The effect of operational sex ratio on sex allocation and neonate phenotype in a viviparous lizard Eremias multiocellata

in Amphibia-Reptilia
Restricted Access
Get Access to Full Text
Rent on DeepDyve

Have an Access Token?

Enter your access token to activate and access content online.

Please login and go to your personal user account to enter your access token.


Have Institutional Access?

Access content through your institution. Any other coaching guidance?


Maternal investment in the production of male versus female neonates was approximately equal in most animal species. However, sex allocation theory predicts that under certain conditions, selection may favor the females’ ability to adjust the sex ratio of their offspring, which females tend to use more for an investment of the rare sex. The mechanism of operational sex ratio (OSR) influence on sex allocation is still unclear, and recent studies conducted on lizards have reached conflicting conclusions. Here, we selected a viviparous lizard Eremias multiocellata to test whether pregnant females could adjust the sex ratio of their offspring in response to OSRs. Our results showed that mothers did not adjust the sex ratios or phenotypes of neonates in the laboratory and field-based experiments, except tail length. However, the OSRs subsequently affected growth in both mass and SVL of the offspring in laboratory experiments; whereas only the mass was affected in the semi-nature field experiments. Our results, thus, contradict the predictions of sex allocation theory and challenge the idea that female investment in the scarcity sex might serve as a mechanism which is used for adjusting the population sex ratio.


Publication of the Societas Europaea Herpetologica



AllsopD.J.WarnerD.A.LangkildeT.DuW.ShineR. (2006): Do operational sex ratios influence sex allocation in viviparous lizards with temperature-dependent sex determination? J. Evolution. Biol. 19: 1175-1182.

AveryH.W.SpotilaJ.R.CongdonJ.D.FischerR.U.StandoraE.A.AveryS.B. (1993): Roles of diet protein and temperature in the growth and nutritional energetics of juvenile slider turtles, Trachemys scripta. Physiol. Zool. 66: 902-925.

BullJ.J. (1987): Temperature-sensitive periods of sex determination in a lizard: similarities with turtles and crocodilians. J. Exp. Zool. 241: 143-148.

CharnovE.L. (1982): The Theory of Sex Allocation. Princeton University Press, Princeton.

ClarkS.J. (1988): The effects of operational sex ratio and food deprivation on copulation duration in the water strider (Gerris remigis Say). Behav. Ecol. and Sociobiol. 23: 317-322.

ClarkeA.L.SaetherB.E.RoskaftE. (1997): Sex biases in avian dispersal: a reappraisal. Oikos 79: 429-438.

Clutton-BrockT. (2007): Sexual selection in males and females. Science 318: 1882-1885.

DonaldP.F. (2007): Adult sex ratios in wild bird populations. Ibis 149: 671-692.

EmlenS.T.OringL.W. (1977): Ecology, sexual selection, and the evolution of mating systems. Science 197: 215-223.

FisherR.A. (1930): The Genetical Theory of Natural Selection. Oxford University Press, Oxford.

FitzeP.S.Le GalliardJ.F. (2008): Operational sex ratio, sexual conflict and the intensity of sexual selection. Ecol. Lett. 11: 432-439.

GageM.J.G. (1991): Risk of sperm competition directly affects ejaculate size in the Mediterranean fruit fly. Anim. Behav. 42: 1036-1037.

HareJ.R.HolmesK.M.WilsonJ.L.CreeA. (2009): Modelling exposure to selected temperature during pregnancy: the limitations of squamate viviparity in a cool-climate environment. Biol. J. Linn. Soc. 96: 541-552.

JiX.LinL.H.LuoL.G.LuH.L.GaoJ.F.HanJ. (2006): Gestation temperature affects sexual phenotype, morphology, locomotor performance, and growth of neonatal brown forest skinks, Sphenomorphus indicus. Biol. J. Linn. Soc. 88: 453-463.

KomdeurJ.DaanS.TinbergenJ.MatemanA.C. (1997): Facultative sex ratio bias in the offspring of Seychelles warblers. Nature 385: 522-525.

KvarnemoC.AhnesjoI. (1996): The dynamics of operational sex ratios and competition for mates. Trends Ecol. Evol. 11: 404-408.

KvarnemoC.AhnesjoI. (2002): Operational sex ratios and mating competition. In: Sex Ratios: Concepts and Research Methods I, p.  366-382. HardyC.W., Ed., Cambridge University Press, Cambridge.

LawrenceW.S. (1986): Male choice and competition in Tetraopes tetraophthalmus: effects of local sex ratio variation. Behav. Ecol. and Sociobiol. 18: 289-296.

Le GalliardJ.-F.FitzeP.S.CoteJ.MassotM.ClobertJ. (2005): Female common lizards (Lacerta vivipara) do not adjust their sex-biased investment in relation to the adult sex ratio. J. Exp. Biol. 18: 1455-1463.

