Evolutionary Ecology

, Volume 24, Issue 6, pp 1339–1351 | Cite as

Plasticity of size and allometry in multiple sexually selected traits in an armed beetle Gnatocerus cornutus

  • Kensuke Okada
  • Takahisa Miyatake
Original Paper


Male-male competition frequently results in the evolution of sexually selected traits used as weapons and ornaments. The expression of these traits often depends on male condition, i.e., condition dependence. Although males often have multiple sexually selected traits, to date many studies have focused on the morphological analysis of one sexual trait whilst ignoring the others. We here report phenotypic plasticity for multiple sexual traits, by manipulating larval diet quality and density, in the broad-horned flour beetle Gnatocerus cornutus. The male beetles possess enlarged mandibles, developed genae and a pair of small horns, but females lack these completely. Larval density significantly affected overall body size but not relative investment in each sexual trait. In contrast, diet quality had no effect on body size but had a significant effect on relative investment in the mandibles and genae. These results indicate that the allometric intercepts of the mandible and genae alter in response to diet quality, i.e., allometric plasticity. However, diet quality had no effect on the growth of the horn. Thus, multiple sexual traits exhibited differences in plasticity as a result of larval nutrient condition in G. cornutus males.


Allometry Condition dependence Exaggerated trait Horned beetle 



We thank Dr Zenobia Lewis for valuable comments. This study was supported by a Grant-in-Aid for Scientific Research (KAKENHI 21870025) to K.O. from Japanese Ministry of Education, Science, Sports, and Culture.


