Evolutionary Ecology

, Volume 24, Issue 5, pp 985–1001 | Cite as

Natural and sexual selection on male behaviour and morphology, and female choice in a wild field cricket population: spatial, temporal and analytical components



We used multiple regression, path analysis and non-parametric selection surface visualisation to investigate natural and sexual selection and, in addition, cross-sectional female choice statistics to analyse female choice in a wild population of the field cricket Gryllus campestris L. in central Germany. Adults (167 males, 75 females) were individually marked and followed daily over the entire adult stage. Two morphological traits (pronotum width, body condition) and two behavioural traits (burrow occupation time, daily displacement) were measured for each male and used in selection analyses. Individuals mated multiply and male mating success was strongly right skewed with less than 6% of the population achieving 50% of the copulations. In males, analysis of natural selection in terms of lifespan revealed positive directional selection on burrow occupation time and stabilising selection on daily displacement. Analysis of sexual selection in term of mating success showed positive directional selection on pronotum width and lifespan. Path analysis confirmed the close association between natural and sexual selection and illustrated indirect effects of the behavioural variables on mating success via their effect on lifespan. Multiple regression analysis further indicated positive quadratic (disruptive) sexual selection on lifespan but the non-parametric cubic spline regression showed this to be an artefact of the quadratic approach. In fact, lifespan was under “threshold selection” i.e. it was not under selection below a threshold and under positive directional sexual selection above the threshold. A positive correlational selection gradient between lifespan and body condition revealed that a high body condition is advantageous among long-lived males. Female choice statistics showed that females chose large and heavy males in the beginning of the season only and that choosiness decreases with increasing distance to potential alternative mates. Our findings highlight the benefits of combining several analytical methods to uncover selection patterns and to avoid misinterpretations based on single methods.


Fitness Gryllidae Male quality Mating success Non-linear selection gradients Selection surface visualization Threshold selection 



K. Reinhardt and an anonymous referee gave us the final push to conduct the detailed selection analysis on our data set. S. K. Sakaluk, R. Brooks and three anonymous referees made helpful comments on the manuscript.


