Reputation-based partner choice is an effective alternative to indirect reciprocity in solving social dilemmas

When group interests clash with individual ones, maintaining cooperation poses a problem. However, cooperation can be facilitated by introducing reputational incentives. Through indirect reciprocity, people who cooperate in a social dilemma are more likely to receive cooperative acts from others. Another mechanism that enhances group cooperation is reputation-based partner choice, or competitive altruism. According to this framework, cooperators benefit via increased access to cooperative partners. Our study compared the effectiveness of indirect reciprocity and competitive altruism in re-establishing cooperation after the typical decline found during repeated public goods games. Twenty groups of four participants first played a series of public goods games, which confirmed the expected decline. Subsequently, public goods games were alternated with either indirect reciprocity games (in which participants had an opportunity to give to another individual from whom they would never receive a direct return) or competitive altruism games (in which they could choose partners for directly reciprocal interactions). We found that public goods game contributions increased when interspersed with competitive altruism games; they were also higher than in public goods games interspersed with indirect reciprocity games. Investing in reputation by increasing contributions to public goods was a profitable strategy in that it increased returns in subsequent competitive altruism and indirect reciprocity games. There was also some evidence that these returns were greater under competitive altruism than indirect reciprocity. Our findings indicate that strategic reputation building through competitive altruism provides an effective alternative to indirect reciprocity as a means for restoring cooperation in social dilemmas.     

Public Information Influences Sperm Transfer to Females in Sailfin Molly Males

In animals, including humans, the social environment can serve as a public information network in which individuals can gather public information about the quality of potential mates by observing conspecifics during sexual interactions. The observing individual itself is also a part of this information network. When recognized by the observed conspecifics as an audience, his/her presence could influence the sexual interaction between those individuals, because the observer might be considered as a potential mate or competitor. One of the most challenging questions in sexual selection to date is how the use of public information in the context of mate choice is linked to the fitness of individuals. Here, we could show that public information influences mate-choice behaviour in sailfin molly males, Poecilia latipinna, and influences the amount of sperm males transfer to a female partner. In the presence of an audience male, males spent less time with the previously preferred, larger of two females and significantly more time with the previously non-preferred, smaller female. When males could physically interact with a female and were faced with an audience male, three audience females or no audience, males transferred significantly more sperm to a female partner in the presence of an audience male than with female audience or no audience and spent less time courting his female partner. This is the first study showing that public information use turns into fitness investment, which is the crucial factor to understand the role of public information in the dynamic processes in sexual selection.     

Female cleaner fish cooperate more with unfamiliar males

Joint group membership is of major importance for cooperation in humans, and close ties or familiarity with a partner are also thought to promote cooperation in other animals. Here, we present the opposite pattern: female cleaner fish, Labroides dimidiatus, behave more cooperatively (by feeding more against their preference) when paired with an unfamiliar male rather than with their social partner. We propose that cooperation based on asymmetric punishment causes this reversed pattern. Males are larger than and dominant to female partners and are more aggressive to unfamiliar than to familiar female partners. In response, females behave more cooperatively with unfamiliar male partners. Our data suggest that in asymmetric interactions, weaker players might behave more cooperatively with out-group members than with in-group members to avoid harsher punishment.     

Extra-pair paternity as the result of reproductive transactions between paired mates

Transactional ('optimal skew' or concessions') models of social evolution emphasize that dominant members of society can be favoured for donating parcels of reproduction to same-sexed subordinates in return for cooperation by the latter. We developed a mathematically similar model in which extra-pair paternity in broods receiving biparental care is viewed as emerging from a reproductive transaction between the paired mates. The model quantitatively predicted the maximum paternity that a male mate can demand before its female mate is favoured to break the pair bond and caring solitarily for a brood sired entirely by a neighbouring male. The model predicts that extra-pair paternity results when the neighbouring male is of sufficiently higher quality than the male mate. In such cases, the exact amount of extra-pair paternity will vary directly with the difference in quality between the two males and inversely with the value (fitness impact) of the male mate's parental care. Importantly, the transactional model provided a unified explanation for experimental and observational evidence that extra-pair paternity rises with decreasing quality of the male mate, increasing genetic variability among breeding males, increasing breeding density, increasing availability of food and decreasing involvement of the male mate in parental care.     

