Evolutionary Ecology

, Volume 22, Issue 6, pp 701–709 | Cite as

Protandry and postandry in two related butterflies: conflicting evidence about sex-specific trade-offs between adult size and emergence time

  • Gabriel NèveEmail author
  • Michael C. Singer
Original Paper


Natural selection acting on timing of metamorphosis can be sex-specific, resulting in differences in timing between males and females. Insects with discrete generations frequently show protandry: males usually mature before females. Both Euphydryas editha and E. aurinia butterflies followed this trend. The present study was motivated by the unusual observation of consistent postandry in addition to protandry. In a single E. editha population observed over 20 years the emergence period of males was longer than that of females, both the first and last emerging individuals being males. Variance of timing among individual E. editha larvae is imposed by spatial patchiness of the snowmelt that releases them from winter diapause. If individual larvae released late from diapause were to compensate for their lateness by shortening their development times, they would be small at maturity. If such compensation were only partial, they would be both late and small. Size and timing would become associated. If females were more prone to such partial compensation than males, the observations of postandry could be explained and the prediction made that any tendency for late individuals to be small should be stronger in females than in males. This was the case: in 1 year late males were the same size as early males, in a second year they were larger. Late females were significantly smaller than early females in both years. In E. aurinia, results were opposite both to theoretical prediction and to the observations from E. editha: although the male emergence period was longer than that of females exactly as in E. editha, late males were smaller than early ones, while late females were not small. The data from E. editha support the hypothesis of a sex-specific trade-off between size and emergence time, the data from E. aurinia do not.


Protandry Postandry Size Euphydryas aurinia Euphydryas editha Lepidoptera-Nymphalidae 



Philippe Goffart, David Boughton and Chris Singer helped gather the data. We thank Philippe Lebrun and Michel Baguette (University of Louvain) and the Ministère de la Région Wallonne for support to the E. aurinia study.


  1. Arnason AN, Schwarz CJ (1999) Using POPAN-5 to analyse banding data. Bird Study 46(suppl):S157–S168CrossRefGoogle Scholar
  2. Arnason AN, Schwarz CJ, Boyer G (1998) POPAN-5: a data maintenance and analysis system for mark-recapture data. Department of Computer Science, WinnipegGoogle Scholar
  3. Benard MF (2004) Predator-induced phenotypic plasticity in organisms with complex life histories. Ann Rev Ecol Syst 35:651–673CrossRefGoogle Scholar
  4. Berger D, Walters R, Gotthard K (2006) What keeps insects small?—Size dependent predation on two species of butterfly larvae. Evol Ecol 20:575–589CrossRefGoogle Scholar
  5. Boggs CL, Freeman KD (2005) Larval food limitation in butterflies: effects on adult resource allocation and fitness. Oecologia 144:353–361PubMedCrossRefGoogle Scholar
  6. Boggs CL, Nieminen M (2004) Checkerspot reproductive biology. In: Ehrlich PR, Hanski I (eds) On the wings of checkerspots. Oxford University Press, Oxford, pp 92–111Google Scholar
  7. Boughton DA (1999) Empirical evidence for complex source-sink dynamics with alternative states in a butterfly metapopulation. Ecology 80:2727–2739Google Scholar
  8. Candolin U, Voigt HR (2003) Size-dependent selection on arrival times in sticklebacks: why small males arrive first. Evolution 57:862–871PubMedGoogle Scholar
  9. Ehrlich AH, Ehrlich PR (1978) Reproductive strategies in the butterflies. I. Mating frequency, plugging, and egg number. J Kansas Ent Soc 51:666–697Google Scholar
  10. Fagerström T, Wiklund C (1982) Why do males emerge before females? Protandry as a mating strategy in male and female butterflies. Oecologia 52:164–166CrossRefGoogle Scholar
  11. Fischer K, Fiedler K (2001a) Sexual differences in life-history traits in the butterfly Lycaena tityrus: a comparison between direct and diapause development. Ent Exp Appl 100:325–330CrossRefGoogle Scholar
  12. Fischer K, Fiedler K (2001b) Effects of larval starvation on adult life-history traits in the butterfly species Lycaena tityrus (Lepidoptera: Lycaenidae). Ent Gen 25:249–254Google Scholar
  13. Fischer K, Zeilstra I, Hetz SK, Fiedler K (2004) Physiological costs of growing fast: does accelerated growth reduce pay-off in adult fitness? Evol Ecol 18:343–353CrossRefGoogle Scholar
  14. Forrest TG (1987) Insect size tactics and developmental strategies. Oecologia 73:178–184CrossRefGoogle Scholar
  15. Garland T Jr, Adolph SC (1994) Why not to do two-species comparative studies. Physiol Zool 67:797–828Google Scholar
  16. Higgins L (2000) The interaction of season length and development time alters size at maturity. Oecologia 122:51–59CrossRefGoogle Scholar
  17. Iwasa Y, Odendaal FJ, Murphy DD, Ehrlich PR, Launer AE (1983) Emergence patterns in male butterflies: a hypothesis and a test. Theor Pop Biol 23:363–379CrossRefGoogle Scholar
  18. Leimar O, Karlsson B, Wiklund C (1994) Unpredictable food and sexual size dimorphism in insects. Proc R Soc Lond B 258:121–125CrossRefGoogle Scholar
  19. Nylin S, Gotthard K (1998) Plasticity in life-history traits. Ann Rev Ent 43:63–83CrossRefGoogle Scholar
  20. R Development Core Team (2004) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Available from (accessed January 2007)
  21. Singer MC (1982) Sexual selection for small size in male butterflies. Am Nat 119:440–443CrossRefGoogle Scholar
  22. Stearns SC (1992) The evolution of life histories. Oxford University Press, OxfordGoogle Scholar
  23. Taylor BW, Anderson CR, Peckarsky BL (1998) Effects of size at metamorphosis on stone fly fecundity, longevity, and reproductive success. Oecologia 114:494–502CrossRefGoogle Scholar
  24. Thomas CD, Singer MC, Boughton DA (1996) Catastrophic extinction of population sources in a butterfly metapopulation. Am Nat 148:957–975CrossRefGoogle Scholar
  25. Vonesh JR, Warkentin KM (2006) Opposite shifts in size at metamorphosis in response to larval and metamorph predators. Ecology 87:556–562PubMedCrossRefGoogle Scholar
  26. Wiklund C, Forsberg J (1991) Sexual size dimorphism in relation to female polygamy and protandry in butterflies: a comparative study of Swedish Pieridae and Satyridae. Oikos 60:373–381CrossRefGoogle Scholar
  27. Wiklund C, Kaitala A (1995) Sexual selection for large male size in a polyandrous butterfly: the effect of body size on male versus female reproductive success in Pieris napi. Behav Ecol 6:6–13CrossRefGoogle Scholar
  28. Zonneveld C (1996) Being big or emerging early? Polyandry and the trade-off between size and emergence time in male butterflies. Am Nat 147:946–965CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media B.V. 2007

Authors and Affiliations

  1. 1.EA 3781 Evolution Génome Environnement, Case 36Université de ProvenceMarseille Cedex 3France
  2. 2.Integrative Biology, Patterson LaboratoriesUniversity of TexasAustinUSA

Personalised recommendations