Table list of papers not included in the final analysis Table S2

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Dougherty & Shuker, 2014 Supplementary material

Dougherty & Shuker, 2014

Supplementary material

Section 1: Search methods

Literature search results

Table S1. List of papers not included in the final analysis

Table S2. Methods used for calculating effect sizes for papers in which statistics were not fully reported
Section 2: Phylogenetic methods

List of studies used to assemble phylogeny

Methods for phylogenetic analysis

Phylogeny in Newick format

Figure S1. Phylogeny used in analysis
Section 3: Effect sizes used in final analysis

Table S3. All effect sizes extracted for analysis
Section 4: Supplementary results.

Results of multivariate meta-analysis models incorporating random effects

Figure S2. Forest plots showing models with and without random effects

Table S4. Results of models incorporating random effects

Figure S3. Scatter plot of effect size against variance

Figure S4. Funnel plot showing trim and fill results

Figure S5: Cumulative forest plot

Section 1: Search methods

Literature search results
Detailed results of all online literature searches performed. An Endnote library of all our search results is also available on request. In all cases we searched across all years and sources available to each database.
All searches, excluding duplicates: 613 results
Google Scholar

Searched 17th June 2013, saved first 100 results (first ten pages)

  • Search: “sequential simultaneous mate choice” 100 results


Searched 17th June 2013, saved first 100 results (first ten pages), removed any results that were not articles

  • Search: “sequential simultaneous mate choice” 68 results


All searches 19th June 2013, in search field: “Article Title, Abstract, Keywords”, all years, subject areas, document types

  • Search: “no choice” AND “multiple choice” 1 result

  • Search: “no choice” AND “two choice” 3 results

  • Search: “no choice” AND “simultaneous” 1 result

  • Search: “sequential” AND “simultaneous” 14 result

  • Search: “sexual* isolat*” AND “no choice” AND “multiple choice” 1 result

Web of Knowledge

All searches 19th June 2013, in search field: “Topic”, all databases, all years

  • Search: “no choice” AND “multiple choice” 28 results

  • Search: “no choice” AND “two choice” 23 results

  • Search: “no choice” AND “simultaneous” 29 results

  • Search: “sequential” AND “simultaneous” 134 results

  • Search: “sexual* isolat*” AND “no choice” AND “multiple choice” 8 results

View all citing articles in all databases

  • WoS, papers citing Coyne et al., (2005) 36 results

  • WoS, papers citing MacLaren & Rowland, (2004) 24 results

  • WoS, papers citing Rowland, (1982) 68 results

  • WoS, papers citing Wagner, (1998) 218 results

Studies excluded from analysis

Table S1. List of relevant studies that were not included in our analysis, and the reasons for them not being included. In all these studies both no-choice and choice tests were used, however they did not meet all of our inclusion criteria. See below for full references.



