non-commercial pur 32: 984-993,whichhasbeenpublishedinfinalformathttps://doi.org/10.1111/jeb.13500.Thisarticlemaybeusedfor c This isthepeerr Accepted Article onstraints innonrandommatingpatter *Corresponding autho Canada This by is protectedarticle reserved. Allrights copyright. doi: 10.1111/jeb.13500 lead to differences between this version and the throughbeen the copyediting, pagination typesetting, which process, may and proofreading hasThis article been accepted publicationfor andundergone peerfull but review not has 3 2 1 Rosalind L Murray* ornaments female with adance for patterns mating non-random in constraints functional role of The typeArticle :ResearchPapers MURRAY 0000-0001-6464-0072)DR ROSALIND (Orcid ID: Email: [email protected] 828 3876 +1905Fax: Phone: +1 647 530 9779 ofUniversity Toronto Mississauga Biology Department Rosalind L Murray BiologicalandEnvironmental UniversitySciences, of Stirling, UK Stirling, FK9 4LA UniversityBiology ofToronto Dept., at ONCanada L5LMississauga,Mississauga, 1C6 andof Ecology Dept. Evolutionary Univ Biology, eviewed versionofthefollowingarticle:Murray,RL,Gwynne,DT,Bussièr poses inaccordancewithWileyTerms 1,2,3 r: , Darryl T Gwynne ns foradanceflywithfemaleornaments.JournalofEv 2 , Luc F, Bussière Version of Record. Please Version as this cite ofRecord. article andConditionsfor ersity ersity of Toronto, Toronto, 3B2 ON,M5S 3

self-archiving. e, LF.Theroleoffunctional olutionary Biology2019; Accepted Article This by is protectedarticle reserved. Allrights copyright. observed in intermediatelyadorned females ismorelikelyto becausingthenon-random patternsmating relationship of assortative mating by mass. We suggest activethat male choice mate for that while Weheavier found males are that heavierfound oftenmore suggesting with mating females determine whethercarried loads bymalesduring the lightercopulation were expected. than male’sthe withcopulation wingloading performed alsoduring permutationWe traits. to tests copulationwere experiencing by comparing limitations load-lift by themasscarried males species. We whether thereforewild-caught tested male previouslyobserved couldproduce size constraint onfemale selection stabilizing inthis the In concertwith directional preferences largefemaleswithmature for a load-lifting eggs, such pairing opportunities for the heaviestfemales,asreportedfornon-ornamented dance . a . andability theory, carryIn to couldnuptial gifts females male’s constrain bearsand mating,weight entire male the of the andthe female giftdurationthe nuptial for of selection. ornamentsunusual thatare amongsexualtraitsbecause under they are stabilizing longicauda for tested functional load-lifting constraints during in aerial copulation can result in functional constraints also can that causenon-random matingWe patterns. display elaboratespecies that However, traits on ornaments. pressures selection conflicting Most hypotheses explainnon-random to invoke patternsmating choice, mate particularly in Abstract Running Title: Load-lift constraints on aerial mating R. longicauda R. longicauda , a speciesdisplays of, dancefly that female-specific multiple extravagant R. longicauda males do not experiencenot malesdoa constraint, a load-lift strong is there males providefemales witha nuptial engaging before gift inaerial . R. longicauda collected during Accepted Article

Partridge, 1983,Crespi, 1989). andevolutionaryconsequencessuch asinbreedingand assortative mating (e.g. see to changeobservedonly mating patterns, could butbringabout also important ecological 2001). Loadingconstraints biologya thereproductiveof impact that species notare likely femalesbecause often are such females heaviest and the fecundmost (Bonduriansky, abilitymating, couldduring load-lifting attractive withlimits tomate to thea male’s constrain engage in aerial copulations (Marden, 1989, Peckarsky et al., 2002). malesIf carry females (Marden,McLachlan 1989,& 1987, SawadogoAllen, especially etal.,2013), that those 1991, 2002). etal., Peckarsky mating patterns, andcan prevent pairingsregardlessof preferences (Schlutermating al., et non-random in also result 2008)can 1989, Almbro &Kullberg, (Marden, mating success 2011).Dubois2003,al., (Candolin, et functionalSimilarly, individual’s an limitations to Funk Candolin,2003, 1998, Tallamy,2000) can & in result choice mate maladaptive example, the inability to accurately evaluate a potential mate’s quality (Endler & Basolo, otherare aspects pair of formation canthat influencethe identity of For partners. mating This by is protectedarticle reserved. Allrights copyright. Introduction constraint, Keywords:dance assortative mating, ornaments,flies, female choice, matemale mating Functional constraints on pairing may be particularly important for flying Hypotheses toexplainanimal patterns mating typicallyinvoke choice, mate but there Rhamphomyia longicauda , sexual selection Accepted Article nuptialthey couldcarrygift (Marden, 1989). male suggesting that approachedbutnever surpassed the predictedtheoretically load-liftingconstraint threshold, genus belongingthe to load-lift constraints haveainobserved been previously species undescribed fly ofdance nuptial gift and durationcopulation the offor (Cumming, 1994).with Patterns consistent nuptialgiftboth givingand must aerialcopulation; carry males the combinedfemale loadof the mass of the nuptial gift they offer and the mass of the female with whom they mate. males who are at the upper limit of their load-carrying capacity may face a trade-off between size (and importantly, the mass) of the gift and female are under selection to be maximized, (toimprove offspringnumber, assuming largest the females arealso the mostfecund). Ifthe offerthat largenuptialgifts nutrition(to improve ofthe mates)intake tothe largestfemales likely tofunctionally constrain reproductive success. Sexual selection should favour males load-liftConsequently, limitations in species with nuptialboth gifts and aerial copulation are giving species because a male must carry the mass of both gift and mating partner. 1998, Loading al.,1990). et Svensson are constraintsin potentially intense nuptial gift anin with insemination advantage successand sperm (Thornhill, competition 1979, Vahed, largercases, thelonger nuptial the and copulationthe gift, longcopulations lasts, covary an aerialduring copulation (Kessel,1955, Marden, 1989,FunkTallamy, 2000).& mostIn 2014). Following nuptial giftexchange, many male support the female and her meal males providewithfemalesa nutritious al.,et 1983,(Thornhill nuptialgift Lewis Alcock, & This by is protectedarticle reserved. Allrights copyright. subfamilyDance flies from (Diptera:the ) with speciesinclude There are many species that whilemate flying, and in some of these taxa Hilara sp. Hilara (Marden, 1989). In that study, wild caught mating pairs were were bythe andfunctionally constrained mass offemale Accepted Article both in evolutionaryorigin oftheirterms (e.g. Funksee Cumming,1994, 2000) & Tallamy, longicauda current In role of the study,investigate wethe functional constraintsin the evolution non-randomof in patternsmating assessed in any ornamented species of dance fly. males during copulation. To aerial hasthis alternative ourknowledge,been never formally largest mating for females the success by mightload-lifting beproduced constraints on al.,2016, et Murray2018).et al., Anintriguing lower isthat alternative thanexpected the (over-) ornamentation by females (Fitzpatrick et al., 1995, Chenoweth et al., 2006, Herridge oncauses, includinghave severalof the constraints elaboratefor malechoiceoperation adorned highest having females mating success.the Thisapparentcould inconsistency 2000), Wheeler(2012) etal. showed intermediately stabilizing selection onornamentswith advantages for femaleslargeornaments with (e.g.LeBas 2003,et al., Funk &Tallamy, 2003, 2008, al., et Bussière al.,etWheeler 2012).somestudieshave While shownmating andformselectionof strength (Sadowski 1999,et Funk al., &Tallamy, 2000,LeBas et al., studieson offlies dancesuccess, sexual selectionfemale report in remarkable variation the In spitestrong of betweencovariance predictedthe and mating expression sexual trait 1961, LeBas etal.,2003,Wheeleral.,et2012,Funk Tallamy, 2000, & Murrayetal., 2018). appearcharacters that improve to(Cumming, to theirattractiveness Collin, males 1994, bymalesprovided have andfemalesa evolved series ofremarkable secondary sex Presumablya consequence, as thefemales inmanyspeciescompeteintenselyfor foodgifts hunt, and must obtain their dietary protein from male giftsnuptial (Hunter & Bussiere, 2018). by presentation the ofnuptial(Cumming, gifts 1994), are butadultfemalesto known not In subfamily,belongs. this (theonlynotmating Empidinae), always almost is accompanied broadtheconsidering range behaviour ofmating subfamily among to the which This by is protectedarticle reserved. Allrights copyright. Theconstraints load-lifting of presence becomes particularlyinterestingwhen display severalornaments,which female-specific elaborate display perplexing remain Rhamphomyia longicauda, the long-tailedthe dancefly. Hilara sp. R. R.

