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.

  • Letter
  • Published:

Ecological and evolutionary processes at expanding range margins


Many animals are regarded as relatively sedentary and specialized in marginal parts of their geographical distributions1,2. They are expected to be slow at colonizing new habitats. Despite this, the cool margins of many species’ distributions have expanded rapidly in association with recent climate warming3,4,5,6,7,8,9,10. We examined four insect species that have expanded their geographical ranges in Britain over the past 20 years. Here we report that two butterfly species have increased the variety of habitat types that they can colonize, and that two bush cricket species show increased fractions of longer-winged (dispersive) individuals in recently founded populations. Both ecological and evolutionary processes are probably responsible for these changes. Increased habitat breadth and dispersal tendencies have resulted in about 3- to 15-fold increases in expansion rates, allowing these insects to cross habitat disjunctions that would have represented major or complete barriers to dispersal before the expansions started. The emergence of dispersive phenotypes will increase the speed at which species invade new environments, and probably underlies the responses of many species to both past11 and future climate change.

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

Figure 1: Habitat and range expansion by H. comma. a, b, Aspects of populated H. comma habitats in 1982 (a), and colonized habitats between 1983–2000 (b).
Figure 2: Expanding distributions of C. discolor (a) and M. roeselii7–9 (b).

Similar content being viewed by others


  1. Thomas, J. A. Holocene climate change and warm man-made refugia may explain why a sixth of British butterflies inhabit unnatural early-successional habitats. Ecography 16, 278–284 (1993).

    Article  Google Scholar 

  2. Thomas, J. A., Rose, R. J., Clarke, R. T., Thomas, C. D. & Webb, N. R. Intraspecific variation in habitat availability among ectothermic animals near their climatic limits and their centres of range. Funct. Ecol. 13 (Suppl.1), 55–64 (1999).

    Article  CAS  Google Scholar 

  3. Hughes, L. Biological consequences of global warming: is the signal already apparent? Trends Ecol. Evol. 15, 56–61 (2000).

    Article  CAS  Google Scholar 

  4. Hill, J. K., Thomas, C. D. & Huntley, B. Climate and habitat availability determine 20th century changes in a butterfly's range margin. Proc. R. Soc. Lond. B 266, 1197–1206 (1999).

    Article  Google Scholar 

  5. Parmesan, C. et al. Poleward shifts in geographical ranges of butterfly species associated with regional warming. Nature 399, 579–583 (1999).

    Article  ADS  CAS  Google Scholar 

  6. Asher, J. et al. The Millennium Atlas of Butterflies in Britain and Ireland: Provisional Maps (Oxford Univ. Press, 2001).

    Google Scholar 

  7. Marshall, J. A. & Haes, E. C. M. Grasshoppers and Allied Insects of Great Britain and Ireland (Harley Books, Colchester, 1988).

    Google Scholar 

  8. Haes, E. C. M. & Harding, P. T. Atlas of the Grasshoppers, Crickets and Allied Insects in Britain and Ireland (The Stationery Office, London, 1997).

    Google Scholar 

  9. Widgery, J. Orthoptera Recording Scheme for Britain and Ireland. Newsletter 26 (BRC, Huntingdon, 2000).

    Google Scholar 

  10. Thomas, C. D. & Lennon, J. J. Birds extend their ranges northwards. Nature 399, 213 (1999).

    Article  ADS  CAS  Google Scholar 

  11. Cwynar, L. C. & MacDonald, G. M. Geographical variation of lodgepole pine in relation to population history. Am. Nat. 129, 463–469 (1987).

    Article  Google Scholar 

  12. Thomas, J. A., Thomas, C. D., Simcox, D. J. & Clarke, R. T. The ecology and declining status of the silver-spotted skipper butterfly (Hesperia comma) in Britain. J. Appl. Ecol. 23, 365–380 (1986).

    Article  Google Scholar 

  13. Hanski, I. A practical model of metapopulation dynamics. J. Anim. Ecol. 63, 151–162 (1994).

    Article  Google Scholar 

  14. Wahlberg, N., Moilanen, A. & Hanski, I. Predicting the occurrence of endangered species in fragmented landscapes. Science 273, 1536–1538 (1996).

    Article  ADS  CAS  Google Scholar 

  15. Moilanen, A. The equilibrium assumption in estimating the parameters of metapopulation models. J. Anim. Ecol. 69, 143–153 (2000).

