Assortative Mating in Animals

Yuexin Jiang, Daniel I. Bolnick and Mark Kirkpatrick
The American Naturalist
Vol. 181, No. 6 (June 2013), pp. E125-E138
DOI: 10.1086/670160
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Page Count: 14
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Assortative Mating in Animals
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AbstractAssortative mating occurs when there is a correlation (positive or negative) between male and female phenotypes or genotypes across mated pairs. To determine the typical strength and direction of assortative mating in animals, we carried out a meta-analysis of published measures of assortative mating for a variety of phenotypic and genotypic traits in a diverse set of animal taxa. We focused on the strength of assortment within populations, excluding reproductively isolated populations and species. We collected 1,116 published correlations between mated pairs from 254 species (360 unique species-trait combinations) in five phyla. The mean correlation between mates was 0.28, showing an overall tendency toward positive assortative mating within populations. Although 19% of the correlations were negative, simulations suggest that these could represent type I error and that negative assortative mating may be rare. We also find significant differences in the strength of assortment among major taxonomic groups and among trait categories. We discuss various possible reasons for the evolution of assortative mating and its implications for speciation.

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This item contains 106 references.

Literature Cited
  • ['Adams, D. C. 2008. Phylogenetic meta-analysis. Evolution 62:567–572.']
  • ['Arak, A. 1983. Male-male competition and mate choice in anuran amphibians. Pages 181–210 in P. Bateson, ed. Mate choice. Cambridge University Press, Cambridge.']
  • ['Arnqvist, G. 2011. Assortative mating by fitness and sexually antagonistic genetic variation. Evolution 65:2111–2116.']
  • ['Arnqvist, G., L. Rowe, J. J. Krupa, and A. Sih. 1996. Assortative mating by size: a meta-analysis of mating patterns in water striders. Evolutionary Ecology 10:265–284.']
  • ['Bank, C., J. Hermisson, and M. Kirkpatrick. 2011. Can reinforcement complete speciation? Evolution 66:229–239.']
  • ['Bastos, R. P., and C. F. B. Haddad. 1996. Breeding activity of the Neotropical treefrog Hyla elegans (Anura, Hylidae). Journal of Herpetology 30:355–360.']
  • ['Bearhop, S., W. Fiedler, R. W. Furness, S. C. Votier, S. Waldron, J. Newton, G. J. Bowen, P. Berthold, and K. Farnsworth. 2005. Assortative mating as a mechanism for rapid evolution of a migratory divide. Science 310:502–504.']
  • ['Beltran, S., F. Cézilly, and J. Boissier. 2008. Genetic dissimilarity between mates, but not male heterozygosity, influences divorce in schistosomes. PLoS ONE 3:e3328.']
  • ['Bernstein, R., and S. Bernstein. 1999. Assortative mating in soldier beetles (Cantharidae: Chaulignathus): test of the mate-choice hypothesis. Western North American Naturalist 59:188–192.']
  • ['———. 2003. Assortative mating in insects: sexual selection vs. sampling artifact. Western North American Naturalist 63:374–381.']
  • ['Birkhead, T., and K. Clarkson. 1980. Mate selection and precopulatory guarding in Gammarus pulex. Zeitschrift für Tierpsychologie 52:365–380.']
  • ['Bollache, L., and F. Cézilly. 2004. Sexual selection on male body size and assortative pairing in Gammarus pulex (Crustacea: Amphipoda): field surveys and laboratory experiments. Journal of Zoology 264:135–141.']
  • ['Bollache, L., G. Gambade, and F. Cézilly. 2000. The influence of micro-habitat segregation on size assortative pairing in Gammarus pulex (L.) (Crustacea, Amphipoda). Archiv für Hydrobiologie 147:547–558.']
  • ['Bolnick, D. I., and M. Kirkpatrick. 2012. The relationship between intraspecific assortative mating and reproductive isolation between divergent populations. Current Zoology 58:484–492.']
  • ['Bonneaud, C., O. Chastel, P. Federici, H. Westerdahl, and G. Sorci. 2006. Complex Mhc-based mate choice in a wild passerine. Proceedings of the Royal Society B: Biological Sciences 273:1111–1116.']