OlssonM.ShineR. (1997): The limits to reproductive output: offspring size versus number in the sand lizard (Lacerta agilis). Am. Nat. 149: 179-188.

OlssonM.ShineR. (2001): Facultative sex allocation in snow skink lizards (Niveoscincus microlepidotus). J. Exp. Biol. 14: 120-128.

QuallsC.P.ShineR. (1996): Reconstructing ancestral reaction norms: an example using the evolution of reptilian viviparity. Funct. Ecol. 10: 688-697.

RobertK.ThompsonM.B. (2003): Reconstructing Thermochron iButtons to reduce size and weight as a new technique in the study of small animal thermal biology. Herpetological Review 34: 130-132.

RobertK.A.ThompsonM.B.SeebacherF. (2003): Facultative sex allocation in the viviparous lizard Eulamprus tympanum, a species with temperature-dependent sex determination. Aust. J. Zool. 51: 367-370.

RoweL. (1992): Convenience polyandry in a water strider: foraging conflicts and female control of copulation frequency and guarding duration. Anim. Behav. 44: 189-202.

ShineR.DownesS.J. (1999): Can pregnant lizards adjust their offspring phenotypes to environmental conditions? Oecologia 119: 1-8.

SwainR.JonesS.M. (2000): Maternal effects associated with gestation conditions in a viviparous lizard, Niveoscincus metallicus. Herpetol. Monogr. 14: 432-440.

TangX.L.YueF.YanX.F.ZhangD.J.XinY.WangC.ChenQ. (2012): Effects of gestation temperature on offspring sex and maternal reproduction in a viviparous lizard (Eremias multiocellata) living at high altitude. J. Therm. Biol. 37: 438-444.

ThomopsonJ. (1981): A study of the sources of nutrients for embryonic development in a viviparous lizard, Sphenomorphus quoyii. Comp. Biochem. Phys. A 70: 509-518.

TriversR.L.WillardD.E. (1973): Natural selection of parental ability to vary the sex ratio of offspring. Science 191: 249-263.

WangT.ZaarM.ArvedsenS.Vedel-SmithC.OvergaardJ. (2002): Effects of temperature on the metabolic response to feeding in Python molurus. Comp. Biochem. Phys. A 133: 519-527.

WapstraE. (2000): Maternal basking opportunity affects juvenile phenotype in a viviparous lizard. Funct. Ecol. 14: 345-352.

WapstraE.SwainR.JonesS.M.O’ReillyJ. (1999): Geographic and annual variation in reproductive cycles in the Tasmanian spotted snow skink, Niveoscincus ocellatus (Squamata: Scincidae). Aust. J. Zool. 47: 539-550.

WarnerD.A.ShineR. (2007): Reproducing lizards modify sex allocation in response to operational sex ratios. Biol. Letters 3: 47-50.

WerrenJ.H. (1980): Sex ratio adaptations to local mate competition in a parasitic wasp. Science 208: 1157-1159.

WerrenJ.H.CharnovE.L. (1978): Facultative sex ratios and population dynamics. Nature 272: 349-350.

WhileG.M.WapstraE. (2009): Effects of basking opportunity on birthing asynchrony in a viviparous lizard. Anim. Behav. 77: 1465-1470.

WilliamsG.C. (1979): The question of adaptive sex ratio in outcrossed vertebrates. Proc. R. Soc. London, Ser. B 205: 567-580.

YueF.TangX.L.ZhangD.J.YanX.F.XinY.ChenQ. (2012): Body temperature and standard metabolic rate of the female viviparous lizard Eremias multiocellata during reproduction. Can. J. Zool. 90: 79-84.

ZhangD.J.TangX.L.YueF.ChenZ.H.LiR.D.ChenQ. (2010): Effect of gestation temperature on sexual and morphological phenotypes of offspring in a viviparous lizard, Eremias multiocellata. J. Therm. Biol. 35: 129-133.

ZhaoE.M.AdlerK. (1993): Herpetology of China, p.  201-204. SSAR, Oxford, Ohio.

ZhaoK.T. (1999): Lacertidae. In: Fauna Sinica, Reptilia, Vol. 2 (Squamata: Lacertilia), p.  219-242. ZhaoE.M.ZhaoK.T.ZhouK.Y., Eds, Science Press, Beijing. [In Chinese.]


  • The effects of OSR treatment on maximum body temperature of the pregnant female lizards in the laboratory conditions.

    View in gallery
  • Mean sex ratio (proportion male) of litters produced in response to OSR treatment in the laboratory and semi-nature field conditions.

    View in gallery


Content Metrics

Content Metrics

All Time Past Year Past 30 Days
Abstract Views 24 24 9
Full Text Views 1 1 1
PDF Downloads 1 1 1
EPUB Downloads 0 0 0