Skip to main content
SpringerLink
Log in

Variation in species richness: Towards a unification of hypotheses

  • Published:
Folia Geobotanica et Phytotaxonomica Aims and scope Submit manuscript

Abstract

The question, “why do areas vary in species richness?” has been important throughout the history of ecology. It is difficult to answer definitively because we have so many (at least 120) plausible hypotheses. This abundance of hypotheses has led to a number of attempts to classify them. Unfortunately, richness hypotheses often defy such categorization. Instead of placing species richness hypotheses into categories, I suggest an alternative approach: to treat species richness hypotheses as violations of the assumptions of Gause’s Competitive Exclusion Principle. This is a very similar approach to the pedagogy of population genetics: evolution occurs if and only if at least one assumption of the Hardy-Weinberg principle is violated. The classification of hypotheses advocated here treats interspecific competition as a central organizing concept in community theory. However, it does not treat competition as an organizing concept in communities: indeed, the relaxation or disruption of competition is considered to be the status quo in the majority of communities.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Log in via an institution

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Institutional subscriptions

Similar content being viewed by others

References

  • Aarssen L.W. (1983): Ecological combining ability and competitive combining ability in plants: toward a general evolutionary theory of coexistence in systems of competition. — Am. Nat. 122: 707–731.

    Article  Google Scholar 

  • Aarssen L.W. (1992): Causes and consequences of variation in competitive ability in plant communities. — J. Veg. Sci. 3: 165–174.

    Article  Google Scholar 

  • Abugov R. (1982): Species diversity and the phasing of disturbance. — Ecology 63: 289–293.

    Article  Google Scholar 

  • Adams J.M. &Woodward F.I. (1989): Patterns in tree species richness as a test of the glacial extinction hypothesis. — Nature 339: 699–701.

    Article  Google Scholar 

  • Atkinson W.D. &Shorrocks B. (1981): Competition on a divided an ephemeral resource: a simulation model. — J. Anim. Ecol. 50: 461–471.

    Article  Google Scholar 

  • Auclair A.N. &Goff F.G. (1971): Diversity relations of upland forests in the western Great Lakes area. — Am. Nat. 205: 499–528.

    Article  Google Scholar 

  • Auerbach M. &Shmida A. (1987): Spatial scale and the determinants of plant species richness. — Tr. Ecol. Evol. 2: 238–242.

    Article  Google Scholar 

  • Barbour M.G., Burk J.H. &Pitts W.D. (1987): Terrestrial plant ecology. — Benjamin Cummings, Menlo Park.

    Google Scholar 

  • Begon M., Harper J.L. &Townsend C.R. (1990): Ecology: individuals, populations, and communities. — Blackwell Scientific, Boston.

    Google Scholar 

  • Bengtsson J., Fagerström T. &Rydin H. (1994): Competition and coexistence in plant communities. — Tr. Ecol. Evol. 9: 246–250.

    Article  Google Scholar 

  • Botkin D.B. (1990): Discordant Harmonies: a new ecology for the twenty-first century. — Oxford University Press, New York.

    Google Scholar 

  • Boughey A.S. (1973): Ecology of populations. — MacMillan, New York.

    Google Scholar 

  • Brewer R. (1988): The science of ecology. — Saunders College Publications, Philadelphia.

    Google Scholar 

  • Brown J.H. (1988): Species diversity. — In:Myers A.A. &Giller P.S. [eds.]: Analytical biogeography, Chapman and Hall, New York, pp. 57–89.

    Google Scholar 

  • Brown J.H. &Davidson D.W. (1977): Competition between seed-eating rodents and ants in desert ecosystems. — Science 196: 880–882.

    Article  PubMed  Google Scholar 

  • Brown J.H. &Kodric-Brown A. (1977): Turnover rates in insular biogeography: effect of immigration on extinction. — Ecology 58: 445–449.

    Article  Google Scholar 

  • Burrough P.A. (1981): Fractal dimensions of landscapes and other environmental data. — Nature 294: 240–242.

