Policing and punishment across the domains of social evolution

Several decades of research in humans, other vertebrates, and social insects have offered fascinating insights into the dynamics of punishment (and its subset, policing), but authors have only rarely addressed whether there are fundamental joint principles underlying the maintenance of these behaviors. Here we present a punisher/bystander approach rooted in inclusive fitness logic to predict which individuals should take on punishing roles in animal societies. We apply our scheme to societies of eusocial Hymenoptera and nonhuman vertebrate social breeders, and we outline potential extensions for understanding conflict regulation among cells in metazoan bodies and unrelated individuals in human societies. We highlight that: 1) no social unit is expected to express punishment behavior unless it collects positive inclusive fitness benefits that surpass alternative benefits of bystanding; 2) punishment with public good benefits can be maintained through either direct fitness benefits (coercion) or indirect fitness benefits (correction) or both; 3) differences across social systems in the distributions of power, relatedness, and reproductive options drive variation in the extent to which individuals actively punish; and 4) inclusive fitness logic captures many punishment‐relevant evolutionary and ecological variables in a single framework that appears to apply across very different types of social arrangements. Synthesis Researchers have long observed that individuals in animal societies punish (and by extension, police) each other, but they have rarely investigated whether general principles underlie this behavior across social arrangements. In this paper, we present a punisher/bystander approach rooted in inclusive fitness logic to predict which individuals should take on punisher roles in animal societies. We apply the approach to eusocial insects and cooperatively breeding vertebrates and outline extensions towards the control of cancer cell lineages and punishment in human groups. We highlight how variation in specific social variables may drive differences in punishing/policing across the social domains.     

Ethnicity,the search for rapists and the press

Media reporting of rape continues to provoke concern. This article focuses on the press reporting of the discovery and search for alleged rapists. On examining nine British newspapers for one complete year (1992), a serious distortion in the media reporting of ethnicity emerges at this early stage of the criminal process. While there is an over‐representation of minority ethnic men accused of rape coming to the notice of the police, the cases identified as involving minority ethnic (particularly black) men as assailants, are much more likely to be reported widely. The distortion is then intensified by associating minority ethnic (and again particularly, black men) with certain types of rape. However, there are important differences between newspapers in their coverage. Furthermore, while media coverage of the search for rapists is a matter of concern, it also needs to be seen as part of the wider process of the social construction of rapists. Nevertheless, the conclusion is that the media coverage of rape tends to distort in ways which may well exacerbate hostility against minority ethnic groups.     

Exact versus heuristic models of kin selection

This paper examines the conditions under which the classical inclusive fitness formulation of Hamilton (1964) provides an adequate approximation to the dynamics of gene frequency change and to conditions for genetic equilibrium, in the “additive” model of altruism between sibs of Uyenoyama and Feldman (1981). It is concluded that the classical formulation is adequate, provided that either the effect of the gene on the probability of behaving altruistically is low or the costs and benefits of altruism are small, unless the benefit/cost ratio k is very close to 2, the value that must be exceeded for altruism to be favoured. In addition, the gene for altruism must be underdominant, recessive or partially recessive in its effect on the probability of behaving altruistically, for the inclusive fitness predictions to break down significantly.     

Pape in Panorpa scorpionflies and a general rape hypothesis

The adaptive significance of heterosexual rape is difficult to demonstrate because (1) female coyness is difficult to distinguish from apparent rape, and (2) male fitness must be enhanced by rape. Male Panorpa scorpionflies inseminate unwilling females by securing the female's wings in an abdominal clamp. Panorpa rapists gain fitness through avoidance of risks of predation by web-building spiders associated with the more typical male behaviours of feeding females a dead insect or salivary secretion during copulation. Males should be most strongly selected to rape in species in which males provide resources important for female reproduction. In such species, the strength of selection on males to rape should exceed the strength of selection on females to prevent rape.     

Male Androphilia in the Ancestral Environment

The kin selection hypothesis posits that male androphilia (male sexual attraction to adult males) evolved because androphilic males invest more in kin, thereby enhancing inclusive fitness. Increased kin-directed altruism has been repeatedly documented among a population of transgendered androphilic males, but never among androphilic males in other cultures who adopt gender identities as men. Thus, the kin selection hypothesis may be viable if male androphilia was expressed in the transgendered form in the ancestral past. Using the Standard Cross-Cultural Sample (SCCS), we examined 46 societies in which male androphilia was expressed in the transgendered form (transgendered societies) and 146 comparison societies (non-transgendered societies). We analyzed SCCS variables pertaining to ancestral sociocultural conditions, access to kin, and societal reactions to homosexuality. Our results show that ancestral sociocultural conditions and bilateral and double descent systems were more common in transgendered than in non-transgendered societies. Across the entire sample, descent systems and residence patterns that would presumably facilitate increased access to kin were associated with the presence of ancestral sociocultural conditions. Among transgendered societies, negative societal attitudes toward homosexuality were unlikely. We conclude that the ancestral human sociocultural environment was likely conducive to the expression of the transgendered form of male androphilia. Descent systems, residence patterns, and societal reactions to homosexuality likely facilitated investments in kin by transgendered males. Given that contemporary transgendered male androphiles appear to exhibit elevated kin-directed altruism, these findings further indicate the viability of the kin selection hypothesis.     

