Partner choice creates competitive altruism in humans

Reciprocal altruism has been the backbone of research on the evolution of altruistic behaviour towards non-kin, but recent research has begun to apply costly signalling theory to this problem. In addition to signalling resources or abilities, public generosity could function as a costly signal of cooperative intent, benefiting altruists in terms of (i) better access to cooperative relationships and (ii) greater cooperation within those relationships. When future interaction partners can choose with whom they wish to interact, this could lead to competition to be more generous than others. Little empirical work has tested for the possible existence of this 'competitive altruism'. Using a cooperative monetary game with and without opportunities for partner choice and signalling cooperative intent, we show here that people actively compete to be more generous than others when they can benefit from being chosen for cooperative partnerships, and the most generous people are correspondingly chosen more often as cooperative partners. We also found evidence for increased scepticism of altruistic signals when the potential reputational benefits for dishonest signalling were high. Thus, this work supports the hypothesis that public generosity can be a signal of cooperative intent, which people sometimes 'fake' when conditions permit it.     

Preferences for symmetry in human faces in two cultures: data from the UK and the Hadza, an isolated group of hunter-gatherers

Many studies show agreement within and between cultures for general judgements of facial attractiveness. Few studies, however, have examined the attractiveness of specific traits and few have examined preferences in hunter-gatherers. The current study examined preferences for symmetry in both the UK and the Hadza, a hunter-gatherer society of Tanzania. We found that symmetry was more attractive than asymmetry across both the cultures and was more strongly preferred by the Hadza than in the UK. The different ecological conditions may play a role in generating this difference. Such variation in preference may be adaptive if it reflects adaptation to local conditions. Symmetry is thought to indicate genetic quality, which may be more important among the Hadza with much higher mortality rates from birth onwards. Hadza men who were more often named as good hunters placed a greater value on symmetry in female faces. These results suggest that high quality Hadza men are more discriminating in their choice of faces. Hadza women had increased preferences for symmetry in men's faces when they were pregnant or nursing, perhaps due to their increased discrimination and sensitivity to foods and disease harmful to a foetus or nursing infant. These results imply that symmetry is an evolutionarily relevant trait and that variation in symmetry preference appears strategic both between cultures and within individuals of a single culture.     

Mate preferences and choices for facial and body hair in heterosexual women and homosexual men: influence of sex,population, homogamy,and imprinting-like effect

Recent research has reported that male body and facial hair influence women's mate preferences. However, it is not clear whether such preferences are typical for women or for individuals who prefer males as sexual partners. Here we explored body and facial hair in preferred and actual partners among men and women who prefer men as sexual partners. Including homosexual individuals provides a unique opportunity to investigate whether evolved mating psychologies are specific to the sex of the individual or sex of the partner. Based on an online survey of 1577 participants from Brazil and the Czech Republic, we found that, on average, homosexual men preferred hairier stimuli than heterosexual women, supporting past findings that homosexual men have strong preferences for masculine traits. Preferences for facial and body hair appear to be influenced less by sex of the preferred partner than sex of the individual, pointing to a possible sex-specific mating psychology. Further, Brazilians preferred bigger beards than Czechs, which was positively associated with the self-reported amount of beardedness in Brazil, suggesting that familiarity effects underpin cross-cultural differences in preferences for facial hair. Moreover, homosexual men preferred a self-similar degree of beardedness, and Czech women preferred a similar degree of beardedness as their fathers had during their childhood. However, these effects were not associated with the level of facial hair in their actual partners; in general, mate preferences and actual mate choices for facial and body hair differed. Thus, individual differences in some self-reported characteristics, cultural factors, and aspects of personal experience may modulate differences in preferences for masculine traits.     

