Journal of Insect Behavior

, Volume 20, Issue 2, pp 215–227 | Cite as

Sex Differences in “Time Out” from Reproductive Activity and Sexual Selection in Male Bushcrickets (Orthoptera: Zaprochilinae: Kawanaphila mirla)

  • Gerlind U. C. Lehmann
  • Arne W. Lehmann

Animals of many species prefer some partners over others. Discriminating among potential mates causes strong sexual selection that shapes characters and behaviors. In bushcrickets the sexes shows different latencies to remate due to differences in investment in production of the nuptial gift by males and the induced refractory period in females. We conducted experiments with the Australian bushcricket Kawanaphila mirla to test the variation in male mating success by female choice.

Male remating intervals under unlimited access to food and mates were around two days, whereas most females did not remate within 12 days. Males had therefore a much shorter “time-out” from mating than females. The adult sex ratio from field samples was near to 1:1. Consequently, the OSR was male-biased with more males than females ready to mate. This male-biased OSR led to mating competition in males and choosiness in females. In a field enclosure with unlimited supply of receptive females the number of matings varied widely between males, with twenty percent of males neglected by the females. The number of matings within this enclosure was neither related to male size nor to song characters, recorded previously in the lab. However, the number of matings by individual males was positively correlated to the size of their spermatophore producing accessory gland. Females appear to prefer males with a large nutritive donation, thereby receiving a direct fitness benefit.


female choice male-male-competition mating success calling spermatophore nuptial gifts 



This study would not have been possible without the help of Winston J. Bailey who invited us to the University of Western Australia, Zoology Department, and provided office and research space during our stay. Brenton Knott generously loaned us camping equipment for the field experiments. We thank Winston J. Bailey and Robert Hickson and three anonymous referees for comments on the manuscript. The research was funded by a postdoctoral grant from the DAAD to GL.