The hierarchy of virtue: mutualism, altruism and signaling in Martu women's cooperative hunting

Cooperative hunting is often assumed to be mutualistic, maintained through returns to scale, where, by working together, foragers can gain higher per capita return rates or harvest sizes than they can by hunting alone. We test this hypothesis among Martu hunters and find that cooperation only provides increased returns to poorer hunters while disadvantaging better hunters. Even so, better hunters still cooperate as frequently as poorer hunters. We ask whether better hunters are advantaged in secondary sharing distributions or whether they bias their partner choice to kin or household members. We find that better hunters are not more likely to pair up with kin and they do not gain consumption benefits from acquiring more. They share a greater proportion of their harvest than poorer hunters: no matter how much one produces — better hunter, worse hunter, cooperator, solitary hunter — all eat the same amount in the end. Such a result suggests the hypothesis that cooperation might be a costly signal of commitment to the public interest on the part of better hunters, which generates trust among camp members and facilitates strong social networks, particularly among women, who cooperate more than men. While some foragers may benefit through cooperation from returns to scale or risk reduction, others may benefit more through signaling commitment and generating trust.     

Eider females form non-kin brood-rearing coalitions

Kin selection is a powerful tool for understanding cooperation among individuals, yet its role as the sole explanation of cooperative societies has recently been challenged on empirical grounds. These studies suggest that direct benefits of cooperation are often overlooked, and that partner choice may be a widespread mechanism of cooperation. Female eider ducks (Somateria mollissima) may rear broods alone, or they may pool their broods and share brood-rearing. Females are philopatric, and it has been suggested that colonies may largely consist of related females, which could promote interactions among relatives. Alternatively, shared brood care could be random with respect to relatedness, either because brood amalgamations are accidental and nonadaptive, or through group augmentation, assuming that the fitness of all group members increases with group size. We tested these alternatives by measuring the relatedness of co-tending eider females in enduring coalitions with microsatellite markers. Females formed enduring brood-rearing coalitions with each other at random with respect to relatedness. However, based on previous data, partner choice is nonrandom and dependent on female body condition. We discuss potential mechanisms underlying eider communal brood-rearing decisions, which may be driven by the specific ecological conditions under which sociality has evolved in this species.     

A simple mechanism for complex social behavior

The evolution of cooperation is a paradox because natural selection should favor exploitative individuals that avoid paying their fair share of any costs. Such conflict between the self-interests of cooperating individuals often results in the evolution of complex, opponent-specific, social strategies and counterstrategies. However, the genetic and biological mechanisms underlying complex social strategies, and therefore the evolution of cooperative behavior, are largely unknown. To address this dearth of empirical data, we combine mathematical modeling, molecular genetic, and developmental approaches to test whether variation in the production of and response to social signals is sufficient to generate the complex partner-specific social success seen in the social amoeba Dictyostelium discoideum. Firstly, we find that the simple model of production of and response to social signals can generate the sort of apparent complex changes in social behavior seen in this system, without the need for partner recognition. Secondly, measurements of signal production and response in a mutant with a change in a single gene that leads to a shift in social behavior provide support for this model. Finally, these simple measurements of social signaling can also explain complex patterns of variation in social behavior generated by the natural genetic diversity found in isolates collected from the wild. Our studies therefore demonstrate a novel and elegantly simple underlying mechanistic basis for natural variation in complex social strategies in D. discoideum. More generally, they suggest that simple rules governing interactions between individuals can be sufficient to generate a diverse array of outcomes that appear complex and unpredictable when those rules are unknown.     

Cooperation as a volunteer’s dilemma and the strategy of conflict in public goods games

Conflict and cooperation for the exploitation of public goods are usually modelled as an N‐person prisoner’s dilemma. Many social dilemmas, however, would be described more properly as a volunteer’s dilemma, in which a certain number of individuals are necessary to produce a public good. If volunteering is costly, but so is failure to produce the public good, cheaters can invade and form a stable mixed equilibrium with cooperators. The dilemma is that the benefit for the group decreases with group size because the larger the group is, the less likely it is that someone volunteers. This problem persists even in the presence of a high degree of relatedness between group members. This model provides precise, testable predictions for the stability of cooperation. It also suggests a counterintuitive but practical solution for this kind of social dilemmas: increasing the damage resulting from the failure to produce the public good increases the probability that the public good is actually produced. Adopting a strategy that entails a deliberate risk (brinkmanship), therefore, can lead to a benefit for the society without being detrimental for the individual.     