    Article  Google Scholar 

  16. Heath, J., Pollard, E. & Thomas, J. A. Atlas of Butterflies in Britain and Ireland (Viking, Harmondsworth, 1984).

    Google Scholar 

  17. Perring, F. H. & Walters, S. M. Atlas of the British Flora 2nd edn (EP Publishing, Wakefield, 1976).

    Google Scholar 

  18. Bourn, N. A. D. & Thomas, J. A. The ecology and conservation of the brown argus butterfly Aricia agestis in Britain. Biol. Conserv. 63, 67–74 (1993).

    Article  Google Scholar 

  19. Wilson, R. J. The Spatiotemporal Dynamics of Three Lepidopteran Herbivores of Helianthemum chamaecistus. Thesis, Univ. Leeds (1999).

    Google Scholar 

  20. Thomas, C. D., Thomas, J. A. & Warren, M. S. Distributions of occupied and vacant butterfly habitats in fragmented landscapes. Oecologia 92, 563–567 (1992).

    Article  ADS  CAS  Google Scholar 

  21. Thomas, C. D. & Hanski, I. in Metapopulation Biology: Ecology, Genetics and Evolution (eds Hanski, I. A. & Gilpin, M. E.) 359–386 (Academic, London, 1997).

    Book  Google Scholar 

  22. Niemela, J & Spence, J. R. Distribution and abundance of an exotic ground-beetle (Carabidae): a test of community impact. Oikos 62, 351–359 (1991).

    Article  Google Scholar 

  23. Hill, J. K., Thomas, C. D. & Blakeley, D. S. Evolution of flight morphology in a butterfly that has recently expanded its geographic range. Oecologia 121, 165–170 (1999).

    Article  ADS  CAS  Google Scholar 

  24. Hill, J. K., Thomas, C. D. & Lewis, O. T. Flight morphology in fragmented populations of a rare British butterfly, Hesperia comma. Biol. Conserv. 87, 277–284 (1999).

    Article  Google Scholar 

  25. Zera, A. J. & Denno, R. F. Physiology and ecology of dispersal poymorphisms in insects. Anu. Rev. Ent. 42, 207–230 (1997).

    Article  CAS  Google Scholar 

  26. Doebeli, M. & Ruxton, R. D. Evolution of dispersal rates in metapopulation models: branching and cyclic dynamics in phenotype space. Evolution 51, 1730–1741 (1997).

    Article  Google Scholar 

  27. Coope, G. R. in Extinction Rates (eds Lawton, J. & May, R.) 55–74 (Oxford Univ. Press, 1995).

    Google Scholar 

  28. Williamson, M. Biological Invasions (Chapman & Hall, London, 1996).

    Google Scholar 

  29. Ratnieks, F. L. W. in The “African” Honey Bee (eds Spivak, M., Fletcher, D. J. C. & Breed, M. D.) 119–135 (Westview, Boulder, Colorado, 1991).

    Google Scholar 

  30. Thomas, C. D. & Jones, T. M. Partial recovery of a skipper butterfly (Hesperia comma) from population refuges: lessons for conservation in a fragmented landscape. J. Anim. Ecol. 62, 472–481 (1993).

    Article  Google Scholar 

Download references


We thank D. Blakeley, H. Burton, C. Dytham, K. Ericson, P. Ewin, R. Fox, C. Gardener, S. Glencross, A. Goodhand, S. Hanna, I. Hanski, S. Hartley, J. Hill, C. Holloway, P. Hoskin, W. Kunin, R. Leaper, G. Jeffcoate, J. Lennon, J. Mellings, A. Moilanen, C. Parmesan, G. Parris, M. Rosenzweig, D. Sazer, J. Silvertown, M. Singer, D. Taneyhill, J. Thomas, J. Travis, J. Widgery, K. Wilson and I. Wynne for discussions, assistance and/or comments on the manuscript. Maps were produced in DMap. This work was funded by The UK Natural Environment Research Council, EU TMR FRAGLAND and English Nature. We also thank English Nature, the National Trust and other landowners for permission to carry out the research, and the amateur butterfly and Orthoptera recorders who recorded 10-km distributions.

Author information

Authors and Affiliations


Corresponding author

Correspondence to C. D. Thomas.

Rights and permissions

Reprints and permissions

About this article

Cite this article

Thomas, C., Bodsworth, E., Wilson, R. et al. Ecological and evolutionary processes at expanding range margins. Nature 411, 577–581 (2001).

Download citation

  • Received:

  • Accepted:

  • Issue Date:

  • DOI:

This article is cited by


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