Male body size and parental relatedness but not nuptial colouration influence paternity success during scramble competition in Rana arvalis

in Behaviour
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Female mate choice promotes the development of male secondary sexual traits such as nuptial colouration, whereas scramble competition favours male traits which enhance their ability for access to females. In the explosively breeding moor frog (Rana arvalis), males express a conspicuous blue colouration during a short reproductive period characterised by scramble competition. In the present study we used controlled mating experiments and genetic markers to disentangle the effects of colouration, body size and pairwise genetic relatedness in determining paternity success. Males observed in amplexus with a female prior to spawning were larger than their competitors but did not differ from them in colouration. Polyandry occurred in 67% of the 18 analysed egg clutches, and amplecting males did not always achieve the highest siring success, most likely due to stray sperm. Successful mating pairs were characterised by higher genetic divergence between them than expected by chance. We confirm previous evidence that male nuptial colouration is not a trait selected by females, and provide evidence that male reproductive success is influenced by male size as well as genetic attributes.

Male body size and parental relatedness but not nuptial colouration influence paternity success during scramble competition in Rana arvalis

in Behaviour



AnderssonM. (1994). Sexual selection. — Princeton University PressPrinceton, NJ.

AnderssonM.IwasaY. (1996). Sexual selection. — Trends Ecol. Evol. 11: 53-58.

ArakA. (1983). Male–male competition and mate choice in anuran amphibians. — In: Mate choice ( BatesonP. ed.). Cambridge University PressCambridge p.  181-210.

ArensP.SluisT.Van’t WestendeW.P.C.VosmanB.VosC.C.SmuldersM.J.M. (2007). Genetic population differentiation and connectivity among fragmented Moor frog (Rana arvalis) populations in The Netherlands. — Landscape Ecol. 22: 1489-1500.

BellR.C.ZamudioK.R. (2012). Sexual dichromatism in frogs: natural selection, sexual selection and unexpected diversity. — Proc. Roy. Soc. Lond. B: Biol. Sci. 279: 4687-4693.

BergerL.RybackiM. (1992). Sperm competition in European water frogs. — Alytes (Paris) 10: 113-116.

BervenA.K. (1981). Mate choice in the wood frog, Rana sylvatica. — Evolution 35: 707-722.

BolkerB.M.BrooksM.E.ClarkC.J.GeangeS.W.PoulsenJ.R.StevensM.H.H.WhiteJ.-S.S. (2009). Generalized linear mixed models: a practical guide for ecology and evolution. — Trends Ecol. Evol. 24: 127-135.

BurnhamK.P.AndersonD.R. (2002). Model selection and inference a practical information-theoretic approach. — SpringerNew York, NY.

ByrneP.G.RobertsJ.D. (2004). Intrasexual selection and group spawning in quacking frogs (Crinia georgiana). — Behav. Ecol. 15: 872-882.

CrawleyM.J. (2007). The R book. — WileyChichester.

DaviesN.B.HallidayT.R. (1977). Optimal mate selection in the toad Bufo bufo. — Nature 269: 56-58.

EdwardsD.L.MahonyM.J.ClulowJ. (2004). Effect of sperm concentration, medium osmolality and oocyte storage on artificial fertilisation success in a myobatrachid frog (Limnodynastes tasmaniensis). — Reprod. Fertil. Dev. 16: 347-354.

GerhardtH.C.HuberF. (2002). Acoustic communication in Insects and Anurans: common problems and diverse solutions. — University of Chicago PressChicago, IL.

GlandtD. (2006). Der Moorfrosch — Einheit und Vielfalt einer Braunfroschart. — Laurenti VerlagBielefeld.

GomezD. (2006). AVICOL a program to analyse spectrometric data. — Free program available online at (accessed 3 August 2012).

GomezD.RichardsonC.LengagneT.PlenetS.JolyP.LénaJ.-P.ThéryM. (2009). The role of nocturnal vision in mate choice: females prefer conspicuous males in the European tree frog (Hyla arborea). — Proc. Roy. Soc. Lond. B: Biol. Sci. 276: 2351-2358.

GrafeT.U.PreiningerD.SztatecsnyM.KasahR.DehlingJ.M.ProkschS.HödlW. (2012). Multimodal communication in a noisy environment: a case study of the Bornean rock frog Staurois parvus. — Plos One 7: e37965.

GreeneA.E.FunkW.C. (2009). Sexual selection on morphology in an explosive breeding amphibian, the Columbia spotted frog (Rana luteiventris). — J. Herpetol. 43: 244-251.

HedengrenI. (1987). Selection of body size arm length and colour in male and female moor frogs (Rana arvalis). — University of StockholmStockholm.