  1. Andersson M (1986) Evolution of condition-dependent sex ornaments and mating preferences: sexual selection based on viability differences. Evolution 40:804–816CrossRefGoogle Scholar
  2. Andersson M (1994) Sexual Selection. Princeton University Press, New JerseyGoogle Scholar
  3. Andersson S, Pryke SR, Ornborg J, Lawes MJ, Andersson M (2002) Multiple receivers, multiple ornaments, and a trade-off between agonistic and epigamic signaling in a widowbird. Am Nat 160:683–691CrossRefPubMedGoogle Scholar
  4. Bonduriansky R (2007) The evolution of condition-dependent sexual dimorphism. Am Nat 169:9–19CrossRefPubMedGoogle Scholar
  5. Bonduriansky R, Rowe L (2005) Intralocus sexual conflict and the genetic architecture of sexually dimorphic traits in Prochyliza xanthostoma (Diptera: Piophilidae). Evolution 59:1965–1975PubMedGoogle Scholar
  6. Cotton S, Fowler K, Pomiankowski A (2004) Do sexual ornaments demonstrate heightened condition-dependent expression as predicted by the handicap hypothesis? Proc R Soc Lond B 271:771–783CrossRefGoogle Scholar
  7. David P, Hingle A, Greig D, Rutherford A, Pomiankowski A, Fowler K (1998) Male sexual ornament size but not asymmetry reflects condition in stalk-eyed flies. Proc R Soc Lond B 265:2211–2216CrossRefGoogle Scholar
  8. Eberhard WG (1979) The function of horns in Podischnus agenor (Dynastinae) and other beetles. In: Blum MS, Blum NA (eds) Sexual and reproductive competition in insects. Academic Press, New York, pp 231–258Google Scholar
  9. Emlen DJ (1994) Environmental control of horn length dimorphism in the beetle Onthophagus acuminatus (Coleoptera: Scarabaeidae). Proc R Soc Lond B 256:131–136CrossRefGoogle Scholar
  10. Emlen DJ (1997) Diet alters male horn allometry in the dung beetle Onthophagus acuminatus. Proc R Soc Lond B 264:567–574CrossRefGoogle Scholar
  11. Emlen DJ (2001) Costs and the diversification of exaggerated animal structures. Science 291:1534–1536CrossRefPubMedGoogle Scholar
  12. Emlen DJ, Nijhout HF (2000) The development and evolution of exaggerated morphologies in insects. Ann Rev Entomol 45:661–708CrossRefGoogle Scholar
  13. Emlen DJ, Szafran Q, Corley LS, Dworkin I (2006) Insulin signaling and limb-patterning: candidate pathways for the origin and evolutionary diversification of beetle ‘horns’. Heredity 97:179–191CrossRefPubMedGoogle Scholar
  14. Emlen DJ, Lavine LC, Ewen-Campen B (2007) On the origin and evolutionary diversification of beetle horns. Proc Natl Acad Sci USA 104:8661–8668CrossRefPubMedGoogle Scholar
  15. Gould SJ (1966) Allometry and size in ontogeny and phylogeny. Biol Rev 41:587–640CrossRefPubMedGoogle Scholar
  16. Grafen A (1990) Sexual selection unhandicapped by the Fisher process. J Theor Biol 144:473–516CrossRefPubMedGoogle Scholar
  17. Grafen A, Hails R (2002) Modern statistics for the life sciences. Oxford University Press, OxfordGoogle Scholar
  18. Green AJ (1992) Positive allometry is likely with mate choice, competitive display and other functions. Anim Behav 43:170–172CrossRefGoogle Scholar
  19. Hongo Y (2003) Appraising behaviour during male–male interaction in the Japanese horned beetle Trypoxylus dichotomus septentrionalis (Kono). Behaviour 140:501–517CrossRefGoogle Scholar
  20. Hunt J, Bussiere LF, Jennions MD, Brooks R (2004) What is genetic quality? Trends Ecol Evol 19:329–333CrossRefPubMedGoogle Scholar
  21. Iguchi Y (1998) Horn dimorphism of Allomyrina dichotoma septentrionalis (Coleoptera: Scarabaeidae) affected by larval nutrition. Ann Entomol Soc Am 91:845–847Google Scholar
  22. Iwasa Y, Pomiankowski A (1994) The evolution of mate preferences for multiple handicaps. Evolution 48:853–867CrossRefGoogle Scholar
  23. Karino K, Seki N, Chiba M (2004) Larval nutritional environment determines adult size in Japanese horned beetles Allomyrina dichotoma. Ecol Res 19:663–668CrossRefGoogle Scholar
  24. Knell RJ, Fruhauf A, Norris KA (1999) Conditional expression of a sexually selected trait in the stalk-eyed fly Diasemopsis aethiopica. Ecol Entomol 24:323–328CrossRefGoogle Scholar
  25. Kodric-Brown A, Sibly RM, Brown JH (2006) The allometry of ornaments and weapons. Proc Nat Acad Sci USA 103:8733–8738CrossRefPubMedGoogle Scholar
  26. Kotiaho JS (2001) Costs of sexual traits: a mismatch between theoretical considerations and empirical evidence. Biol Rev 76:365–376CrossRefPubMedGoogle Scholar
  27. Kotiaho JS, Simmons LW, Tomkins JL (2001) Towards a resolution of the lek paradox. Nature 410:684–686CrossRefPubMedGoogle Scholar
  28. Miller CW, Emlen DJ (2010) Dynamic effects of oviposition site on offspring sexually-selected traits and scaling relationships. Evol Ecol 24:375–390CrossRefGoogle Scholar