  1. Ahnesjö, I., Kvarnemo, C., and Merilaita, S. (2001). Using potential reproductive rates to predict mating competition among individuals qualified to mate. Behav. Ecol. 12: 397–401.CrossRefGoogle Scholar
  2. Andersson, M. B. (1994). Sexual Selection, Princeton University Press, Princeton.Google Scholar
  3. Andersson, M., and Iwasa, Y. (1996). Sexual selection. Trends Ecol. Evol. 11: 53–58.CrossRefGoogle Scholar
  4. Andersson, M., and Simmons, L. W. (2006). Sexual selection and mate choice. Trends Ecol. Evol. 21: 296–302.CrossRefGoogle Scholar
  5. Bussière, L. F. (2002). A model of the interaction between ‘good genes’ and direct benefits in courtship-feeding animals: When do males of high genetic quality invest less? Phil. Trans. R. Soc. Lond. B 357: 309–317.CrossRefGoogle Scholar
  6. Clutton-Brock, T. H., and Parker, G. A. (1992). Potential reproductive rates and the operation of sexual selection. Quart. Rev. Biol. 67: 437–456.CrossRefGoogle Scholar
  7. Clutton-Brock, T. H., and Vincent, A. C. J. (1991). Sexual selection and the potential reproductive rates of males and females. Nature 351: 58–60.CrossRefGoogle Scholar
  8. Emlen, S. T., and Oring, L. W. (1977). Ecology, sexual selection and the evolution of mating systems. Science 197: 215–223.CrossRefGoogle Scholar
  9. Gerhardt, H. C. (1991). Female mate choice in treefrogs: Static and dynamic acoustic criteria. Anim. Behav. 42: 615–635.CrossRefGoogle Scholar
  10. Gerhardt, H. C., and Huber, F. (2002). Acoustic Communication in Insects and Anurans—Common Problems and Diverse Solutions, The University of Chicago Press, Chicago.Google Scholar
  11. Greenfield, M. D., Tourtellot, M. K., and Snedden, W. A. (1997). Precedence effects and the evolution of chorusing. Proc. R. Soc. Lond. B 264: 1355–1361.CrossRefGoogle Scholar
  12. Gwynne, D. T. (1982). Mate selection by female katydids (Orthoptera: Tettigoniidae, Conocephalus nigropleurum). Anim. Behav. 30: 734–738.CrossRefGoogle Scholar
  13. Gwynne, D. T. (1990). Testing parental investment and the control of sexual selection in katydids: The operational sex ratio. Am. Nat. 136: 474–484.CrossRefGoogle Scholar
  14. Gwynne, D. T. (1993). Food quality controls sexual selection in mormon crickets by altering male mating investment. Ecology 74: 1406–1413.CrossRefGoogle Scholar
  15. Gwynne, D. T. (1995). Phylogeny of the Ensifera (Orthoptera): A hypothesis supporting multiple origins of acoustical signalling, complex spermatophores and maternal care in crickets, katydids, and weta. J. Orth. Res. 4: 203–218.CrossRefGoogle Scholar
  16. Gwynne, D. T. (1997). The evolution of edible ‘sperm sacs’ and other forms of courtship feeding in crickets, katydids and their kin (Orthoptera: Ensifera). In Choe, J. C., and Crespi, B. J. (eds.), The Evolution of Mating Systems in Insects and Arachnids, Cambridge University Press, Cambridge, pp. 110–129.Google Scholar
  17. Gwynne, D. T. (2001). Katydids and Bush-Crickets: Reproductive Behavior and the Evolution of the Tettigoniidae, Cornell University Press, Ithaka.Google Scholar
  18. Gwynne, D. T. (2004). Sexual differences in response to larval food stress in two nuptial feeding orthopterans—implications for sexual selection. Oikos 105: 619–625.CrossRefGoogle Scholar
  19. Gwynne, D. T., and Bailey, W. J. (1988). Mating system, mate choice and ultrasonic calling in a zaprochiline katydid (Orthoptera: Tettigoniidae). Behaviour 105: 202–223.Google Scholar
  20. Gwynne, D. T., and Bailey, W. J. (1999). Female-female competition in katydids: Sexual selection for increased sensitivity to a male signal? Evolution 53: 546–551.CrossRefGoogle Scholar
  21. Gwynne, D. T., and Simmons, L. W. (1990). Experimental reversal of courtship roles in an insect. Nature 346: 172–174.CrossRefGoogle Scholar
  22. Gwynne, D. T., Bailey, W. T., and Annels, A. (1998). The sex in short supply for matings varies over small spatial scales in a katydid (Kawanaphila nartee, Orthopetra: Tettigoniidae). Behav. Ecol. Sociobiol. 42: 157–162.CrossRefGoogle Scholar
  23. Hartbauer, M., Kratzer, S., Steiner, K., and Römer, H. (2005). Mechanisms for synchrony and alternation in song interactions of the bushcricket Mecopoda elongate (Tettigoniidae: Orthoptera). J. Comp. Physiol. A 191: 175–188.CrossRefGoogle Scholar
  24. Hartbauer, M., Kratzer, S., and Römer, H. (2006). Chirp rate is independent of male condition in a synchronising bushcricket. J. Ins. Physiol. 52: 221–230.CrossRefGoogle Scholar
  25. Heller, K.-G., and Reinhold, K. (1994). Mating effort function of the spermatophore in the bushcricket Poecilimon veluchianus (Orthoptera, Phaneropteridae): Support from a comparison of the mating behaviour of two subspecies. Biol. J Linn. Soc. 53: 153– 163.CrossRefGoogle Scholar
