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.

General patterns of taxonomic and biomass partitioning in extant and fossil plant communities


A central goal of evolutionary ecology is to identify the general features maintaining the diversity of species assemblages1,2,3. Understanding the taxonomic and ecological characteristics of ecological communities provides a means to develop and test theories about the processes that regulate species coexistence and diversity. Here, using data from woody plant communities from different biogeographic regions, continents and geologic time periods, we show that the number of higher taxa is a general power-function of species richness that is significantly different from randomized assemblages. In general, we find that local communities are characterized by fewer higher taxa than would be expected by chance. The degree of taxonomic diversity is influenced by modes of dispersal and potential biotic interactions. Further, changes in local diversity are accompanied by regular changes in the partitioning of community biomass between taxa that are also described by a power function. Our results indicate that local and regional processes2 have consistently regulated community diversity and biomass partitioning for millions of years.

This is a preview of subscription content, access via your institution

Relevant articles

Open Access articles citing this article.

Access options

Buy article

Get time limited or full article access on ReadCube.


All prices are NET prices.

Figure 1: Three graphical hypotheses for the relationship between species richness and number of higher taxa within a local community.
Figure 2: Relationship between the number of species and higher taxa across 227 0.1-ha sites from around the world.
Figure 3: Relationship between the number of species and higher taxa across 28 local palaeoflora sites ranging from 4.5 to 45 Myr ago.
Figure 4: Relationship between species pool size and the results of randomization experiments using the total number of sampled sites from South America.
Figure 5: Biomass partitioning between species (open squares), genera (open circles), and families (filled diamonds) across global woody plant communities.


  1. Hutchinson, G. E. Homage to Santa Rosalia, or why are there so many kinds of animals? Am. Nat. 93, 145–159 (1959)

    Article  Google Scholar 

  2. Hubbell, S. P. A Unified Theory of Biodiversity and Biogeography (Princeton Univ. Press, Princeton, 2001)

    Google Scholar 

  3. Ricklefs, R. E. & Schluter, D. (eds) Species Diversity in Ecological Communities (Univ. Chicago Press, Chicago, 1993)

  4. Elton, J. Competition and the structure of ecological communities. Anim. Ecol. 15, 54–68 (1946)

    Article  Google Scholar 

  5. Williams, C. B. Patterns in the Balance of Nature (Academic, New York, 1964)

    Google Scholar 

  6. Kinzig, A. P., Levin, S. A., Dushoff, J. & Pacala, S. Limiting similarity, species packing, and system stability for hierarchical competition-colonization models. Am. Nat. 153, 371–383 (1999)

    CAS  PubMed  Google Scholar 

  7. MacArthur, R. H. & Levins, R. The limiting similarity, convergence, and divergence of coexisting species. Am. Nat. 101, 377–385 (1967)

    Article  Google Scholar 

  8. Simberloff, D. S. Taxonomic diversity of island biotas. Evolution 24, 23–47 (1970)

    Article  Google Scholar 

  9. Gentry, A. H. Changes in plant community diversity and floristic composition on environmental and geographic gradients. Ann. Missouri Bot. Gard. 75, 1–34 (1988)

    Article  Google Scholar 

  10. Gentry, A. H. Biological Relationships Between Africa and South America (ed. Goldblatt, P.) 500–547 (Yale Univ. Press, New Haven, 1993)

    Book  Google Scholar 

  11. Webb, C. O. Exploring the phylogenetic structure of ecological communities: An example for rain forest trees. Am. Nat. 156, 145–155 (2000)

    Article  Google Scholar 

  12. Williams, P. H., Humphries, C. J. & Gaston, K. J. Centers of seed-plant diversity—the family way. Proc. R. Soc. Lond. B 256, 67–70 (1994)

    ADS  Article  Google Scholar 

  13. Roy, K., Jablonski, D. & Valentine, J. W. Higher taxa in biodiversity studies: patterns from eastern Pacific marine mollusks. Phil. Trans. R. Soc. Lond. B 351, 1605–1613 (1996)