Accepted Article might be mediated bymight be mediated behavioural initialattractiveness that interactions stage. the follow R. longicauda large ornaments could be important for initially attracting males potentialto mates within the This by is protectedarticle reserved. Allrights copyright. in selection on acting separatethe episodes two of females displaying intermediate-sized ornaments could also reflect differences in (Chenoweth al., et 2006). (2012) al. et Wheeler acknowledgedheightenedthat the success ornamented femaleswere beingrejected because ornament costreduces fecundity that ornaments were under stabilizing selection,consistent with suppositionthe that over- traits (Wheeler etal.,2012). Wheeler Instead, (2012) et al. supportfound forthehypothesis (unpaired) females found no evidence for directional on selection either of two ornamental of sectional comparison the phenotypictraitsand ofsuccessful (paired) unsuccessful onsexual selection directionalsuggesting positive However, ornamentation. a cross- are likelytoattractmale more (Funka & Tallamy, 2000, Wheeler, 2008, Murray etal.,2018) nuptialgifthe hasprovided (Funk &Tallamy, 2000). carrying pattern inafigure-8 female the orwhile circular she stereotypically feeds onthe pairs that transfercopulationnuptial gift,successfully themalethewing, with onthe occurs complete, and each returningto the swarmmating (R.Murray personalobservation). For transfersare unsuccessful, withthemaleandfemale separating pair before is formation involvestransfera lossforaltitudeof typically and the somepair, initialattemptsgiftat transferring nuptial the themale giftfromfemale the to astheytheexit mating This swarm. from below. When mated pairs first form, they engage in the apparently cumbersome task of Males enter mating the swarm preycarrying a asanuptial item approachgift, and females pinnateleg scalesabdominaland inflatable sacs (Newkirk, Funk 1970, 2000). &Tallamy, mating dusk atandswarms dawninwhich females displaytwo sex-specific ornaments: andfor theirrelationshipmating success with (Wheeler etal.,2012). Previous studies have shown that mating swarm (Murray swarm et actualmating al.,2018), the matingsuccess ofa female R. longicauda R. longicauda females with the largest ornaments mate pairing process. While R. longicauda R. forms Accepted Article This by is protectedarticle reserved. Allrights copyright. randommating. assumes observe alower load-lift ourdata ratiofrom compared simulated nullthe to that distribution constraint contributesnon-randomthat to we hypothesized matingpatterns,would we that ratios (total wingarea). masscarried/male If a load-lift wildexperience inthe males females.we Third, performed permutation aacreateto load-lift ofmean nulldistribution test wholargeload (maleswing heavy relative are lighter wingsize)wouldpairwith their to wingsto would largerwith females,with pairbe able massivewith andthat more a males aerialcopulationduringwing hypothesized we traits.that and amale’sown Here, males correlation.negative we a tested forSecond, relationship between the total masscarried and hypothesizedmassmassand nuptialconstraintload-lift that a gift would a in result constraints loss ofaltitudeduringpair andaerialformation) copulation. massfemales be may carry moreduring difficultto prey transferobserveoften (whenwe a covary,tend albeitto imperfectly,with increased mass;Wheeler, attractive,high- 2008), femalesIf displayingtheornaments largest heaviest(because arealsothe largeornaments range of possible partners to a subset of males (Bussière et al., 2008, Wheeler et 2012).al., individuals,and despitebeingmoreattractive (Murray etal.,2018), their mass the constrains ornamented females(Wheeler al., 2012), etare thesefemalesis that oftenheavier out ona large fractionof thepaternity share (Parker, 1970). beforemale’s ejaculatelong ovipositionand fertilization causing potentially amaletolose obtain toandThesenuptialfurthersubsequent meals. amatings byfemales coulddisplace knowing that such females are likely to mate repeatedly both because they are attractive maturity, ovarian of stages early at females ornate overly avoid to prefer might Males In the currentIn study, wethe performedtestto analyses three load-lift for functional An additional alternative depressed the successfor hypothesis ofmating over- Rhamphomyia longicauda . First, we tested. First, arelationshipbetween we for female Accepted Article This by is protectedarticle reserved. Allrights copyright. near Glen Williams (Halton Co., Ontario, Canada: 43 Sample Collection Methods mass and/or eggmassand/or number/size. not consumed (or mated nuptial gift) captureto priora because itchange could a female’s inspermspermatheca;account we wanted the potential to having effect any for offemales measures. also We recorded each formating status presence asthe female or absence of could relate measures of fecundity (egg number) and mass to female ornamentation eggs.mature We ornamentation measured legscales)(pinnate wesofemale eachfor that eggato measure length(asproxy for egg quality)andcount of number the having more, or higher quality, eggs (e.g. Bonduriansky, 2001), we dissected each female’s Morphological Measurements within hoursfreezer two following collection. sometimes dropped theupon gift capture. Allsampleswere frozenatlaboratory -20C ina weretubes. unable We allnuptialto retrieve giftseach from pairbecause mated the female net. sweep entomological placedpairs We gifts andtheircollection in individually nuptial 2018).swarming eachcollected At we event, an pairsfromusing mated nuptial flight al., 2007, Gwynne & 2002,Bussière, Wheeler et al., 2012, Murray et al., 2017, Murray et al., been used for several previous studies on We collected collected We maleBecause choice mate largerforis females typicallythose also linkedto females R. longicauda samples onthebanksRiver inJune Credit ofthe 2012 R. longicauda o 41’117”N, 79 41’117”N, (Bussière Gwynne et etal., 2008, o 55’34”W). has Thissite Accepted Article regression ofoocyte countsize against and female mass. mass was an accurate measure of fecundity in abdominalornamentdeflatedwas measure.impossible to and Toassess whether female onornamentation because our mass;female specimens frozen, were theinflatable covary with ornaments.female We performed a linear regression of female leg on pattern Female ornamentation and mating status one damageof side, to onlyundamagedthewe measured side. measured and right both sides and left took mean.When the thiswas not possible because summedleg area of mid andhindfore,the legs Foreach individual. of pairedcharacters we frozen, adult females.We measured female ornamentation (pinnateleg scales) asthe abdomen.dissected the Theeggsare immediatelydissectedin the obvious abdomensof acountobtain number ofthe eggs within ofeach (as female proxy afecundity) for we measurements were collected as proxies for body size and/or ornamentation. To to assess male fly traits withassociated lift and flight endurance (see below), while leg and of length egg eggsperfive (see individual Figure Wingwere measurementsS1). collected pinnateleg scalesin the female, see area, details), belowfor and number wing the ofeggs of(centre theneck abdominal-thorax centre of the to the area connection), leg (including together),lengthpoint wing (the longest theaxillary apex), area from the to thorax length measurements:the leglength following morphological (femora andtibialengthsummed dissected each frozen specimen and photographed only the body part of interest. We took softwareusingImageJ (Abràmoff For etal.,2004). we morphologicalmeasurements, microscopecameradissecting attached a to and microscope measured digitalimages the This by is protectedarticle reserved. Allrights copyright. In order a for loadingconstraintcontributeto to stabilizingobserved the selection photographedWe morphologicaltraits usinganAmScope digitalMA500-CK R. longicauda ornaments (Wheeleret2012),female al., mass must positively R. longicauda , we also performedlinear we a , Accepted Article copulation (to improve sperm transfer success). improve pairingbut success), endurance mightflight be important sustainedaerial for might expect maneuverability tobe more important during initial nuptial gift transfer (e.g., to maneuverabilitya 2002).in tomales lowaspect with relative Dudley, (reviewed ratio We a high AR (relatively long, narrow wings) will have better flight endurance but lower byconstrainedto beof and the theycandisplaying females with. copulatemass carry Males comparedDudley,(reviewed withalowWL to and in are 2002)males therefore likely more displaying2014). Males ahighWLlift (relativelymass largewillsize)winghave less to wingspan squared of the (e.g.wing area to seeBerwaerts Gyulavári 2002, al., et etal., mass: wingload(WL),the ratio ofmalemasswing to and aspect area, (AR), theratio ratio Male loadingmeasures nuptialgift mass possible, when frozen specimenshours within 24offrom collection. underestimatingnumber the females. virgin of We recorded male and, mass, female mass sperm from the male she is paired with during collection, and consequently we may be potentialaccount for this of We source variation. note thatafemalehave may received change their behaviour when they are hungry (e.g. Sandhu et 2018),al., we wanted to female had(and matednuptial consumeda gift)previously. Because individuals can (paleopaquesubstance) orabsence(empty) of tospermwhether determine not or the dissected the spermatheca (sperm storage organ) from females and recorded the presence This by is protectedarticle reserved. Allrights copyright. computedWe two insect’s loadon indicesfrom ofthe its an wings arising own body To assess a female’s previous mating behaviour (and nuptial gift consumption), we Accepted Article the total aerialthe masstotal (g)(mm unit per combined mass male, ofthe and female gift.theterm nuptialFinally, ‘load-lift ratio’ to refers ourWithin permutation (see tests below),weuseterm the aerial ‘total refer to mass’ the to mass of the female and, where possible, the nuptial gift) by a male during aerial copulation. This by is protectedarticle reserved. Allrights copyright. models we examinedmodels we variance inflationfactors ‘vif’using (VIF)carthe function in the one of several alternative indices of underlying phenomena. To test for collinearity in our estimates estimates over strict partial of effects, of some especially since our traitsrepresent (Morrissey Ruxton,& 2018), but we favoured model simplicity and confidence in parameter which coefficients.of Theproblematic to model extent varianceinflation is iscontroversial of aretraits related, functionally and such correlations can pose problems for the estimation come from fixedmodels. little variation among observations (<10%). For simplicity, the results we report here all resultsfrom mixed and models same; were fixed date qualitativelythe account totendedfor as“date”each thatcollected arandomsample was a in effect mixed foundbut model, the requires disrupting theswarm,setmodels statistical of each we effectoffor tested an for can vary (Wheeler,2008),temporally and tomeasure trying a covariate composition as interpretability of coefficients (Schielzeth, 2010). Because independentvariables werealso beforez-transformed improve analyses to the measurementswereroot cube transformed statisticalpriormodels. All toinclusioninany Statistical Analyses Below weuse the carried’ ‘totalmass term (TMC)torefertothemasslifted (i.e. the Area were andwings)measurements square root (legs transformed and mass High positive correlations betweenHighpositive correlations phenotypic traitsare expectedwhenever groups 2 ) of wingareaduring ourpermutation tests. R. longicauda swarm composition Accepted Article This by is protectedarticle reserved. Allrights copyright. area), AR (wingspan andresponse, morphological male measures forWLthorax length, mass/wing (male male Gyulavári etal., 2014). We fit the total mass carried during aerial copulation as the insect andconstraints flightperformance (Berwaerts etal.,2002, Peckarsky 2002,al., et measures), predictorfitting variables previouslythat have been shown to influence loading subset of observations including a nuptial gift as well as thelarger dataset with only female aerial copulation and male wing traits. We performed linear models on both datasets (for the giftinvestigated mass. Second,we therelationship betweenthea loadmalecarries during longicauda limit correlations betweenphenotypic possible.traits as much as wantedbetween for potential used we predictor wing variables, the collinearity to as traits three.noteis We that this aconservative cut-offVIF (Zuur et al.,2010), but giventhe variable highestwith value and VIFthe all reran modeluntilvaluesthe below VIF were package (Fox, 2011). our IfanyVIF in linear weexceeded model removed three, the pairing total the carried for (TMC)mmmass per resampling generate methods null theexpectationsto assumption underthe random of copulation wasless than expected by chance (i.e. if mating was random). We used hypothesis. Third, we tested whether observed the carriesa mass aerialduringmale that superior of measure by totalmass estimated carriedthe testing male forthe constraintthe she feeds. Therefore, the combined mass of both the female and the nuptial gift is a proportion how to has thefemale fed. much mass Similarly,thefemale’s willincrease as dancenuptialthe fliesfeedon gift duringcopulation, willgift massoftheinthe decrease carried to male wing area. This exercise produced 9999 means from the null distribution, afemalemasswith andmass, nuptial gift andthenthe recomputed of mean mass total ratio procedure,our we resampling randomly pairedareamale eachdataset inour measure wing We performedWe aseriestest analysesto of evidence for constraintsloading of in mated pairs. wetested First, mated arelationship for mass between femaleand nuptial 2 /wing area) and area)/wing as predictor winglength variables. female Because 2 of male wing area.male of For every iteration of R. R. Accepted Article of previousof activityinanymating analyses. further fractionourof sampleshad no spermthat inthe proxywe did spermatheca, notinclude this foursure femaleswere that collected during their first matingevent. Becauseof small the andspermatheca,while we recognize that underestimate,an be this might wecan onlybe traits can be found in Table 1. Only 5% of the collectedfemales had no sperm in their gift during collection. Summaries of our morphological measurements for female and male collected theirnuptial wealsogifts.those pairs Thepairs remaining dropped 55 nuptial the This by is protectedarticle reserved. Allrights copyright. Results underexpected random pairing. actual TMC by wild random distributionofloading lowerratios. than A that wouldindicate expected ratio the 2.5%actualwouldthe lower datasetfall aof (for off cut within significance p<0.05) of the mating patterns observedin constraintsload-lifting nuptialcollected gifts.If without important are non-random the for procedure butomittedof advantage mass nuptialgiftnumerous themore take to samples couldhavecollected samples through arisen pairingrandom We innature.repeated this assessedand itsrankamong of10000 astotal distribution the probabilitythe our that hadno bearingpair on formation. We observed added then meanour these to 9999 trials, femalegiftareaandwhere mass, malewing mass, assigned wererandomly and therefore We collected 112

