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Productivity–biodiversity relationships depend on the history of community assembly


Identification of the causes of productivity–species diversity relationships remains a central topic of ecological research1,2. Different relations have been attributed to the influence of disturbance3,4, consumers5,6, niche specialization7 and spatial scale8,9,10,11,12,13,14. One unexplored cause is the history of community assembly, the partly stochastic sequential arrival of species from a regional pool of potential community members. The sequence of species arrival can greatly affect community structure15,16,17,18,19. If assembly sequence interacts with productivity to influence diversity, different sequences can contribute to variation in productivity–diversity relationships. Here we report a test of this hypothesis by assembling aquatic microbial communities at five productivity levels using four assembly sequences. About 30 generations after assembly, productivity–diversity relationships took various forms, including a positive, a hump-shaped, a U-shaped and a non-significant pattern, depending on assembly sequence. This variation resulted from idiosyncratic joint effects of assembly sequence, productivity and species identity on species abundances. We suggest that the history of community assembly should be added to the growing list of factors that influence productivity–biodiversity patterns.

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Figure 1: Response of species diversity to productivity.
Figure 2: Response of the abundance of Uronema sp. to productivity and assembly sequence on day 25.


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

    Article  Google Scholar 

  2. Morin, P. J. Biodiversity's ups and downs. Nature 406, 463–464 (2000)

    CAS  Article  Google Scholar 

  3. Huston, M. A. Biological Diversity: The Coexistence of Species on Changing Landscapes (Cambridge Univ. Press, 1994)

    Google Scholar 

  4. Kondoh, M. Unifying the relationships of species richness to productivity and disturbance. Proc. R. Soc. Lond. B 268, 269–271 (2001)

    CAS  Article  Google Scholar 

  5. Worm, B. et al. Consumer versus resource control of species diversity and ecosystem functioning. Nature 417, 848–851 (2002)

    ADS  CAS  Article  Google Scholar 

  6. Leibold, M. A. et al. Species turnover and the regulation of trophic structure. Annu. Rev. Ecol. Syst. 28, 467–494 (1997)

    Article  Google Scholar 

  7. Kassen, R. et al. Diversity peaks at intermediate productivity in a laboratory microcosm. Nature 406, 508–512 (2000)

    ADS  CAS  Article  Google Scholar 

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

    Article  Google Scholar 

  9. Wright, D. H., Currie, D. J. & Maurer, B. A. in Species Diversity in Ecological Communities: Historical and Geographical Perspectives (eds Ricklefs, R. & Schluter, D.) 66–74 (Univ. Chicago Press, 1993)

    Google Scholar 

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

    Article  Google Scholar 

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

    Article  Google Scholar 

  12. Scheiner, S. M. et al. Species richness, species–area curves and Simpson's paradox. Evol. Ecol. Res. 2, 791–802 (2000)

    Google Scholar 

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

    Article  Google Scholar 

  14. Chase, J. M. & Leibold, M. A. Spatial scale dictates the productivity–biodiversity relationship. Nature 416, 427–430 (2002)

    ADS  CAS  Article  Google Scholar 

  15. Gilpin, M. E. & Case, T. J. Multiple domains of attraction in competition-communities. Nature 261, 40–42 (1976)

    ADS  CAS  Article  Google Scholar 

  16. Post, W. M. & Pimm, S. L. Community assembly and food web stability. Math. Biosci. 64, 169–192 (1983)

    Article  Google Scholar 

  17. Drake, J. A. Community–assembly mechanics and the structure of experimental species ensemble. Am. Nat. 137, 1–26 (1991)

    Article  Google Scholar 

  18. Wilson, D. S. Complex interactions in metacommunities, with implications for biodiversity and higher levels of selection. Ecology 73, 1984–2000 (1992)

    Article  Google Scholar 

  19. Law, R. & Morton, R. D. Alternative permanent states of ecological communities. Ecology 74, 1347–1361 (1993)

    Article  Google Scholar 

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

    Article  Google Scholar 

  21. Leigh, E. G. Jr On the relationship between productivity, biomass, diversity and stability of a community. Proc. Natl Acad. Sci. USA 53, 777–783 (1965)

    ADS  CAS  Article  Google Scholar 

  22. Pianka, E. R. Latitudinal gradients in species diversity: a review of concepts. Am. Nat. 100, 33–46 (1966)

    Article  Google Scholar 

  23. Rosenzweig, M. L. Species diversity gradients: We know more and less than we thought. J. Mammal. 73, 715–730 (1992)

    Article  Google Scholar 

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

    Book  Google Scholar 

  25. Tilman, D. & Pacala, S. in Species Diversity in Ecological Communities: Historical and Geographical Perspectives (eds Ricklefs, R. & Schluter, D.) 13–25 (Univ. Chicago Press, 1993)

    Google Scholar 

  26. Van de Koppel, J. et al. Patterns of herbivory along a productivity gradient: An empirical and theoretical investigation. Ecology 77, 736–745 (1996)

    Article  Google Scholar 

  27. Holt, R. D. & Polis, G. A. A theoretical framework for intraguild predation. Am. Nat. 149, 745–764 (1997)

    Article  Google Scholar 

  28. Chase, J. M. Food web effects of prey size refugia: Variable interactions and alternative stable equilibria. Am. Nat. 154, 559–570 (1999)

    Article  Google Scholar 

  29. Simpson, E. H. Measurement of diversity. Nature 163, 688 (1949)

    ADS  Article  Google Scholar 

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

    Article  Google Scholar 

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We thank members of the Morin laboratory for discussion, and J. A. Drake, C. M. K. Kaunzinger, M. A. Leibold, Z. T. Long, P. B. Rainey and D. Simberloff for comments. The National Science Foundation and the Department of Ecology and Evolutionary Biology at the University of Tennessee supported this research.

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Correspondence to Tadashi Fukami.

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The authors declare that they have no competing financial interests.

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Supplementary Information: This file contains supplementary information on the following: regression analysis, MOS test, species richness, population dynamics, comparison of assembly in our study and assembly in nature, and local and regional diversity. (DOC 220 kb)

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Fukami, T., Morin, P. Productivity–biodiversity relationships depend on the history of community assembly. Nature 424, 423–426 (2003).

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