquisition and cultural institutions. We conclude by urging careful, theoretically motivated study of behavioral flexibility in intergroup encounters in humans and the nonhuman great apes.     

Dispersal and fighting in male pollinating fig wasps

For more than two decades, it has been the dogma that the males of pollinating fig wasps do not fight and that they only mate in their native fig. Their extreme degree of local mating leads to highly female biased sex ratios that should eliminate the benefits of fighting and dispersal by males. Furthermore, males sharing a fig are often brothers, and fighting may be barred by kin selection. Therefore, theory supported the presumed absence of fighting and dispersal in pollinating fig wasp males. However, we report here that in pollinating fig wasps, fighting between brothers evolved at least four and possibly six time, and dispersal by males at least twice. This finding supports the idea that competition between relatives can cancel the ameliorating effects of relatedness. The explanation to this evolutionary puzzle, as well as the consequences of male dispersal and fighting, opens the doors to exciting new research.     

The implicit rules of combat reflect the evolved function of combat: An evolutionary-psychological analysis of fairness and honor in human aggression

Aggression was a powerful selection pressure for our human ancestors. Aggressive contests would have selected not only for anatomical features (e.g., superior upper-body strength in adult males), but also for behavior-regulation adaptations for the conditional and cost-effective deployment of aggression. Here, we explore two concepts that appear to be part of the neurocognitive suite that regulates aggression. The first is the concept of a “fair fight”, which tags whether an aggressive bout can provide an unbiased estimate of the relative fighting ability of the combatants. The second concept is the “honorability” of a fighter, which tags whether an individual can be trusted to not inflict aggression in excess of what is needed to establish the relative fighting ability of the combatants. Eight predictions were derived from these hypotheses and tested experimentally with vignettes that varied relevant aspects of a fight. The results support the predictions. Fights are perceived as “fair” to the extent that the fighters avoid biased tactics that improve their probability of winning absent superior fighting ability (e.g., not attacking when the opponent is unprepared). Further, fighters are perceived as “honorable” when they minimize the danger to both combatants (e.g., not attacking after the opponent surrenders). We conclude by listing additional implications of the theory.     

What can animal aggression research tell us about human aggression?

Research on endocrinological correlates of aggression in laboratory animals is implicitly motivated by an expectation that the results of such studies may be applicable to human aggression as well. Research with a focus on the stimulus antecedents of aggression, its response characteristics, and its outcomes suggests a number of detailed correspondences between offensive aggression in laboratory rodents and human angry aggression. These include resource (including status and territory) competition as motives that are particularly elicited by conspecific challenge situations and, when the aggression is successful, outcomes of reduction of challenge and enhancement of resource control and status. Although the response characteristics of human aggression have been dramatically altered by human verbal, technological, and social advancements, there is some evidence for targeting of blows, similar to a well-established pattern for offensive aggression in many nonhuman mammals. Finally, for people as well as for nonhuman mammals, fear of defeat or punishment is a major factor inhibiting the expression of offensive aggression. While defensive aggression has been very little researched in people, it may represent a different phenomenon than angry aggression, again providing a parallel to the offense-defense distinction of laboratory rodent studies.     

Defence, intrusion and the evolutionary stability of territoriality

Territorial behaviour can only be adaptive if its costs are outweighed by its benefits. Territorial individuals incur costs by defending their territories against intruders. Usually these intruders are assumed to be non-territorial floaters attempting to take over the whole territory or neighbours trying to extend the borders of their own territory. We instead investigate how costs and benefits of territorial behaviour are affected by neighbours which invade to steal resources on a territory.We show analytically that in the absence of defence intrusion into neighbouring territories always pays and that even if territories are defended intrusion levels can still be high. Using a more detailed simulation model we find that territory defence usually disappears from the population even if owners have a strong advantage over intruders in terms of fighting costs or foraging efficiency. Defence and thus territoriality can only be evolutionarily stable if fighting costs for the intruder relative to the productivity of the territory are very high or if crossing the borders between territories carries additional costs.Our results show that stealing of resources by neighbours can have a considerable effect on the evolutionary stability of territory defence and thus territoriality itself. A more mechanistic model of territorial behaviour is needed to incorporate these kinds of mechanisms into a general theory on the evolution of territoriality.     

