You are on page 1of 7

PSYCHOLOGICAL SCIENCE

Research Report
HUMAN FLUCTUATING ASYMMETRY AND SEXUAL BEHAVIOR
Randy Thornhili and Steven W. Gangestad
University of New Mexico Abstract—This report presents evidence that sexual selection may favor developmental stability (i.e., the absence of fluctuating asymmetry) in humans. Subtle, heritable asymmetries in seven nonfacial human body traits correlated negatively with number of self-reported, lifetime sex partners and correlated positively with self-reported age at first copulation in a college student sample. These relationships remained statistically significant when age. marital status, body height, ethnicity, physical anomalies associated with early prenatal development, and physical attractiveness were statistically controlled. The strength of the relationships did not differ significantly as a function of sex. It is unlikely that the relationships are generated by false self-reporting of sexual experience. Fluctuating asymmetry (FA) is deviation from symmetry in morphological traits for which signed differences between the left and right sides have a mean of zero and are near-normally distributed within a population (Van Valen, 1962). A long-standing interest in FA (Lerner, 1954; Ludwig, 1932) derives from the notion that perfect bilateral symmetry in such traits is the optimal and asymmetry results from imprecise expression of developmental design. FA increases with exposure to environmental perturbations such as parasites (Bailit, Workman, Niswander, & MacLean, 1970; M0Uer, 1992b; Polak, in press), pollutants (Parsons, 1990), extreme temperatures (Parsons, 1990), and other physical conditions (Parsons, 1990) during ontogeny, and with exposure to genetic perturbations such as deleterious recessives (Lerner, 1954; Parsons, 1990), homozygosity (Lerner, 1954; Mitton, in press), and episodes of directional selecAddress correspondence to Randy Thornhill, Department of Biology, The University of New Mexico, Albuquerque, NM 871311091. VOL. 5, NO. 5, SEPTEMBER 1994 tion (Clark & McKenzie, 1987). Within populations, FA can vary considerably across individual organisms. One explanation of this fact is that individuals differ in the extent to which they are able to resist developmental perturbations. In a range of species, individuals' FA negatively predicts their fecundity, growth rate, and survival (Mitton & Grant, 1984; Palmer & Strobeck, 1986; Parsons, 1990; Thornhili, 1992; Thornhili & Sauer, 1992). Recently, behavioral ecologists have studied the role of FA in sexual selection, that is, in the individual variation in offspring production that is the consequence of individual differences in traits that affect access to number and quality of mates. In sexual selection, differential access of individuals to mates arises from mate choice or direct competition for mates (e.g., searching or fighting). Three main models of sexual selection currently exist (Cronin, 1991), one nonfunctional and two functional models. According to the arbitrary, or aesthetic, model, mate choice of one sex favors certain features in the other sex solely because they are attractive and not because they indicate heritable viability that enhances the survival of offspring. The good-genes view of sexual selection proposes that sexual selection involving both mate choice and competition favors features that correlate with the presence of genes that enhance offspring viability. The good-provider model posits that traits favored by sexual selection in species with male-provided material benefits to a mate or offspring, such as H. sapiens, advertise the bearer's ability to provide, independent of his heritable viability. Because asymmetry in normally bilaterally symmetrical traits apparently reflects the inability of the developing individual to cope with environmental and genetic perturbations during development, it has been argued that evidence of sexual selection favoring body symmetry lends support to the functional hypotheses of sexual selection (i.e., the goodgenes and good-provider models; M0ller, 1990; M0ller & Pomiankowski, 1993; Thornhili, 1992; Watson & Thornhili. 