R. longicauda R. longicauda R. longicauda males during males aerialis lesswouldthanbe copulation mated pairs from the wild in June 2012. For 57 of , we might, weexpect ratios load-lift the that our from Accepted Article This by is protectedarticle reserved. Allrights copyright. mass pairsin mated F (estimate=0.31±0.12, TableWe ±0.97%;14.26 nuptial 1).found the that massgift positively covariedwith female 1A; estimate=0.12±0.016 F ornamentation.or We that foundornamentation masspositivelyfemale predicts leg (Figure massFemaleornamentation fecundity and correlates with 9999 iterations without resampling and found a null distribution of load-lift ratios hadthat a Third, we performed a permutation test determineto nullthe distribution of mean loads a male might carry if foraging and mating were completely random. We performed the absencenuptial of gift F (Figuremeasure estimate=0.62±0.12, 2; S1).and We alsofound highhad athat males WLheavierpaired with in both females male’s WL (male area)mass/wingdatasets both for 2 nuptialTableswithout (withand gift; onlythethat morphologicalpredicted that trait TMC the duringaerial copulationwas a found that the nuptial gift accounts for 4-40% of the total mass carried by a male (mean: Testing for load-lift constraints positively covaries with positively fecundity covariesfemale with (Figure 6.029±1.579,estimate= 1B; F estimate=0.86±0.19, F and for the subset of data where nuptial gift masses were available (Figure S2; P<0.001) oocyte butnot (estimate= F 8.45±8.31, size 2We tested whether or not or2We whether covaried tested mass female fecundity, oocyte female with size,

