Suspending disbelief—of Wynne-Edwards and his reception

Critique of Wynne-Edwards' views on population regulation and sociality suppose a population of discrete, mutually exclusive groups essential to his thought. Yet both his past and present work focus on continually distributed, philopatric populations; his critics have argued the untenability of a position never his own. Wynne-Edwardsian ‘group selection’ focuses on local population productivity under philopatry. A ‘group’ is a local confluence of genotypes which need not be reified, and group selection consists of the differential replication (hence heritability) of the local social environment in which a genotype is embedded. Differential productivity contingent on social environment can eliminate some relational structures on genotypes in favor of others, creating an expanding wave of population productivity as in Wright's shifting balance metaphor. Such a process is inherent in the evolution of reciprocity, where cooperators must cluster to successfully invade a population of defectors. Regulation of resource exploitation in continuously distributed populations may be modeled as overlapping n-person Prisoner's Dilemmas, where each individual participates in several distinct commons and defection represents local over-exploitation of resources.     

Weaving a tight social net: Allogrooming in free-ranging female langurs (Presbytis entellus)

We studied grooming among adults of a one-male multifemale troop of free-ranging Hanuman langurs (Presbytis entellus)living near Jodhpur, India, for 9 years. The 11–13 females devoted about 6% of their day to allogrooming. Adult males, whose tenures averaged 2.2 years, were transient figures in the troop's history, as reflected by their rather peripheral role in the grooming network. Females groomed males 4–40 times more frequently (1006 episodes) than vice versa- (176 episodes). Adult females received 97% of all grooming from other adult females (6655 episodes). Although females exhibited an age- inversed dominance hierarchy, they did not compete for grooming access to particular troop mates. Dyads of all possible rank differences occurred as frequently as expected: 51% of grooming was directed up the hierarchy and 49% down it. Young, high- ranking individuals gave and received significantly more grooming than the oldest, low- ranking females did. The pattern seemed to be influenced by kin selection because of the presumably high degree of female relatedness. They invested most in troopmates with the highest reproductive value, i.e., the youngest individuals. This trend was coupled with a preference of closest kin (mothers and daughters). Reciprocity was the outstanding feature since all adult females groomed and were groomed by all others. Such a tight social net might establish the necessary cohesion during frequent territorial disputes with neighboring troops.     

Reputation,a universal currency for human social interactions

Decision rules of reciprocity include ‘I help those who helped me’ (direct reciprocity) and ‘I help those who have helped others’ (indirect reciprocity), i.e. I help those who have a reputation to care for others. A person''s reputation is a score that members of a social group update whenever they see the person interacting or hear at best multiple gossip about the person''s social interactions. Reputation is the current standing the person has gained from previous investments or refusal of investments in helping others. Is he a good guy, can I trust him or should I better avoid him as a social partner? A good reputation pays off by attracting help from others, even from strangers or members from another group, if the recipient''s reputation is known. Any costly investment in others, i.e. direct help, donations to charity, investment in averting climate change, etc. increases a person''s reputation. I shall argue and illustrate with examples that a person''s known reputation functions like money that can be used whenever the person needs help. Whenever possible I will present tests of predictions of evolutionary theory, i.e. fitness maximizing strategies, mostly by economic experiments with humans.     

Different patterns of allopreening in the same-sex and opposite-sex interactions of juvenile large-billed crows (Corvus macrorhynchos)

Allogrooming, where an individual grooms another, has been extensively studied in various social animals to understand its role in the evolution of cooperation/prosociality. In existing studies in mammals, allogrooming has been suggested to exhibit not only a hygiene but also a social function. Allopreening, a topic of increasing interest in mammals but recently also in birds, has been studied mostly with mature animals. However, in some species immature individuals also show allopreening and its function remains poorly understood. Crows, Corvus spp., are an ideal model to study this phenomenon, because juveniles form year-round aggregates during their long juvenile stage (e.g., throughout 3–4 years). Here, we investigated the function of allopreening in juvenile groups of wild-caught large-billed crows (C. macrorhynchos). Allopreening frequency and duration for three groups of wild-caught juveniles were analysed to determine whether there was a symmetrical (i.e., reciprocal) or asymmetrical allopreening pattern, and if sex composition of the dyad and/or relative dominance of donor and recipient had an effect. We found that both the frequency and duration of male allopreening correlated with frequency of aggression. Allopreening between both males and females occurred unidirectionally from dominants to subordinates but not in the opposite direction. On the contrary, allopreening between a male and a female was found to be reciprocated, though the absolute frequency and duration were both greater in males than in females. These results suggest that the social function of allopreening in juvenile crows differs depending on the sex composition of the dyad, functioning as a dominance signal for same-sex dyads, and serving a social bonding function for opposite-sex dyads. These findings may reflect the potentially crucial roles of allopreening in within-sex competition and opposite-sex attraction during the 3 year-long juvenile stage affecting future mate choice in lifelong monogamy.     