Parental antagonism,relatedness asymmetries,and genomic imprinting.

The theory of inclusive fitness can be modified to consider separate coefficients of relatedness for an individual''s maternal and paternal alleles. A gene is said to have parentally antagonistic effects if it has an inclusive fitness benefit when maternally derived, but an inclusive fitness cost when paternally derived (or vice versa). Parental antagonism favours the evolution of alleles that are expressed only when maternally derived or only when paternally derived (genomic imprinting).     

A reconciliation of inclusive fitness and personal fitness approaches: a proposed correcting term for the inclusive fitness formula

Hamilton implicitly defined the inclusive fitness of an individual as the number of genomes, identical by descent to its own, but not in its own body, which owe their existence to expression of genes in said individual. Hamilton regarded inclusive fitness as the true metric of evolutionary success and the thin- maximized by selection. Williams, Stern and Orlove either claimed this property for mean reproductive success, or stated that expected reproductive success equals expected inclusive fitness. These statements are reconciled if a correcting term is added to Hamilton's inclusive fitness formula.This change completely accounts for inclusive fitness in personal fitness terminology. The use of ? in place of r renders the new formula exact. This has less numerical impact than the addition of the correcting term to begin with, but helps show inclusive fitness theory holds exactly.     

The validity and value of inclusive fitness theory

Social evolution is a central topic in evolutionary biology, with the evolution of eusociality (societies with altruistic, non-reproductive helpers) representing a long-standing evolutionary conundrum. Recent critiques have questioned the validity of the leading theory for explaining social evolution and eusociality, namely inclusive fitness (kin selection) theory. I review recent and past literature to argue that these critiques do not succeed. Inclusive fitness theory has added fundamental insights to natural selection theory. These are the realization that selection on a gene for social behaviour depends on its effects on co-bearers, the explanation of social behaviours as unalike as altruism and selfishness using the same underlying parameters, and the explanation of within-group conflict in terms of non-coinciding inclusive fitness optima. A proposed alternative theory for eusocial evolution assumes mistakenly that workers' interests are subordinate to the queen's, contains no new elements and fails to make novel predictions. The haplodiploidy hypothesis has yet to be rigorously tested and positive relatedness within diploid eusocial societies supports inclusive fitness theory. The theory has made unique, falsifiable predictions that have been confirmed, and its evidence base is extensive and robust. Hence, inclusive fitness theory deserves to keep its position as the leading theory for social evolution.     

Longevity suppresses conflict in animal societies

Models of social conflict in animal societies generally assume that within-group conflict reduces the value of a communal resource. For many animals, however, the primary cost of conflict is increased mortality. We develop a simple inclusive fitness model of social conflict that takes this cost into account. We show that longevity substantially reduces the level of within-group conflict, which can lead to the evolution of peaceful animal societies if relatedness among group members is high. By contrast, peaceful outcomes are never possible in models where the primary cost of social conflict is resource depletion. Incorporating mortality costs into models of social conflict can explain why many animal societies are so remarkably peaceful despite great potential for conflict.     

Social dynamics of mammals: Reproductive success, kinship and individual fitness

Apparent altruism, in which an individual seemingly decreases its evolutionary fitness by assisting others, can confer benefits if the individual assists kin. Thus, an animal can increase its total or inclusive fitness by producing offspring (direct fitness) and/or helping kin to reproduce (indirect fitness). Although kin selection has been suggested as the mechanism underlying the formation of mammalian societies, many species act as if they attempt to maximize the direct fitness component of their inclusive fitness.     

Optimal flight initiation distance

Decisions regarding flight initiation distance have received scant theoretical attention. A graphical model by Ydenberg and Dill (1986. The economics of fleeing from predators. Adv. Stud. Behav. 16, 229-249) that has guided research for the past 20 years specifies when escape begins. In the model, a prey detects a predator, monitors its approach until costs of escape and of remaining are equal, and then flees. The distance between predator and prey when escape is initiated (approach distance = flight initiation distance) occurs where decreasing cost of remaining and increasing cost of fleeing intersect. We argue that prey fleeing as predicted cannot maximize fitness because the best prey can do is break even during an encounter. We develop two optimality models, one applying when all expected future contribution to fitness (residual reproductive value) is lost if the prey dies, the other when any fitness gained (increase in expected RRV) during the encounter is retained after death. Both models predict optimal flight initiation distance from initial expected fitness, benefits obtainable during encounters, costs of escaping, and probability of being killed. Predictions match extensively verified predictions of Ydenberg and Dill's (1986) model. Our main conclusion is that optimality models are preferable to break-even models because they permit fitness maximization, offer many new testable predictions, and allow assessment of prey decisions in many naturally occurring situations through modification of benefit, escape cost, and risk functions.     

