Your PDF has successfully downloaded.

You may be interested in finding more content on these topics:


You are not currently logged in.

Access JSTOR through your library or other institution:


Log in through your institution.

Journal Article

The Combined Influence of the Local Environment and Regional Enrichment on Bird Species Richness

Ethan P. White and Allen H. Hurlbert
The American Naturalist
Vol. 175, No. 2 (February 2010), pp. E35-E43
DOI: 10.1086/649578
Stable URL:
Page Count: 9
Were these topics helpful?
See somethings inaccurate? Let us know!

Select the topics that are inaccurate.

  • Download PDF
  • Add to My Lists
  • Cite this Item
We're having trouble loading this content. Download PDF instead.


Abstract: It is generally accepted that local species richness at a site reflects the combined influence of local and regional processes. However, most empirical studies evaluate the influence of either local environmental variables or regional enrichment but not both simultaneously. Here we demonstrate the importance of combining these processes to understand continental‐scale richness patterns in breeding birds. We show that neither regional enrichment nor the local environment in isolation is sufficient to characterize observed patterns of species richness. Combining both sets of variables into a single model results in improved model fit and the removal of residual spatial autocorrelation. At short timescales, local processes are most important for determining local richness, but as the timescale of analysis increases, regional enrichment becomes increasingly important. These results emphasize the need for increased integration of multiple scales of processes into models of species richness.

Notes and References

This item contains 59 references.