  • ['Borenstein, M., L. V. Hedges, J. P. T. Higgins, and H. R. Rothstein. 2009. Introduction to meta-analysis. Wiley, Chichester.']
  • ['Bürger, R., K. A. Schneider, and M. Willensdorfer. 2006. The conditions for speciation through intraspecific competition. Evolution 60:2185–2206.']
  • ['Burley, N. 1983. The meaning of assortative mating. Ethology and Sociobiology 4:191–203.']
  • ['Cézilly, F. 2004. Assortative mating. Pages 876–881 in M. Bekoff, ed. Encyclopedia of animal behavior. Greenwood, Westport, CT.']
  • ['Cézilly, F., and A. R. Johnson. 1995. Re-mating between and within breeding seasons in the greater flamingo Phoenicopterus ruber roseus. Ibis 137:543–546.']
  • ['Chamberlain, N. L., R. I. Hill, D. D. Kapan, L. E. Gilbert, and M. R. Kronforst. 2009. Polymorphic butterfly reveals the missing link in ecological speciation. Science 326:847–850.']
  • ['Cook, T. 1994. Meta-analysis for explanation: a casebook. Russell Sage Foundation, New York.']
  • ['Cooper, H. M., L. V. Hedges, and J. C. Valentine. 2009. The handbook of research synthesis and meta-analysis. Russell Sage Foundation, New York.']
  • ['Courtiol, A., M. Raymond, B. Godelle, and J.-B. Ferdy. 2010. Mate choice and human stature: homogamy as a unified framework for understanding mating preferences. Evolution 64:2189–2203.']
  • ['Coyne, J., and H. Orr. 2004. Speciation. Sinauer, Sunderland, MA.']
  • ['Coyne, J., and T. Price. 2000. Little evidence for sympatric speciation in island birds. Evolution 54:2166–2171.']
  • ['Crespi, B. J. 1989. Causes of assortative mating in arthropods. Animal Behaviour 38:980–1000.']
  • ['Crozier, W. J. 1917. Evidence of assortative mating in a nudibranch. Proceedings of the National Academy of Sciences of the USA 3:519–522.']
  • ['———. 1918. Assortative mating in a nudibranch, Chromodoris zebra Heilprin. Journal of Experimental Zoology 27:247–292.']
  • ['De Cara, M. A. R., N. H. Barton, and M. Kirkpatrick. 2008. A model for the evolution of assortative mating. American Naturalist 171:580–596.']
  • ['Dekkers, J. 1994. Optimal breeding strategies for calving ease. Journal of Dairy Science 77:3441–3453.']
  • ['Dick, J., and R. Elwood. 1996. Effects of natural variation in sex ratio and habitat structure on mate-guarding decisions in amphipods (Crustacea). Behaviour 133:985–996.']
  • ['Dieckmann, U., and M. Doebeli. 1999. On the origin of species by sympatric speciation. Nature 400:354–357.']
  • ['Dieckmann, U., M. Doebeli, J. A. J. Metz, and D. Tautz. 2004. Adaptive speciation. Cambridge University Press, Cambridge.']
  • ['Doebeli, M. 1996. A quantitative genetic competition model for sympatric speciation. Journal of Evolutionary Biology 9:893–909.']
  • ['Drès, M., and J. Mallet. 2002. Host races in plant–feeding insects and their importance in sympatric speciation. Philosophical Transactions of the Royal Society B: Biological Sciences 357:471–492.']
  • ['Duenez-Guzman, E. A., J. Mavárez, M. D. Vose, and S. Gavrilets. 2009. Case studies and mathematical models of ecological speciation. 4. Hybrid speciation in butterflies in a jungle. Evolution 63:2611–2626.']
  • ['Edelaar, P., A. M. Siepielski, and J. Clobert. 2008. Matching habitat choice causes directed gene flow: a neglected dimension in evolution and ecology. Evolution 62:2462–2472.']
  • ['Egger, M., G. D. Smith, M. Schneider, and C. Minder. 1997. Bias in meta-analysis detected by a simple, graphical test. British Medical Journal 315:629–634.']
  • ['Endler, J. A. 1986. Natural selection in the wild. Princeton University Press, Princeton, NJ.']