Sexual preferences during artificial menstrual cycles in social groups of rhesus monkeys (Macaca mulatta)

Eight groups of rhesus monkeys each consisting of one male and four ovariectomized females were observed while two of the females were treated with hormones to produce artificial menstrual cycles. These were either synchronized or offset by 7-day increments. Sexually preferred females, defined by the numbers of ejaculations per test, received almost twice as many ejaculations as did non-preferred females during all synchronized and offset cycles and during all cycle phases. However, short-term changes in partner preference occurred when the midcycle phase of non-preferred females coincided with the middle or late progesterone phase of preferred females, suggesting a negative effect of progesterone on behavior during the menstrual cycle. There were highly significant differences between preferred and non-preferred partners for almost all of their sexual and social interactions, and preferred partners showed longer proximity and grooming times as well as higher levels of sexual activity. Partner preferences accounted for more of the behavioral variance between pairs than did female dominance, although males sought the proximity of dominant females independently of their partner preferences. Thus, in a setting uncomplicated by male mate competition, sexual preference by male rhesus monkeys is a robust phenomenon depending on complex interactions between dominance, hormonal status, and the individual behavior of female partners.     

Men''s preferences for women''s profile waist-to-hip ratio in two societies

Women's waist-to-hip ratio (WHR) varies with age, and a lower WHR is associated with a higher estrogen-to-androgen ratio and possibly higher fecundity, at least in some populations. Consequently, it has been argued that selection has favored a universal male preference for a low female WHR. In previous studies using frontal pictures, men in the United States preferred a low WHR of 0.7, but men among Hadza hunter–gatherers and a few other small-scale societies preferred higher ratios. Unlike the actual WHR of women, measured with a tape around the waist and the hips and buttocks, the WHR in frontal pictures excludes the buttocks. Because frontal WHR gives only a partial picture, we used profile views of women to measure men's preferences for the profile WHR. Hadza men preferred a lower profile WHR (more protruding buttocks) than American men. Since Hadza men preferred higher frontal WHR but lower profile WHR, and since both contribute to the actual WHR, these results imply there is less disparity between American and Hadza preferences for the actual WHR of real women. We suggest men's preferences vary with the geographic variation in the shape of women who have wider hips in some populations and more protruding buttocks in others.     

Self-interested partner selection can lead to the emergence of fairness

The theory of biological markets and competitive altruism contends that competitive partner selection is favorable to the selection of prosocial behaviors in social evolution. The current study provides an empirical assessment of this theory based on a laboratory experiment with human subjects using the Ultimatum game. The experimental results show that more generous proposers and more tolerant responders are preferred as partners. This indicates that subjects tend to choose partners in a manner that coincides with their own interests. In competitive partner selection, partner preferences driven by self-interest nevertheless generate an assortative pairing structure that prompts players to behave fairly in the game. The study shows that a free market of partner selection, plus the type of partner preferences driven by self-interests, can facilitate the emergence of fairness in social exchange.     

Populational differences in attractiveness judgements of male and female faces: Comparing British and Jamaican samples

In the UK and Japan, both men and women prefer somewhat feminised opposite-sex faces, especially when choosing a long-term partner. Such faces are perceived as more honest, caring, and sensitive; traits that may be associated with successful male parental investment. By contrast, women prefer less feminised faces for short-term relationships and when they are near ovulation. As genetic quality may be associated with facial masculinity, women may ‘trade-off’ cues between genetic quality and paternal investment in potential partners. No analogous trade-off has been suggested to influence men's preferences, as both attributions of prosociality and potential cues to biological quality are associated with facial femininity in female faces. Ecological and cultural factors may influence the balance of trade-offs leading to populational differences in preferences. We predicted that Jamaican women would prefer more masculine faces than British women do because parasite load is higher in Jamaica, medical care less common (historically and currently), and male parental investment less pronounced. Male preferences, however, were predicted to vary less cross-culturally, as no trade-off has been identified in female facial characteristics. We constructed masculinised and feminised digital male and female face stimuli of three populations (Jamaican, Japanese, and British) and presented them to men and women in Jamaica and in Britain. The results demonstrated that Jamaican women preferred more masculine male faces than their British counterparts did. Jamaican men tended to prefer more masculine female faces than did British men did, but this effect was complicated by an interaction suggesting that more feminised faces were preferred within culture.     