Literature Cited
  • ['Algar, A. C., J. T. Kerr, and D. J. Currie. 2007. A test of metabolic theory as the mechanism underlying broad‐scale species‐richness gradients. Global Ecology and Biogeography 16:170–178.']
  • ['Allen, A. P., J. H. Brown, and J. F. Gillooly. 2002. Global biodiversity, biochemical kinetics, and the energetic‐equivalence rule. Science 297:1545–1548.']
  • ['Angermeier, P. L., and M. R. Winston. 1998. Local vs. regional influences on local diversity in stream fish communities of Virginia. Ecology 79:911–927.']
  • ['Belmaker, J. 2009. Species richness of resident and transient coral‐dwelling fish responds differentially to regional diversity. Global Ecology and Biogeography 18:426–436.']
  • ['Boulinier, T., J. D. Nichols, J. R. Sauer, J. E. Hines, and K. H. Pollock. 1998. Estimating species richness: the importance of heterogeneity in species detectability. Ecology 79:1018–1028.']
  • ['Brown, J. H. 1981. Two decades of homage to Santa‐Rosalia: toward a general theory of diversity. American Zoologist 21:877–888.']
  • ['Brown, J. H., T. G. Whitham, S. K. M. Ernest, and C. A. Gehring. 2001. Complex species interactions and the dynamics of ecological systems: long‐term experiments. Science 293:643–650.']
  • ['Burnham, K. P., and D. R. Anderson. 2002. Model selection and multimodel inference: a practical information‐theoretic approach. Springer, New York.']
  • ['Chong, D. L. S., E. Mougin, and J. P. Gastellu‐Etchegorry. 1993. Relating the global vegetation index to net primary productivity and actual evapotranspiration over Africa. International Journal of Remote Sensing 14:1517–1546.']
  • ['Cornell, H. V., and R. H. Karlson. 1996. Species richness of reef‐building corals determined by local and regional processes. Journal of Animal Ecology 65:233–241.']
  • ['Cornell, H. V., and J. H. Lawton. 1992. Species interactions, local and regional processes, and limits to the richness of ecological communities: a theoretical perspective. Journal of Animal Ecology 61:1–12.']
  • ['Currie, D. J., G. G. Mittelbach, H. V. Cornell, R. Field, J. F. Guegan, B. A. Hawkins, D. M. Kaufman, et al. 2004. Predictions and tests of climate‐based hypotheses of broad‐scale variation in taxonomic richness. Ecology Letters 7:1121–1134.']
  • ['Diniz‐Filho, J. A., L. M. Bini, and B. A. Hawkins. 2003. Spatial autocorrelation and red herrings in geographical ecology. Global Ecology and Biogeography 12:53–64.']
  • ['Evans, K. L., S. E. Newson, D. Storch, J. J. D. Greenwood, and K. J. Gaston. 2008. Spatial scale, abundance and the species‐energy relationship in British birds. Journal of Animal Ecology 77:395–405.']
  • ['Freestone, A. L., and S. Harrison. 2006. Regional enrichment of local assemblages is robust to variation in local productivity, abiotic gradients, and heterogeneity. Ecology Letters 9:95–102.']
  • ['Gough, L., C. W. Osenberg, K. L. Gross, and S. L. Collins. 2000. Fertilization effects on species density and primary productivity in herbaceous plant communities. Oikos 89:428–439.']
  • ['Graham, M. H. 2003. Confronting multicollinearity in ecological multiple regression. Ecology 84:2809–2815.']
  • ['Griffiths, D. 1999. On investigating local‐regional species richness relationships. Journal of Animal Ecology 68:1051–1055.']
  • ['Grinnell, J. 1922. The role of the “accidental.” Auk 39:373–380.']
  • ['Harrison, S., and H. Cornell. 2008. Toward a better understanding of the regional causes of local community richness. Ecology Letters 11:969–979.']
  • ['Harrison, S., H. D. Safford, J. B. Grace, J. H. Viers, and K. F. Davies. 2006. Regional and local species richness in an insular environment: serpentine plants in California. Ecological Monographs 76:41–56.']
  • ['Hawkins, B., R. Field, H. Cornell, D. Currie, J. Guegan, D. Kaufman, J. Kerr, et al. 2003a. Energy, water, and broad‐scale geographic patterns of species richness. Ecology 84:3105–3117.']
  • ['Hawkins, B. A., E. E. Porter, and J. A. F. Diniz‐Filho. 2003b. Productivity and history as predictors of the latitudinal diversity gradient of terrestrial birds. Ecology 84:1608–1623.']
  • ['Hawkins, B. A., F. S. Albuquerque, M. B. Araujo, J. Beck, L. M. Bini, F. J. Cabrero‐Sanudo, I. Castro‐Parga, et al. 2007. A global evaluation of metabolic theory as an explanation for terrestrial species richness gradients. Ecology 88:1877–1888.']
  • ['He, F. L., K. J. Gaston, E. F. Connor, and D. S. Srivastava. 2005. The local‐regional relationship: immigration, extinction, and scale. Ecology 86:360–365.']
  • ['Holling, C. S. 1992. Cross‐scale morphology, geometry, and dynamics of ecosystems. Ecological Monographs 62:447–502.']
  • ['Hortal, J., J. Rodriguez, M. Nieto‐Diaz, and J. M. Lobo. 2008. Regional and environmental effects on the species richness of mammal assemblages. Journal of Biogeography 35:1202–1214.']
  • ['Hurlbert, A. H., and J. P. Haskell. 2003. The effect of energy and seasonality on avian species richness and community composition. American Naturalist 161:83–97.']
  • ['Hurlbert, A. H., and E. P. White. 2005. Disparity between range map‐ and survey‐based analyses of species richness: patterns, processes and implications. Ecology Letters 8:319–327.']