    Article  Google Scholar 

  • Bush E.O., Aho J.M. &Kennedy C.R. (1990): Ecological versus phylogenetic determinants of helminth parasite community richness. — Evol. Ecol. 4: 1–20.

    Article  Google Scholar 

  • Butterfield H. (1957): The origins of modern science. — Collier Books, New York.

    Google Scholar 

  • Case T.J. (1991): Invasion resistance, species build-up and community collapse in metapopulation models with interspecies competition. — Biol. J. Linn. Soc. 42: 239–266.

    Article  Google Scholar 

  • Coleman B.D., Mares M.A., Willig M.R. &Hsieh Y.-H. (1982): Randomness, area, and species richness. — Ecology 63: 1121–1133.

    Article  Google Scholar 

  • Collier B.D., Cox G.W., Johnson A.W. &Miller P.C. (1973): Dynamic ecology. — Prentice-Hall, Englewood Cliffs.

    Google Scholar 

  • Collins S.L. (1992): Fire frequency and community heterogeneity in tallgrass prairie vegetation. — Ecology 73: 2001–2006.

    Article  Google Scholar 

  • Condit R., Hubbell S.P. &Foster R.B. (1992): Recruitment near conspecific adults and the maintenance of tree and shrub diversity in a neotropical forest. — Am. Nat. 140: 261–286.

    Article  PubMed  CAS  Google Scholar 

  • Connell J.H. (1978): Diversity in tropical rain forests and coral reefs. — Science 199: 1302–1309.

    Article  PubMed  Google Scholar 

  • Connell J.H. &Lowman M.D. (1989): Low-diversity tropical rain forests: some possible mechanisms for their existence. — Am. Nat. 134: 88–119.

    Article  Google Scholar 

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

    Article  Google Scholar 

  • Cornell H.V. &Lawton J.H. (1992): Species interactions, local and regional processes, and limits to the richness of ecological communities: a theoretical perspective. — J. Anim. Ecol. 61: 1–12.

    Article  Google Scholar 

  • Cousins S.H. (1989): Species richness and the energy theory. — Nature 340: 350–351.

    Article  Google Scholar 

  • Crawley M.J. (1986): The structure of plant communities. — In:Crawley M.J. [ed.]: Plant Ecology, Blackwell, Oxford, pp. 1–50.

    Google Scholar 

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

    Article  Google Scholar 

  • DeAngelis D.L. &Waterhouse J.C. (1987): Equilibrium and nonequilibrium concepts in ecological models. — Ecol. Mon. 57: 1–21.

    Article  Google Scholar 

  • Diamond J. (1988): Factors controlling species diversity: overview and synthesis. — Ann. Miss. Bot. Gard. 75: 117–129.

    Article  Google Scholar 

  • Dobzhansky T. (1950): Evolution in the tropics. — Am. Sci. 38: 209–221.

    Google Scholar 

  • Fagerström T. (1988): Lotteries in communities of sessile organisms. — Tr. Ecol. Evol. 3: 303–306.

    Article  Google Scholar 

  • Fox J.F. (1977): Alternation and coexistence of tree species. — Am. Nat. 111: 69–89.

    Article  Google Scholar 

  • Gause G.F. (1937): Experimental populations of microscopic organisms. — Ecology 18: 173–179.

    Article  Google Scholar 

  • Gentry A.H. (1988): Changes in plant community diversity and floristic composition on environmental and geographical gradients. — Ann. Miss. Bot. Gard. 75: 1–34.

    Article  Google Scholar 

  • Giller P.S. (1984): Community structure and the niche. — Chapman and Hall, London.

    Google Scholar 

  • Gleason H.A. (1925): Species and area. — Ecology 6: 66–74.

    Article  Google Scholar 

  • Grime J.P. (1973): Control of species density in herbaceous vegetation. — J. Env. Manage. 1: 151–167.

    Google Scholar 

  • Grime J.P., Mackey J.M.L., Hillier S.H. &Read D.J. (1987): Floristic diversity in a model system using experimental microcosms. — Nature 328: 420–422.