Of Fighting Flies,Mice, and Men: Are Some of the Molecular and Neuronal Mechanisms of Aggression Universal in the Animal Kingdom?

Aggressive behavior is widespread in the animal kingdom, but the degree of molecular conservation between distantly related species is still unclear. Recent reports suggest that at least some of the molecular mechanisms underlying this complex behavior in flies show remarkable similarities with such mechanisms in mice and even humans. Surprisingly, some aspects of neuronal control of aggression also show remarkable similarity between these distantly related species. We will review these recent findings, address the evolutionary implications, and discuss the potential impact for our understanding of human diseases characterized by excessive aggression.     

Sex,gender, and difference

Empirical research has demonstrated that women’s aggressive behavior is widespread and displays regularities across societies. Until recently, however, discussions about the aggressive behavior of women and gender differences in aggressive behavior have been based largely on data from nonhuman primates, children, or laboratory experiments. Using a unique corpus of naturalistic data on aggressive human interactions both between and among men and women, I explore the complexity of our questions about sex differences in aggression and further illuminate the ways in which men and women may use aggression in human interactions. In this paper I compare the aggressive behavior of men and women in an Australian Aboriginal community. In doing so I argue for the continuing use of a “sex differences” framework for organizing our understanding of gender relations and gender hierarchy. I believe, however, that this form of analysis benefits from, if not requires, a sensitivity to the most taken-for-granted aspects of our gender ideology and a commitment to attend to evidence that challenges our convictions about men and women.     

Proximate perspectives on the evolution of female aggression: good for the gander,good for the goose?

Female–female aggression often functions in competition over reproductive or social benefits, but the proximate mechanisms of this apparently adaptive behaviour are not well understood. The sex steroid hormone testosterone (T) and its metabolites are well-established mediators of male–male aggression, and several lines of evidence suggest that T-mediated mechanisms may apply to females as well. However, a key question is whether mechanisms of female aggression primarily reflect correlated evolutionary responses to selection acting on males, or whether direct selection acting on females has made modifications to these mechanisms that are adaptive in light of female life history. Here, I examine the degree to which female aggression is mediated at the level of T production, target tissue sensitivity to T, or downstream genomic responses in order to test the hypothesis that selection favours mechanisms that facilitate female aggression while minimizing the costs of systemically elevated T. I draw heavily from avian systems, including the dark-eyed junco (Junco hyemalis), as well as other organisms in which these mechanisms have been well studied from an evolutionary/ecological perspective in both sexes. Findings reveal that the sexes share many behavioural and hormonal mechanisms, though several patterns also suggest sex-specific adaptation. I argue that greater attention to multiple levels of analysis—from hormone to receptor to gene network, including analyses of individual variation that represents the raw material of evolutionary change—will be a fruitful path for understanding mechanisms of behavioural regulation and intersexual coevolution.     

Evolution of the human menopause

Menopause is an evolutionary puzzle since an early end to reproduction seems contrary to maximising Darwinian fitness. Several theories have been proposed to explain why menopause might have evolved, all based on unusual aspects of the human life history. One theory is that menopause follows from the extreme altriciality of human babies, coupled with the difficulty in giving birth due to the large neonatal brain size and the growing risk of child-bearing at older ages. There may be little advantage for an older mother in running the increased risk of a further pregnancy when existing offspring depend critically on her survival. An alternative theory is that within kin groups menopause enhances fitness by producing post-reproductive grandmothers who can assist their adult daughters. Such theories need careful quantitative assessment to see whether the fitness benefits are sufficient to outweigh the costs, particularly in circumstances of relatively high background mortality typical of ancestral environments. We show that individual theories fail this test, but that a combined model incorporating both hypotheses can explain why menopause may have evolved.     

