Skip to main content

Thank you for visiting You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

  • Brief Communications Arising
  • Published:

Wild, Gardner & West reply


Replying to: M. J. Wade et al. Nature 463, 10.1038/nature08809 (2010)

We previously showed how inclusive-fitness theory separates various components of selection on parasite virulence1. Wade et al.2 do not seem to dispute our results or make new predictions. Instead, they state that insufficient attention was given to multilevel-selection theory2. However, we pointed out the links to multi-level selection1, and we believe that a misunderstanding has arisen because they have fundamentally conflated selection and adaptation.

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

Access options

Buy this article

Prices may be subject to local taxes which are calculated during checkout

Similar content being viewed by others


  1. Wild, G., Gardner, A. & West, S. A. Adaptation and the evolution of parasite virulence in a connected world. Nature 459, 983–986 (2009)

    Article  ADS  CAS  Google Scholar 

  2. Wade, M. J. et al. Multilevel and kin selection in a connected world. Nature 463 10.1038/nature08809 (2010)

  3. West, S. A., Griffin, A. S. & Gardner, A. Social semantics: how useful has group selection been? J. Evol. Biol. 21, 374–385 (2008)

    Article  Google Scholar 

  4. Wilson, D. S. & Wilson, E. O. Rethinking the theoretical foundation of sociobiology. Q. Rev. Biol. 82, 327–348 (2007)

    Article  Google Scholar 

  5. Goodnight, C. et al. Evolution in spatial predator-prey models and the “prudent predator”: the inadequacy of steady-state organism fitness and the concept of individual and group selection. Complexity 13, 23–44 (2008)

    Article  MathSciNet  Google Scholar 

  6. Wilson, D. S. Social semantics: toward a genuine pluralism in the study of social behaviour. J. Evol. Biol. 21, 368–373 (2008)

    Article  CAS  Google Scholar 

  7. Werfel, J. & Bar-Yam, Y. The evolution of reproductive restraint through social communication. Proc. Natl Acad. Sci. USA 101, 11019–11024 (2004)

    Article  ADS  CAS  Google Scholar 

  8. Kohn, M. Darwin 200: the needs of the many. Nature 456, 296–299 (2008)

    Article  CAS  Google Scholar 

  9. Wilson, D. S. & O’Brien, D. T. in Games, Groups, and the Global Good (ed. Levin, S. A.) 115–168 (Springer-Verlag, 2009)

    Google Scholar 

  10. Gardner, A. Adaptation as organism design. Biol. Lett. 5, 861–864 (2009)

    Article  Google Scholar 

  11. Hamilton, W. D. in Biosocial Anthropology (ed. Fox, R.) 133–155 (Wiley, 1975)

    Google Scholar 

  12. Hamilton, W. D. The genetical theory of social behaviour I and II. J. Theor. Biol. 7, 1–16 (1964)

    Article  CAS  Google Scholar 

  13. Grafen, A. Optimization and inclusive fitness. J. Theor. Biol. 238, 541–563 (2006)

    Article  MathSciNet  Google Scholar 

  14. Gardner, A. & Grafen, A. Capturing the superorganism: a formal theory of group adaptation. J. Evol. Biol. 22, 659–671 (2009)

    Article  CAS  Google Scholar 

  15. Krebs, J. R. & Davies, N. B. Introduction to Behavioural Ecology 3rd edn (Blackwell, 1993)

    Google Scholar 

Download references

Author information

Authors and Affiliations


Ethics declarations

Competing interests

The authors declare no competing financial interests

Rights and permissions

Reprints and permissions

About this article

Cite this article

Wild, G., Gardner, A. & West, S. Wild, Gardner & West reply . Nature 463, E9–E10 (2010).

Download citation

  • Issue Date:

  • DOI:


By submitting a comment you agree to abide by our Terms and Community Guidelines. If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate.


Quick links

Nature Briefing

Sign up for the Nature Briefing newsletter — what matters in science, free to your inbox daily.

Get the most important science stories of the day, free in your inbox. Sign up for Nature Briefing