    Article  Google Scholar 

  • Grinnell J. (1917): The niche-relationships of the California Thrasher. — Auk 34: 427–433.

    Google Scholar 

  • Grubb P. (1977): The maintenance of species richness in plant communities: the importance of the regeneration niche. — Biol. Rev. 52: 107–145.

    Article  Google Scholar 

  • Hairston N.G., Allan J.D., Colwell R.K., Futuyma D.J., Howell J., Lubin M.D., Mathias J. &Van der Meer J.H. (1968): The relationship between species diversity and stability: an experimental approach with protozoa and bacteria. — Ecology 49: 1091–1101.

    Article  Google Scholar 

  • Hara T. (1993): Effects of variation in individual growth on plant species coexistence. — J. Veg. Sci. 4: 409–416.

    Article  Google Scholar 

  • Hardy G.H. (1908): Mendelian proportions in a mixed population. — Science 28: 49–50.

    Article  PubMed  Google Scholar 

  • Harman W.N. (1972): Benthic substrates: their effect on freshwater molluscs. — Ecology 53: 271–272.

    Article  Google Scholar 

  • Harrer J.L. (1977): Population biology of plants. — Academic Press, London.

    Google Scholar 

  • Hart T.B. (1990): Monospecific dominance in tropical rain forests. — Tr. Ecol. Evol. 5: 6–11.

    Article  Google Scholar 

  • Haukisalmi V. &Henttonen H. (1993): Coexistence in helminths of the bank voleClethrionomys glareolus. II. Intestinal distribution and interspecific interactions. — J. Anim. Ecol. 62: 230–238.

    Article  Google Scholar 

  • Hawkins B.A., Shaw M.R. &Askew R.R. (1992): Relations among assemblage size, host specialization, and climatic variability in North American parasitoid communities. — Am. Nat. 139: 58–79.

    Article  Google Scholar 

  • Heatwole H. (1991): Factors affecting the number of species of plants on islands of the Great Barrier Reef, Australia. — J. Biogeogr. 18: 213–221.

    Article  Google Scholar 

  • Hofstadter D.R. (1980): Gödel, Escher, Bach: an eternal golden braid. — Random House, New York.

    Google Scholar 

  • Holt R.D. (1992): A neglected facet of island biogeography: the role of internal spatial dynamics in area effects. — Theor. Popul. Biol. 41: 354–371.

    Article  Google Scholar 

  • Huston M. (1979): A general hypothesis of species diversity. — Am. Nat. 113: 81–101.

    Article  Google Scholar 

  • Huston M.A. (1985): Patterns of species diversity on coral reefs. — Ann. Rev. Ecol. Syst. 16: 149–177.

    Article  Google Scholar 

  • Hutchinson G.E. (1958): Concluding remarks. — Cold Spring Harbor Symp. Quant. Biol. 22: 415–427.

    Google Scholar 

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

    Article  Google Scholar 

  • Hutchinson G.E. &Deevey E.S. (1949): Ecological studies on populations. — Surv. Biol. Prog. 1: 325–359.

    Google Scholar 

  • Janos D.P. (1983): Tropical mycorrhizas, nutrient cycles and plant growth. — In:Sutton S.L., Whitmore T.C. &Chadwick A.C. [eds.]: Tropical rain forest: ecology and management, Blackwell Scientific, Oxford, pp. 327–345.

    Google Scholar 

  • Janzen D.H. (1970): Herbivores and the number of tree species in tropical forests. — Am. Nat. 104: 501–528.

    Article  Google Scholar 

  • Jenkins J.B. (1979): Genetics. — Houghton Mifflin Company, Boston.

    Google Scholar 

  • Jones C.G. &Lawton J.H. (1991): Plant chemistry and insect species richness of British umbellifers. — J. Anim. Ecol. 60: 767–777.

    Article  Google Scholar 

  • Kelly B.J., Wilson J.B. &Mark A.F. (1989): Causes of the species-area relation: a study of islands in Lake Manapouri, New Zealand. — J. Ecol. 77: 1021–1028.