Second, we tested for a relationship between TMC and male wing traits. We found First, we First, a fortested femalethe relationship between mass andthenuptial gift. We (1,56) =19.6, P<0.001). =19.6, (1, 105) =58.35, P<0.001). We also found that female mass mass female that found also We P<0.001). =58.35, (1, 56) =6.85, P=0.01). (1, 105) =1.035, P=0.31).=1.035, (1, 105) =26.95, P<0.001) P<0.001) =26.95, (1, 105) =14.33, =14.33, Accepted Article longicauda functional constraints arenon-randomcontributing tothe patternsmating inobserved differratio didnot thenulldistributionfrom (Figure 3) providingnoevidence load-lifting that ratios when random foraging and mating are assumed. We found that our observed load-lift also copulation. performedWe assessnulldistribution permutation teststo the of load-lift covaried positivelymass withtheof and nuptial female bya carried gift male aerialduring non-random patterns.mating We nowing that but found load, wing othermale traits, male longicauda cumbersome for duringmales tocarry aerial copulation. We tested wild-caught areclass ofornamentsor are isthat heavy too less frequently mated females these ornaments (Wheeler al., et 2012). Onehypothesisforwhydisplayinglargest females the suggested that femalesmorelikely were they tomate displayed intermediate-sizedif This by is protectedarticle reserved. Allrights copyright. Discussion 0.607mean of retrieved the same result: the null distribution of wing loads had a mean of 0.538 inthe loadmeasurement thepotential that (so sampletodraw was larger) from and distribution0.24). (P = performed We asimilar test includedpermutation without nuptial gifts

is 0.606 mg/mm observed Previously patternsselection ofsexual in 2 (Figure 3B) while our data had a mean of 0.535 ± 0.022 mg/mm . pairs mated foraload-lifting functionalthatconstraintbe might to contributing ± 0.016mg/mm 2 , which was not significantly smaller than expected by our null 2 (Figure 3A). The load-lift mean ratiowe observedin nature ± 0.013mg/mm Rhamphomyia longicauda 2 (P=0.13). (P=0.13). R. ± 0.014 0.014