1994). In a range of nonhuman species, males who exhibit low FA tend to obtain more mates than do more asymmetrical males, through either advantage in malemale competition or female choice (M0ller, 1990; M0ller & Pomiankowski, 1993; Thornhili, 1992; Watson & Thornhili, 1994) (including female preference for symmetry itself; M0ller, 1992a). According to A. M0ller and R. Thornhill's unpublished meta-anaiysis of 26 published studies, the overall heritabOity of FA is .25 {SE = 0.09), which differs significantly from zero {p = .007) in a onesample t test with log (x -t- 1) transformed heritabilities. This heritability of FA is consistent with good-genes hypotheses of sexual selection. It is unclear, however, whether heritable FA is due to genetic variation in the machinery of development per se or to genetic variation in resistance to disease or some other aspect of viability that then affects FA variation among individuals. Recent evidence suggests that FA may play a role in human sexual selection as well. In one study, nonfacial body FA of male university students correlated significantly and negatively with ratings of their facial attractiveness (Gangestad, Thornhili, & Yeo, 1994). In another study, opposite-sex attractiveness ratings of facial photographs of men and women correlated positively with the measured bilateral symmetry of the faces of each sex (Grammer & Thornhili, in press). Human body symmetry is heritable (Livshits & Kobyliansky, 1991), suggesting that sexual selection acting on men may favor individuals with heritable propensities for body symmetry. In the present study, we examined the relations of FA with self-reported number of lifetime copulatory partners and age at first sexual intercourse within a U.S. student population in an attempt to determine whether FA affects human 297

Copyright © 1994 American Psychological Society

62 women) who participated in return for credit toward their grade or course research requirement.65 16. 1989..64 . Waldrop. NO.21 SD 5. p < . p < .22 5.07-.83 2. 1991) for each subject. 4% btack. Livshits & Kobyiiansky. ear breadth. and 3% Native American (1 subject did not report ethnicity).5] (Palmer & Strobeck. 5% Oriental. FAI is expressed as z scores of residuals (removing age and agesquared) times 100. & Shetterly. each adjusted for the effects of age and age-squared. The interrater reliability for FAI was modest (r = . and data for men are at the bottom (partial r = .PSYCHOLOGICAL SCIENCE Fluctuating Asymmetry and Sexual Behavior sured 30% of the subjects.47 . & Bell. data for women are at the top (partial r = . & Shetterly. 14% married. 1968) was used in this study as a covariate. and a composite of them (Waldrop.36. Fifty-nine percent were Caucasian.11-.97 6. • ^ 1 * • • 1 20 5 :: • 10 0 •• • -200 -100 0 100 200 300 Fluctuating asymmetry index Fig. low ears) result from developmental disruption during the first trimester of gestation.08 10. Waldrop. VOL.75 16. Halverson. wrist breadth. and 5% divorced. elbow breadth. ankle breadth.51 Men Age Lifetime number of sex partners Age at first sex Fluctuating asymmetry index 24. 1.92 . hand breadth. p < . The authors mea298 • •• Women 5 2 ^ o i 15 Ifei c • a. !986). Subjects signed a consent form after being introduced to the general nature ofthe project and the methodology used. Pederson.10 . wide eyes. 1989. Individual trait asymmetries were then summed to yield a composite fluctuating asymmetry index (FAI. The FAs of individual traits were calculated by the absolute difference between left (L) and right (R) sizes divided by the mean size of the left and dght sides for the subject: individual trait FA = \R . and ear length). 23 METHOD Subjects were 122 undergraduate students (60 men.01). Each subject privately filled out a questionnaire that asked his or her age.g. Table L Descriptive statistics of the sample Variable Age Lifetime number of sex partners Age at first sex Fluctuating asymmetry index Mean Women 23.02). correlation across measures by two researchers on 47 subjects). These traits were chosen because they show FA that can be measured reliably and has moderate heritability (Livshits & Kobyiiansky. & Bell. the left and right sides were measured independently to the nearest 0. 5. Waldrop.1 cm using steel calipers. o 1 10 5 0 • # 40 • Men 1 30 r S. 81% were single. Pederson.L\t{iR + L) x . 1968) was also assessed. three assistants measured the others. SEPTEMBER 1994 . 29% Hispanic. all rights of the subjects were protected. A set of minor physical anomalies (MPAs..75 5. Plot of the fluctuating asymmetry index (FAI) and lifetime number of sex partners.63.46 mating pattems that may directly relate to the component of sexual selection involving mate number.27 9.80 2. 1991).32. Subjects were assessed in groups of 1 to 4. For each of seven traits (feet breadth.001. The Une is the least-squares linear regression line.. 5. MPAs (e. Halverson.08 N 62 60 60 62 Range 17-53 0-20 12-28 . All probabilities reported here are twoUiled.08 60 59 59 60 18-42 1-45 12-25 .