  29. Moczek AP (2002) Allometric plasticity in a polyphenic beetle. Ecol Entomol 27:58–67CrossRefGoogle Scholar
  30. Moczek AP (2003) The behavioral ecology of threshold evolution in a polyphenic beetle. Behav Ecol 14:841–854CrossRefGoogle Scholar
  31. Moczek AP, Nijhout HF (2003) Rapid evolution of a polyphonic threshold. Evol Dev 5:259–268CrossRefPubMedGoogle Scholar
  32. Moczek AP, Hunt J, Emlen DJ, Simmons LW (2002) Threshold evolution in exotic populations of a polyphenic beetle. Evol Ecol Res 4:587–601Google Scholar
  33. Moczek AP, Bruehl CB, Krell FTK (2004) Linear and threshold dependent expression of secondary sexual traits in the same individual: insights from a horned beetle. Biol J Linn Soc 83:473–480CrossRefGoogle Scholar
  34. Nijhout HF, Grunert LW (2002) Bombyxin is a growth factor for wing imaginal disks in Lepidoptera. Proc Nat Acad Sci USA 99:15446–15450CrossRefPubMedGoogle Scholar
  35. Okada K, Miyatake T (2009) Genetic correlations between weapons, body shape and fighting behaviour in the horned beetle Gnatocerus cornutus. Anim Behav 77:1057–1065CrossRefGoogle Scholar
  36. Okada K, Miyanoshita A, Miyatake T (2006) Intra-sexual dimorphism in male mandibles and male aggressive behavior in the broad-horned flour beetle Gnatocerus cornutus (Coleoptera: Tenebrionidae). J Insect Behav 19:457–467CrossRefGoogle Scholar
  37. Okada K, Nomura Y, Miyatake T (2007) Relations between allometry, male-male interactions and dispersal in a sap beetle, Librodor japonicus. Anim Behav 74:749–755CrossRefGoogle Scholar
  38. Okada K, Miyatake T, Nomura Y, Kuroda K (2008) Fighting, dispersing, and sneaking: body-size dependent mating tactics by male Librodor japonicus beetles. Ecol Entomol 33:269–275CrossRefGoogle Scholar
  39. Pomiankowski A (1987) Sexual selection: the handicap principle does work—sometimes. Proc R Soc Lond B 231:123–145CrossRefGoogle Scholar
  40. SAS Institute (2005) JMP 6.0. SAS Institute Inc, CaryGoogle Scholar
  41. Simmons LW, Emlen DJ (2006) Evolutionary trade-off between weapons and testes. Proc Natl Acad Sci USA 103:16346–16351CrossRefPubMedGoogle Scholar
  42. Siva-Jothy MT (1987) Mate securing tactics and the cost of fighting in the Japanese horned beetle, Allomyrina dichotoma L. (Coleoptera Scarabaeidae). J Ethol 5:165–172CrossRefGoogle Scholar
  43. Snedecor GW, Cochran MJ (1980) Statistical methods. Iowa State University Press, AmesGoogle Scholar
  44. Tomkins JL, Brown GS (2004) Population density drives the local evolution of a threshold dimorphism. Nature 431:1099–1103CrossRefPubMedGoogle Scholar
  45. Tomkins JL, Simmons LW (2002) Measuring relative investment: a case study of testes investment in species with alternative male reproductive tactics. Anim Behav 63:1009–1016CrossRefGoogle Scholar
  46. Tomkins JL, Radwan J, Kotiaho JS, Tregenza T (2004) Genic capture and resolving the lek paradox. Trends Ecol Evol 19:323–328CrossRefPubMedGoogle Scholar
  47. Tomkins JL, Kotiaho JS, LeBas NR (2005a) Matters of scale: positive allometry and the evolution of male dimorphisms. Am Nat 165:389–402CrossRefPubMedGoogle Scholar
  48. Tomkins JL, Kotiaho JS, LeBas NR (2005b) Phenotypic plasticity in the developmental integration of morphological trade-offs and secondary sexual trait compensation. Proc R Soc Lond B 272:543–551CrossRefGoogle Scholar
  49. Tsuda Y, Yoshida T (1984) Population biology of the broad-horned flour beetle Gnathocerus cornutus (F) (Coleopteran Tenebrionidae) I: life table and population parameters. Appl Entomol Zool 19:129–131Google Scholar
  50. Tsuda Y, Yoshida T (1985) Population biology of the broad-horned flour beetle, Gnathocerus cornutus (F.), II. Crowding effects of larvae on their survival and development. Res Popul Ecol 27:77–85CrossRefGoogle Scholar
  51. Unrug J, Tomkins JL, Radwan J (2004) Alternative phenotypes and sexual selection: can dichotomous handicaps honestly signal quality? Proc R Soc Lond B 271:1401–1406CrossRefGoogle Scholar
  52. Wilkinson GS, Taper M (1999) Evolution of genetic variation for condition-dependent traits in stalk-eyed flies. Proc R Soc Lond B 266:1685–1690CrossRefGoogle Scholar
  53. Zakladnoi GA, Ratanova VF (1987) Stored-grain pests and their control (Russian translations series). AA Balkema Publishers, RotterdamGoogle Scholar

Copyright information

© Springer Science+Business Media B.V. 2010

Authors and Affiliations

  1. 1.Laboratory of Evolutionary Ecology, Graduate School of Environmental ScienceOkayama UniversityOkayamaJapan

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