  • ['Karlson, R. H., and H. V. Cornell. 1999. Integration of local and regional perspectives on the species richness of coral assemblages. American Zoologist 39:104–112.']
  • ['Karlson, R. H., H. V. Cornell, and T. P. Hughes. 2004. Coral communities are regionally enriched along an oceanic biodiversity gradient. Nature 429:867–870.']
  • ['Kaspari, M., S. O’Donnell, and J. Kercher. 2000. Energy, density, and constraints to species richness: ant assemblages along a productivity gradient. American Naturalist 155:280–293.']
  • ['Legendre, P., and L. Legendre. 1998. Numerical ecology. Elsevier, Amsterdam.']
  • ['Lennon, J. J. 2000. Red‐shifts and red herrings in geographical ecology. Ecography 23:101–113.']
  • ['Lichstein, J. W., T. R. Simons, S. A. Shriner, and K. E. Franzreb. 2002. Spatial autocorrelation and autoregressive models in ecology. Ecological Monographs 72:445–463.']
  • ['MacArthur, R. 1960. On the relative abundance of species. American Naturalist 94:25–36.']
  • ['MacArthur, R. H. 1964. Environmental factors affecting bird species diversity. American Naturalist 98:387–396.']
  • ['Magurran, A., and P. Henderson. 2003. Explaining the excess of rare species in natural species abundance distributions. Nature 422:714–716.']
  • ['McGill, B. 2003. A test of the unified neutral theory of biodiversity. Nature 422:881–885.']
  • ['Nichols, J. D., T. Boulinier, J. E. Hines, K. H. Pollock, and J. R. Sauer. 1998. Inference methods for spatial variation in species richness and community composition when not all species are detected. Conservation Biology 12:1390–1398.']
  • ['Paruelo, J. M., H. E. Epstein, W. K. Lauenroth, and I. C. Burke. 1997. ANPP estimates from NDVI for the central grassland region of the United States. Ecology 78:953–958.']
  • ['Qian, H., P. S. White, and J. S. Song. 2007. Effects of regional vs. ecological factors on plant species richness: an intercontinental analysis. Ecology 88:1440–1453.']
  • ['Rahbek, C., and G. R. Graves. 2001. Multiscale assessment of patterns of avian species richness. Proceedings of the National Academy of Sciences of the USA 98:4534–4539.']
  • ['Rahbek, C., N. J. Gotelli, R. K. Colwell, G. L. Entsminger, T. F. L. V. B. Rangel, and G. R. Graves. 2007. Predicting continental‐scale patterns of bird species richness with spatially explicit models. Proceedings of the Royal Society B: Biological Sciences 274:165–174.']
  • ['Rangel, T., J. A. F. Diniz‐Filho, and L. M. Bini. 2006. Towards an integrated computational tool for spatial analysis in macroecology and biogeography. Global Ecology and Biogeography 15:321–327.']
  • ['Ricklefs, R. E. 1987. Community diversity: relative roles of local and regional processes. Science 235:167–171.']
  • ['———. 2000. The relationship between local and regional species richness in birds of the Caribbean Basin. Journal of Animal Ecology 69:1111–1116.']
  • ['———. 2007. History and diversity: explorations at the intersection of ecology and evolution. American Naturalist 170(suppl.):S56–S70.']
  • ['Ridgely, R. S., T. F. Allnutt, T. Brooks, D. K. McNicol, D. W. Mehlman, B. E. Young, and J. R. Zook. 2003. Digital distribution maps of the birds of the Western Hemisphere. Version 1.0. NatureServe, Arlington, VA.']
  • ['Sauer, J. R., J. E. Hines, and J. Fallon. 2007. The North American breeding bird survey: results and analysis, 1966–2006. Version 10.13.2007. U.S. Geological Survey Patuxent Wildlife Research Center, Laurel, MD.']
  • ['Schmida, A., and M. V. Wilson. 1985. Biological determinants of species diversity. Journal of Biogeography 12:1–20.']
  • ['Schweiger, O., M. Musche, D. Bailey, R. Billeter, T. Diekötter, F. Hendrickx, F. Herzog, et al. 2007. Functional richness of local hoverfly communities (Diptera, Syrphidae) in response to land use across temperate Europe. Oikos 116:461–472.']
  • ['Srivastava, D. S. 1999. Using local‐regional richness plots to test for species saturation: pitfalls and potentials. Journal of Animal Ecology 68:1–16.']
  • ['Stohlgren, T. J., D. T. Barnett, C. S. Jarnevich, C. Flather, and J. Kartesz. 2008. The myth of plant species saturation. Ecology Letters 11:313–322.']
  • ['Tilman, D. 2004. Niche tradeoffs, neutrality, and community structure: a stochastic theory of resource competition, invasion, and community assembly. Proceedings of the National Academy of Sciences of the USA 101:10854–10861.']
  • ['White, E. P. 2004. Two‐phase species‐time relationships in North American land birds. Ecology Letters 7:329–336.']
  • ['White, E. P., P. B. Adler, W. K. Lauenroth, R. A. Gill, D. Greenberg, D. M. Kaufman, A. Rassweiler, et al. 2006. A comparison of the species‐time relationship across ecosystems and taxonomic groups. Oikos 112:185–195.']
  • ['Witman, J. D., R. J. Etter, and F. Smith. 2004. The relationship between regional and local species diversity in marine benthic communities: a global perspective. Proceedings of the National Academy of Sciences of the USA 101:15664–15669.']
  • ['Wright, D. H. 1983. Species‐energy theory: an extension of species‐area theory. Oikos 41:496–506.']


Part of Sustainability