We included only subjects who reported themselves to be heterosexual. Additionally.22 -.32. 1981). 1966) indicated a single Facial Attractiveness factor within each sample.23 < < < < . Gangestad height. the correlation between FAI and lifetime number of partners was significant and negative for both men ( .. exclusion of any subset of the additional potential confounding variables did not critically affect the results. NO. and marital status. The tests do not assume any particular distributional characteristics of variables.no -0. n = 60. ethnicity.44).). men: r = + .004 -0.0001 n. we regressed lifetime number of partners on FAI with the effects of sex. Error terms for F statistics have from 115 (at Step 1) to 93 (at Step 7) df.0003 and .93 < .045 0.37 1 .78 and . 5. /[94] = -4. 1). FAI significantly predicted lifetime number of partners (beta = -0. Parametric test statistics are robust with regard to deviations from normality so long as sample size is not small and deviations from normality are not extreme (Hays. ethnicity..02. For the two correlations.000 random pairings of variables were created. 2 homosexual and 2 bisexual individuals were excluded (see Table 1). the p-value estimates from the randomization tests were within .25. p = . VOL. the randomization test estimated a more extreme value than that estimated from a standard t table).10.00 3.64 0.02) and FAI (women: r = + . df = number of variables entered at that step.10 22. the questionnaire inquired about the subject's sexual history. respectively (in the former case.33. Because number of sexual partners across a lifetime can be expected to increase with age. p < . Consistent with previous research (Gangestad RESULTS Does FA in humans affect lifetime number of mates in men or in women? The answer is yes. Ethnicity and marital status are nominal variables that were dummy-coded for these analyses.01. ethnicity. For each correlation.21 Age-squared 1 . female = 0) 1 .p = . Scree tests (Cattell. Using the full sample of men and women. (For all regression analyses reported here. To assess the appropriateness of parametric tests for assessing the significance of the primary relationships in this research. partial r = .PSYCHOLOGICAL SCIENCE Randy Thornhill and Steven W. With the effects of age and age-squared statistically controlled.116 anomalies 10 . partial r = correlation between the variable entered at that step and the criterion variable with ail variables entered prior to entry of that variable partialed out.86 for ratings of men and women. age-squared. FAI and lifetime number of sex partners were somewhat positively skewed.22. height.76.35 5. Hierarchical multiple regression analyses of regression of lifetime number of sex partners on the fluctuating asymmetry index iFAI) Step 1: 2: 3: 4: df change Beta Partial r . beta = standardized regression coefficient for the variable entered at that step.37. n. body height.023 -0. Factor scores estimated by weighted composites of the ratings served as measures of facial attractiveness (Cronbach's alphas = . These results justify the use of parametric tests in this study. men: r = +.16 7.06 < . but rather estimate probability levels of sample statistics under a null hypothesis from observed distributions.p < .0008 of the p value derived from use of a standard t table. Long.44 -. we partialed age and age-squared out of all correlations involving number of sex partners. and the interactions of all variables with sex removed (Table 2).192 5: Interactions of age. 1974) controlled for a number of additional potential confounds. Exclusion of any variables entered at Steps 4 and 5 has little effect on results at Steps 6 and 7. p < .01. (women: r = +.) The attractiveness of facial photos of subjects were rated by eight raters (four men. & Sokal. minor physical 8 .36.s. Note.23 Steps prior to entry of fluctuating asymmetry Sex (male = 1.24 Age 1 . Hierarchical regression analyses (Neter & Wasserman. it is not possible to report a specific beta or partial r. and minor physical anomalies with sex Steps involving fluctuating asymmetry 1 .88 Height. age-squared. marital status. p = .03 2.060 0. MPAs.09 . One subject had never had sexual intercourse. for both sexes. FAI and sex did not interact to predict number of partners it[93] = -0. marital status. 1986) on the zero-order correlations of FAI with lifetime number of partners and age at first sex. 5. 3 did not respond to the latter.005 6: FAI 7: Interaction of FAI with sex -. In all cases in which multiple degrees of freedom are associated with variables entered. we performed randomization tests (Smouse. Ratings were factor-analyzed separately for each sex. F = p statistic for the increase in multiple correlation associated with entry of the variable(s) at that step. Age correlated positively with lifetime number of partners Table 2.71 3. 5. SEPTEMBER 1994 299 .s. four women) on a scale from 1 (extremely unattractive) to 10 (extremely attractive).24 . see Fig. respectively).96. age.06).01 . marital status.02 . ethnicity.31. including the variables of interest here: "What age were you when you first had sexual intercourse?" and "With how many partners have you had sexual intercourse in your lifetime?" Two subjects did not answer the former question. sexual orientation.0001. p < . n = 59) and women (-.