  • ['Epinat, G., and T. Lenormand. 2009. The evolution of assortative mating and selfing with in- and outbreeding depression. Evolution 63:2047–2060.']
  • ['Felsenstein, J. 1981. Skepticism towards Santa Rosalia, or why are there so few kinds of animals? Evolution 35:124–138.']
  • ['Fitzpatrick, B., J. Fordyce, and S. Gavrilets. 2008. What, if anything, is sympatric speciation? Journal of Evolutionary Biology 21:1452–1459.']
  • ['Follett, P. A., F. Calvert, and M. Golden. 2007. Genetic studies using the orange body color type of Nezara viridula (Hemiptera: Pentatomidae): inheritance, sperm precedence, and disassortative mating. Annals of the Entomological Society of America 100:433–438.']
  • ['Gavrilets, S. 2004. Fitness landscapes and the origin of species. Princeton University Press, Princeton, NJ.']
  • ['Gavrilets, S., and A. Vose. 2007. Case studies and mathematical models of ecological speciation. 2. Palms on an oceanic island. Molecular Ecology 16:2910–2921.']
  • ['Gavrilets, S., A. Vose, M. Barluenga, W. Salzburger, and A. Meyer. 2007. Case studies and mathematical models of ecological speciation. 1. Cichlids in a crater lake. Molecular Ecology 16:2893–2909.']
  • ['Gimelfarb, A. 1986. Is offspring-midparent regression affected by assortative mating of parents? Genetical Research 47:71–75.']
  • ['Harari, A. R., A. M. Handler, and P. J. Landolt. 1999. Size-assortative mating, male choice and female choice in the curculionid beetle Diaprepes abbreviatus. Animal Behaviour 58:1191–1200.']
  • ['Hedges, L. V., I. Olkin, and M. Statistiker. 1985. Statistical methods for meta-analysis. Academic Press, Orlando, FL.']
  • ['Helfenstein, F., E. Danchin, and R. Wagner. 2004. Assortative mating and sexual size dimorphism in black-legged kittiwakes. Waterbirds 27:350–354.']
  • ['Henry, L. 2008. Assortative mating and the role of phenotypic plasticity in male competition: implications for gene flow among host-associated parasitoid populations. Biology Letters 4:508–511.']
  • ['Hooper, P. L., and G. F. Miller. 2008. Mutual mate choice can drive costly signaling even under perfect monogamy. Adaptive Behavior 16:53–70.']
  • ['Hume, K., R. Elwood, J. Dick, and K. Connaghan. 2002. Size-assortative pairing in Gammarus pulex (Crustacea: Amphipoda): a test of the timing hypothesis. Animal Behaviour 64:239–244.']
  • ['Hunter, J. E., and F. L. Schmidt. 2004. Methods of meta-analysis: correcting error and bias in research findings. Sage, Newbury Park, CA.']
  • ['Johannesson, K., E. Rolan-Alvarez, and A. Ekendahl. 1995. Incipient reproductive isolation between two sympatric morphs of the intertidal snail Littorina saxatilis. Evolution:1180–1190.']
  • ['Johnston, R. F., and S. G. Johnson. 1989. Nonrandom mating in feral pigeons. Condor 91:23–29.']
  • ['Jones, A., G. Moore, C. Kvarnemo, D. Walker, and J. Avise. 2003. Sympatric speciation as a consequence of male pregnancy in seahorses. Proceedings of the National Academy of Sciences of the USA 100:6598–6603.']
  • ['Kingsolver, J. G., H. E. Hoekstra, J. M. Hoekstra, D. Berrigan, S. N. Vignieri, C. E. Hill, A. Hoang, et al. 2001. The strength of phenotypic selection in natural populations. American Naturalist 157:245–261.']
  • ['Kirkpatrick, M. 2000. Reinforcement and divergence under assortative mating. Proceedings of the Royal Society B: Biological Sciences 267:1649–1655.']
  • ['Kirkpatrick, M., and S. L. Nuismer. 2004. Sexual selection can constrain sympatric speciation. Proceedings of the Royal Society B: Biological Sciences 271:687–693.']
  • ['Kirkpatrick, M., and V. Ravigné. 2002. Speciation by natural and sexual selection: models and experiments. American Naturalist 159(suppl.):S22–S35.']