R. Accepted Article This by is protectedarticle reserved. Allrights copyright. patterns of mating wherein other carried the taxa male the duringcopulation.female Our Marden (1989)concludeda similarfunctional that mightbeimportant constraint an influence differenceobservedin load-liftthe compared ratio asimulation to of pairings.random between relationshipnegative nuptial and mass female giftmass, significant anda males doexperienceload-lift a during in-flightconstraint copulation.Marden(1989) a found longicauda load-lift of mean found ratios. weagain, load-liftingevidence no Once a in constraint for expected we copulation, our observed ratio load-liftmean less than tobe nulldistributionthe mating andforaging. If amales experienceload-liftconstraint functional during aerial distributionload-lift of mean ratios(total mass carried/malearea) wing assumingrandom correlatedThird, with performed mass. female permutation we a test tocreatea null loaddidnot trade-offwith massof positively was instead the butwith, hepairedfemale the females. We found that female mass did not covary with male wing length and male wing experiencinghaveconstrainta load-lift might smaller and wouldcarrywings, smaller by amaleduring copulation wing and ahypothesized We male’s traits. that male’s relationship. negative we Second,a tested for relationship between masstotal the carried constraint havemight totrade-off femalemasswith nuptialmassgift andexpecteda male during aerialcopulation.We hypothesized loading thatamaleexperiencing a we tested arelationshipfor thefemale between by massandnuptialmassgift carried a evidence that a load-lifting constraint during aerial copulation exists in 1989). performed We test analysesto three forload-lift functional constraints andfound no copulation, duringbearflight maleof to leavingthe load entire the weight (Marden, the he onshe feeds is provides. that nuptial gift It unlikely the contributeis ablethe female to to Testing for load-lift constraints Previousworkload-liftingdifferent constraintsina on fly dancespecies showed that R. longicauda ; theobservedload-lift; ratios differ didnot null fromdistributions (Figure 3). mated pairs copulate in-flight with with in-flight copulate pairs the malecarrying mated female the while R. longicauda . First,. R. Accepted Article ornaments, female-biased OSRmeasures) and ofsexual andload-lift competition Direction constraints 1989)suggests males that competestrongly for to access mates. swarms of the non-ornamented ornamentation (Funk Tallamy,2000, & et2018)Murray al., whereas themajority-male 2016).et Herridge al.,2012, et In al., bequite differentOring, (Emlen & 1977,Clutton-Brock,2009, Hare2018, &Simmons, Kokko cause higher energyexpenditurehasbecause from carry he male the ofto the mass the a siresoffspringmale (Parker 2010).Longer& Pizzari, copulation in-flight durationwill Simmons & Siva-Jothy, 1998, Simmons,2001,etal., Schluter 1991) and the number of limitsflight in-flight copulationduration,number thus sperm of transferred (Thornhill, 1979, measures;1989). Marden,One hypothetical effort explanationisthatthe load aincarry to operational sex ratios, sexual selection and mate choice in the two species et al., 2008, Murray et al., 2017). Given variationthe in ornamentation and contrasting longicauda ornaments. Second,of swarms to (where load-liftingobserved) were functionalconstraints in ways:indiffer First, two contrast This by is protectedarticle reserved. Allrights copyright. in mass by no providedata support wefor but hypothesis,evidence this didfind assortative for mating Marden,1989, Funk 2000).However, &Tallamy, matingin mating gift-giving,swarms,(Collin, copulationCumming,1961, andin-flight nuptial 1994, displayelaborate that mating lek-likeincluding iscommonin danceflyspecies,behaviour heavierfemales (Table 2 Figure2). Both R. longicauda We found contrasting patternsof load-liftconstraints between are consistently female-biased (Funk & Tallamy, 2000, Gwynne, 1991, Bussière R. longicauda (Funk & Tallamy,2000), ; males with a high winghighwith load(heavy a males ; theirpaired wingarea)for with Hilara Hilara sp. R. longicauda

sp . experiencesthat constraints. load-lifting (Marden, Hilara are male-biased (Marden, 1989) while those of R. longicauda

Hilara sp. Hilara sp . does. not have female-specific there ismale there choicemate andfemale and (no ornaments, male-biased OSR OSR male-biased ornaments, (no R. longicauda Hilara sp. R. longicauda (Marden, 1989) and

are expected to Hilara (female

sp.

R. Accepted Article fecund female. fecund female. mate quality), on average choosing an ornamented female will also result in a heavier, more andornamentationregressed on fecundity mass female (arguably twomeasures of female (Bonduriansky, 2001).variation Whilelinear thereis aroundestimates the forboth in with covaries(Figureshow that positively ornamentation1A)mass andfecundity (Figure 1B) ischoicesuggest mate ofresponsible observed for matingin patterns assortative-mating patterns than mate choice. Our data align well with this conclusion and (1989)functional concludedthatconstraintswereexplain less far likely to observed comparativeafemales. However, studyacrossdiverse Crespitaxa, following proposed as a possible tomechanism explain this pattern in species where males carry Thompson, Ridley 1987, 1979,& Speakman et 2007)al., andloadingconstraints have been Assortative-mating by body mass has been observed in diverse taxa (Alcock & Gwynne, be heavier but not larger in linear dimensions than unpaired females (Woodhead, 1981). and S2). However, the two findings need not be contradictory because paired females can al.,2008),et results assortative-mating ourby mass acrosssuggest pairs(Figuresmated 2 by mating Assortative mass bettera not formale thanmatingat all. copulationdurationtransferring (and potentially of asmallnumber undoubtedlysperm) is This by is protectedarticle reserved. Allrights copyright. asinopportunities limited,such are paternity.to ensure However,inspecieswhere choosier arefemales sexandmating the unableabilitiestheir load-lifting areif itmeans transferto they as much necessarysperm as female-biased swarms, mating males mightreject choosetoupperthe females at of limits female and the nuptial gift for the duration (Marden, 1989). Therefore, in species with R. longicauda. While a previousWhile a size-assortative studyfound in no mating Therefore, high-condition maleschoosing couldbe mates fecund more Hilara sp ., aheavymating with female for ashorter R. longicauda R. R. longicauda (Bussière . We Accepted Article not always reveal underlying tradeoffs becausetradeoffs always revealunderlyingnot vary animalsin bothacquisition resource Wheeler,Funk 2008,Tallamy,& 2000). However,correlationsamong phenotypic do traits aconsistently positive see relationship between ornamentation and fecundity (Figure 1B; associated with ornamentation in importantnote, however,to itcurrently that is unclearwhethercost therea isfecundity (Arnqvist Rowe,& 2005), and preferences for more modestly adorned females. It is resistance in ornaments, resulting antagonisticallyfemale seductive to male characters Females may be deceiving males about their potential fecundity by displaying larger ornamented mating inobservational studies. success However, value. signalbe expression ornament correlated easily might previous with of of actual mating success rather than ornaments high of risks sperm competition). preferencesSuch would more sensibly be based on cues potentialAnother is explanation males avoidparticularlypopularthat (who pose females atornamentsexpense the offecundity (Fitzpatrickal., etal.,1995,Chenoweth2006). et is females avoid moderate which with males predicted if ornaments, in overinvestthat observationremains unclear, choicemateactive butonepossibility bymales is forfemales This by is protectedarticle reserved. Allrights copyright. highly-ornamented female thereforeexplainconstraintsfunctional conclude donot thereduced that ofmating success limitedmales are during load they by carry massofthecanthe aerial and copulation, were contributing non-randomto matingpatterns. foundevidence We no that R. longicauda Male mateornamentsfemale-specific choice for An alternative explanation for why we observe selection intermediatelyfor We set outsetWe totesthypothesis by the raised sexualselectionin two studiesmeasuring R. longicauda (Bussière et al., 2008, (Bussière etal., Wheeler 2012) et thatfunctionalal., loadingconstraints females is antagonistic 2005).Rowe, females coevolution (Arnqvist & R. longicauda R. R. longicauda (Wheeler 2012). al., et What doesexplainthis females. Here, and in previous studies we per se per , lestundermine, they ornamental the R. longicauda