Reason for non-inclusion

Doherty, 1985

Gryllus bimaculatus

No data reported

Bissell & Martins, 2006

Sceloporus graciosus

No data reported

Kostarakos et al., 2008

Gryllus bimaculatus

No data reported

Allison & Cardé, 2008

Cadra cautella

No data reported

Yukilevich & True, 2008

Drosophila melanogaster

Do not report results of no-choice tests

Kozak et al., 2011

Gasterosteus aculeatus

Do not report results of no-choice tests

Meffert & Regan, 2012

Musca domestica

Do not report results of no-choice tests

Taborsky et al., 2009

Eretmodus cyanostictus

Do not report results of choice tests

Gupta & Sarandan, 1994

Drosophila kikkawai

Not true no-choice test, Do not report results of no-choice tests

Wu et al., 1995

Drosophila melanogaster

Not true no-choice test

Singh & Sisodia, 1999

Drosophila bipectinata

Not true no-choice test

Tomaru & Oguma, 2000

Drosophila melanogaster

Not true no-choice test

Drosophila sechellia

Not true no-choice test

Zhao et al., 2008

Helicoverpa armigera

Not true no-choice test

Kumaran et al., 2013

Bactrocera tryoni

Not true no-choice test

Ryan & Rand, 1993

Physalaemus pustulosus

No conspecific calls tested in no-choice tests

Willis et al., 2011

Xiphophorus birchmanni

No conspecifics tested in no-choice trials

Bee & Schwartz, 2009

Hyla chrysoscelis

No use of heterospecific call in no-choice tests

Basolo, 1995

Priapella olmecae

Did not test for preference for no sword in no-choice trials

Basolo, 2002

Heterandria bimaculata

Did not test for preference for no sword in no-choice trials

Walling et al., 2010

Xiphophorus helleri

Females only tested with one type of male in no-choice trials

Havens & Etges, 2013

Drosophila mojavensis

No comparison between mated and unmated males in no-choice trials

Uetz & Norton, 2007

Schizocosa ocreata

Preference for different traits tested in choice an no-choice tests

Sharma et al., 2010

Drosophila simulans

Preference for different traits tested in choice an no-choice tests

Deb et al., 2012

Oecanthus henryi

Preference for different traits tested in choice an no-choice tests

Wade et al., 1995

Tribolium confusum

No mate choice: recorded offspring number from different matings

Grant et al., 1995

Oryzias latipes

No mate choice: measured male mating success

Wiegmann, 1999

Gryllus integer

No mate choice: ability of female to locate speakers

Muller & Robert, 2002

Ormia ochracea

No mate choice: ability of parasite to locate host

Silva et al., 2007

Syngnathus abaster

No mate choice in no-choice tests: female associations with females

Kozak & Boughamn, 2009

Gasterosteus aculeatus

Females tested with no-choice tests, males tested with choice tests

Havens et al., 2011

Drosophila mojavensis

No mate choice: recorded offspring fitness of matings


Allison JD, Cardé RT, 2008. Male pheromone blend preference function measured in choice and no-choice wind tunnel trials with almond moths, Cadra cautella. Anim Behav. 75:259-266.

Basolo AL, 1995. Phylogenetic evidence for the role of a pre-existing bias in sexual selection. Proc Biol Sci. 259:307-311.

Basolo AL, 2002. Female discrimination against sworded males in a poeciliid fish. Anim Behav. 63:463-468.

Bee MA, Schwartz JJ, 2009. Behavioral measures of signal recognition thresholds in frogs in the presence and absence of chorus-shaped noise. J Acoust Soc Am. 126:2788-2801.

Bissell AN, Martins EP, 2006. Male approach and female avoidance as mechanisms of population discrimination in sagebrush lizards. Behav Ecol Sociobiol. 60:655-662.

Deb R, Bhattacharya M, Balakrishnan R, 2012. Females of a tree cricket prefer larger males but not the lower frequency male calls that indicate large body size. Anim Behav. 84:137-149.

Doherty JA, 1985. Phonotaxis in the cricket, Gryllus bimaculatus De Geer: comparisons of choice and no-choice paradigms. J Comp Physiol A. 157:279-289.

Grant JWA, Bryant MJ, Soos CE, 1995. Operational sex ratio, mediated by synchrony of female arrival, alters the variance of male mating success in Japanese Medaka. Anim Behav. 49:367-375.

Gupta J, Sundaran A, 1994. Some evidence of incipient speciation in Drosophila kikkawai. Genome 37:1041-1044.

Havens J, Orzack S, Etges W, 2011. Mate choice opportunity leads to shorter offspring development time in a desert insect. J Evolution Biol. 24:1317-1324.

Havens JA, Etges WJ, 2013. Premating isolation is determined by larval rearing substrates in cactophilic Drosophila mojavensis. IX. Host plant and population specific epicuticular hydrocarbon expression influences mate choice and sexual selection. J Evolution Biol. 26:562-576.

Kostarakos K, Hartbauer M, Römer H, 2008. Matched filters, mate choice and the evolution of sexually selected traits. Plos One 3.

Kozak GM, Head ML, Boughman JW, 2011. Sexual imprinting on ecologically divergent traits leads to sexual isolation in sticklebacks. Proc Biol Sci. 278:2604-2610.

Kozak GM, Reisland M, Boughmann JW, 2009. Sex differences in mate recognition and conspecific preference in species with mutual mate choice. Evolution 63:353-365.