Structure and function in mammalian societies

Traditional interpretations of the evolution of animal societies have suggested that their structure is a consequence of attempts by individuals to maximize their inclusive fitness within constraints imposed by their social and physical environments. In contrast, some recent re-interpretations have argued that many aspects of social organization should be interpreted as group-level adaptations maintained by selection operating between groups or populations. Here, I review our current understanding of the evolution of mammalian societies, focusing, in particular, on the evolution of reproductive strategies in societies where one dominant female monopolizes reproduction in each group and her offspring are reared by other group members. Recent studies of the life histories of females in these species show that dispersing females often have little chance of establishing new breeding groups and so are likely to maximize their inclusive fitness by helping related dominants to rear their offspring. As in eusocial insects, increasing group size can lead to a progressive divergence in the selection pressures operating on breeders and helpers and to increasing specialization in their behaviour and life histories. As yet, there is little need to invoke group-level adaptations in order to account for the behaviour of individuals or the structure of mammalian groups.     

Comparative statics of games between relatives

According to Hamilton's theory of kin selection, species tend to evolve behavior such that each organism appears to be attempting to maximize its inclusive fitness. In particular, two neighbors are likely to help each other if the cost of doing so is less than the benefit multiplied by r, their coefficient of relatedness. Since the latter is less than unity, mutual altruism benefits both neighbors. However, is it theoretically possible that acting so as to maximize the inclusive, rather than personal, fitness may harm both parties. This may occur in strategic symmetric pairwise interactions (more specifically, nxn games), in which the outcome depends on both sides' actions. In this case, the equilibrium outcome may be less favorable to the interactants' personal fitness than if each of them acted so as to maximize the latter. This paper shows, however, that such negative effect of relatedness on fitness is incompatible with evolutionary stability. If the symmetric equilibrium strategies are evolutionarily stable, a higher coefficient of relatedness can only entail higher personal fitness for the two neighbors. This suggests that negative comparative statics as above are not likely to occur in nature.     

Policing effectiveness depends on relatedness and group size

Cohesion of social groups requires the suppression of individual selfishness. Indeed, worker egg laying in insect societies is usually suppressed or punished through aggression and egg removal. The effectiveness of such "policing" is expected to increase with decreasing relatedness, as inclusive fitness of group members is more strongly affected by selfish worker reproduction when group members are less closely related to each other. As inclusive fitness is also influenced by the costs and benefits of helping, the effectiveness of policing should decrease with increasing colony size, because the costs for the whole colony from selfish worker reproduction are proportionally reduced in large groups. Here, we show that policing effectiveness in colonies of the ant Temnothorax unifasciatus is low in large groups and high in small groups when relatedness is high. When we experimentally decreased the relatedness in groups, the policing effectiveness reached the same high level as in small, highly related groups, irrespective of group size. Therefore, our results indicate that policing effectiveness is simultaneously shaped by relatedness and group size, that is, an ecological factor. This may have major implications for testing policing across species of animals.     

Direct assessment of queen quality and lack of worker suppression in a paper wasp

Assessing a conspecific's potential is often crucial to increase one's fitness, e.g. in female choice, contests with rivals or reproductive conflicts in animal societies. In the latter, helpers benefit from accurately assessing the fertility of the breeder as an indication of inclusive fitness. There is evidence that this can be achieved using chemical correlates of reproductive activity. Here, we show that queen quality can be assessed by directly monitoring her reproductive output. In the paper wasp Polistes dominulus, we mimicked a decrease in queen fertility by regularly removing brood. This triggered ovarian development and egg-laying by many workers, which strongly suggests that brood abundance is a reliable cue of queen quality. Brood abundance can be monitored when workers perform regular brood care in small size societies where each brood element is kept in a separate cell. Our results also show that although the queen was not manipulated, and thus remained healthy and fully fertile, she did not control worker egg-laying. Nevertheless, when workers laid eggs, the queen secured a near reproductive monopoly by selectively destroying these eggs, a mechanism known as 'queen policing'. By contrast, workers destroyed comparatively few queen-laid eggs, but did destroy each other's eggs.     