Social Environment Affects Peripheral Oxytocin and Cortisol during Stress Responses in Guinea-Pigs

Numerous studies have demonstrated interactions between oxytocin (OT) secretion, adrenal activity, an animal's social environment, and stress responses. In the present study, we hypothesized that partner preference and pair bonding cause increased peripheral OT, which down‐regulates the adrenal and behavioral response in stress. In addition, we tested whether these interactions depended on sex, the social environment of the individual, or the type of stressor. Experiments were carried out on guinea‐pigs held in sexual‐pairs or stressed by isolation. Among the paired individuals choice tests were carried out to document partner preference. Female preference for a male was expressed by spatial cohesion. Stress responses to abiotic stimuli were examined and compared between isolated and cohabited animals with or without preference. Results show that OT of cohabited animals was significantly higher than during isolation. OT levels were further increased in males preferred by females. Cortisol (CORT) levels were elevated in isolated animals. There were no significant differences between pairs with and without female preference. Behaviorally, partner preferences were expressed by high amounts of tactile contact. The stress‐induced behavioral immobility response after exposure to a noise stressor was significantly reduced in preferred males and to a lesser extent in their female partners. Only females with preferences showed an endocrine stress response. Their levels of OT increased. There was no consistent post‐stressor release of CORT. The data indicate that the social environment of an individual, here expressed as preference or isolation, influences peripheral OT secretion. Behavioral stress responses were similarly affected by social factors without a clear involvement of peripheral CORT or OT.     

The effects of female control of resources on sex-differentiated mate preferences

A number of sex differences in mate preferences have been reported across cultures. Women prefer partners who are older than them whereas men prefer partners who are younger than them. Women have stronger preferences for resource-acquisition characteristics whereas men have stronger preferences for physical attractiveness. Recently, studies have reported shifts in female preferences with increasing female empowerment and associated female attitudes. Other studies, however, report opposite effects of female wealth and income. In this study, we investigated the effects of female control of the resources necessary to raise offspring successfully on mate preferences. We developed measures of resource control at the level of the individual and investigated relationships between these and mate preferences using an internet survey. Resource control was associated with preferences for physical attractiveness over good financial prospects and greater maximum partner age tolerated. Resource control, however, was also associated with tolerance of younger partners. The results implicate the role of constraints on female access to and control of resources in sex-differentiated mate preferences and highlight differences between resource control and wealth.     

Parental age difference and offspring count in humans

Preferences for certain age characteristics of partners are reported across cultures: men prefer mates who are younger and women prefer mates older than themselves. To examine whether these age preferences entail fitness effects for men and women, we investigated the association among age differences between partners and offspring count. On the basis of a sample of approximately 10,000 post-reproductive Swedish men and women who did not change their partner between the birth of their first and last child, we find maximum offspring count in men if their partner is approximately 6 years younger, and in women if their partner is approximately 4 years older. We further find that after separation, on average, both men and women shift to a partner younger than the first, albeit in women the new partner is still older than the female herself. We conclude that the age preference for the partner yields fitness benefits for both men and women and may thus be an evolutionarily acquired trait.     

How Universal Are Preferences for Female Waist-to-Hip Ratios? Evidence from the Hadza of Tanzania

Female waist-to-hip ratio (WHR) has been proposed by evolutionary psychologists to be an important component of human male mate choice, because this trait is thought to provide a reliable cue to a woman's reproductive value. Based largely on work conducted in industrialized societies, the claim has been made that preferences for low WHR are culturally invariant. Presumably, the preferences evolved before the advent of agriculture, making foraging populations the best place to test the hypothesis. This was done with the Hadza of Tanzania, who were shown figures of females that varied by weight and waist-to-hip ratio. Low WHR was not preferred. Hadza men did not consider waist-to-hip ratio when expressing preferences for mates. Instead, they were most interested in the weight of potential partners. Research by others with subjects who practice swidden agriculture also revealed that low WHR was not preferred. The data from the Hadza coupled with the information derived from this horticultural group bring into question whether preferences for low WHR are culturally invariant.     