In such species. SEPTEMBER 1994 .23. If symmetry correlates with maturation rate (see Mitton & Grant.s. 10.08. women: r = + . it is reasonable to conclude that sexual selection favors individuals of high developmental competence or health and thus probably individuals of high viability.s. data for women are at the top (partial r = +.s.30) and no interaction between FAI and sex (r[90] = -0. facial attractiveness does not fuUy mediate the association between FAI and number of partners. partial r = +. Regression analyses controlling for potential confounding variables (see above) as well as physical attractiveness and its interaction with sex revealed a significant predictive effect of FAI on lifetime number of partners (beta = -0. women: . then.19.38. p < . (See Table 3.42. Indeed.16. 2 3 .29.s. the correlation did not reach statistical significance (+ .). 1993. In our sample. n. p < . FAI correlated significantly and positively with men's age at first sexual intercourse (+ . agesquared. age correlated with age at first sex (men: r = +. Multiple regression analyses controlling for potential confounding variables (see above) as well as age of flrst intercourse and its interaction with sex yielded a significant predictive effect of FAI on number of partners (beta = -0. We assumed that it is not highly variable in the students studied. f[91] = 2. p < . n. height. n. p < . p < . 1994). n = 60). n. marital status. 300 Women 25 ID 25 Men 10 -200 -100 0 100 200 300 Fluctuating asymmetry index Fig.96.PSYCHOLOGICAL SCIENCE Fluctuating Asymmetry and Sexual Behavior et al.42 for men ip < .. With age and age-squared partialed out.0001. 1994). the correlation was small and nonsignificant (. f[94] = 4.82. 1993. Hence. age. Thornhill.24.15).. Watson & ThomhiU.47) and no interactive effect of sex and FAI (/[86] = -0.. n = 59). DISCUSSION This report provides evidence that lifetime number of sexual partners in both sexes and age of first sex in men depend signiflcantly on an individual's developmental stability as measured by FA.09. Thornhill & Sauer. it is difficult to imagine that false reporting would correlate positively with body symmetry. and data for men are at the bottom (partial r = + . p < .s. NO.005. 5. suggest that humans be added to the growing list of species in which FA plays a role in sexual selection (for reviews.60).). p < . n .) Age at flrst sex correlated negatively with lifetime number of partners (men: . 2). 1984. and the interactions of all variables with sex yielded a significant predictive effect of FAI on age at first sexual intercourse (beta = 0.23. The relations of FAI with age at first intercourse did not rehably differ across the sexes (f[93] = 0. We did not assess the socioeconomic status of our subjects.).93. p < .s. more research will be necessary to determine if human sexual VOL. It seems unlikely that false selfreporting of number of partners or age of first sex could account for the relationship reported. the association between FAI and number of partners is not fully mediated by timing of flrst intercourse. FAI is expressed as z scores of residuals (removing age and age-squared) times 100. partial r = .005) and +.27. 2. in males of certain nonhuman species. M0ller & Pomiankowski.).0001). providing partial control.005). Watson & Thornhill. Although some level of false reporting is expected to occur.16.0001. p = .. FAI and age of flrst intercourse correlated + .40). The line is the least-squares linear regression line.05..) (Fig. n.. both viability and success in mate competition are correlated with body symmetry (Mitton & Grant.35. MPAs. The results. see M0ller & Pomiankowski. n = 56). p < . Clearly.003. ethnicity./[87] = -4. n. 1992. For women. Plot of the fluctuating asymmetry index (FAI) and age of flrst sexual intercourse. 5. For women. men's facial attractiveness correlated significantly with FAI with age and age-squared partialed out ( .s. 4 7 . 1984). each adjusted for the effects of age and age-squared. the association between number of partners and FAI may be partly mediated by maturation rate. 10 for women (n. partial r = . The relations between socioeconomic status and FAI are unknown for humans in general and for our subject population...0005. Hierarchical regression analyses controlling for the effects of sex. However. 1994). 1992.45.

