Sex Differences in Aggression: What Does Evolutionary Theory Predict?
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BEHAVIORAL AND BRAIN SCIENCES (2009) 32, 249–311 Printed in the United States of America doi:10.1017/S0140525X09990951 Does sexual selection explain human sex differences in aggression? John Archer School of Psychology, University of Central Lancashire, Preston PR1 2HE, United Kingdom [email protected] http://www.uclan.ac.uk/scitech/research/rae2008/psychology/ staff_profiles/jarcher.php Abstract: I argue that the magnitude and nature of sex differences in aggression, their development, causation, and variability, can be better explained by sexual selection than by the alternative biosocial version of social role theory. Thus, sex differences in physical aggression increase with the degree of risk, occur early in life, peak in young adulthood, and are likely to be mediated by greater male impulsiveness, and greater female fear of physical danger. Male variability in physical aggression is consistent with an alternative life history perspective, and context-dependent variability with responses to reproductive competition, although some variability follows the internal and external influences of social roles. Other sex differences, in variance in reproductive output, threat displays, size and strength, maturation rates, and mortality and conception rates, all indicate that male aggression is part of a sexually selected adaptive complex. Physical aggression between partners can be explained using different evolutionary principles, arising from the conflicts of interest between males and females entering a reproductive alliance, combined with variability following differences in societal gender roles. In this case, social roles are particularly important since they enable both the relatively equality in physical aggression between partners from Western nations, and the considerable cross-national variability, to be explained. Keywords: aggression; partner violence; sex differences; sexual selection; social role theory 1. Introduction partners in this article, so as to provide a comprehensive explanation of sex differences in aggression. Nevertheless, Darwin (1859/1911; 1871/1901) regarded the greater most of the discussion is devoted to within-sex differences, proneness to physical aggression by men than women as the main concern of sexual selection and social role part of a general mammalian pattern, which can be accounts. In assessing the evidence, I address the issues explained by one aspect of sexual selection, inter-male of ultimate origins, development, mediating variables, and competition. Within the social sciences there is a long tra- sources of individual differences, in relation to predictions dition of explaining sex differences in social behavior, derived from the two theories. I argue that sexual selection including aggression, in human-centered terms, as a con- provides the more complete explanation of the origins of sex sequence of social influences. In this article, I argue that differences in aggression, and that these differences are sexual selection provides a better explanation for human linked to a range of other features indicting the operation sex differences in aggression than the main contemporary of sexual selection in humans. Although reformulations of social science approach, the social role theory. social role theory do encompass some evolved differences, Aggression is typically defined as behavior intended to they are restricted to physical attributes, which form only harm another individual, and its study was originally part of the sexual selection view. restricted to direct physical and verbal forms. Psychologists began systematically documenting human sex differences in these types of aggression in the 1920s, and studies have continued to the present time. The forms of aggression JOHN ARCHER, Professor of Psychology at the Univer- 1 now include indirect (or “relational” ) aggression, which sity of Central Lancashire, Preston, United Kingdom, is Feshbach (1969) and Lagerspetz et al. (1988) found to be the author of more than 100 articles, in a wide range of more typical of girls than boys. They suggested that the journals, in the areas of animal aggression and emotion- sex difference, hitherto characterized as involving “aggres- ality, testosterone and behavior, human sex differences, sion,” was more accurately viewed as involving its form. human aggression, grief and loss, and evolutionary psy- Contemporary studies include both direct and indirect chology. He is also the author of several books, includ- aggression. A separate extensive literature documents sex ing The Behavioural Biology of Aggression (1988), The Nature of Grief (1999), and Sex and Gender (2nd differences (or similarities) in physical aggression between edition, 2002). He is a former President of the Inter- partners, although this is not usually considered in discus- national Society for Research on Aggression (ISRA), sions of sex differences in aggression. I do consider the evol- and is a Fellow of the British Psychological Society. utionary and social forces underlying aggression between # 2009 Cambridge University Press 0140-525X/09 $40.00 249 Archer: Sex differences in aggression 2. Aggression between same-sex individuals distributed, the potential for reproductive competition is less, and monogamy is more likely. These principles also 2.1. Sexual selection as an explanatory framework apply to variation within a species. For example, the for human aggression mating system of the dunnock (Prunella vulgaris) varies 2.1.1. Principles of sexual selection. Sexual selection from polygyny to monogamy to polyandry (Davies & involves the choice of members of one sex by the other, Hartley 1996; Davies & Lundberg 1984), depending on and competition by members of one sex for access to the the food distribution, which affects female range size. other (Darwin 1871/1901). The more competitive and Underlying these associations is the ability of males to aggressive sex is usually the male, particularly in control access to females, or more precisely, the concept mammals. Sexual selection forms a basis for understanding of operational sex ratio (OSR; Emlen & Oring sex differences in aggression in animals, although it is not 1977) – the ratio of sexually active males to fertilizable the only explanation (Ralls 1976; Selander 1972). Trivers females at any given time and place. This will be biased (1972) proposed that the reason why males are typically in the female direction as a consequence of males being the more competitive sex is their lower parental investment, excluded from the breeding group by higher mortality, defined as any contribution by a parent that increases an or being ejected as a result of competition, or maturing offspring’s chances of survival and reproduction, while more slowly (Clutton-Brock & Harvey 1977), and in the reducing the parents’ ability to produce further offspring. male direction if females only gradually become fertile Examples include the energy expended in gamete pro- or available to males (e.g., Smuts 1987b). The OSR is duction (Bateman 1948), and feeding or guarding the essentially an index of the degree of male competition young. The initial greater contribution by the female to for mates, and is associated with greater variance in male the gametes will make desertion to seek access to other than female reproductive success. mates a more beneficial option for males than for females, Pomiankowski and Møller (1995) found greater variabil- providing that one parent can care for the offspring.2 The ity in sexually selected than non-sexually selected traits in sex showing higher parental investment becomes a limiting males than females from a range of species, and they resource, the other sex competing for reproductive access. attributed this to sexual selection producing traits exagger- Competition can take forms other than direct aggression, ated beyond those that were optimal for survival. An for example, sperm competition, or features that aid mate alternative explanation in species where there is some attraction, or securing a dispersed resource (Andersson degree of paternal care is that males of many monogamous 1994, pp. 10–13; Archer 1988, pp. 106–107). species retain traits associated with polygynous mating, When parental care by both sexes is required, inter-male and can therefore be regarded as facultatively polygynous competition will be countered by the male’s greater (Andersson 1994, pp. 157–58; Trivers 1972). The degree paternal investment, as occurs in monogamous birds. to which this occurs depends both on the context (e.g., Where the female can leave the male to brood the eggs the availability of alternative mates) and the individual and care for the young alone, as occurs in polyandrous (e.g., the ability to attract alternative mates). Such individ- wading birds, females will be the more competitive, and ual differences are said to arise from the extent to which the larger and more aggressive, sex (Jenni 1974). This rever- reproductive effort is concentrated on parental or mating sal of the usual sex differences provides crucial support for effort, which is termed a conditional reproductive strategy Trivers’ theory. In mammals, fertilization is internal and the (Gross 1996). It would lead to greater variation in sexually female feeds the developing offspring, so that parental selected traits among males than among females. investment is further female-biased, and polygyny is the usual outcome. In terrestrial mammals, this is typically 2.1.2. Sexual selection applied to human aggression. associated