Kumaran N, Balagawi S, Schutze MK, Clarke AR, 2013. Evolution of lure response in tephritid fruit flies: phytochemicals as drivers of sexual selection. Anim Behav. 85:781-789.

Meffert LM, Regan JL, 2002. A test of speciation via sexual selection on female preferences. Anim Behav. 64:955-965.

Muller P, Robert D, 2002. Death comes suddenly to the unprepared: singing crickets, call fragmentation, and parasitoid flies. Behav Ecol. 13:598-606.

Ryan MJ, Rand AS, 1993. Species recognition and sexual selection as a unitary problem in animal communication. Evolution 47:647-657.

Sharma MD, Tregenza T, Hosken DJ, 2010. Female mate preferences in Drosophila simulans: evolution and costs. J Evolution Biol. 23:1672-1679.

Silva K, Vieira MN, Almada VC, Monteiro NM, 2007. The effect of temperature on mate preferences and female-female interactions in Syngnathus abaster. Anim Behav. 74:1525-1533.

Singh BN, Sisodia S, 1999. Mating propensity in Drosophila bipectinata under different sex-ratios and choice situations. Curr Sci India. 76:222-225.

Taborsky B, Guyer L, Taborsky M, 2009. Size-assortative mating in the absence of mate choice. Anim Behav. 77:439-448.

Tomaru M, Oguma Y, 2000. Mate choice in Drosophila melanogaster and D. sechellia: criteria and their variation depending on courtship song. Anim Behav. 60:797-804.

Uetz GW, Norton S, 2007. Preference for male traits in female wolf spiders varies with the choice of available males, female age and reproductive state. Behav Ecol Sociobiol. 61:631-641.

Wade MJ, Chang NW, McNaughton M, 1995. Incipient speciation in the flour beetle, Tribolium confusum: premating isolation between natural populations. Heredity 75:453-459.

Walling CA, Royle NJ, Lindstrom J, Metcalfe NB, 2010. Do female association preferences predict the likelihood of reproduction? Behav Ecol Sociobiol. 64:541-548.

Wiegmann DD, 1999. Search behaviour and mate choice by female field crickets, Gryllus integer. Anim Behav. 58:1293-1298.

Willis PM, Ryan MJ, Rosenthal GG, 2011. Encounter rates with conspecific males influence female mate choice in a naturally hybridizing fish. Behav Ecol. 22:1234-1240.

Wu C-I, Hollocher H, Begun DJ, Aquadro CF, Xu Y, Wu M-L, 1995. Sexual isolation in Drosophila melanogaster: a possible case of incipient speciation. P Natl Acad Sci USA. 92:2519-2523.

Yukilevich R, True JR, 2008. Incipient sexual isolation among cosmopolitan Drosophila melanogaster populations. Evolution 62:2112-2121.

Zhao XC, Wu KM, Liang GM, Guo YY, 2008. Altered mating behaviour in a Cry1Ac-resistant strain of Helicoverpa armigera (Lepidoptera : Noctuidae). J Appl Entomol. 132:360-365.

Methods for calculating effect sizes not presented in papers

For 25 out of the 38 studies present in the analysis all useful statistics were not fully reported in the text; statistics were obtained from information in figures and tables. Details of all methods are presented in Figure S2.
Table S2: Methods used for calculating effect sizes for papers in which statistics were not fully reported. Data were extracted from figures using the image analysis software Digitize It 2010.



Wagner et al., 1995

No-choice effect size calculated from data extracted from Figure 3, converted to eta-squared using sums of squares, then converted to r. Was unable to account for fact that data are from repeated-measures. Choice test: calculated Hedge's d using data from Table 1

Jang & Gerhardt, 2006

Performed t tests using data from Figure 3

Gershmann & Sakaluk, 2009

Repeated χ2 test using data in text

Lehmann & Lehmann, 2008

Performed χ2 test using data from Figure 2

Coyne et al., 2005

Statistics for both mating frequency and mating latency taken from data in table 1

Wood & Ringo, 1980

Mating frequency data: repeated analysis in order to obtain χ2 statistics (Tables 1 & 2). Correlation of 1 for some measures, set at 2 when converted to Zr