. Hierarchical multiple regression analyses of regression of age at first sexual intercourse on the fluctuating asymmetry index (FAI) R' Step 1: 2: 3: 4: df change Beta Partial r -. 89.P. de Lisio. 65. Nature. and minor physical anomalies with sex 6: FAI 7: Interaction of FAI with sex Steps involving fluctuating asymmetry 1 . & Sokal.s. Multiple regression and correlation extensions 301 .. (1974).20 -. The negative relationship between number of partners and FAI in women is perhaps the most intriguing result. age-squared. & Kobyiiansky.. T. REFERENCES Bailit. 63.L. SEPTEMBER 1994 sexual selection in additional human populations and in samples of older adults. (1990). Roots for help with measurements and logistics.. 391. Human Biology. (1992a). Polak. The evolution of desire: Strategies of human mating. M.L. marital status. Ethnicity and marital status are nominal variables that were dummy-coded for these analyses. H. Cronin. J. Thornhill.07 3.008 -0.P. & Wasserman. Fluctuating asymmetry and sexual selection. see ThomhiU & Gangestad. (in press). R.R. analysis. A.M.B. and hence may not be cues directly responsible for the associations we observed. Homewood. 479-499. E. The ant and the peacock: Altruism and sexual selection from Darwin to today. Hays. More research is needed. Symons.2i .004 0. 267-279.. Genetica. Annual Review of Ecology & Systematic!. minor physical 8 . 42. selection is usually driven by good genes or good provisions.T. Body symmetry and facial symmetry may correlate. & MacLean. (1994). 245-276. Animal Behaviour.. P. (1932).0001 n. ethnicity. Smouse. Facial attractiveness..09 Age 1 . I. df = number of variables entered at that step. M011er.05 < . & Yeo.C. partial r = correlation between the variable entered at that step and the criterion variable with all variables entered prior to entry of that variable partialed out. R. IL: Irwin. The asymmetries we measured are subtle. 325. 238-240. G. Fluctuating asymmetry: Measurement. (in press)..04 1 . (1986).. Pawlak. (in press). Fluctuating asymmetry: An epigenetic measure of stress.40 18. Berlin: Springer. Genetic a. cannot be readily observed. (1966). Rinehan and Winston. The scree test for the number of factors. & Strobeck. to examine the role of symmetry in VOL. Human Biology. M0ller. Gangestad. and. I. and E. Parsons. 1979). ethnicity.B. The heritability of FA in huniians suggests the possibility that human sexual selection may be driven. for women favored by men as mates should not be assumed to convert the advantage into a high number of sexual partners (Buss. R. as weil. J. Genetica.19 0. Palmer. Exclusion of any variables entered at Steps 4 and 5 has little effect on results at Steps 6 and 7. Multivariate Behavioral Research. C.14 1. Workman. facial symmetry would mark overall developmental stability.67 < . Ludwig. Buss. Developmental stability of insecticide resistance phenotypes in the blowfly.101 0. female = 0) 1 . W. P. 1185-1187. developmental stability andfluctuatingasymmetry. Edinburgh: Oliver and Boyd. (1994). Parasites increase fluctuating asymmetry of male Drosophila nigrospiracula: Implications for sexual selection. J. 5.R.86 n. M011er. It seems unlikely to be due to male choice alone.J. A. marital status. (1986). Cosmides and R.177 anomalies 10 .35 1 .42 0. We suggest that men who evidence low FA may manifest behaviors or physical features other than body symmetry that are advantaged in male-male competition or females' choice of partners (for full discussion.005 < .A. Cattell.PSYCHOLOGICAL SCIENCE Randy Thornhill and Steven W.94 4. In all cases in which multiple degrees of freedom are associated with variables entered. Genetic homeostasis. Acknowledgments—We thank T. in part. J. Grammer. by genetic variation affecting offspring developmental quality and hence survival. New York: Holt. L. W. Dental asymmetry as an indicator of genetic and environmental con. Note. growth rate and developmental homeostasis. M.s. A. W. it is not possible to report a specific beta or partial r.07 Steps prior to entry of fluctuating asymmetry Sex (male = 1. pattem. 