Male chimpanzees' grooming rates vary by female age, parity, and fertility status

Copulation preferences in our closest living relative, the chimpanzee, suggest that males prefer older females who have had previous offspring. However, this finding is counter to some behavioral models, which predict that chimpanzee males, as promiscuous breeders with minimal costs to mating, should show little or no preference when choosing mating partners (e.g. should mate indiscriminately). To determine if the preferences indicated by copulations appear in other contexts as well as how they interact, we examined how male chimpanzees' grooming patterns varied amongst females. We found that males' preferences were based on interactions among females' fertility status, age, and parity. First, grooming increased with increasing female parity. We further found an effect of the estrous cycle on grooming; when females were at the lowest point of their cycle, males preferentially groomed parous females at peak reproductive age, but during maximal tumescence, males preferred the oldest multiparous females. Nulliparous females received relatively little grooming regardless of age or fertility. Thus, male chimpanzees apparently chose grooming partners based on both female's experience and fertility, possibly indicating a two-pronged social investment strategy. Male selectivity seems to have evolved to effectively distribute costly social resources in a pattern which may increase their overall reproductive success.     

Partner preferences in the context of concurrency: What Himba want in formal and informal partners

Research on human mate preferences that distinguishes between short- and long-term partners has been conducted mainly in industrialized societies, where multiple mating and concurrent partnerships are stigmatized. However, cross-culturally, there is significant variation in the frequency and the level of acceptance of such relationships. Furthermore, the dichotomy between short- and long-term partnerships does not fully describe the diversity in actual extra-pair behavior, which ranges from single sexual encounters to multi-year love affairs. Here we present another comparison, between formal (marital) and informal (non-marital) partners, which we feel better captures this diversity. We assess the traits that men and women prefer in each type of partner among Himba pastoralists, where concurrent partnerships are common and accepted for both sexes. We situate our findings in relation to three main explanations for concurrent partnerships: dual-mating, trading-up and multiple investors. We find some similarities with the existing literature in the traits that are listed as important, including physical attractiveness, wealth and intelligence. Our evidence suggest that Himba men follow a dual strategy, preferring hard-working wives and attractive girlfriends. Women's preferences align most strongly with a multiple investors explanation, most clearly articulated through their preferences for wealthy husbands and generous boyfriends. Limited support is also found for a dual-mating strategy in women. These findings suggest that local cultural norms and ecologies modulate mate preferences in important ways.     

What people prefer and what they think they prefer in short- and long-term partners. The effects of the phase of the menstrual cycle,hormonal contraception,pregnancy, and the marital and the parenthood status on partner preferences

The issue with most studies concerned with mate selection preferences in humans is that they rely on declarations and rational actions of experimental subjects, which are affected by their pre-conceived opinions and prejudices. Moreover, current research suggests that subcortical structures and processes, rather than the neocortex, play the principal role in actual partner choice behaviour. Consequently, we have only limited information on how relevant our current knowledge is in relation to real-life human ethology. To address these issues, we surveyed 2718 women and 4073 men between the ages of 16–50 and compared their declared and observed preferences for various properties in short-term and long-term partners. We found differences between what the subjects declared to prefer and what they preferred in reality: for example, men declared that wealth was the second least desirable property out of eleven in short-term partners, but we observed that in reality, they considered wealth the third most important factor after charisma and sense of humour. Similarly, while women declared that dominance and masculinity were desirable properties in short-term partners, in the observational part of the study, they showed little preference for these traits. Furthermore, we investigated the effects of the phase of the menstrual cycle, hormonal contraception, pregnancy, and partnership and parenthood status on these preferences. We found some support for the good parents hypothesis and no support for the good genes and the immunocompetence handicap hypotheses when observed, rather than declared preferences were considered. We also detected that hormonal contraception, and parenthood and partnership status influenced declared preferences in considerable ways, but had only a small, if any, impact on observed preferences. We suggest interpreting the results of studies reliant on declarations and rational actions of experimental subjects with great caution.     