    ADS  Article  Google Scholar 

  14. Valentine, J. W. How many marine invertebrate fossil species? A new approximation. J. Paleontol. 44, 410–415 (1970)

    Google Scholar 

  15. Sepkoski, J. J. Jr The Unity of Evolutionary Biology (ed. Dudley, E. C.) 210–236 (Dioscorides, Portland, 1991)

    Google Scholar 

  16. Robeck, H., Maley, C. C. & Donoghue, M. Taxonomy and temporal diversity patterns. Paleobiology 26, 171–187 (2000)

    Article  Google Scholar 

  17. Manly, B. F. J. Randomization, Bootstrap and Monte Carlo Methods in Biology (Chapman & Hall, New York, 1997)

    MATH  Google Scholar 

  18. Gotelli, N. J. & McCabe, D. J. Species co-occurrence: a meta-analysis of J.M. Diamond's assembly rules model. Ecology 83, 2091–2096 (2002)

    Article  Google Scholar 

  19. MacArthur, R. H. & Wilson, E. O. The Theory of Island Biogeography (Princeton Univ. Press, Princeton, 1967)

    Google Scholar 

  20. Darwin, C. The Origin of Species (John Murray, London, 1859)

    Google Scholar 

  21. Karr, J. R. & James, F. C. Ecology and Evolution of Communities (eds Cody, M. L. & Diamond, J. M.) 258–291 (Harvard Univ. Press, Cambridge, 1975)

    Google Scholar 

  22. Ricklefs, R. E. & O'Rourke, K. Aspect diversity in moths: a temperate-tropical comparison. Evolution 29, 313–324 (1975)

    Article  Google Scholar 

  23. Hubbell, S. P. et al. Light-gap disturbances, recruitment limitation, and tree diversity in a neotropical forest. Science 283, 554–557 (1999)

    ADS  CAS  Article  Google Scholar 

  24. Tilman, D., Lehman, C. L. & Thompson, K. T. Plant diversity and ecosystem productivity: Theoretical considerations. Proc. Natl Acad. Sci. USA 94, 1857–1861 (1997)

    ADS  CAS  Article  Google Scholar 

  25. Hughes, L. et al. Predicting dispersal spectra—a minimal set of hypotheses based on plant attributes. J. Ecol. 82, 933–950 (1994)

    Article  Google Scholar 

  26. Vazquez., J. A. G. & Givnish, T. J. Altitudinal gradients in tropical forest composition, structure, and diversity in the Sierra de Manantlan. J. Ecol. 86, 999–1020 (1998)

    Article  Google Scholar 

  27. Ribbens, E., Silander, J. A. & Pacala, S. W. Seedling recruitment in forests: Calibrating models to predict patterns of tree seedling dispersion. Ecology 75, 1794–1806 (1994)

    Article  Google Scholar 

  28. Enquist, B. J. & Niklas, K. J. Invariant scaling relations across tree-dominated communities. Nature 401, 655–660 (2001)

    ADS  Article  Google Scholar 

Download references


We thank J. H. Brown, S. Collins, R. Colwell, M. J. Donoghue, N. J. Gotelli, D. Post, S. P. Hubbell, C. J. Humphries, W. P. Maddison, K. J. Niklas, N. Pittman, F. A. Smith, M. Weiser, J. Williams, R. Whittaker and the members of the NCEAS Body Size Working Group and Phylogenies and Community Ecology Working Group for critical discussions and/or comments on earlier drafts. In particular, C. O. Webb provided valuable comments. This work stems in part from the Body Size in Ecology and Evolution Working Group (F.A. Smith, Principal Investigator) sponsored by The National Center for Ecological Analysis and Synthesis (NCEAS, a national centre funded by the NSF, the University of California Santa Barbara and the State of California). B.J.E. was supported by the NSF and NCEAS. J.P.H. was supported by a student internship from NCEAS. Computer resources for the simulations were provided by the UNM Department of Biology, Sevilleta Long Term Ecological Research site and NCEAS.

Author information

Authors and Affiliations


Corresponding author

Correspondence to Brian J. Enquist.

Ethics declarations

Competing interests

The authors declare that they have no competing financial interests.

Supplementary information

Rights and permissions

Reprints and Permissions

About this article

Cite this article

Enquist, B., Haskell, J. & Tiffney, B. General patterns of taxonomic and biomass partitioning in extant and fossil plant communities. Nature 419, 610–613 (2002).

Download citation

  • Received:

  • Accepted:

  • Issue Date:

  • DOI:

Further reading


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