  26. Johnstone, R. A. (1995). Sexual selection, honest advertisement and the handicap principle: Reviewing the evidence. Biol. Rev. 70: 1–65.CrossRefGoogle Scholar
  27. Kokko, H., Brooks, R., Jennions, M. D., and Morley, J. (2003). The evolution of mate choice and mating biases. Proc. R. Soc. Lond. B 270: 653–664.CrossRefGoogle Scholar
  28. Kvarnemo, C., and Ahnesjö, I. (1996). The dynamics of operational sex ratios and competition for mates. Trends Ecol. Evol. 11: 404–408.CrossRefGoogle Scholar
  29. Lehmann, G. U. C., and Lehmann, A. W. (2000a). Spermatophore characteristics in bushcrickets vary with parasitism and remating interval. Behav. Ecol. Sociobiol. 47: 393– 399.CrossRefGoogle Scholar
  30. Lehmann, G. U. C., and Lehmann, A. W. (2000b). Female bushcrickets mated with parasitized males show rapid remating and reduced fecundity (Orthoptera: Phaneropteridae: Poecilimon mariannae). Naturwissenschaften 87: 404–407.CrossRefGoogle Scholar
  31. Lehmann, G. U. C., and Lehmann, A. W. (accepted). Bushcrickets song is essential for female mate choice of heavier males, who provide larger direct (material) benefits. Behav. Ecol. Sociobiol. Google Scholar
  32. Mason, A. C., and Bailey, W. J. (1998). Ultrasound hearing and male-male communication in Australian katydids (Tettigoniidae: Zaprochilinae) with sexually dimorphic ears. Phys. Entomol. 23: 139–149.CrossRefGoogle Scholar
  33. Møller, A. P., and Jennions, M. D. (2001). How important are direct fitness benefits of sexual selection? Naturwissenschaften 88: 401–415.CrossRefGoogle Scholar
  34. Parker, G. A., and Simmons, L. W. (1996). Parental investment and the control of sexual selection: Predicting the direction of sexual competition. Proc. R. Soc. Lond. B 263: 315–321.CrossRefGoogle Scholar
  35. Reid, M. L., and Stamps, J. A. (1997). Female mate choice tactics in a resource-based mating system: Field tests of alternative models. Am. Nat. 150: 98–121.CrossRefGoogle Scholar
  36. Reinhold, K., and von Helversen, D. (1997). Sperm number, spermatophore weight and remating in the bushcricket Poecilimon veluchianus. Ethology 103: 12–18.CrossRefGoogle Scholar
  37. Rentz, D. C. F. (1993). A monograph of the Tettigoniidae of Australia. Vol. 2, The Phasmodinae, Zaprochilinae and Austrosaginae, CSIRO Australia, Victoria.Google Scholar
  38. Robinson, D. J., and Hall, M. J. (2002). Sound signaling in Orthoptera. Adv. Insect Physiol. 29: 151–278.CrossRefGoogle Scholar
  39. Simmons, L. W. (1990). Nuptial feeding in tettigoniids: Male costs and the rates of fecundity increase. Behav. Ecol. Sociobiol. 27: 43–47.CrossRefGoogle Scholar
  40. Simmons, L. W. (1992). Quantification of role reversal in relative parental investment in a bush cricket. Nature 358: 61–63.CrossRefGoogle Scholar
  41. Simmons, L. W. (1995a). Relative parental expenditure, potential reproductive rates, and the control of sexual selection in katydids. Am. Nat. 145: 797–808.CrossRefGoogle Scholar
  42. Simmons, L. W. (1995b). Male bushcrickets tailor spermatophores in relation to their remating intervals. Funct. Ecol. 9: 881–886.CrossRefGoogle Scholar
  43. Simmons, L. W., and Bailey, W. J. (1990). Resource influenced sex roles of zaprochiline tettigoniids (Orthoptera: Tettigoniidae). Evolution 44: 1853–1868.CrossRefGoogle Scholar
  44. Simmons, L. W., and Gwynne, D. T. (1991). The refractory period of female katydids (Orthoptera: Tettigoniidae): Sexual conflict over the remating interval? Behav. Ecol. 2: 276–282.CrossRefGoogle Scholar
  45. Trivers, R. L. (1972). Parental investment and sexual selection. In Campbell, B. (ed.), Sexual Selection and the Descent of Man, 1871–1971, Heinemann, London, pp. 136–179.Google Scholar
  46. Vahed, K. (1998). The function of nuptial feeding in insects: Review of empirical-studies. Biol. Rev. 73: 43–78.CrossRefGoogle Scholar
  47. Vahed, K., and Gilbert, F. S. (1996). Differences across taxa in nuptial gift size correlate with differences in sperm number and ejaculate volume in bushcrickets (Orthoptera: Tettigoniidae). Proc. R. Soc. Lond. B 263: 1255–1263.CrossRefGoogle Scholar
  48. Voigt, C. C., Lehmann, G. U. C., Michener, R. H., and Joachimski, M. M. (2006). Nuptial feeding is reflected in tissue nitrogen isotope ratios of female katydids. Funct. Ecol. 20: 656–661.CrossRefGoogle Scholar
  49. Wedell, N. (1993). Spermatophore size in bushcrickets: Comparative evidence for nuptial gifts as a sperm protection device. Evolution 47: 1203–1212.CrossRefGoogle Scholar
  50. Wedell, N. (1994). Dual function of the bushcricket spermatophore. Proc. R. Soc. Lond. B 258: 181–185.CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media, LLC 2007

Authors and Affiliations

  1. 1.School of Animal BiologyUniversity of Western AustraliaPerthWestern Australia
  2. 2.Institut für ZoologieFreie Universität BerlinBerlinGermany

Personalised recommendations