Jennings et al., 2011

Performed χ2 tests using data from Table 1

Hoikkala & Aspi, 1993

Performed χ2 test on proportion data extracted from Figure 3. Exact sample sizes not presented (only range), took largest sample size presented

Xu & Wang, 2009

Performed χ2 tests on data from Figure 2

Schofl et al., 2011

Performed χ2 tests using data from Table 2 and Figure 3

Cook et al., 1994

Performed χ2 test on data in Table 1 & 2. Had to assume individuals were not encountered more than once

Barry et al., 2010

Performed χ2 tests on data from text

King et al., 2005

Performed χ2 tests on data from Tables 1 & 4

McNamara et al., 2004

Performed χ2 test on mating frequency data from Figure 1

Dougherty & Shuker, 2013

Re-analysed own data, excluding mutual choice treatment

Jordan & Brooks, 2011

Performed t tests performed using data from Figures 1 & 2

Hurt et al., 2004

No-choice t statistic calculated from data extracted from Table 1. Choice test: calculated Hedge's d using data from Table 4

Rowland, 1982

Performed Wilcoxon test using data from Tables 1 & 2

Jamieson & Colgan, 1989

Performed χ2 test from data in text

Belle-Isles, 1990

Performed χ2 test using data from Table 2, combined results from all males

Owen et al., 2012

Statistics obtained by contacting authors

Itzkowitz et al., 1998

z scores back-converted from presented P values

Suk & Choe, 2002

t statistic for no-choice test calculated from P value and N

Phelps et al., 2006

Performed χ2 tests on data from Figure 5

Verrell, 1995

Performed χ2 test from data in text

Section 2: Phylogenetic methods

Obtaining the phylogeny
The species in the meta-analysis are very widespread taxonomically and many are not commonly studied, thus there is no single phylogeny available. Therefore a supertree was constructed by combining multiple trees from several different sources.
For the basal node between vertebrates and arthropods we used Glenner et al. (2004). For the relationship among arthropods we used Regier & Shultz (1998) and Giribet et al. (2001). For the relationship among insect orders we used Gaunt & Miles (2002) and Kjer et al. (2006). For the relationship among Orthoptera we used Huang et al. (2000) and Jost & Shaw (2006). For the relationship among Lepidoptera we used Regier et al. (2009) and Mutanen et al. (2010). For the relationship among Drosophila we used Lachaise et al. (2000), Spicer & Bell (2002) and van der Linde & Houle (2008).
For the relationships among vertebrates we used Ureta-Vidal et al. (2003) and Xia et al. (2003). For the relationships among fish orders we used Miya et al. (2003) and Near et al. (2012), though the position of these nodes are particularly unresolved and may be subject to change. For the relationships among the Cyprinodontiformes we used Meyer & Lydeard (1993) and Ghedotti (2000).
Gaunt MW, Miles MA, 2002. An insect molecular clock dates the origin of the insects and accords with palaeontological and biogeographic landmarks. Mol Biol Evol. 19:748-761.

Ghedotti MJ, 2000. Phylogenetic analysis and taxonomy of the poecilioid fishes (Teleostei: Cyprinodontiformes). Zool J Linn Soc-Lon. 130:1-53.

Giribet G, Edgecombe GD, Wheeler WC, 2001. Arthropod phylogeny based on eight molecular loci and morphology. Nature 413:157-161.

Glenner H, Hansen AJ, Sørensen MV, Ronquist F, Huelsenbeck JP, Willerslev E, 2004. Bayesian inference of the metazoan phylogeny: a combined molecular and morphological approach. Curr Biol. 14:1644-1649.

Huang Y, Ortı́ G, Sutherlin M, Duhachek A, Zera A, 2000. Phylogenetic relationships of North American field crickets inferred from mitochondrial DNA data. Mol phylogenet evol. 17:48-57.

Jost M, Shaw K, 2006. Phylogeny of Ensifera (Hexapoda: Orthoptera) using three ribosomal loci, with implications for the evolution of acoustic communication. Mol phylogenet evol. 38:510-530.