Fairness versus favoritism in children

Children respond positively to individuals who favor them and also to individuals who are fair. The present studies examine the interaction between these two preferences by presenting children with two distributors who share resources with the child participant and another recipient. Children are asked whom they like better: the distributor who was unfair but favored the child participant or the distributor who was fair and showed no (or reduced) favoritism. In Study 1, we find that when fairness and favoritism are in conflict, children are split on whom they prefer. In Study 2, we find that placing children in a competitive context leads to a stronger preference for the distributor who favored the child participant. In Study 3, we examine whether children's preference for favoritism persists when both distributors gave the child the same number of rewards, but one distributor gave the child participant relatively more than the other recipient. In this situation, we find that children prefer the fair distributor. However, we again find that creating a competitive context reduces children's preference for the fair distributor. Finally we find that in a third-party context, children value fairness over generosity. Taken together, these results show how children balance competing concerns for fairness and favoritism.     

Habitat preferences of ptarmigan hunters in Norway

Hunting for willow ptarmigan (Lagopus lagopus) is a popular recreational activity in Norway, but studies of the hunters are limited. While large game management in Norway is based on sound models and research, management of small game hunters and hunting have not been subject to the same rigour. Compared to the extensive body of knowledge about ptarmigan behaviour and habitat preferences, the knowledge base on ptarmigan hunters is very limited. We surveyed the habitat preferences of 3,056 hunters to identify preferred landscape categories using pictures covering a range of landscape types. We also examined to which extent residence types and forms of hunting were related to habitat preferences. Through factor analysis, we identified three categories of hunting habitat; mountain forest, low alpine and high alpine. Mode of hunting is more important than residence status in terms of preferred habitat. Hunters using dogs preferred mountain forest and low alpine habitats more than hunters without dogs. Hunters without dogs had a higher preference than dog hunters for high alpine habitats. Hunters with mixed modes of hunting have wider habitat preferences. Residence status only affects the perception of the mountain forest habitat. The results may have implications for management as land use and place-based meanings are currently changing in many natural and rural landscapes, and hunting needs to be integrated with other recreational uses.     

Cooperators benefit through reputation-based partner choice in economic games

Explaining unconditional cooperation, such as donations to charities or contributions to public goods, continues to present a problem. One possibility is that cooperation can pay through developing a reputation that makes one more likely to be chosen for a profitable cooperative partnership, a process termed competitive altruism (CA) or reputation-based partner choice. Here, we show, to our knowledge, for the first time, that investing in a cooperative reputation can bring net benefits through access to more cooperative partners. Participants played a public goods game (PGG) followed by an opportunity to select a partner for a second cooperative game. We found that those who gave more in the PGG were more often selected as desired partners and received more in the paired cooperative game. Reputational competition was even stronger when it was possible for participants to receive a higher payoff from partner choice. The benefits of being selected by a more cooperative partner outweighed the costs of cooperation in the reputation building phase. CA therefore provides an alternative to indirect reciprocity as an explanation for reputation-building behaviour. Furthermore, while indirect reciprocity depends upon individuals giving preference to those of good standing, CA can explain unconditional cooperation.     