5.A. Lerner. C. New York: Basic Books. Mitton. A. Future research may identify the role of symmetry in affecting women's number of sexual partners. Parasites differentially increase the degree of fluctuating asymmetry in secondary sexual characters. Mitton. 131-145. K.. however. (1993). Enzyme heterozygosity. R. F = F statistic for the increase in multiple correlation associated with entry of the variable(sj at that step. Human (Homo sapiens) facial attractiveness and sexual selection: The role of symmetry and averageness. Neter.041 0. Females prefer large and symmetrical ornaments. D. D. (1991). (1981).20 Age-squared -1.116 5: Interactions of age. Journal of Comparative Psychology. (1984). & Grant. Ethology & Sociobiology.P. 73-«5. & McKenzie. New York: Cambridge tJniversity Press. Annual Review of Ecology &. A. Gangestad Table 3. 15.05 < . ditions in human populations. M01ler. Livshits. P. 1994.. J. beta = standardized regression coefficient for the variable entered at that step. (1991). J45-346. S. 5. Statisiics. Clark.L. 15. J.09 . J. (1987). 441^166. Biological Review. & Thornhill. G. & Pomiankowski. R. metaboiism and developmental stability. 626-638. A. 17. A. Fluctuating asymmetry as a possible measure of developmental homeostasis in humans: A review. (1992b).M. Nature. height. Dos rechis-links problem in tierreich und beim menschen.94 4.. 1993).P. NO. a result of canalizing natural selectioti. Applied linear statistical models. 40. < . if so. (1954). Trivers provided useful suggestions on the manuscript.D. 691-699. (1990). Niswander. 357. (1970). Associations among protein heterozygosity. Journal of Evolutionary Biology. Systematic!. Error terms for F statistics have from 115 (at Step 1) to 93 (at Step 7) df.C. Eggert. Long.B.033 Height.. Ingram. Fluctuating asymmetry in male sexual ornaments may reliably reveal male quality.

There is a short.A. (1989). M. select audience of academic. Human Nature. Department of Psychology. Minor physical anomalies and behavior in preschool children..F. 391-400. Suite 1100 Washington. 35.Lee Herring. (1993). ThomhiU. R.F. Trends in Ecology & Evolution. & Bell. Child Development. SEPTEMBER 1994 . NW.. Evolution. Editor 1010 Vermont Ave. 202-783-2077 • Fax 202-783-20S3 302 VOL. Readers include a large. The evolution of human sexuality.. K. & Gangestad. Unpublished manuscript.. DC 20005-4907 Here's why you should . Waldrop. Fluctuating asymmetry and sexual selection. • Fax: 202-783-2083 (ATTN: Observer) • Emat'f (Bitnet or Intemet. Suite 1 iOO • Washington. Include PO# with order. 43.. & Shetteriy.W. (1962). (1994). Systematic Zoology. Manual for assessing minor physical anomalies. 125-142. D. & Sauer. & Thornhill. 255-264.W. R. Human facial beauty: Averageness. ThomhiU. NO. 4. (RECEIVED II2AI9A. (1992). or research position openings? Advertise in the American Psychological Society's APS Observer Employment Bulletin Three easy ways to place your ad in the bulletin . _ ? employment line-ads are $5.. ^ ^ ^^ ^^ * The Observer has the most competitive ad rates. Symons. Pederson. Halverson. 5. 21-25. Watson. applied. Waldrop. and research psychologists distributed across all psychology's scientific subdiscipiines. R. S. APS • 1010 Vermont Ave. (1992). symmetry and parasite resistance. Oxford: Oxford University Press. Van Valen.P. 9. NW. There are no typesetting charges for display ads. DC 20005-4907 • Tel. C. Readers easily find ads by referring to the one-of-a-i<ind |ob subject index In each issue.. 16.PSYCHOLOGICAL SCIENCE Fluctuating Asymmetry and Sexual Behavior of the mantel test of matrix correspondence.Q. L. A study of fluctuating asymmetry... ACCEPTED 3/24/94) AMERICAN PSYCHOLOGlCAl. 867879. K. applied. 627-632. Athens. 5. ThomhiU. P. Animal Behaviour. Want the best candidates for your academic. three-week iead lime to get your ad published. 44. R. respectivelviLHERRING@APS LHERRiNG@BITNiC.50 per line (34 characters fit on a line and there is a 6I rates and a publication calendar are available on reqtiest. F. R..EDUCOM. (1979). M. University of Georgia. Panorpajaponica. (1968). Fluctuating asymmetry and the mating system of the Japanese scorpionfly...F. 237269.. Genetic sire effects on the fighting ability of sons and daughters and mating success of sons in a scorpionfly.EDU • Postal malh ^^n?^^^^"^^^. 39. Animal Behaviour.