HettyeyA.RobertsJ.D. (2006). Sperm traits of the quacking frog, Crinia georgiana: intra- and interpopulation variation in a species with a high risk of sperm competition. — Behav. Ecol. Sociobiol. 59: 389-396.

HettyeyA.RobertsJ.D. (2007). Sperm traits in the quacking frog (Crinia georgiana), a species with plastic alternative mating tactics. — Behav. Ecol. Sociobiol. 61: 1303-1310.

HettyeyA.BaksayS.VagiB.HoiH. (2009a). Counterstrategies by female frogs to sexual coercion by heterospecifics. — Anim. Behav. 78: 1365-1372.

HettyeyA.HerczegG.HoiH. (2009b). Testing the phenotype-linked fertility hypothesis in male moor frogs (Rana arvalis) exhibiting a conspicuous nuptial colouration. — Amphibia-Reptilia 30: 581-586.

HettyeyA.HerczegG.LaurilaA.CrochetP.-A.MerilaJ. (2009c). Body temperature, size, nuptial colouration and mating success in male moor frogs (Rana arvalis). — Amphibia-Reptilia 30: 37-43.

HillG.E. (1991). Plumage coloration is a sexually selected indicator of male quality. — Nature 350: 337-339.

HöglundJ. (1989). Pairing and spawning patterns in the common toad, Bufo bufo — the effects of sex-ratios and the time available for male–male competition. — Anim. Behav. 38: 423-429.

HöglundJ.RobertsonJ.G.M. (1987). Random mating by size in a population of common toads Bufo bufo. — Amphibia-Reptilia 8: 321-330.

HöglundJ.RobertsonJ.G.M. (1988). Chorusing behavior, a density-dependent alternative mating strategy in male common toads (Bufo bufo). — Ethology 79: 324-332.

HöglundJ.SäterbergL. (1989). Sexual selection in common toads: correlates with age and body size. — J. Evol. Biol. 2: 367-372.

HowardR.D. (1988). Sexual selection on male body size and mating behaviour in American toads, Bufo americanus. — Anim. Behav. 36: 1796-1808.

JehleR.SztatecsnyM.WolfJ.B.W.WhitlockA.HödlW.BurkeT. (2007). Genetic dissimilarity predicts paternity in the smooth newt (Lissotriton vulgaris). — Biol. Lett. 3: 526-528.

KalinowskiS.T.TaperM.L.MarshallT.C. (2007). Revising how the computer program CERVUS accommodates genotyping error increases success in paternity assignment. — Mol. Ecol. Notes 16: 1099-1106.

KnoppT.MerilaJ. (2009). Multiple paternity in the moor frog, Rana arvalis. — Amphibia-Reptilia 30: 515-521.

KrupaJ.J. (1994). Breeding biology of the great-plains toad in Oklahoma. — J. Herpetol. 28: 217-224.

KvarnemoC.SimmonsL.W. (2013). Polyandry as a mediator of sexual selection before and after mating. — Philos. Trans. Roy. Soc. Lond. B: Biol. Sci. 368: 20120042.

LaneJ.E.BoutinS.GunnM.R.ColtmanD.W. (2009). Sexually selected behaviour: red squirrel males search for reproductive success. — J. Anim. Ecol. 78: 296-304.

LaurilaA.SeppäP. (1998). Multiple paternity in the common frog (Rana temporaria): genetic evidence from tadpole kin groups. — Biol. J. Linn. Soc. 63: 221-232.

LiaoW.B.LuX. (2011a). Male mating success in the Omei treefrog (Rhacophorus omeimontis): the influence of body size and age. — Belg. J. Zool. 141: 3-12.

LiaoW.B.LuX. (2011b). Proximate mechanisms leading to large male-mating advantage in the Andrew’s toad Bufo andrewsi. — Behaviour 148: 1087-1102.

LomanJ.MadsenT. (1986). Reproductive tactics of large and small male toads Bufo bufo. — Oikos 46: 57-61.

MarmetJ.PisanuB.ChapuisJ.-L.JacobG.BaudryE. (2012). Factors affecting male and female reproductive success in a chipmunk (Tamias sibiricus) with a scramble competition mating system. — Behav. Ecol. Sociobiol. 66: 1449-1457.

MilinskiM.BakkerT.C.M. (1990). Female sticklebacks use male coloration in mate choice and hence avoid parasitized males. — Nature 344: 330-333.

MurphyC.G. (1998). Interaction-independent sexual selection and the mechanisms of sexual selection. — Evolution 52: 8-18.

NarinsP.M.HödlW.GrabulD.S. (2003). Bimodal signal requisite for agonistic behavior in a dart-poison frog, Epipedobates femoralis. — Proc. Natl. Acad. Sci. USA 100: 577-580.