  1. Arnold SJ, Wade MJ (1984a) On the measurement of natural and sexual selection: applications. Evolution 38:720–734CrossRefGoogle Scholar
  2. Arnold SJ, Wade MJ (1984b) On the measurement of natural and sexual selection: theory. Evolution 38:709–719CrossRefGoogle Scholar
  3. Bateman PW, Gilson LN, Ferguson JWH (2001) Male size and sequential mate preference in the cricket Gryllus bimaculatus. Anim Behav 61:631–637CrossRefGoogle Scholar
  4. Benjamini Y, Hochberg Y (1995) Controlling the false discovery rate—a practical and powerful approach to multiple testing. J R Stat Soc B 57:289–300Google Scholar
  5. Bentsen CL, Hunt J, Jennions MD, Brooks R (2006) Complex multivariate sexual selection on male acoustic signaling in a wild population of Teleogryllus commodus. Am Nat 167:E102–E116CrossRefPubMedGoogle Scholar
  6. Blows MW, Brooks R (2003) Measuring nonlinear selection. Am Nat 162:815–820CrossRefPubMedGoogle Scholar
  7. Brodie ED, Janzen FJ (1996) On the assignment of fitness values in statistical analyses of selection. Evolution 50:437–442CrossRefGoogle Scholar
  8. Brodie ED, Moore AJ, Janzen FJ (1995) Visualizing and quantifying natural selection. Trends Ecol Evol 10:313–318CrossRefGoogle Scholar
  9. Cade WH (1981) Field cricket spacing, and the phonotaxis of crickets and parasitoid flies to clumped and isolated cricket songs. Zeitschrift für Tierpsychologie 55:365–375Google Scholar
  10. Conner JK (2001) How strong is natural selection? Trends Ecol Evol 16:215CrossRefPubMedGoogle Scholar
  11. Gray DA (1999) Intrinsic factors affecting female choice in house crickets: time cost, female age, nutritional condition, body size, and size-relative reproductive investment. J Insect Behav 12:691–700CrossRefGoogle Scholar
  12. Green PJ, Silverman BW (1994) Nonparametric regression and generalized linear models. Chapman and Hall, LondonGoogle Scholar
  13. Gwynne DT (2001) Katydids and bush-crickets: reproductive behavior and evolution of the Tettigoniidae. Cornell University Press, New YorkGoogle Scholar
  14. Hedrick AV, Dill LM (1993) Mate choice by female crickets is influenced by predation risk. Anim Behav 46:193–196CrossRefGoogle Scholar
  15. Hereford J, Hansen TF, Houle D (2004) Comparing strengths of directional selection: how strong is strong? Evolution 58:2133–2143PubMedGoogle Scholar
  16. Hockham LR, Vahed K (1997) The function of mate guarding in a field cricket (Orthoptera: Gryllidae; Teleogryllus natalensis Otte and Cade). J Insect Behav 10:247–256CrossRefGoogle Scholar
  17. Hoekstra HE, Hoekstra JM, Berrigan D, Vignieri SN, Hoang A, Hill CE, Beerli P, Kingsolver JG (2001) Strength and tempo of directional selection in the wild. Proc Natl Acad Sci USA 98:9157–9160CrossRefPubMedGoogle Scholar
  18. Huber F, Moore TE, Loher WE (1989) Cricket behavior and neurobiology. Cornell University Press, IthacaGoogle Scholar
  19. Hunt J, Brooks R, Jennions MD, Smith MJ, Bentsen CL, Bussiere LF (2004) High-quality male field crickets invest heavily in sexual display but die young. Nature 432:1024–1027CrossRefPubMedGoogle Scholar
  20. Kiflawi M, Gray DA (2000) Size-dependent response to conspecific mating calls by male crickets. Proc R Soc Lond B 267:2157–2161CrossRefGoogle Scholar
  21. Kingsolver JG, Pfennig DW (2007) Patterns and power of phenotypic selection in nature. Bioscience 57:561–572CrossRefGoogle Scholar
  22. Kingsolver JG, Hoekstra HE, Hoekstra JM, Berrigan D, Vignieri SN, Hill CE, Hoang A, Gibert P, Beerli P (2001) The strength of phenotypic selection in natural populations. Am Nat 157:245–261CrossRefPubMedGoogle Scholar
  23. Kline RB (2004) Principles and practice of structural equation modelling. Guilford Press, New YorkGoogle Scholar
  24. Köhler G, Reinhardt K (1992) Beitrag zur Kenntnis der Feldgrille (Gryllus campestris L.) in Thüringen. Articulata 7:63–76Google Scholar
  25. Kokko H, Mappes J (2005) Sexual selection when fertilization is not guaranteed. Evolution 59:1876–1885PubMedGoogle Scholar
  26. Lande R, Arnold SJ (1983) The measurement of selection on correlated characters. Evolution 37:1210–1226CrossRefGoogle Scholar
  27. Mitchell-Olds T, Bergelson J (1990) Statistical genetics of an annual plant, Impatiens capensis. II. Natural selection. Genetics 124:417–421Google Scholar
  28. Mitchell-Olds T, Shaw RG (1987) Regression analysis of natural selection: statistical inference and biological interpretation. Evolution 41:1149–1161CrossRefGoogle Scholar
  29. Petraitis PS, Dunham AE, Niewiarowski PH (1996) Inferring multiple causality: the limitations of path analysis. Funct Ecol 10:421–431CrossRefGoogle Scholar
  30. Phillips PC, Arnold SJ (1989) Visualizing multivariate selection. Evolution 43:1209–1222CrossRefGoogle Scholar
  31. Popov AV (1975) Acoustic behaviour and migrations of field crickets Gryllus campestris (in russian). Zoologichesky Zhurnal 54:1803–1809Google Scholar
  32. Popov AV, Shuvalov VF (1977) Phonotactic behavior of crickets. J Comp Physiol A 119:111–126CrossRefGoogle Scholar
  33. Prosser MR, Murray A-M, Cade WH (1997) The influence of female age on phonotaxis during single and multiple song presentations in the field cricket, Gryllus integer (Orthoptera: Gryllidae). J Insect Behav 10:437–449CrossRefGoogle Scholar
  34. R Development Core Team (2006) R—a language and environment for statistical computing. R Foundation for Statistical Computing, ViennaGoogle Scholar