Accepted Article the previously observed non-random matingpatterns in findings suggest male mate choice for ornamented females as a probable explanation for theassortative pattern: opposite we saw positive matingforconclude mass. We our that mating pairs andtestsandno permutation found constraint. aload-lift evidencefor Instead, dance fly Conclusions 2000,Tallamy, Murray etal.,2018). 2012) over the large-ornamented females they are initially attractedmost to (Funk & mating process malesare choosingintermediately-ornamented (Wheeler females al.,et Throughapartteasewe couldalso male trials when thetimingof during choicemate the directlywouldallowus to assess choiceornamentsfor mate male female in successfullaboratory-controlled However,future,in the species incaptivity. if trials mating species. couldtherethatfecundity bea ornamented costthat payrelativetofemales non-ornamented unornamented in fecundityof inflatable abdomenornament, orother female is the size abdomen amuchbetter predictor This by is protectedarticle reserved. Allrights copyright. comparison between acquisition andallocationdirectly(Houslay & 2012).Bussière, We in athat though, note, deallocationand (VanNoordwijk Jong, 1986), & we are unableto and measure resource

We aload-lift for tested constraint inshaping non-randomin patterns mating the At present we have not been successful in rearing or maintaining any dance fly Rhamphomyia longicauda. R. longicauda R. R.

sociabilis and We used We morphological wild-caught measures from Rhamphomyia sociabilis (Figure 9 in Funk & Tallamy, 2000) suggesting R. longicauda , a, dance flywith no . R. longicauda . Accepted Article This by is protectedarticle reserved. Allrights copyright. *N=57 and males (N=112). TableMeans andstandard 1. errorsmorphological for traits ofpaired Tables offromfunding University the Stirling HorizonScholarship RLM to andNatural Science ResearchEngineeringof Canada Council PGS as toRLM wellas providing useful feedback on this study. This research was supported by funding from the K. Vahed,two P.Lee,anonymous reviewer Acknowledgements Trait wing length (mm) length wing (mm) area wing squareroot thorax length (mm) females win (mm) area wing squareroot thorax length (mm) (mm) size oocyte count) (oocyte fecundity squareroot hindleg ornament area (mm) mm) tibia: and (femora length leg hind mass (mg) nuptial gift mass (mg)* mass gift nuptial squareroot hindleg area (mm) mm) tibia: and (femora length leg hind mass (mg) males g len

g

th (mm) th

s and the Bussière and Gwynne labs for Mean 1.57 66.3 2.23 6.46 4.63 6.74 1.60 0.80 1.29 5.77 2.68 6.31 0.31 71 98 Standard Error R. longicaudaR. 0.023 0.012 0.014 0.040 0.021 0.012 0.068 0.008 0.033 0.085 0.039 1.67 0.36 0.13 0.01

females Accepted Article This by is protectedarticle reserved. Allrights copyright. (mg). female mass longicauda Male Tablebya wingtraits predict 2. that mass carried thecopulation maleduring in (wingspan ratio aspect (mass/wing area) load wing male length wing male intercept Trait

2 /wing area) . Estimates are standardised coefficients of a multiple regression. Mass carried is

Estimate -0.056 0.62 0.18 5.6 Standard Error 0.13 0.13 0.12 0.19 28.82 2.39 4.76 1.15 z

<0.001 <0.001 0.68 0.14 R. R. p

Accepted Article Figures Rhamphomyia longicauda Figure 1.Female body mass correlates andornamentationpositively with in andfecundity (egg count) square-root female leg ornament area (mm)

20 40 Fecundity60 80

squarerooted2.50 2.75 female 3.00 leg ornament3.25 area (mm)3.50 B A 2− 2 1 0 −1 −2 2− 2 1 0 −1 −2 standardised cube-rootstandardised (g offemalemass standardised cuberooted femalestandardised mass (g^1/3) . A) Linear. A) regressionofsquare-rooted female leg ornament area Standardised mass (g) female 1/3 )

Accepted Article This by is protectedarticle reserved. Allrights copyright. (mm) onstandardised cube-rooted femalemass(g standardised body female mass 6.029±1.579,(estimate= F P<0.001). B) Linear regression of female fecundity (measured as oocyte count) against red). shaded area represents the standard error measure around linearthe estimate (shown in 1/3 ; estimate=0.12±0.016, F estimate=0.12±0.016, ; (1, 105) =14.33, P<0.001).=14.33, The (1, 105) =58.35, Accepted Article This by is protectedarticle reserved. Allrights copyright. inred). (estimate=0.00062± F 0.00012, Thearea shaded represents thestandarderror measurearound thelinearestimate (shown longicauda Figure 2. Linear regression of the female mass (g) on standardised male wing load in

0.002 Female0.004 mass (g)0.006 0.008 mated pairs.isareaas calculated Wing mated load details). mass/wing fortext (see 2− 2 1 0 −1 −2 (1, 105) Male wing load =26.95, P<0.001). =26.95, R.