Polo-CaviaN.LópezP.MartínJ. (2013). Head coloration reflects health state in the red-eared slider Trachemys scripta elegans. — Behav. Ecol. Sociobiol. 67: 153-162.

PreiningerD.BoeckleM.FreudmannA.StarnbergerI.SztatecsnyM.HödlW. (2013). Multimodal signaling in the small torrent frog (Micrixalus saxicola) in a complex acoustic environment. — Behav. Ecol. Sociobiol. 67: 1449-1456.

R Development Core Team (2014). R: a language and environment for statistical computing 3.1.0. — R Foundation for Statistical ComputingVienna available online at: (accessed 10 April 2014).

RiesC.SpaetheJ.SztatecsnyM.StrondlC.HödlW. (2008). Turning blue and ultraviolet: sex-specific colour change during mating season in the Balkan moor frog. — J. Zool. (Lond.) 276: 229-236.

RobertsJ.D.ByrneP.G. (2011). Polyandry, sperm competition, and the evolution of anuran amphibians. — Adv. Stud. Behav. 43: 1-53.

RyserJ. (1989). Weight loss, reproductive output, and the cost of reproduction in the common frog, Rana temporaria. — Oecologia 78: 264-268.

SchwagmeyerP.L.WoontnerS.J. (1986). Scramble competition polygyny in thirteen-lined ground squirrels: the relative contributions of overt conflict and competitive mate searching. — Behav. Ecol. Sociobiol. 19: 359-364.

SeehausenO.van AlphenJ.J.M. (1998). The effect of male coloration on female mate choice in closely related Lake Victoria cichlids (Haplochromis nyererei complex). — Behav. Ecol. Sociobiol. 42: 1-8.

SheldonB.C.ArponenH.LaurilaA.CrochetP.A.MeriläJ. (2003). Sire coloration influences offspring survival under predation risk in the moorfrog. — J. Evol. Biol. 16: 1288-1295.

ShermanC.D.H.UllerT.WapstraE.OlssonM. (2008). Within-population variation in ejaculate characteristics in a prolonged breeder, Peron’s tree frog, Litoria peronii. — Naturwissenschaften 95: 1055-1061.

ShermanC.D.H.SagvikJ.OlssonM. (2010). Female choice for males with greater fertilisation success in the Swedish moor frog, Rana arvalis. — Plos One 5: e13634.

SztatecsnyM.JehleR.BurkeT.HödlW. (2006). Female polyandry under male harassment: the case of the common toad (Bufo bufo). — J. Zool. (Lond.) 270: 517-522.

SztatecsnyM.StrondlC.BaierlA.RiesC.HödlW. (2010). Chin up: are the bright throats of male common frogs a condition-independent visual cue?Anim. Behav. 79: 779-786.

SztatecsnyM.PreiningerD.FreudmannA.LorettoM.-C.MaierF.HödlW. (2012). Don’t get the blues: conspicuous nuptial colouration of male moor frogs (Rana arvalis) supports visual mate recognition during scramble competition in large breeding aggregations. — Behav. Ecol. Sociobiol. 66: 1587-1593.

TaylorR.C.BuchananB.W.DohertyJ.L. (2007). Sexual selection in the squirrel treefrog Hyla squirella: the role of multimodal cue assessment in female choice. — Anim. Behav. 74: 1753-1763.

VieitesD.R.Nieto-RomaS.BarluengaM.PalancaA.VencesM.MeyerA. (2004). Post-mating clutch piracy in an amphibian. — Nature 431: 305-307.

WangJ. (2002). An estimator for pairwise relatedness using molecular markers. — Genetics 160: 1203-1215.

WangJ. (2011). COANCESTRY: a program for simulating, estimating and analysing relatedness and inbreeding coefficients. — Mol. Ecol. Resour. 11: 141-145.

WangJ.SantureA.W. (2009). Parentage and sibship inference from multilocus genotype data under polygamy. — Genetics 181: 1579-1594.

WellsK.D. (2007). The ecology and behavior of amphibians. — University of Chicago PressChicago, IL.

ZamudioK.R.ChanL.M. (2008). Alternative reproductive tactics in amphibians. — In: Alternative reproductive tactics: an integrative approach ( OliveiraR.F.TaborskyM.BrockmannH.J. eds). Cambridge University PressCambridge.


  • View in gallery

    A female moor frog (Rana avails, white arrow) within a group of bluely coloured males. This figure is published in colour in the online edition of this journal, which can be accessed via

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    Relationship between inferred share of offspring sired and pairwise relatedness between mating partners for each of 7 experimental moor frog mating tanks. The pairwise relatedness coefficients are tank-specific, and we therefore refrained from overall correlational analyses.


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