Community Ecology

, Volume 1, Issue 2, pp 171–179 | Cite as

Do assembly rules for bird communities operate in small, fragmented woodlands in an agricultural landscape?

  • R. J. PakemanEmail author
  • S. A. Hinsley
  • P. E. Bellamy


The favoured state approach sensu Fox (1987) was used to investigate the existence of assembly rules for woodland bird communities in an agricultural landscape. When birds were classified according to gross breeding habitat requirements, year-round resident, ‘true’ woodland species showed an excess of favoured states suggesting a possible assembly rule. There was weaker evidence for a similar assembly rule governed by foraging requirements. This pattern was shown for all woods together, and for most categories of woods, grouped according to size, shape or size and shape together. Summer migrants did not show such patterns, and their arrival appeared to mask any patterns established by year-round resident species. The statistical significance of the excess of favoured states was highest in 1990, when bird population densities were considerably higher than in 1991 and 1992. Interspecific competition appears to be a factor in structuring woodland bird communities within the area sampled. Some implications for the action of these assembly rules on the results of further habitat fragmentation are discussed.


Favoured state Guild Landscape Ecology Null models Species distribution 


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Bellamy, P.E., S. A. Hinsley and I. Newton. 1996. Local extinctions and recolonisations of passerine bird populations in small woods. Oecologia 108: 64–71.CrossRefGoogle Scholar
  2. Belyea, L.R. and J. Lancaster. 1999. Assembly rules within a contingent ecology. Oikos 86: 402–416.CrossRefGoogle Scholar
  3. British Ornithologists’ Union. 2000. The British List, 3rd edition. Tring, UK.Google Scholar
  4. Diamond, J.M. 1975. Assembly of species communities. In: M.L. Cody and J. M. Diamond (eds), Ecology and Evolution of Communities. Harvard University Press, Cambridge MA., pp. 342–444.Google Scholar
  5. Fox, B.J. 1987. Species assembly and the evolution of community structure. Evolutionary Ecology 1: 201–213.CrossRefGoogle Scholar
  6. Fox, B.J. 1999. Genesis of an assembly rule. In: E. Weiher and P.A. Keddy (eds), Ecological Assembly Rules. Perspectives, Advances, Retreats. Cambridge University Press, Cambridge, pp. 23–57.Google Scholar
  7. Fox, B.J. and J. H. Brown. 1993. Assembly rules for functional groups in North American desert rodent communities. Oikos 67: 358–370.CrossRefGoogle Scholar
  8. Gotelli, N.J. 2000. Null model analysis of species co-occurrence patterns. Ecology 81: 2606–2621.CrossRefGoogle Scholar
  9. Gotelli, N.J. and G.R. Graves. 1996. Null Models in Ecology. Smithsonian Institution Press, Washington.Google Scholar
  10. Gotelli, N.J., N.J. Buckley, and J.A. Wiens. 1997. Co-occurrence of Australian land birds: Diamond’s assembly rules revisited. Oikos 80: 311–324.CrossRefGoogle Scholar
  11. Graves, G.R. and N.J. Gotelli. 1983. Neotropical land-bridge avifaunas: new appraoches to null hypotheses in biogeography. Oikos 41: 322–333.CrossRefGoogle Scholar
  12. Haila, Y., I.K. Hanski and S. Raivio. 1993. Turnover of breeding birds in small forest fragments: the “sampling” colonization hypothesis corroborated. Ecology 74: 714–725.CrossRefGoogle Scholar
  13. Hinsley, S.A., P.E. Bellamy and I. Newton. 1995. Bird species turnover and stochastic extinction in woodland fragments. Ecography 18: 41–50.CrossRefGoogle Scholar
  14. Hinsley, S.A., P.E. Bellamy, I. Newton, I. and T. H. Sparks. 1996a. Influences of population size and woodland area on bird species distributions in small woods. Oecologia 105: 100–106.CrossRefGoogle Scholar
  15. Hinsley, S.A., R.J. Pakeman, P.E. Bellamy and I. Newton. 1996b. Influences of habitat fragmentation on bird species distributions and regional population sizes. Proceedings of the Royal Society of London, B. 263: 307–313.CrossRefGoogle Scholar
  16. Kelt, D.A., M.L. Taper and P.L. Meserve. 1995. Assessing the impact of competition on community assembly: a case study using small mammals. Ecology 76: 1283–1296.CrossRefGoogle Scholar
  17. Marchant, J. 1983. BTO Common Bird Census Instructions. British Trust for Ornithology, Tring, Hertfordshire.Google Scholar
  18. Marchant, J. and L. Musty. 1992. Common bird census 1990–91 index report. British Trust for Ornithology News 182: 9–12.Google Scholar
  19. Simberloff, D., L. Stone and T. Dayan. 1999. Ruling out a community assembly rule: the method of favored states. In: E. Weiher and P.A. Keddy (eds), Ecological Assembly Rules. Perspectives. Advances, Retreats. Cambridge University Press, Cambridge, pp. 58–74.CrossRefGoogle Scholar
  20. Stone, L., T. Dayan and D. Simberloff. 1996. Community-wide assembly patterns unmasked: the importance of species’ differing geographic ranges. The American Naturalist 148: 997–1015.CrossRefGoogle Scholar
  21. Timonen, S., M. Mönkkönen and M. Orell. 1994. Does competition with residents affect the distribution of migrant territories? Ornis Fennica 71: 55–60.Google Scholar
  22. Weiher, E. and P. A. Keddy. 1999. Ecological Assembly Rules. Perspectives, Advances, Retreats. Cambridge University Press, Cambridge.CrossRefGoogle Scholar
  23. Weiher, E., G.D.P Clarke and P.A. Keddy. 1998. Community assembly rules, morphological dispersion, and the coexistence of plant species. Orkos 81: 309–322.Google Scholar
  24. Wiens, J.A. 1989. The Ecology of Bird Communities. Volume 1. Foundations and Pattern. Cambridge University Press, Cambridge.CrossRefGoogle Scholar
  25. Wilson, J.B. 1995. Null models for assembly rules: the Jack Horner effect is more insidious than the Narcissus effect. Oikos 72: 139–144.CrossRefGoogle Scholar
  26. Wilson, J.B. 1999. Guilds, functional types and ecological groups. Oikos 86: 507–522.CrossRefGoogle Scholar
  27. Wilson, J.B. and S. H. Roxburgh. 1994. A demonstration of guild-based assembly rules for a plant community, and determination of intrinsic guilds. Oikos 69: 267–276.CrossRefGoogle Scholar

Copyright information

© Akadémiai Kiadó, Budapest 2000

Authors and Affiliations

  • R. J. Pakeman
    • 1
    Email author
  • S. A. Hinsley
    • 2
  • P. E. Bellamy
    • 2
  1. 1.Macaulay Land Use Research InstituteCraigiebuckler, AberdeenUK
  2. 2.Centre for Ecology and Hydrology, Monks Wood, Abbots RiptonHuntingdon, CambridgeshireUK

Personalised recommendations