Free mutual mate preferences in house mice affect reproductive success and offspring performance

When social constraints on the expression of mate preferences are absent, variation in offspring viability is predicted to favour females and males that display mate preferences. Earlier studies showed that female and male house mice, Mus domesticus, tested individually and mated with preferred (P) partners had higher reproductive success and better progeny performance than individuals mated with nonpreferred (NP) partners. Here we tested the effects of mutual mate preferences on reproductive success, offspring viability and performance. We conducted mate preference tests and created four types of reproductive pairings. One involved females and males that preferred each other (P-P); the second type had females that preferred the male but the male did not prefer the female (P-NP); the third had females that did not prefer the male but the male did prefer the female (NP-P). The last set consisted of females and males that did not prefer each other (NP-NP). We measured components of fitness for breeders (reproductive success) and offspring viability (birth-to-weaning viability and weight variation) as well as measures of offspring performance. There were no statistical differences in reproductive success of breeders or offspring viability and quality (weight variation) among the four types of pairings. There were, however, consistent differences between P-P versus NP-NP matings. The number of pups weaned, time to first litter, birth-to-weaning viability, pup body weight at birth and weaning, and the growth rates for pups of both sexes were consistently greater for progeny from P-P matings than NP-NP matings. Significant differences occurred among the four mating types in dominance of sons during aggression trials, nest construction and predator avoidance. Progeny from P-P matings displayed behaviour associated with higher fitness more often than progeny from NP-NP matings. These data show that breeders produce more highly competent progeny, most likely to survive, when social constraints on the expression of mate preferences in both sexes are relaxed.Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Body Weight and Rearing Conditions of Males, Female Choice and Paternities in a Small Mammal, Cavia aperea

Patterns of mate choice may be important determinants of a species' social organisation and mating system. At least two different aspects of female mate choice can be distinguished: choice of a social partner and choice of the genetic father of the offspring. Different characteristics of males can qualify them for these two roles. Although social and reproductive partners have been shown to differ in many species, social association times are often used in laboratory choice tests to infer reproductive preferences. The traits for which females may choose partners are diverse. Body size can correlate with the male's strength in defending resources or other abilities benefiting the female and her offspring. In species living in social groups, social skills learned from group members during infancy can be important for later reproductive success. In this laboratory study, we conducted choice tests with wild cavies, Cavia aperea , a harem-living species of South American rodents, to determine social preferences of females towards two simultaneously available males. For offspring sired during these tests, paternities were determined by microsatellite DNA profiling. Males used in the tests differed in body weight and in rearing conditions: Half of the males had been reared in the presence or absence of their father, respectively. Male rearing conditions had no effect on either female social preferences or paternities. Females significantly preferred heavier males as social partners. In five of six tests, the heavier male also sired the offspring. Sires were in most cases but not consistently socially preferred. Heavier males may be preferable as social partners because they are better able to provide females with resources or have more experience in paternal care or predator avoidance as weight correlates with age. When choosing reproductive partners, females may prefer other male traits and the distribution of paternities may also be influenced by sperm competition.     

Cryptic preference for MHC-dissimilar females in male red junglefowl,Gallus gallus

An increasing number of studies test the idea that females increase offspring fitness by biasing fertilization in favour of genetically compatible partners; however, few have investigated or controlled for corresponding preferences in males. Here, we experimentally test whether male red junglefowl, Gallus gallus, prefer genetically compatible females, measured by similarity at the major histocompatibility complex (MHC), a key gene complex in vertebrate immune function. Theory predicts that because some degree of MHC heterozygosity favours viability, individuals should prefer partners that carry MHC alleles different from their own. While male fowl showed no preference when simultaneously presented with an MHC-similar and an MHC-dissimilar female, they showed a ‘cryptic’ preference, by allocating more sperm to the most MHC-dissimilar of two sequentially presented females. These results provide the first experimental evidence that males might respond to the MHC similarity of a female through differential ejaculate expenditure. By revealing that cryptic male behaviours may bias fertilization success in favour of genetically compatible partners, this study demonstrates the need to experimentally disentangle male and female effects when studying preferences for genetically compatible partners.