    Article  Google Scholar 

  • Kingsland S.E. (1991): Defining ecology as a science. — In:Real L.A. &Brown J.H. [eds.]: Foundations of ecology, University of Chicago Press, Chicago, pp. 1–13.

    Google Scholar 

  • Koetsier P., Dey P., Mladenka G. &Check J. (1990): Rejecting ecological theory — a cautionary note. — Bull. Ecol. Soc. Am. 71: 229–230.

    Google Scholar 

  • Kohyama T. (1993): Size-structured tree populations in gap-dynamic forests — the forest architecture hypothesis for the stable coexistence of species. — J. Ecol. 81: 131–143.

    Article  Google Scholar 

  • Kuhn T.S. (1970): The structure of scientific revolutions. — University of Chicago Press, Chicago.

    Google Scholar 

  • Latham R.E. (1992): Co-occurring tree species change rank in seedling performance with resources varied experimentally. — Ecology 73: 2129–2144.

    Article  Google Scholar 

  • Latham R.E. &Ricklefs R.E. (1993): Global patterns of tree species richness in moist forests: energy-diversity theory does not account for variation in species richness. — Oikos 67: 325–333.

    Article  Google Scholar 

  • Lavorel S. &Lebreton J.D. (1992): Evidence for lottery recruitment in Mediterranean old fields. — J. Veg. Sci. 3: 91–100.

    Article  Google Scholar 

  • Lechowicz M. &Bell G. (1991): The ecology and genetics of fitness in forest plants. II. Microspatial heterogeneity of the edaphic environment. — J. Ecol. 79: 687–696.

    Article  Google Scholar 

  • Levin S.A. (1974): Dispersion and population interactions. — Am. Nat. 108: 207–228.

    Article  Google Scholar 

  • Loreau M. (1992): Time scale of resource dynamics and coexistence through time partitioning. — Theor. Popul. Biol. 41: 401–412.

    Article  Google Scholar 

  • Lubchenco J. (1978): Plant species diversity in a marine intertidal community: importance of herbivore food preference and algal competitive abilities. — Am. Nat. 112: 23–39.

    Article  Google Scholar 

  • Luescher A. &Jacquard P. (1991): Coevolution between interspecific plant competitors? — Tr. Ecol. Evol. 6: 355–358.

    Article  Google Scholar 

  • MacArthur R.H. (1965): Pattern of species diversity. — Biol. Rev. 40: 510–533.

    Article  Google Scholar 

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

    Article  Google Scholar 

  • MacArthur R.H. &Wilson E.O. (1967): The theory of island biogeography. — Princeton University Press, Princeton.

    Google Scholar 

  • May R.M. (1988): How many species are there on Earth? — Science 241: 1441–1449.

    Article  PubMed  Google Scholar 

  • McIntosh R.P. (1985): The background of ecology. — Cambridge University Press, Cambridge.

    Google Scholar 

  • Milligan B.G. (1985): Evolutionary divergence and character displacement in two phenotypically-variable, competing species. — Evol. 39: 1207–1222.

    Article  Google Scholar 

  • Milne B.T. (1991): Lessons from applying fractal models to landscape patterns. — In:Turner M.G. &Gardner R.H. [eds.]: Quantitative methods in landscape ecology. Springer-Verlag, Berlin, pp. 199–235.

    Google Scholar 

  • Milne B.T. &Forman R.T.T. (1986): Peninsulas in Maine: woody plant diversity, distance, and environmental patterns. — Ecology 67: 967–974.

    Article  Google Scholar 

  • Mohler C.L. (1990): Co-occurrence of oak subgenera: implications for niche differentiation. — Bull. Torr. Bot. Club 117: 247–255.

    Article  Google Scholar 

  • Murdoch W.W. (1991): Equilibrium and non-equilibrium paradigms. — Bull. Ecol. Soc. Am. 72: 49–51.