Accepted Article References References wing area (mm assuming random mating andmaleforaging. Load-lift ratiois the totalaerial(g) mass male/ Figure oftheload-lift 3.Nulldistribution ratio from permutation with9999tests iterations, mean of load-liftmean of the ratiofromdatadisplayed ourdashed is as lines (A: P=0.24; B: P=0.13). randomised female values excludingmass (seetext gift nuptialThe details). for observed male mass values and (A) randomised gift andfemale nuptial massvaluesor (B) B A 2 ). The totalcarried aerialmass is by up male the non-randomisedmade of

Accepted Article This by is protectedarticle reserved. Allrights copyright. an selection, 1977.sexual L.W. Ecology, Emlen, S. &Oring, T. 2002. R. Dudley, Alcock,J. & Gwynne,D. T.1987. Abràmoff,M. D., Magalhães,P. & J. Ram, S. 2004.J. Image proc Gwynne, D. T., Bussière, L. F. & Ivy, T. M. 2007. Female orname Female 2007. M. T. Ivy, F. & L. Bussière, T., D. Gwynne, Funk,D. H.& Tallamy, D.W. 2000.Courtship role reversal and Fox, J. 2011. SanfordW., Fitzpatrick, S., & Berglund, A. Endler, J. A.& Basolo, A.L.1998. Sensory ecology, receiverb Dubois, F., Drullion, D. & Witte, K. 2011. Social informationu 2011. & K. Witte, Social D. Dubois, F.,Drullion, danc of evolution the and selection Sexual 1994. M. J. Cumming, . matingin assortative of Causes 1989. J. Crespi,B. L.2005. G.& Arnqvist, Rowe, Almbro, M. Kullberg,& C. 2008. downfall The of mating: the eff Gwynne, D. T. & Buss Gwynne,D. Gwynne, T.D. 1991.Sexual competition among females: What caus Collin, J. E. 1961. Clutton-Brock, T.2009. Sexual selection in females. P.Chenoweth,& Doughty, S.F., Candolin, U. 2003. Theuse of multiple cuesin mate choice. Bonduriansky, evolutionThe 2001.R. ofmale mate choice inins Berwaerts, &K.,Ae H. VanDyck, Bussière, L. F., Gwynne, D. T. & Brooks, R. 2008. Contrasting s Contrasting 2008. R. Brooks, & D.T. Gwynne, F., L. Bussière, systems. systems. University Press,Princeton, New Jersey. (Hymenoptera, Tiphiidae). international Biophotonics Trends in Ecology &Evolution spider webs arole-reversedin swarmingfly. dance long-tailed dance fly, Thousand Oaks, CA. Linnean Society reproductive success restrict sexual selection processes. decisions: a game model. theoretic Empidinae). Empididae; (Diptera: 980-1000. UK. Cambridge, Press, University Cambridge Sociobiology and flight muscle ratio on the escape abilityof aPierid butterfly 1082. 1082. reversal? preferencesfacilitate honest female ornamentation? 595. Loew Empididae).(Diptera: ideasand evidence. Functional Ecology USA. and females in a role-reversed swarming dance fly, performance? An experimental test with the butterfly Science Science The biomechanics of insect fight: form, function, evolution Trends inEcology & Evolution British Flies VI: Empididae Part 2: Hybotinae, Empidinae (except Flies VI:Empididae (except Hilara) Part British Empidinae 2: Hybotinae, ière, L. T. 2002. Femal T. ière, 2002. L. 55 197 63 An R Companion to Applied Regression, : 251-260. 16 : 413-420. Biological Reviews : 215-223. 215-223. : Rhamphomyia : : 484-491. Sexual Conflict Sexual Rosenqvist, G. 1995. Ornaments Ornaments G. 1995. Rosenqvist, Kokko, H. 2006. Cannon-directi 2006.Kokko, H. Courtship feeding matechoand rts, P. 2002. Does flight morph flight Does rts, P.2002. 11 Australian Journal of Zoology Journal ofEvolutionaryBiology : 36-43. 13 : 415-420. : The Canadian Entomologist Canadian The Behavioral Ecology

e mating swarms incr e mating swarms longicauda .

PrincetonUniversityPress, Princeton, NJ, 76 6 : 118-121. : 118-121. : 305-339. Behaviour Behaviour Animal . Animal Behaviour Behaviour Animal Rhamphomyia longicauda Rhamphomyia Animal Behaviour Behaviour Animal Biological Reviews Ecology Letters iases and sexual selection. se maylead to maladaptive d the evolutionof mating exual selection maleson

essingImageJ. with Pararge 23 Animal Behaviour or offspring: costs to to costs offspring: or nts hinder escape from deceptive signals in the inthe deceptive signals ect of mate-carrying and ology relate to flight onal male onalmating male Second ed. e fly mating systems systems mating fly e 35 ice in Thynnine wasps Biological Journal of the ects: a synthesis of of synthesis a ects: : 225-231. ease inrisk a : 451-458. . 21 es es courtship-role Behavioral Ecology 126

: 1683-1691. : 77 aegeria : 3-11. . : 907-920. 59

Princeton Princeton Sage, 9 : 411-421. : 73 : 179-184. : . 78 : 1077-: 38 : 575- : :

.

Accepted Article Marden, J. H. 1989. Effects of loa Marden,Effects H.1989. J. Koene,J Vahed,K., M., S. Lewis, L. M. 2003. Hockham, LeBas, &G. R. R., N.a Ritchie, Nonlinear McLachlan, A. J. & Allen, D. F. 1987. Male mating success in Di in success mating Male 1987. F. D. Allen, & J. A. McLachlan, & Kokko,H. H.,Jennions, Klug, E.Kessel, mating L.1955. The Murray, R. L., Herridge, E. J., Ness, R. W. & Bussiere, L. F. 2 F. L. Bussiere, & W. R. Ness, E.J., Herridge, L., Murray,R. D. G. Ruxton, & B. M. Morrissey, Hunter,F.D. L. & Bussiere,F. L. 2018. Comparative evidence s Houslay, T.M. & Bussière, L.F. (2012) Sexual selection and li F. L. Bussiere, & T. D. Gwynne, L., R. Murray, E.Herridge,J., evo its and selection Sexual 2018. L. W. Simmons, & R. M. Hare, L., A., Therry, H. Gyulavári, This by is protectedarticle reserved. Allrights copyright. Gwyn J., L.,Wheeler, Murray,R. Peckarsky, B. L., McIntosh, A. R., Caudill, C. C. & Dahl, J. 20 I fitness. offspring mating and L.(1983)Non-random Partridge, 2010. T. Pizzari, & A. G. Parker, cons competitionevolutionary its and Parker,Sperm 1970. G. A. Newkirk, M. R. 1970. Biology of the longtailed dance fly, Ecology Ecology Biology Letters Lehmann, G. U. C. 2014.Emerging issues theevolutionin of ani Biological Sciences selection on honest ornamentation. female size. size. investment. operational sexratio, Bateman gradient and scopethe for compe Entomology Behaviour (Diptera: Empidinae)? Empidinae)? (Diptera: dan among variation mating for system responsible endosymbionts Philosophy, Theory, andPractice in Biology The Meaning Multipleof Regression and Non-Problemthe ofColli reproductive allocation in the evolution offemale ornament div Encyclopedia of Life Sciences. andparental sex roles. in animals.female Evolutionary Ecology endurance, flight-related morphology and physiologyin a scramb Sciences Sciences flies. dance in ornaments multiple Empididae: (Diptera: fly role-reversed dance size. size. a of behavior (Bateson,P., ed.) Reviews Reviews of Philosophical the Cambridge Society 1412. Empididae). (Diptera: Oikos Behavioral Ecology and Sociobiology 70 85 285 48 139 : 496-502. : 897-934. . Ecology Letters : 20181525. : 11-14. : 1425-1430. 10 . pp. : 20140336. : Biological Reviews Biological 270 Dévai, G. & Stoks, R. 2014.Sexua &Stoks, R. Dévai, G.