A Transactional Theory of Within-Group Conflict

Transactional models of social evolution emphasize that dominant members of the society can be favored to donate parcels of reproduction to subordinate members in return for cooperation. I construct a formal theory of intragroup conflict within the framework of transactional models by determining the maximum extent to which colony members can be selfish without destabilizing the group. The difference between the maximum value of the subordinate's fraction of group reproduction that the dominant can tolerate before ejecting the subordinate and the minimum value required by the subordinate to stay and cooperate peacefully in the group defines the "window of selfishness," which in turn predicts the frequency of within-group conflict. The window of selfishness tends to increase with increasing group reproductive output, increasingly harsh ecological constraints on solitary breeding, and, counterintuitively, increasing relatedness between subordinate and dominant. Increasing fighting ability of the subordinate can either widen or narrow the window of selfishness, the latter being most likely when ecological constraints on group living are strong. Although increasing relatedness is predicted to increase the rate of within-group aggression, the mean intensity of an aggressive act should decline, as predicted by the general theory of honest signaling between relatives and the tug-of-war models of within-group selfishness. In the bidding game, in which multiple dominants bid for the services of a subordinate, the window of selfishness is predicted to have zero width. A zero-width window of selfishness and low conflict also are predicted for saturated N-person groups, that is, groups whose total output is a concave function of group size and in which the dominant is not favored to admit additional subordinates. The model's predictions are compared to empirical evidence and to predictions of alternative models of intragroup aggression, including the value-aggression model and the pure tug-of-war model.     

Negotiations over Grooming in Wild Vervet Monkeys (Chlorocebus pygerythrus)

Mutual grooming plays a central role in the establishment and maintenance of social relationships in primates. Allogrooming has two main functions: hygiene and bonding with partners. The duration of grooming bouts is commonly used in studies of the functional aspects of grooming, but few reflect on the proximate mechanisms that determine grooming bout lengths. As it is highly unlikely that groomer and groomee prefer exactly the same bout length, we are likely to observe the result of some form of negotiation. We currently lack information about the signals that primates employ to inform others about their intentions and desires concerning grooming interactions. From October 2006 until April 2007 we studied three behaviors shown in grooming interactions that could potentially have a signaling function in the negotiation process over the initiation and length of grooming bouts among adult females of two vervet groups freely ranging in the Loskop Dam Nature Reserve, South Africa: approaching another individual as far as that resulted in a grooming session, changing of the body position by the groomed individual, and lip smacking. We found that “approach” did not reliably predict which individual would receive grooming first, although approaching individuals groomed significantly more than those approached. Thus, in the context of grooming interactions, moving toward a group member may signal the willingness to invest. Body part presentations appeared to be the main signal used to demand a prolongation of the grooming by the partner. Finally, lip smacking was used under potentially stressful circumstances, notably shortly before using the mouth to groom the partner or an attempt to touch a mother’s infant. Our exploratory study hopefully inspires colleagues to start looking at the role of communication during cooperative interactions for a better appreciation of how animals manage cooperation and negotiate exchange rates.     

The evolution of strong reciprocity: cooperation in heterogeneous populations

How do human groups maintain a high level of cooperation despite a low level of genetic relatedness among group members? We suggest that many humans have a predisposition to punish those who violate group-beneficial norms, even when this imposes a fitness cost on the punisher. Such altruistic punishment is widely observed to sustain high levels of cooperation in behavioral experiments and in natural settings. We offer a model of cooperation and punishment that we call STRONG RECIPROCITY: where members of a group benefit from mutual adherence to a social norm, strong reciprocators obey the norm and punish its violators, even though as a result they receive lower payoffs than other group members, such as selfish agents who violate the norm and do not punish, and pure cooperators who adhere to the norm but free-ride by never punishing. Our agent-based simulations show that, under assumptions approximating likely human environments over the 100000 years prior to the domestication of animals and plants, the proliferation of strong reciprocators when initially rare is highly likely, and that substantial frequencies of all three behavioral types can be sustained in a population. As a result, high levels of cooperation are sustained. Our results do not require that group members be related or that group extinctions occur.     

Charitable giving as a signal of trustworthiness: Disentangling the signaling benefits of altruistic acts

It has been shown that psychological predispositions to benefit others can motivate human cooperation and the evolution of such social preferences can be explained with kin or multi-level selection models. It has also been shown that cooperation can evolve as a costly signal of an unobservable quality that makes a person more attractive with regard to other types of social interactions. Here we show that if a proportion of individuals with social preferences is maintained in the population through kin or multi-level selection, cooperative acts that are truly altruistic can be a costly signal of social preferences and make altruistic individuals more trustworthy interaction partners in social exchange. In a computerized laboratory experiment, we test whether altruistic behavior in the form of charitable giving is indeed correlated with trustworthiness and whether a charitable donation increases the observing agents' trust in the donor. Our results support these hypotheses and show that, apart from trust, responses to altruistic acts can have a rewarding or outcome-equalizing purpose. Our findings corroborate that the signaling benefits of altruistic acts that accrue in social exchange can ease the conditions for the evolution of social preferences.     

