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Spatial scale dictates the productivity–biodiversity relationship


The diversity of life is heterogeneously distributed across the Earth. A primary cause for this pattern is the heterogeneity in the amount of energy, or primary productivity (the rate of carbon fixed through photosynthesis), available to the biota in a given location1,2,3,4,5,6,7,8,9,10,11,12. But the shape of the relationship between productivity and species diversity is highly variable10,11,12,13,14. In many cases, the relationship is ‘hump-shaped’, where diversity peaks at intermediate productivity7,9,10,12,15,16,17,18. In other cases, diversity increases linearly with productivity4,5,6,10,11,12. A possible reason for this discrepancy is that data are often collected at different spatial scales10,12,14. If the mechanisms that determine species diversity vary with spatial scale, then so would the shape of the productivity–diversity relationship. Here, we present evidence for scale-dependent productivity–diversity patterns in ponds. When the data were viewed at a local scale (among ponds), the relationship was hump-shaped, whereas when the same data were viewed at a regional scale (among watersheds), the relationship was positively linear. This dependence on scale results because dissimilarity in local species composition within regions increased with productivity.

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  1. 1

    Lotka, A. J. Elements of Mathematical Biology (Dover, New York, 1956).

  2. 2

    Connell, J. H. & Orias, E. The ecological regulation of species diversity. Am. Nat. 98, 399–414 (1964).

  3. 3

    MacArthur, R. H. Patterns of species diversity. Biol. Rev. 40, 510–533 (1965).

  4. 4

    Brown, J. H. Two decades of homage to Santa Rosalia: Towards a general theory of biodiversity. Am. Zool. 21, 877–888 (1981).

  5. 5

    Currie, D. J. & Paquin, V. Large-scale biogeographical patterns of species richness of trees. Nature 329, 326–327 (1987).

  6. 6

    Currie, D. J. Energy and large-scale patterns of animal and plant species richness. Am. Nat. 137, 27–49 (1991).

  7. 7

    Tilman, D. & Pacala, S. in Species Diversity in Ecological Communities: Historical and Geographical Perspectives (eds Ricklefs, R. E. & Schluter, D.) 12–25 (Univ. Chicago Press, Chicago, 1993).

  8. 8

    Wright, D. H., Currie, D. J. & Maurer, B. A. in Species Diversity in Ecological Communities: Historical and Geographical Perspectives (eds Ricklefs, R. E. & Schluter, D.) 66–76 (Univ. Chicago Press, Chicago, 1993).

  9. 9

    Rosenzweig, M. L. Species Diversity in Space and Time (Cambridge Univ. Press, Cambridge, 1995).

  10. 10

    Waide, R. B. et al. The relationship between primary productivity and species richness. Annu. Rev. Ecol. Syst. 30, 257–300 (1999).

  11. 11

    Gaston, K. J. Global patterns in biodiversity. Nature 405, 220–227 (2000).

  12. 12

    Mittelbach, G. G. et al. What is the observed relationship between productivity and diversity? Ecology 82, 2381–2396 (2001).

  13. 13

    Abrams, P. A. Monotonic or unimodal diversity–productivity gradients: What does competition theory predict? Ecology 76, 2019–2027 (1995).

  14. 14

    Gross, K. L. et al. Patterns of species density and productivity at different spatial scales in herbaceous plant communities. Oikos 89, 417–427 (2000).

  15. 15

    Leibold, M. A. Biodiversity and nutrient enrichment in pond plankton communities. Evol. Ecol. Res. 1, 73–95 (1999).

  16. 16

    Ritchie, M. E. & Olff, H. Spatial scaling laws yield a synthetic theory of biodiversity. Nature 400, 557–560 (1999).

  17. 17

    Abramsky, Z. & Rosenzweig, M. L. Tilman's predicted productivity–diversity relationship shown by desert rodents. Nature 309, 150–151 (1984).

  18. 18

    Dodson, S. I., Arnott, S. E. & Cottingham, K. L. The relationship in lake communities between primary productivity and species richness. Ecology 81, 2662–2679 (2000).

  19. 19

    Whittaker, R. H. Evolution and the measurement of species diversity. Taxon 21, 213–251 (1972).

  20. 20

    Lande, R. Statistics and partitioning of species diversity and similarity among multiple communities. Oikos 76, 393–401 (1996).

  21. 21

    Loreau, M. Are communities saturated? On the relationship between α, β, and γ diversity. Ecol. Lett. 3, 73–76 (2000).

  22. 22

    Law, R. in Advances in Ecological Theory: Principles and Applications (ed. McGlade, J.) 141–171 (Blackwell, Oxford, 1999).

  23. 23

    Vitousek, P. M. et al. Human Alteration of the Global Nitrogen Cycle: Causes and Consequences. Issues in Ecology Vol. 1 (Ecological Society of America, Washington DC, 1997).

  24. 24

    Carpenter, S. et al. Nonpoint Pollution of Surface Waters with Phosphorus and Nitrogen. Issues in Ecology Vol. 3 (Ecological Society of America, Washington DC, 1998).

  25. 25

    Clesceri, L. S., Greeberg, A. E. & Eaton, A. D. (eds) Standard Methods for the Examination of Water and Wastewater 20th edn (American Public Health Association, Washington DC, 1998).

  26. 26

    Mitchell-Olds, T. & Shaw, R. E. Regression analysis of natural selection: statistical inference and biological interpretation. Evolution 41, 1149–1161 (1987).

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We thank J. Shurin, M. Willig, M. Vandermuelen, P. Lorch and especially T. Knight for discussions and comments. A. Downing, J. Shurin and T. Leibold helped with the pond survey. This research was supported by the Kellogg Biological Station (Michigan State University), the University of Chicago, the University of California—Davis, the University of Pittsburgh and the NSF.

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The authors declare that they have no competing financial interests

Correspondence to Jonathan M. Chase.

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Figure 1: Results from the survey of pond species diversity relative to in situ primary productivity at local and regional scales.
Figure 2: The relative dissimilarity in species composition among local ponds within the watersheds (calculated as one minus Jaccard's index of similarity).


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