HUMAN COOPERATION BASED ON PUNISHMENT REPUTATION

The threat of punishment usually promotes cooperation. However, punishing itself is costly, rare in nonhuman animals, and humans who punish often finish with low payoffs in economic experiments. The evolution of punishment has therefore been unclear. Recent theoretical developments suggest that punishment has evolved in the context of reputation games. We tested this idea in a simple helping game with observers and with punishment and punishment reputation (experimentally controlling for other possible reputational effects). We show that punishers fully compensate their costs as they receive help more often. The more likely defection is punished within a group, the higher the level of within‐group cooperation. These beneficial effects perish if the punishment reputation is removed. We conclude that reputation is key to the evolution of punishment.     

Personal experience and reputation interact in human decisions to help reciprocally

There is ample evidence that human cooperative behaviour towards other individuals is often conditioned on information about previous interactions. This information derives both from personal experience (direct reciprocity) and from experience of others (i.e. reputation; indirect reciprocity). Direct and indirect reciprocity have been studied separately, but humans often have access to both types of information. Here, we experimentally investigate information use in a repeated helping game. When acting as donor, subjects can condition their decisions to help recipients with both types of information at a small cost to access such information. We find that information from direct interactions weighs more heavily in decisions to help, and participants tend to react less forgivingly to negative personal experience than to negative reputation. Moreover, effects of personal experience and reputation interact in decisions to help. If a recipient''s reputation is positive, the personal experience of the donor has a weak effect on the decision to help, and vice versa. Yet if the two types of information indicate conflicting signatures of helpfulness, most decisions to help follow personal experience. To understand the roles of direct and indirect reciprocity in human cooperation, they should be studied in concert, not in isolation.     

Indirect reciprocity can overcome free-rider problems on costly moral assessment

Indirect reciprocity is one of the major mechanisms of the evolution of cooperation. Because constant monitoring and accurate evaluation in moral assessments tend to be costly, indirect reciprocity can be exploited by cost evaders. A recent study crucially showed that a cooperative state achieved by indirect reciprocators is easily destabilized by cost evaders in the case with no supportive mechanism. Here, we present a simple and widely applicable solution that considers pre-assessment of cost evaders. In the pre-assessment, those who fail to pay for costly assessment systems are assigned a nasty image that leads to them being rejected by discriminators. We demonstrate that considering the pre-assessment can crucially stabilize reciprocal cooperation for a broad range of indirect reciprocity models. In particular for the most leading social norms, we analyse the conditions under which a prosocial state becomes locally stable.     

Transforming the dilemma

How does natural selection lead to cooperation between competing individuals? The Prisoner's Dilemma captures the essence of this problem. Two players can either cooperate or defect. The payoff for mutual cooperation, R, is greater than the payoff for mutual defection, P. But a defector versus a cooperator receives the highest payoff, T, where as the cooperator obtains the lowest payoff, S. Hence, the Prisoner's Dilemma is defined by the payoff ranking T > R > P > S . In a well‐mixed population, defectors always have a higher expected payoff than cooperators, and therefore natural selection favors defectors. The evolution of cooperation requires specific mechanisms. Here we discuss five mechanisms for the evolution of cooperation: direct reciprocity, indirect reciprocity, kin selection, group selection, and network reciprocity (or graph selection). Each mechanism leads to a transformation of the Prisoner's Dilemma payoff matrix. From the transformed matrices, we derive the fundamental conditions for the evolution of cooperation. The transformed matrices can be used in standard frameworks of evolutionary dynamics such as the replicator equation or stochastic processes of game dynamics in finite populations.     

Food sharing and affiliation: An experimental and longitudinal study in cockatiels (Nymphicus hollandicus)

Food sharing has attracted much attention because of its apparently altruistic nature and its link to prosociality. However, food sharing has been mostly studied in a reproductive context, during courtship and parental care, where the fitness benefits are obvious. We still lack a clear understanding of the functions of food sharing outside any reproductive context and within social groups of same‐aged peers. Previous studies suggest that cofeeding, the action to let another animal feed from the same monopolizable food source, may be used to build and strengthen bonds between individuals. This may be particularly crucial in social birds forming long‐term associations between mates or siblings such as psittacids and corvids. Here, we investigated food sharing and affiliative behaviors such as allopreening in a psittacine species, namely in a group of captive juvenile cockatiels (Nymphicus hollandicus) consisting of five siblings and five unrelated birds. Our main objective was to study the developmental pattern of food sharing over time and its implication in social bonding depending on kinship, affiliation, and sex. Studying cockatiels in this context is providing many new information since most of the studies on food sharing in birds focused on corvids. We found that, contrary to jackdaws, cockatiels continued to share food with multiple individuals, although the frequency of cofeeding as well as the number of cofeeding partners decreased over time. Cockatiels shared more food with their siblings than with other conspecifics but they were not more likely to do cofeeding with birds of the opposite sex. We also found evidence that young cockatiels exchanged more food with those from whom they received food (reciprocity) and, to a lesser extent, allopreening (interchange), than from other cockatiels. Our findings suggest that in cockatiels, food sharing within social groups serves the formation (and maintenance) of affiliative bonds, especially between siblings, rather than pair bonds, but might additionally be explained by reciprocity, interchange, and harassment avoidance.