  • ['Kondrashov, A. S., and M. Shpak. 1998. On the origin of species by means of assortative mating. Proceedings of the Royal Society B: Biological Sciences 265:2273–2278.']
  • ['Kuramoto, M. 1978. Correlations of quantitative parameters of fecundity in amphibians. Evolution 32:287–296.']
  • ['Lewontin, R., D. Kirk, and J. Crow. 1968. Selective mating, assortative mating, and inbreeding: definitions and implications. Eugenics Quarterly 15:141–143.']
  • ['Lipsey, M. W., and D. B. Wilson. 2001. Practical meta-analysis. Sage, Thousand Oaks, CA.']
  • ['Lu, X., X. Ma, Y. Li, and L. Fan. 2009. Breeding behavior and mating system in relation to body size in Rana chensinensis, a temperate frog endemic to northern China. Journal of Ethology 27:391–400.']
  • ['Lynch, M., and B. Walsh. 1998. Genetics and analysis of quantitative traits. Sinauer, Sunderland, MA.']
  • ['Malausa, T., M. Bethenod, A. Bontemps, D. Bourguet, J. Cornuet, and S. Ponsard. 2005. Assortative mating in sympatric host races of the European corn borer. Science 308:258–260.']
  • ['Maynard Smith, J. 1966. Sympatric speciation. American Naturalist 100:637–650.']
  • ['Mays, H. L., Jr., and G. E. Hill. 2004. Choosing mates: good genes versus genes that are a good fit. Trends in Ecology and Evolution 19:554–559.']
  • ['McLain, D. K. 1982. Density dependent sexual selection and positive phenotypic assortative mating in natural populations of the soldier beetle, Chauliognathus pennsylvanicus. Evolution 36:1227–1235.']
  • ['McLain, D. K., and R. D. Boromisa. 1987. Male choice, fighting ability, assortative mating and the intensity of sexual selection in the milkweed longhorn beetle, Tetraopes tetraophthalmus (Coleoptera, Cerambycidae). Behavioral Ecology and Sociobiology 20:239–246.']
  • ['McNamara, J. M., and E. J. Collins. 1990. The secretary problem as an employer-candidate game. Journal of Applied Probability 27:815–827.']
  • ['Merikangas, K. 1982. Assortative mating for psychiatric disorders and psychological traits. Archives of General Psychiatry 39:1173–1180.']
  • ['Olson, D. H., A. R. Blaustein, and R. K. O’Hara. 1986. Mating pattern variability among western toad (Bufo boreas) populations. Oecologia (Berlin) 70:351–356.']
  • ['Ortego, J., G. Calabuig, R. Bonal, A. Muñoz, J. M. Aparicio, and P. J. Cordero. 2009. Temporal variation of heterozygosity-based assortative mating and related benefits in a lesser kestrel population. Journal of Evolutionary Biology 22:2488–2495.']
  • ['Otto, S. P., M. R. Servedio, and S. L. Nuismer. 2008. Frequency-dependent selection and the evolution of assortative mating. Genetics 179:2091–2112.']
  • ['Palmer, A. R. 1999. Detecting publication bias in meta-analyses: a case study of fluctuating asymmetry and sexual selection. American Naturalist 154:220–233.']
  • ['Partridge, L. 1983. Non-random mating and offspring fitness. Pages 227–256 in P. Bateson, ed. Mate choice. Cambridge University Press, Cambridge.']
  • ['Piertney, S. B., and M. K. Oliver. 2006. The evolutionary ecology of the major histocompatibility complex. Heredity 96:7–21.']
  • ['Pryke, S. R., and S. C. Griffth. 2007. The relative role of male vs. female mate choice in maintaining assortative pairing among discrete colour morphs. Journal of Evolutionary Biology 20:1512–1521.']
  • ['Pusey, A., and M. Wolf. 1996. Inbreeding avoidance in animals. Trends in Ecology and Evolution 11:201–206.']
  • ['R Development Core Team. 2009. R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna.']
  • ['Redden, D., and D. Allison. 2006. The effect of assortative mating upon genetic association studies: spurious associations and population substructure in the absence of admixture. Behavior Genetics 36:678–686.']
