Skip to main content

Costly signalling theories: beyond the handicap principle


Two recent overviews of costly signalling theory—Maynard-Smith and Harper (2003) and Searcy and Nowicki (2005)—both refuse to count signals kept honest by punishment of dishonesty, as costly signals, because (1) honest signals must be costly in cases of costly signalling, and (2) punishment of dishonesty itself requires explanation. I argue that both pairs of researchers are mistaken: (2) is not a reason to discount signals kept honest by punishment of dishonesty as cases of costly signalling, and (1) betrays too narrow a focus on certain versions of costly signalling theory. In the course of so arguing, I propose a new schema for classifying signal costs, which suggests productive research questions for future conceptual and empirical work on costly signalling.

This is a preview of subscription content, access via your institution.


  1. Thanks to an anonymous referee for urging me to emphasise this point.

  2. And, as Kim Sterelny has pointed out to me, punishment need not impose any extra costs on signal receivers. If known liars are punished via ostracism—not being chosen as partners for mutually profitable interactions—then punishment takes place as a side-effect of partner choice, and involves no costs that discriminating individuals were not paying anyway.

  3. Thanks to Richard Joyce for this point.


  • Boyd R, Richerson P (1992) Punishment allows the evolution of cooperation (or anything else) in sizable groups. Ethol Sociobiol 13:171–195

    Article  Google Scholar 

  • Caro T (1986a) The functions of stotting: a review of the hypotheses. Anim Behav 34:649–662

    Article  Google Scholar 

  • Caro T (1986b) The functions of stotting in Thompson’s gazelle: some tests of the predictions. Anim Behav 34:663–684

    Article  Google Scholar 

  • Enquist M, Plane E, Roed J (1985) Aggressive communication in fulmars (Fulmarus glacialis) competing for food. Anim Behav 33:1007–1020

    Article  Google Scholar 

  • Grafen A (1990a) Biological signals as handicaps. J Theor Biol 144:517–546

    Article  Google Scholar 

  • Grafen A (1990b) Sexual selection unhandicapped by the Fisher process. J Theor Biol 144:473–516

    Article  Google Scholar 

  • Johnstone R (1995) Sexual selection, honest advertisement and the handicap principle. Biol Rev 70:1–65

    Article  Google Scholar 

  • Kotiaho J (2001) Costs of sexual traits. Biol Rev 76:365–376

    Article  Google Scholar 

  • Lachmann M, Szamado S, Bergstrom C (2001) Cost and conflict in animal signals and human language. Proc Natl Acad Sci USA 98(23):13189–13194

    Article  Google Scholar 

  • Maynard-Smith J (1976) Sexual selection and the handicap principle. J Theor Biol 57:239–242

    Article  Google Scholar 

  • Maynard-Smith J, Harper D (2003) Animal signals. Oxford University Press, Oxford

    Google Scholar 

  • Miller G (2007) Sexual selection for moral virtues. Q Rev Biol 82:97–121

    Article  Google Scholar 

  • Mueno-Rueda G, Redondo T (2011) Begging at high level simultaneously impairs growth and immune response in southern shrike (Lanius meridionalis) nestlings. J Evol Biol 24:1091–1098

    Article  Google Scholar 

  • Saunders S (2009) Costly signalling: a work in progress. Biol Philos 24(3):405–416

    Article  Google Scholar 

  • Searcy W, Nowicki S (2005) The evolution of animal communication. Princeton University Press, Princeton

    Google Scholar 

  • Smith E, Bliege-Bird R (2005) Costly signalling and cooperative behaviour. In: Gintis H, Bowles S, Boyd R, Fehr E (eds) Moral sentiments and material interests. MIT Press, Cambridge

    Google Scholar 

  • Vehrencamp S (2000) Handicap, index, and conventional signal elements of bird song. In: Espmark Y, Amundsen T, Rosenqvist G (eds) Animal signals: signalling and signal design in animal communication. Tapir Academic Press, Trondheim

    Google Scholar 

  • Zahavi A (1975) Mate selection: a selection for handicap. J Theor Biol 53:205–214

    Article  Google Scholar 

  • Zahavi A (1977) The cost of honest (further remarks on the handicap principle). J Theor Biol 67(3):603–605

    Article  Google Scholar 

  • Zahavi A, Zahavi A (1997) The handicap principle: a missing piece of Darwin’s puzzle. Oxford University Press, Oxford

    Google Scholar 

Download references

Author information

Authors and Affiliations


Corresponding author

Correspondence to Ben Fraser.

Rights and permissions

Reprints and Permissions

About this article

Cite this article

Fraser, B. Costly signalling theories: beyond the handicap principle. Biol Philos 27, 263–278 (2012).

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: