, Volume 7, Issue 3, pp 296–310 | Cite as

Species Diversity Across Nutrient Gradients: An Analysis of Resource Competition in Model Ecosystems

  • Darrell A. Herbert
  • Edward B. Rastetter
  • Laura Gough
  • Gaius R. Shaver


The capture and efficient use of limiting resources influence the competitive success of individual plant species as well as species diversity across resource gradients. In simulations, efficient nutrient acquisition or nutrient retention by species were key predictors of success when nutrients were limiting. Increased nutrient supply favored species with characteristics that improved light interception or light use. Ecological theory suggests that low diversity on fertile sites may be a consequence of competitive exclusion by one or a few species with superior light-interception characteristics. On infertile sites, competitive exclusion may be a function of superior nutrient-acquisition characteristics in species. At intermediate fertility, a shift from single-resource specialization to a balanced effort in the acquisition of multiple resources should allow for greater species diversity. Thus, a unimodal relationship between diversity and nutrient supply, vegetation biomass, or productivity is predicted. However, simulations demonstrated alternate relationships depending on the ecosystem characteristic to which diversity was compared. Diversity was greatest at intermediate total biomass but increased monotonically with net primary production and nitrogen (N) supply. The highest diversity occurred midrange on a scale of community-level leaf area to fine-root length ratios, which in the context of the model indicates that the vegetation as a whole was simultaneously limited by both N and light and that effort toward the acquisition of both resources is distributed in such a way that both resources are equally exploited. Diversity was lowered by the presence of species with a superior ability to sequester resources.


diversity competition nutrient use light use biogeochemical model 



National Science Foundation grant DEB95-09613 provided funding support for this research. We thank Bonnie Kwaitkowski for assistance in development of the MEL model and Jim Grace for substantive comments on the manuscript.


  1. 1.
    Abrams, PA 1988How should resources be counted?Theor Popul Biol3322642Google Scholar
  2. 2.
    Abrams, PA 1995Monotonic or unimodal diversity–productivity gradients: what does competition theory predict?Ecology76201927Google Scholar
  3. 3.
    Aerts, RE 1990Nutrient use efficiency in evergreen and deciduous species from heathlands.Oecologia (Berl)843917Google Scholar
  4. 4.
    Aerts, RE, Chapin III, FS 2000The mineral nutrition of wild plants revisited : a re-evaluation of processes and patterns.Adv Ecol Res30167Google Scholar
  5. 5.
    Al-Mufti, MM, Sydes, CL, Furness, SB, Grime, JP, Band, SR 1977A quantitative analysis of shoot phenology and dominance in herbaceous vegetation.J Ecol6575991Google Scholar
  6. 6.
    Bloom, AJ, Chapin III, FS, Mooney, HA 1985Resource limitation in plants: an economic analogy.Annu Rev Ecol Syst1636392Google Scholar
  7. 7.
    Chapin III, FS 1980The mineral nutrition of wild plants.Annu Rev Ecol Syst1123360Google Scholar
  8. 8.
    Connell, JE, Orias, E 1964Ecological regulation of species diversity.Am Nat98399414CrossRefGoogle Scholar
  9. 9.
    Connell, JE, Slatyer, RO 1977Mechanisms of succession in natural communities and their role in community stability and organization.Am Nat111111944CrossRefGoogle Scholar
  10. 10.
    Currie, DJ 1991Energy and large-scale patterns of animal- and plant-species richness.Am Nat1372749CrossRefGoogle Scholar
  11. 11.
    Darwin C. 1872. The origin of species. 6th London ed. Chicago: Thomas and Thomas.Google Scholar
  12. 12.
    Elton, CS 1958The ecology of invasions by animals and plants.MethuenLondonGoogle Scholar
  13. 13.
    Giblin, AE, Nadelhoffer, KJ, Shaver, GR, Laundre, JA, McKerrow, AJ 1991Biogeochemical diversity along a riverside toposequence in arctic Alaska.Ecol Monogr6141535Google Scholar
  14. 14.
    Gough, L, Grace, JB, Taylor, KL 1994The relationship between species richness and community biomass: the importance of environmental variables.Oikos702719Google Scholar
  15. 15.
    Grace, JB 1993The effect of habitat productivity on competition intensity.Trends Ecol Evol822930CrossRefGoogle Scholar
  16. 16.
    Grace, JB 1999The factors controlling species density in herbaceous plant communities: an assessment.Perspect Plant Ecol Evol Syst2128Google Scholar
  17. 17.
    Grime, JP 1973aCompetition and diversity in herbaceous vegetation: a reply.Nature24431011Google Scholar
  18. 18.
    Grime, JP 1973bControl of species diversity in herbaceous vegetation.J Environ Manage115167Google Scholar
  19. 19.
    Grime, JP 1979Plant strategies and vegetation processes.John Wiley and SonsChichester (UK)Google Scholar
  20. 20.
    Guo, Q, Berry, WL 1998Species richness and biomass: dissection of the hump-shaped relationships.Ecology7925559Google Scholar
  21. 21.
    Hedin, LO, Armesto, JJ, Johnson, AH 1995Patterns of nutrient loss from unpolluted, old–growth temperate forests: evaluation of biogeochemical theory.Ecology76493509Google Scholar
  22. 22.
    Herbert, DA, Rastetter, EB, Shaver, GR, Ågren, GI 1999Effects of plant growth characteristics on biogeochemistry and community composition in a changing climate.Ecosystems236782CrossRefGoogle Scholar
  23. 23.
    Huston, MA 1994Biological diversityCambridge University PressCambridgeGoogle Scholar
  24. 24.
    Huston, MA, DeAngelis, DL 1994Competition and coexistence: the effects of resource transport and supply rates.Am Nat14495477CrossRefGoogle Scholar
  25. 25.
    Lesack, FW, Melack, JM 1991The deposition, composition and potential sources of ionic solutes in rain of the central Amazon basin.Water Resour Res27295377CrossRefGoogle Scholar
  26. 26.
    Lovett, GM, Weathers, KC, Sobczak, W 2000Nitrogen saturation and retention in forested watersheds of the Catskill Mountains, NY.Ecol Appl107384Google Scholar
  27. 27.
    MacArthur, RH 1955Fluctuation of animal populations and a measure of community stability.Ecology365336Google Scholar
  28. 28.
    Margalef, R 1963On certain unifying principles in ecology.Am Nat9735774CrossRefGoogle Scholar
  29. 29.
    Magurran, A 1988Ecological diversity and its measurementPrinceton University PressPrincetonGoogle Scholar
  30. 30.
    Marrs, RH, Grace, JB, Gough, L 1996On the relationship between plant species diversity and biomass: a comment on a paper by Gough, Grace and Taylor.Oikos753236Google Scholar
  31. 31.
    Mittelbach, GG, Steiner, CF, Scheiner, SM, Gross, KL, Reynolds, HL, Waide, RB, Willig, MR, Dodson, SI, Gough, L 2001What is the observed relationship between species richness and productivity?Ecology82238196Google Scholar
  32. 32.
    Monsi, M, Saeki, T 1953Über den Lichtfaktor in den Pflanzengesellschaften und seine Bedeutung für die Stoffproduktion.Jpn J Bot142252Google Scholar
  33. 33.
    Olff, H, Huisman, J, Van Tooren, BF 1993Species dynamics and nutrient accumulation during early primary succession in coastal sand dunesJ Ecol81693706Google Scholar
  34. 34.
    Preston, FW 1962The canonical distribution of commonness and rarity.Ecology43185215Google Scholar
  35. 35.
    Rastetter, EB, Ågren, GI 2002Changes in individual allometry can lead to species coexistence without niche separation.Ecosystems5789801CrossRefGoogle Scholar
  36. 36.
    Rastetter, EB, Ågren, GI, Shaver, GR 1997Responses of N-limited ecosystems to increased CO2: a balanced-nutrition, coupled-element-cycles model.Ecol Appl744460Google Scholar
  37. 37.
    Rastetter, EB, Shaver, GR 1992A model of multiple-element limitation for acclimating vegetation.Ecology73115774Google Scholar
  38. 38.
    Rastetter, EB, Vitouse, PM, Fiel, C, Shave, GR, Herbert, D, Ågren, GI 2001Resource optimization and symbiotic N fixation.Ecosystems436988Google Scholar
  39. 39.
    Reader, RJ, Wilson, SD, Belcher, JW, Wisheau, I, Keddy, PA, Tilman, D, Morris, EC, Grace, JB, McGraw, JB, Olff, H,  et al. 1994Plant competition in relation to neighbor mass: an intercontinental study with Poa pratensis.Ecology75175360Google Scholar
  40. 40.
    Rosenzweig, ML, Abramski, Z 1993How are diversity and productivity related?Ricklefs, RESchluter, E eds. Species diversity in ecological communities: historical and geographical perspectivesUniversity of Chicago PressChicago5265Google Scholar
  41. 41.
    Schoener, TW 1976Alternatives to Lotka–Volterra competition: models of intermediate complexity.Theor Popul Biol1030933PubMedGoogle Scholar
  42. 42.
    Sokal, RR, Rohlf, FJ 1980Biometry: the principles and practice of statistics in biological research. 2nd ed.WH FreemanNew YorkGoogle Scholar
  43. 43.
    Tilman, D 1982Resource competition and community structurePrinceton University PressPrincetonGoogle Scholar
  44. 44.
    Tilman, D 1987Secondary succession and the pattern of plant dominance along experimental nitrogen gradients.Ecol Monogr57189214Google Scholar
  45. 45.
    Tilman, D 1990Constraints and tradeoffs: toward a predictive theory of competition and succession.Oikos58315Google Scholar
  46. 46.
    Tilman, D 1997Distinguishing between the effects of species diversity and species composition.Oikos80185Google Scholar
  47. 47.
    Tilman, D, Pacala, S 1993The maintenance of species richness in plant communities.Ricklefs, RESchluter, D eds. Species diversity in ecological communities: historical and geographical perspectivesUniversity of Chicago PressChicago1325Google Scholar
  48. 48.
    Tilman, D, Wedin, D 1991Plant traits and resource reduction for five grasses growing on a nitrogen gradient.Ecology72685700Google Scholar
  49. 49.
    Vitousek, PM 1982Nutrient cycling and nutrient use efficiency.Am Nat11955372CrossRefGoogle Scholar
  50. 50.
    Waide, RB, Willig, MR, Steiner, CF, Mittelbach, GG, Gough, L, Dodson, SI, Juday, GP, Parmenter, R 1999The relationship between primary productivity and species richness.Annu Rev Ecol Syst30257300CrossRefGoogle Scholar
  51. 51.
    Wedin, D, Tilman, D 1993Competition among grasses along a nitrogen gradient: initial conditions and mechanisms of competition.Ecol Monogr63199229Google Scholar
  52. 52.
    Wilson, SD, Tilman, D 1991Components of plant competition along an experimental gradient of nitrogen availability.Ecology7210508Google Scholar
  53. 53.
    Wilson, SD, Tilman, D 1995Competitive responses of eight old-field plant species in four environments.Ecology76116980Google Scholar
  54. 54.
    Zobel, K, Liira, J 1997A scale-independent approach to the richness vs biomass relationship in ground-layer plant communities.Oikos8032532Google Scholar

Copyright information

© Springer-Verlag New York, Inc. 2004

Authors and Affiliations

  • Darrell A. Herbert
    • 1
  • Edward B. Rastetter
    • 1
  • Laura Gough
    • 2
  • Gaius R. Shaver
    • 1
  1. 1.Ecosystems CenterMarine Biological Laboratory, Woods Hole, Massachusetts 02543USA
  2. 2.Department of Biological SciencesUniversity of Texas, Arlington, Texas 76019USA

Personalised recommendations