Kjer KM, Carle FL, Litman J, Ware J, 2006. A molecular phylogeny of Hexapoda. Arthropod Syst Phylo. 64:35-44.

Lachaise D, Harry M, Solignac M, Lemeunier F, Benassi V, Cariou M-L, 2000. Evolutionary novelties in islands: Drosophila santomea, a new melanogaster sister species from Sao Tome. Proc Biol Sci. 267:1487-1495.

Meyer A, Lydeard C, 1993. The evolution of copulatory organs, internal fertilization, placentae and viviparity in killifishes (Cyprinodontiformes) inferred from a DNA phylogeny of the tyrosine kinase gene X-src. Proc Biol Sci. 254:153-162.

Miya M, Takeshima H, Endo H, Ishiguro NB, Inoue JG, Mukai T, Satoh TP, Yamaguchi M, Kawaguchi A, Mabuchi K, 2003. Major patterns of higher teleostean phylogenies: a new perspective based on 100 complete mitochondrial DNA sequences. Mol phylogenet evol. 26:121-138.

Mutanen M, Wahlberg N, Kaila L, 2010. Comprehensive gene and taxon coverage elucidates radiation patterns in moths and butterflies. Proc Biol Sci. 277:2839-2848.

Near TJ, Eytan RI, Dornburg A, Kuhn KL, Moore JA, Davis MP, Wainwright PC, Friedman M, Smith WL, 2012. Resolution of ray-finned fish phylogeny and timing of diversification. P Natl Acad Sci USA. 109:13698-13703.

Paradis E, Claude J, Strimmer K, 2004. APE: analyses of phylogenetics and evolution in R language. Bioinformatics 20:289-290.

Regier JC, Shultz JW, 1998. Molecular phylogeny of arthropods and the significance of the Cambrian “explosion” for molecular systematics. Am Zool. 38:918-928.

Regier JC, Zwick A, Cummings MP, Kawahara AY, Cho S, Weller S, Roe A, Baixeras J, Brown JW, Parr C, 2009. Toward reconstructing the evolution of advanced moths and butterflies (Lepidoptera: Ditrysia): an initial molecular study. Bmc Evol Biol. 9:280.

Spicer GS, Bell C, 2002. Molecular phylogeny of the Drosophila virilis species group (Diptera: Drosophilidae) inferred from mitochondrial 12S and 16S ribosomal RNA genes. Ann Entomol Soc Am. 95:156-161.

Team RDC, 2012. R: A language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing.

Ureta-Vidal A, Ettwiller L, Birney E, 2003. Comparative genomics: genome-wide analysis in metazoan eukaryotes. Nat Rev Gen. 4:251-262.

van der Linde K, Houle D, 2008. A supertree analysis and literature review of the genus Drosophila and closely related genera (Diptera, Drosophilidae). Insect Syst Evol. 39:241-267.

Xia X, Xie Z, Kjer KM, 2003. 18S ribosomal RNA and tetrapod phylogeny. Syst Biol. 52:283-295.

Phylogenetic analysis methods
Once the supertree was obtained it was manually converted into Newick format (see below), with branch lengths all set to one. This standard tree is not ultrametric (tips do not all line up): lineages with more branches (i.e. with more species) are longer. In order to obtain an ultrametric tree the standard tree was run in FigTree v1.4 (Andrew Rambaut, 2012) and transformed using the cladogram option to obtain the final tree seen below (Figure 1). Branch lengths are thus transformed so that all tips are contemporaneous. The total length of the tree is 12 units. Note that in this tree branching events only roughly correspond to taxonomic groupings; distortion is apparent especially for basal nodes and lineages with several species (such as the Drosophila genus). The ultrametric tree was then imported into R 3.0.1 (R Development Core Team, 2014), and a correlation matrix obtained using the vcv function in the package ape v3.1.1 (Paradis et al., 2004). This gives a standardised matrix with diagonal values equal to 1 (each tip is fully correlated with itself). This phylogeny was then incorporated into a mixed-effects meta-analysis model using the function in the package Metafor v1.9-2 (Viechtbauer 2010).

Phylogeny in Newick format:


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