title: Facultative adjustment of the offspring sex ratio and male attractiveness: a systematic review and meta-analysis.
abstract: Females can benefit from mate choice for male traits (e.g. sexual ornaments or body condition) that reliably signal the effect that mating will have on mean offspring fitness. These male-derived benefits can be due to material and/or genetic effects. The latter include an increase in the attractiveness, hence likely mating success, of sons. Females can potentially enhance any sex-biased benefits of mating with certain males by adjusting the offspring sex ratio depending on their mate's phenotype. One hypothesis is that females should produce mainly sons when mating with more attractive or higher quality males. Here we perform a meta-analysis of the empirical literature that has accumulated to test this hypothesis. The mean effect size was small (r = 0.064-0.095; i.e. explaining <1% of variation in offspring sex ratios) but statistically significant in the predicted direction. It was, however, not robust to correction for an apparent publication bias towards significantly positive results. We also examined the strength of the relationship using different indices of male attractiveness/quality that have been invoked by researchers (ornaments, behavioural displays, female preference scores, body condition, male age, body size, and whether a male is a within-pair or extra-pair mate). Only ornamentation and body size significantly predicted the proportion of sons produced. We obtained similar results regardless of whether we ran a standard random-effects meta-analysis, or a multi-level, Bayesian model that included a correction for phylogenetic non-independence. A moderate proportion of the variance in effect sizes (51.6-56.2%) was due to variation that was not attributable to sampling error (i.e. sample size). Much of this non-sampling error variance was not attributable to phylogenetic effects or high repeatability of effect sizes among species. It was approximately equally attributable to differences (occurring for unknown reasons) in effect sizes among and within studies (25.3, 22.9% of the total variance). There were no significant effects of year of publication or two aspects of study design (experimental/observational or field/laboratory) on reported effect sizes. We discuss various practical reasons and theoretical arguments as to why small effect sizes should be expected, and why there might be relatively high variation among studies. Currently, there are no species where replicated, experimental studies show that mothers adjust the offspring sex ratio in response to a generally preferred male phenotype. Ultimately, we need more experimental studies that test directly whether females produce more sons when mated to relatively more attractive males, and that provide the requisite evidence that their sons have higher mean fitness than their daughters.
authors.affiliation: Division of Evolution, Ecology & Genetics, Research School of Biology, The Australian National University, Daley road, Canberra, Australian Capital Territory, 2601, Australia.|Department of Animal Ecology, Evolutionary Biology Centre, Uppsala University, Norbyvägen 18D, SE-75236, Uppsala, Sweden.|Institute of Evolutionary Biology and Environmental Studies, University of Zürich, Winterthurerstrasse 190, CH-8057, Zürich, Switzerland.
authors.forename: Isobel
authors.initials: I
authors.lastname: Booksmythe
authors.affiliation: Division of Evolution, Ecology & Genetics, Research School of Biology, The Australian National University, Daley road, Canberra, Australian Capital Territory, 2601, Australia.|Department of Animal Ecology, Evolutionary Biology Centre, Uppsala University, Norbyvägen 18D, SE-75236, Uppsala, Sweden.
authors.forename: Brian
authors.initials: B
authors.lastname: Mautz
authors.affiliation: Division of Evolution, Ecology & Genetics, Research School of Biology, The Australian National University, Daley road, Canberra, Australian Capital Territory, 2601, Australia.|Department of Psychology, University of Cambridge, Downing Street, CB2 3EB, Cambridge, U.K.
authors.forename: Jacqueline
authors.initials: J
authors.lastname: Davis
authors.affiliation: Department of Zoology, University of Otago, PO Box 56, Dunedin 9054, New Zealand.|Evolution & Ecology Research Centre, School of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney, New South Wales, 2052, Australia.
authors.forename: Shinichi
authors.identifier: http://orcid.org/0000-0002-7765-5182
authors.initials: S
authors.lastname: Nakagawa
authors.affiliation: Division of Evolution, Ecology & Genetics, Research School of Biology, The Australian National University, Daley road, Canberra, Australian Capital Territory, 2601, Australia.
authors.forename: Michael D
authors.initials: MD
authors.lastname: Jennions
mesh_terms: D000818:Animals; D001499:Bayes Theorem; D005260:Female; D008297:Male; D008954:Models, Biological; D010802:Phylogeny; D012098:Reproduction; D012744:Sex Ratio; D012726:Sexual Behavior, Animal / Q000502:physiology*
keywords: female choice; mate choice; maternal effects; sex allocation; sex ratio adjustment; sexual selection