  35. Ritz MS (2000) Verhalten der Männchen in einer Wildpopulation der Feldgrille (Gryllus campestris L.). Articulata 15:87–98Google Scholar
  36. Ritz MS, Köhler G (2007) Male behaviour over the season in a wild population of the field cricket Gryllus campestris L. Ecol Entomol 32:384–392CrossRefGoogle Scholar
  37. Ritz MS, Sakaluk SK (2002) The role of the male’s cerci in copulation and mate guarding in decorated crickets (Gryllodes sigillatus). J Zool 257:519–523CrossRefGoogle Scholar
  38. Rost R, Honegger HW (1987) The timing of premating and mating behavior in a field population of the cricket Gryllus campestris L. Behav Ecol Sociobiol 21:279–289CrossRefGoogle Scholar
  39. Ryder JJ, Siva-Jothy MT (2000) Male calling song provides a reliable signal of immune function in a cricket. Proc R Soc Lond B 267:1171–1175CrossRefGoogle Scholar
  40. Sakaluk SK (1991) Post-copulatory mate guarding in decorated crickets. Anim Behav 41:207–216CrossRefGoogle Scholar
  41. Sakaluk SK, Cade WH (1983) The adaptive significance of female multiple matings in house and field crickets. In: Gwyne DT, Morris GK (eds) Orthopteran mating systems, sexual competition in a diverse group of insects. Westview Press, Boulder, pp 319–336Google Scholar
  42. Sakaluk SK, Schaus JM, Eggert A-K, Snedden WA, Brady PL (2002) Polyandry and fitness of offspring reared under varying nutritional stress in decorated crickets. Evolution 56:1999–2007PubMedGoogle Scholar
  43. Scheiner SM, Mitchell RJ, Callahan HS (2000) Using path analysis to measure natural selection. J Evol Biol 13:423–433CrossRefGoogle Scholar
  44. Schluter D (1988) Estimating the form of natural selection on a quantitative trait. Evolution 42:849–861CrossRefGoogle Scholar
  45. Schluter D, Nychka D (1994) Exploring fitness surfaces. Am Nat 143:597–616CrossRefGoogle Scholar
  46. Shipley B (2000) Cause and correlation in biology: a user’s guide to path analysis, structural equations and causal inference. Cambridge University Press, CambridgeCrossRefGoogle Scholar
  47. Shuvalov VF, Popov AV (1973) The importance of the calling song rhythmic pattern of males of the genus Gryllus for phonotaxis of females. Zoologichesky Zhurnal 52:1179–1185Google Scholar
  48. Simmons LW (1987) Sperm competition as a mechanism of female choice in the field cricket, Gryllus bimaculatus (De Geer). Behav Ecol Sociobiol 21:197–202CrossRefGoogle Scholar
  49. Simmons LW (1988a) The calling song of the field cricket, Gryllus bimaculatus (De Geer): constraints on transmission and its role in intermale competition and female choice. Anim Behav 36:380–394CrossRefGoogle Scholar
  50. Simmons LW (1988b) Male size, mating potential and lifetime reproductive success in the field cricket, Gryllus bimaculatus (De Geer). Anim Behav 36:372–379CrossRefGoogle Scholar
  51. Simmons LW (1992) Sexual selection and body size in a natural population of the field cricket, Gryllus campestris (L.). Journal of Orthoptera Research 1:12–13CrossRefGoogle Scholar
  52. Simmons LW (1995) Correlates of male quality in the field cricket, Gryllus campestris L.: age, size, and symmetry determine pairing success in field populations. Behav Ecol 6:376–381CrossRefGoogle Scholar
  53. Simmons LW, Ritchie MG (1996) Symmetry in the songs of crickets. Proc R Soc Lond B 263:1308–1311Google Scholar
  54. Simmons LW, Zuk M (1992) Variability in call structure and pairing success of male field crickets, Gryllus bimaculatus: the effect of age, size and parasite load. Anim Behav 44:1145–1152CrossRefGoogle Scholar
  55. Stinchcombe JR, Agrawal AF, Hohenlohe PA, Arnold SJ, Blows MW (2008) Estimating nonlinear selection gradients using quadratic regression coefficients: Double or nothing? Evolution 62:2435–2440CrossRefPubMedGoogle Scholar
  56. Tregenza T, Wedell N (1998) Benefits of multiple mates in the cricket Gryllus bimaculatus. Evolution 52:1726–1730CrossRefGoogle Scholar
  57. Verhoeven KJF, Simonsen KL, McIntyre LM (2005) Implementing false discovery rate control: increasing your power. Oikos 108:643–647CrossRefGoogle Scholar
  58. Vieth E (1989) Fitting piecewise linear regression functions to biological responses. J Appl Physiol 67:390–396PubMedGoogle Scholar
  59. Wagner WEJ, Hoback WW (1999) Nutritional effects on male calling behaviour in the variable field cricket. Anim Behav 57:89–95CrossRefPubMedGoogle Scholar
  60. Wright S (1920) The relative importance of heredity and environment in determining the piebald pattern of guinea pigs. Proc Natl Acad Sci USA 6:320–332CrossRefPubMedGoogle Scholar
  61. Zar JH (1999) Biostatistical analysis, 4th edn. Prentice-Hall, New JerseyGoogle Scholar
  62. Zippelius H-M (1949) Die Paarungsbiologie einiger Orthopteren-Arten. Zeitschrift für Tierpsychologie 6:372–390Google Scholar
  63. Zuk M (1987) Variability in attractiveness of male field crickets (Orthoptera: Gryllidae) to females. Anim Behav 35:1240–1248CrossRefGoogle Scholar
  64. Zuk M (1988) Parasite load, body size, and age of wild-caught male field crickets (Orthoptera: Gryllidae): effects on sexual selection. Evolution 42:969–976CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media B.V. 2009

Authors and Affiliations

  1. 1.Institute of EcologyFriedrich Schiller University JenaJenaGermany

Personalised recommendations