Stress hormones and mate choice

A few recent studies have suggested that glucocorticoid stress hormones can play a role in sexual selection. In terms of mate choice, these studies have shown that individuals can exhibit preferences for mates with either low baseline or peak glucocorticoid levels. This appears to occur because stress hormones can be key mediators of many condition-dependent, sexually selected traits that serve as honest signals of mate quality. We suggest that our understanding of how sexual selection can influence the glucocorticoid stress response will be expanded by studying the interactions among glucocorticoid stress hormones, sex steroids and body condition.     

It's all relative: allometry and variation in the baculum (os penis) of the harp seal, Pagophilus groenlandicus (Carnivora: Phocidae)

We compared allometry and variation in the baculum (os penis), mandible, and humerus of the harp seal, Pagophilus groenlandicus. This species is presumed to have a promiscuous mating system in which choice of mate by females during intromission with different males is likely. The baculum is large and grows throughout life so may be an honest indicator of males' quality (size) or viability (age). We predicted that bacular size would exhibit stronger allometry relative to body size than mandibles or humeri. The baculum is less functionally (mechanically) constrained than mandibles or humeri so we also predicted it would be more variable, though less variable than sexually selected traits which do not function as honest indicators. Our sample (N=67 seals) represented broad ranges of size and age (0–35 yr) so we compared variation using residuals from allometric regressions of skeletal measurements on body length. Bacular size was isometric to body length until ∼ 137 cm (when some seals enter puberty) in body length then was highly positively allometric; mandibular and humeral size were negatively allometric to body length throughout growth. Bacula were more variable than mandibles or humeri. Bacular size in large specimens (>137 cm in body length) was related strongly to body length and weakly to age. We interpret bacular size to be an uncheatable honest indicator of male quality and viability. High bacular variation conforms with theoretical predictions of females' asymmetrical choice of mate and choice of extremes, and may reflect corresponding anatomical variation among females. Some bacular variation may also result incidentally from positive allometry coupled with lifelong bacular growth, which can amplify early differences between reproductive and somatic growth, enabled by weak selection on bacular form in relation to function.     

Behavioural phenotype affects social interactions in an animal network

Animal social networks can be extremely complex and are characterized by highly non-random interactions between group members. However, very little is known about the underlying factors affecting interaction preferences, and hence network structure. One possibility is that behavioural differences between individuals, such as how bold or shy they are, can affect the frequency and distribution of their interactions within a network. We tested this using individually marked three-spined sticklebacks (Gasterosteus aculeatus), and found that bold individuals had fewer overall interactions than shy fish, but tended to distribute their interactions more evenly across all group members. Shy fish, on the other hand, tended to associate preferentially with a small number of other group members, leading to a highly skewed distribution of interactions. This was mediated by the reduced tendency of shy fish to move to a new location within the tank when they were interacting with another individual; bold fish showed no such tendency and were equally likely to move irrespective of whether they were interacting or not. The results show that animal social network structure can be affected by the behavioural composition of group members and have important implications for understanding the spread of information and disease in social groups.     

Cannibalism as an interacting phenotype: precannibalistic aggression is influenced by social partners in the endangered Socorro Isopod (Thermosphaeroma thermophilum)

Models for the evolution of cannibalism highlight the importance of asymmetries between individuals in initiating cannibalistic attacks. Studies may include measures of body size but typically group individuals into size/age classes or compare populations. Such broad comparisons may obscure the details of interactions that ultimately determine how socially contingent characteristics evolve. We propose that understanding cannibalism is facilitated by using an interacting phenotypes perspective that includes the influences of the phenotype of a social partner on the behaviour of a focal individual and focuses on variation in individual pairwise interactions. We investigated how relative body size, a composite trait between a focal individual and its social partner, and the sex of the partners influenced precannibalistic aggression in the endangered Socorro isopod, Thermosphaeroma thermophilum. We also investigated whether differences in mating interest among males and females influenced cannibalism in mixed sex pairs. We studied these questions in three populations that differ markedly in range of body size and opportunities for interactions among individuals. We found that relative body size influences the probability of and latency to attack. We observed differences in the likelihood of and latency to attack based on both an individual's sex and the sex of its partner but found no evidence of sexual conflict. The instigation of precannibalistic aggression in these isopods is therefore a property of both an individual and its social partner. Our results suggest that interacting phenotype models would be improved by incorporating a new conditional ψ, which describes the strength of a social partner's influence on focal behaviour.     