Baetis Cambridge University Press, Cambridge, UK. UK. Cambridge, Press, University Cambridge 28 Annals Annals Entomologicalof Society of the America Plos One Plos M. D. 2012. Unifying cornerston M. 2012.Unifying D. activities of balloon ofballoon flies. activities EncyclopediaEvolutionary of Biology : 2159-2165. . M., Engqvist, L., Bussière, Bussière, L., Engqvist, . M., Sperm competition and ejacula and competition Sperm ne, D. T. & Bussiere, L. F. 201 F. L. Bussiere, & T. ne, D. : 639-654. 2018. Multiple Regression Is N Is Regression Multiple 2018. d-lifting constraints on the constraints d-lifting

bicaudatus 15 pp. : 1340-1351. 1340-1351. : 12

Wiley. Proceedingsof the Royal Society B-Biological : e0178364. : 000-000. suggest stabilizing selection for male body male for selection stabilizing suggest 51 10 Proceedings ofProceedings the Royal Society B: : 530-537. 530-537. : Rhamphomyia longicauda Rhamphomyia . 45 Systematic Zoology Rhamphomyia longicauda longicauda Rhamphomyia : 525-567. 02. Swarming and mating 017. Are distorting ratio 017. Are sex L. F., Perry, J.C., Gwynne, D. & 2016. Diversity in mating inmating 2016.Diversity l selectionl on flight mating system of adance fly. 8. Sexual selection on fe allocation.history fe In: ptera: ptera: advantages of small nd sexual correlational upportsa role for es ofes selection:sexual te economics. economics. te ot Multiple Regressions: lutionary consequences n: n: equences. Mate Choice Mate : 453-458. mal nuptial gifts. ersity. titive titive ling damselfly. nearity. nearity. Biological ce flies flies ce 4 63 : 97-104. : Biological Ecological , Loew). : 1407- Accepted Article Wheeler, J.(2008) selection Sexual andfemale ornamentation in Svensson, B. G., Petersson, E. Frisk,& M. 1990.Nuptial gift s 20 M.D. & Jackson, J. Stewart, K., Djafarian, R., J. Speakman, 1998. T. M. Siva-Jothy, & W. L. Simmons, Sandhu, P., Shura,O., Murray, R. L. & Guy, C.2018. Worms make Zuur, A. F., Ieno, E. N. & Elphick, C. S. 2010. A protocol for C. Zuur,protocol 2010. S. &Elphick, E. F., A. A Ieno, N. Woodhead, dry Female P.A. 1981. Wheeler, Gwynne, J., D This by is protectedarticle reserved. Allrights copyright. Van JongJ. & Noordwijk, de A. Vahed, K.1998. Thefunctionnuptialof feeding in insects: rev 1983. J. R.&Alcock, Thornhill, 1979. R. Thornhill, Simmons, L.W. 2001. Schluter, D.,Price,& Row D. T. Sadowski,J. A.,Moore, A.J. & Brodie, E.D. 1999. Theevoluti Ridley,M. Thompson,& D. J. 1979. and Size mating in Schielzeth, H. 2010. Simple means to improve the interpretabili Toé, A., Diabaté, P., S. Sawadogo, 142. 142. influence on variation in life history tactics. tactics. history life in variation on influence durationthe in dance fly obesity. selection for potential the Princeton University Press, Princeton, New Jersey, USA. ( earthworm common the in foraging on starvation of effect ornaments indance a fly. In: fly. common statistical problems. problems. statistical common pennsylvanicus Biological Reviews of the Cambridge Philosophical Society UniversityPress, Cambridge, Massachusetts & London, England. insects in strategies trade-offs. Zoology of Journal Canadian BehavioralEcology and Sociobiology dance EmpisBigger snoddyi (Diptera: isn't fly,Empididae): alw Isopoda). coefficients. coefficients. 285-293. gambiae S.& Sinkins, G., F., Gibson, Ecology andEvolutionary Biology. The American journal of clinical nutrition s.s. male mating success. matingsuccess. mosquito male s.s. Journal of Comparative Ethology Ethology Comparative of Journal Proceedings of the Royal Society B-Biological Sciences male and female sexual selection and the evolution of mating mating of evolution the and selection sexual and female male Methods Ecologyin and Evolution . Sperm Sperm competition in and evolutionary consequences its the insects . T. & Bussière, L. F. 2012. Stabilizing Stabilizing F. 2012. L.T. Bussière, & . Evolution Evolution . , G. 1986. Acquisition and alloca and 1986.Acquisition , G. e, L. Conflicting1991. selecti The Evolution of Insect Mating Systems Mating Insect of Evolution The . H. K., Sanon, A., Lefevre, T

Journal ofEvolutionaryBiology 35 Academic Press. Dabiré, R. K. 2013. effects ag Dabiré,of 2013. R. effects K. -weight and female choice in choice female and -weight 96 : 192-193. MethodsEcology in andEvolution

: 1278-1283. : borealis Sperm competition competition Sperm in mechanisms insects: and 45 : 161-166. : . Ecological entomology pp.87. 51 The American Naturalist Naturalist American The Journal of medical entomology : : 380-397. 1 Asellus aquaticus Asellus

: 103-113. : University ofToronto. 86 data exploration to avoid ., Baldet, T., J., Gilles, Simard, on pressures and life history history life and pressures on 07. Assortative mating for for mating Assortative 07. ize prolongsize copulation : 316-323. : on of empty nuptial gifts in a a in gifts nuptial empty of on sexual selection forfemale iew of empirical studies. of empirical iew tion ofresources: their Chauliognathus Chauliognathus ty of regression risky choices too:the 25 e and on size 73 arole-reversed dance Lumbricus terrestris Lumbricus : 1233-1242. : : 43-78. .

Harvard 15 ays better.ays (Crustacea, 1 246 : 3-14. : 225-229. : 11-17. 11-17. : 128 Anopheles : 137- 50 : ). .