  • ['Reid, W. V. 1988. Age correlations within pairs of breeding birds. Auk 105:278–285.']
  • ['Rice, W. R., and E. E. Hostert. 1993. Laboratory experiments on speciation: what have we learned in 40 years? Evolution 47:1637–1653.']
  • ['Roberts, S. C., M. L. Hale, and M. Petire. 2006. Correlations between heterozygosity and measures of genetic similarity: implications for understanding mate choice. Journal of Evolutionary Biology 19:558–569.']
  • ['Roulin, A. 2004. The evolution, maintenance and adaptive function of genetic colour polymorphism in birds. Biological Reviews 79:815–848.']
  • ['Rowe, L., and G. Arnqvist. 1996. Analysis of the causal components of assortative mating in water striders. Behavioral Ecology and Sociobiology 38:279–286.']
  • ['Sadedin, S., J. Hollander, M. Panova, K. Johannesson, and S. Gavrilets. 2009. Case studies and mathematical models of ecological speciation. 3. Ecotype formation in a Swedish snail. Molecular Ecology 18:4006–4023.']
  • ['Salthe, S. N., and W. E. Duellman. 1973. Quantitative constraints associated with reproductive mode in anurans. Pages 229–249 in J. Vial, ed. Evolutionary biology of the anurans: contemporary research on major problems. University of Missouri Press, Columbia.']
  • ['Seehausen, O., J. J. M. Alphen, and F. Witte. 1997. Cichlid fish diversity threatened by eutrophication that curbs sexual selection. Science 277:1808–1811.']
  • ['Shine, R., D. O’Connor, M. LeMaster, and R. Mason. 2001. Pick on someone your own size: ontogenetic shifts in mate choice by male garter snakes result in size-assortative mating. Animal Behaviour 61:1133–1141.']
  • ['Snowberg, L. K., and D. I. Bolnick. 2008. Assortative mating by diet in a phenotypically unimodal but ecologically variable population of stickleback. American Naturalist 172:733–739.']
  • ['Spuhler, J. 1968. Assortative mating with respect to physical characteristics. Biodemography and Social Biology 15:128–140.']
  • ['Udovic, D. 1980. Frequency-dependent selection, disruptive selection, and the evolution of reproductive isolation. American Naturalist 116:621–641.']
  • ['Viechtbauer, W. 2010. Conducting meta-analyses in R with the metafor package. Journal of Statistical Software 36:1–48.']
  • ['Waser, N. M. 1993. Population structure, optimal outbreeding, and assortative mating in angiosperms. Page 173–199 in N. W. Thornhill, ed. The natural history of inbreeding and outbreeding: theoretical and empirical perspectives. University of Chicago Press, Chicago.']
  • ['Wedekind, C., T. Seebeck, F. Bettens, and A. J. Paepke. 1995. MHC-dependent mate preferences in humans. Proceedings of the Royal Society B: Biological Sciences 260:245–249.']
  • ['Weis, A. 2005. Direct and indirect assortative mating: a multivariate approach to plant flowering schedules. Journal of Evolutionary Biology 18:536–546.']
  • ['Weis, A., J. Winterer, C. Vacher, T. Kossler, C. Young, and G. LeBuhn. 2005. Phenological assortative mating in flowering plants: the nature and consequences of its frequency dependence. Evolutionary Ecology Research 7:161–181.']
  • ['Wogel, H., P. A. Abrunhosa, and J. P. Pombal. 2005. Breeding behaviour and mating success of Phyllomedusa rohdei (Anura, Hylidae) in south-eastern Brazil. Journal of Natural History 39:2035–2045.']
  • ['Wolański, N. 1994. Assortative mating in somatic traits and its consequences. Studies in Human Ecology 11:73–111.']
  • ['Wright, S. 1921. Systems of mating. III. Assortative mating based on somatic resemblance. Genetics 6:144–161.']
  • ['Yamazaki, K., E. A. Boyse, V. Mike, H. T. Thaler, B. J. Mathieson, J. Abbott, J. Boyse, et al. 1976. Control of mating preferences in mice by genes in the major histocompatibility complex. Journal of Experimental Medicine 144:1324–1335.']