Human mate choice and the wedding ring effect

Individuals are often restricted to indirect cues when assessing the mate value of a potential partner. Females of some species have been shown to copy each other’s choice; in other words, the probability of a female choosing a particular male increases if he has already been chosen by other females. Recently it has been suggested that mate-choice copying could be an important aspect of human mate choice as well. We tested one of the hypotheses, the so-called wedding ring effect—that women would prefer men who are already engaged or married—in a series of live interactions between men and women. The results show that women do not find men signaling engagement, or being perceived as having a partner, more attractive or higher in socioeconomic status. Furthermore, signs of engagement did not influence the women’s reported willingness to engage in short-term or long-term relationships with the men. Thus, this study casts doubt on some simplified theories of human mate-choice copying, and alternative, more complex scenarios are outlined and discussed. Tobias Uller works on broad issues in evolutionary biology, such as life-history evolution and evolutionary genetics. Christoffer Johansson recently received his Ph.D. with a dissertation on biomechanics of swimming birds. Their collaborative work on humans is focused on mate choice.     

从合作的进化探讨植物与传粉者的相互作用

合作的进化为研究植物–传粉者相互关系提供了新的视角。植物与传粉者通过"报酬换服务"建立种间合作关系。这一合作关系从建立、维持到解体面临着3个关键问题:(1)在植物和传粉者不了解对方质量信息时,双方如何选择出最适伙伴,进而建立合作关系;(2)合作方如何限制欺骗策略(比如,盗蜜和欺骗性传粉)的扩散以维持合作关系;(3)什么过程可导致传粉合作关系的解体。植物与传粉者间信号博弈或筛选博弈可促进二者合作关系的建立。面对欺骗策略,传粉者和植物分别采用伙伴选择机制和防御机制加以应对。合作者与欺骗者的稳定共存也有助于植物–传粉者合作的维持。从合作转向对抗、转向新的伙伴和合作放弃3个过程可导致植物–传粉者的合作关系的解体。植物与传粉者合作关系的理论预期已经得到了部分实验结果支持,深化了我们对植物与传粉者合作过程中关键机制的理解。在今后的研究中,需要进一步探讨以下问题:(1)传粉者对植物信号诚实性的选择作用和植物对传粉者的筛选作用;(2)植物与传粉者各自应对欺骗策略的可能机制及其相对重要性;(3)合作者与欺骗者稳定共存的机制;(4)植物与传粉者合作系统对全球变化的响应。     

Same-Sex Gaze Attraction Influences Mate-Choice Copying in Humans

Mate-choice copying occurs when animals rely on the mating choices of others to inform their own mating decisions. The proximate mechanisms underlying mate-choice copying remain unknown. To address this question, we tracked the gaze of men and women as they viewed a series of photographs in which a potential mate was pictured beside an opposite-sex partner; the participants then indicated their willingness to engage in a long-term relationship with each potential mate. We found that both men and women expressed more interest in engaging in a relationship with a potential mate if that mate was paired with an attractive partner. Men and women''s attention to partners varied with partner attractiveness and this gaze attraction influenced their subsequent mate choices. These results highlight the prevalence of non-independent mate choice in humans and implicate social attention and reward circuitry in these decisions.     

Pair Swimming and Mutual Partner Guarding in Monogamous Butterflyfish (Pisces,Chaetodontidae): a Joint Advertisement for Territory

The ecology and social systems of Red Sea butterflyfish were studied along the Sinai coast. Distribution, population density and sociography indicate a preponderance of monogamy in all species, independent of reef zonation and seasonal or long-term changes of the reef environment. The social and mating systems are mutual monogamy; partners remain together for up to 6 years. Links between the monogamy and benthic habits of these fish are indicated by their feeding ecology and by a comparison with Indopacific plankton-feeding species, which usually occur in groups. Territories are defended pairwise. Pair partners nearly always remain within visual contact, but a species-typical distance apart. Pair swimming is independent of season or time of day. Experimental partner removals led to speedy replacement of mates. Possible causes of high partner permanency with little partner fidelity are discussed. In good habitats, single territorial adults do not occur. They would be unable to defend and maintain a territory over long periods. While cooperating in defence and maintenance of a territory, each individual can exploit the resources of the territory including availability of a mate. Pair swimming with mutual partner guarding is considered a joint territorial advertisement similar to duetting in birds; it minimizes agonistic interactions between territory holders. As butterflyfish have planktonic larvae without any broodcare, monogamy could not have evolved in response to broodcare activities as in many other vertebrate groups.