Models of the evolution of altruism

There are two ways of calculating the spread of a gene for altruism. One, originally proposed by Hamilton, is to allow for the effects of the gene on the survival and reproduction of collateral relatives of the individual carrying it (i.e., “inclusive fitness”); this leads to the condition k > 1/r for the spread of the gene, where k is a benefit/cost ratio. The other is to count only the direct offspring of a carrier, but to allow for the altruistic acts performed toward the carrier by its relatives (“neighbour modulated fitness” or “personal fitness”). A recent personal fitness model (L. L. Cavalli Sforza and M. W. Feldman, 1978, Theor. Pop. Biol.14, 268–280) analyses parent-offspring and sib-sib altruism and concludes that k > 1/r is applicable only when fitness components are combined additively. The present paper analyses some simple models in which the phenotypic effects are carefully specified. It is concluded that it is sometimes, but not always, appropriate to combine fitness components additively. The relative roles of inclusive and personal fitness models are compared. The former have the virtue of being easier to think about in causal terms; and the latter of incorporating the evolution of altruism into the corpus of population genetics as an example of frequency-dependent selection.     

Rape and Rape Avoidance in Ethno-National Conflicts: Sexual Violence in Liminalized States

Mass rape is a common but not universal occurrence in ethnic or nationalist conflicts. Using South Asian and Bosnian data, in this article I argue that mass rape is likely when such conflicts take place during the partition of a territory and its population, when the state itself is liminal, both its territory and control over it uncertain. In conflicts in which the state is not itself threatened, and thus groups feel that they will continue to coexist, there is some evidence that rape is avoided, even when murder is accepted. However, such instances of rape avoidance are largely unstudied, in large part because of the focus on the violence of mass rape. Further, this focus on violence tends toward classifying all sexual relations between groups whose members have participated in mass rape as improper, thus depriving women who may not wish to rejoin their natal groups of agency, [rape, genocide, violence, India, Yugoslavia/Bosnia]     

How Biochemical Constraints of Cellular Growth Shape Evolutionary Adaptations in Metabolism

Evolutionary adaptations in metabolic networks are fundamental to evolution of microbial growth. Studies on unneeded-protein synthesis indicate reductions in fitness upon nonfunctional protein synthesis, showing that cell growth is limited by constraints acting on cellular protein content. Here, we present a theory for optimal metabolic enzyme activity when cells are selected for maximal growth rate given such growth-limiting biochemical constraints. We show how optimal enzyme levels can be understood to result from an enzyme benefit minus cost optimization. The constraints we consider originate from different biochemical aspects of microbial growth, such as competition for limiting amounts of ribosomes or RNA polymerases, or limitations in available energy. Enzyme benefit is related to its kinetics and its importance for fitness, while enzyme cost expresses to what extent resource consumption reduces fitness through constraint-induced reductions of other enzyme levels. A metabolic fitness landscape is introduced to define the fitness potential of an enzyme. This concept is related to the selection coefficient of the enzyme and can be expressed in terms of its fitness benefit and cost.     

Inclusive Fitness from Multitype Branching Processes

I use multitype branching processes to study genetic models for the evolution of social behaviour, i.e. behaviours that, when acted out, affect the success of the actor’s neighbours. Here, I suppose an individual bearing a mutant copy of a gene influences the reproductive success of a neighbour by altering its own competitive ability. Approximations based on assumptions about the rareness of the mutant allele and the strength of selection allow me to formulate statements concerning the probability of mutant extinction in terms of inclusive fitness. Inclusive fitness is an idea well known to biologists and can be thought of as a sum of an individual’s fitness and the fitness of each of its relatives, weighted by some measure of genetic relatedness. Previous work has led to some confusion surrounding the definition of the inclusive-fitness effect of a mutant allele when individuals carrying that allele experience demographic conditions that fluctuate randomly. In this paper, I emphasise the link between inclusive fitness and the probability of mutant extinction. I recover standard results for populations of constant size, and I show that inclusive fitness can be used to determine the short-term fate of mutants in the face of stochastic demographic fluctuations. Overall, then, I provide a connection between certain inclusive-fitness-based approaches routinely applied in theoretical studies of social evolution.     

Hamilton's rule,inclusive fitness maximization,and the goal of individual behaviour in symmetric two‐player games

Hamilton's original work on inclusive fitness theory assumed additivity of costs and benefits. Recently, it has been argued that an exact version of Hamilton's rule for the spread of a pro‐social allele (rb > c) holds under nonadditive pay‐offs, so long as the cost and benefit terms are defined as partial regression coefficients rather than pay‐off parameters. This article examines whether one of the key components of Hamilton's original theory can be preserved when the rule is generalized to the nonadditive case in this way, namely that evolved organisms will behave as if trying to maximize their inclusive fitness in social encounters.