Male Siamese fighting fish use gill flaring as the first display towards territorial intruders

The Siamese fighting fish (Betta splendens) is well known as an aggressive fish with unique spawning and parental care behavior. During reproduction, male fish construct a bubble nest, court females, protect the brood, and defend the territory through aggressive displays. Aggression in male Siamese fighting fish has long been the subject of investigation; however, the kinematics of aggression during contests have been largely overlooked. Here we investigated how nest-holding, male Siamese fighting fish use two different types of displays, gill flaring and fin spreading, towards intruders during various reproductive phases; before (BB) and after bubble nest building, and after spawning (AS), and hatching (AH). Males were more aggressive towards male than female intruders and the level of aggression changed significantly between reproductive phases. Gill flaring, the more energetically costly display, was the dominant initial display towards male and female intruders in BB, AS, AH phases. However, defending males switched to fin spreading after prolonged exposure to intruders. The results suggest that Siamese fighting fish use gill flaring as an acute response in order to defend their territory; this response may be replaced by fin spreading as a chronic response, probably to reduce the energetic costs during the contest.     

Agonistic Displays and the Benefits of Fighting in the Field Cricket, Gryllus bimaculatus

Fighting is often composed of discrete agonistic displays. Few studies have partitioned fighting behavior into its component agonistic displays and evaluated the relationships between the frequency of the displays and the potential benefits of fighting, particularly mating success. In this study, we quantified the frequency of male field cricket, Gryllus bimaculatus, agonistic displays. The displays were quantified under three social environments which varied in the potential benefits of fighting: males with other males only, males with other males and female scents, and males with other males and females. We found that (1) the presence of females elicited an increase in agonistic displays characteristic of intermediate levels of escalation, (2) female scents did not produce a similar increase in the frequency of agonistic displays, and (3) in the presence of females, the frequency of agonistic displays was positively correlated with mating success. Aggressive stridulation, an energetically low-cost display, was more strongly associated with mating success than were more costly displays. The results are discussed in the context of the evolutionary theory of aggression and in the context of cricket mating systems.     

Costly punishment prevails in intergroup conflict

Understanding how societies resolve conflicts between individual and common interests remains one of the most fundamental issues across disciplines. The observation that humans readily incur costs to sanction uncooperative individuals without tangible individual benefits has attracted considerable attention as a proximate cause as to why cooperative behaviours might evolve. However, the proliferation of individually costly punishment has been difficult to explain. Several studies over the last decade employing experimental designs with isolated groups have found clear evidence that the costs of punishment often nullify the benefits of increased cooperation, rendering the strong human tendency to punish a thorny evolutionary puzzle. Here, we show that group competition enhances the effectiveness of punishment so that when groups are in direct competition, individuals belonging to a group with punishment opportunity prevail over individuals in a group without this opportunity. In addition to competitive superiority in between-group competition, punishment reduces within-group variation in success, creating circumstances that are highly favourable for the evolution of accompanying group-functional behaviours. We find that the individual willingness to engage in costly punishment increases with tightening competitive pressure between groups. Our results suggest the importance of intergroup conflict behind the emergence of costly punishment and human cooperation.     

Aggression Levels Affect Social Interaction in the Non‐Breeding Territorial Aggression of the Weakly Electric Fish,Gymnotus omarorum

Animals typically decide whether to fight or retreat from conspecifics based on their individual estimates of the costs and benefits of fighting. Theoretical models predict how contenders solve a conflict, but the evaluation processes involved in these decisions depend on multiple factors that are difficult to explore experimentally. We addressed these questions using the non‐breeding territorial aggression of Gymnotus omarorum, in which subordinates make three distinctive decisions to signal their submission during a fight: (1) interruption of their electric discharges to hide from the dominant, (2) stop attacking and retreat, and (3) emission of ‘chirps’, transient submissive electric signals. We confirmed that subordinates take into account the aggressive performance of dominants to shape their own agonistic decisions and performance. The intensity of aggression is highly correlated with an agonistic dyad, and the decision of subordinates to retreat is influenced by the attack rates of dominants. When we lowered the aggression of expected dominants with a 5‐HT_1A receptor agonist, the correlation between the two contenders' aggression levels was lost and subordinates completely stopped emitting electric chirp signals. The aforementioned results contribute to the understanding of the decision‐making strategies driven by social challenge inherent to agonistic encounters.