    Google Scholar 

  • Nilsson C. &Wilson S.D. (1991): Convergence in plant community structure along disparate gradients: are lakeshores inverted mountainsides? — Am. Nat. 137: 774–790.

    Article  Google Scholar 

  • Nilsson C., Grelsson G., Dynesius M., Johansson M.E. &Sperens U. (1991): Small rivers behave like large rivers: effects of postglacial history on plant species richness along riverbanks. — J. Biogeogr. 18: 533–541.

    Article  Google Scholar 

  • Nilsson S.G., Bengtsson J. &Ás S. (1988): Habitat diversity or area per se? Species richness of woody plants, carabid beetles, and land snails on islands. — J. Anim. Ecol. 57: 685–704.

    Article  Google Scholar 

  • Noss R.F. (1991): Effects of edge and internal patchiness on avian habitat use in an old-growth Florida hammock. — Nat. Areas J. 11: 34–47.

    Google Scholar 

  • Odum E.P. (1959): Fundamentals of ecology. — Saunders, Philadelphia.

    Google Scholar 

  • Odum E.P. (1975): Ecology. — Holt, Rinehart and Winston, New York.

    Google Scholar 

  • Osborne L.L. &Wiley M.J. (1992): Influence of tributary spatial position on the structure of warmwater fish communities. — Can. J. Fish. Aquat. Sci. 49: 671–681.

    Article  Google Scholar 

  • Pagel M.D., May R.M. &Collie A.R. (1991): Ecological aspects of the geographical distribution and diversity of mammalian species. — Am. Nat. 137: 791–815.

    Article  Google Scholar 

  • Paine R.T. (1966): Food web complexity and species diversity. — Am. Nat. 100: 65–75.

    Article  Google Scholar 

  • Palmer M.W. (1987): Variability in species richness within Minnesota oldfields: a use of the variance test. — Vegetatio 70: 61–64.

    Google Scholar 

  • Palmer M.W. (1988): Fractal geometry: a tool for describing spatial patterns of plant communities. — Vegetatio 75: 91–102.

    Article  Google Scholar 

  • Palmer M.W. (1990a): Spatial scale and patterns of species-environment relationships in hardwood forests of the North Carolina piedmont. — Coenoses 5: 79–87.

    Google Scholar 

  • Palmer M.W. (1990b): Spatial scale and patterns of vegetation, flora and species richness in hardwood forests of the North Carolina piedmont. — Coenoses 5: 89–96.

    Google Scholar 

  • Palmer M.W. (1991): Patterns of species richness among North Carolina hardwood forests: tests of two Hypotheses. — J. Veg. Sci. 2: 361–366.

    Article  Google Scholar 

  • Palmer M.W. (1992): The coexistence of species in fractal landscapes. — Am. Nat. 139: 375–397.

    Article  Google Scholar 

  • Palmer M.W. &Dixon P.M. (1990): Small scale environmental variability and the analysis of species distributions along gradients. — J. Veg. Sci. 1: 57–65.

    Article  Google Scholar 

  • Peet R.K. &Christensen N.L. (1980): Hardwood forest vegetation of the North Carolina Piedmont. — Veröff. Geobot. Inst. ETH Stift. Rübel, Zürich 69: 14–39.

    Google Scholar 

  • Petraitis P.S., Latham R.E. &Niesenbaum R.A. (1989): The maintenance of species diversity by disturbance. — Quart. Rev. Biol. 64: 393–418.

    Article  Google Scholar 

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

    Article  Google Scholar 

  • Pianka E.R. (1973): The structure of lizard communities. — Ann. Rev. Ecol. Syst. 4: 53–74.

    Article  Google Scholar 

  • Pickett S.T.A. (1980): Non-equilibrium coexistence of plants. — Bull. Torr. Bot. Club. 107: 238–248.

    Article  Google Scholar 

  • Pimentel D. (1968): Population regulation and genetic feedback. — Science 159: 1432–1437.

    Article  PubMed  CAS  Google Scholar 

  • Pimm S.L. (1991): The balance of nature? — The University of Chicago Press, Chicago.

    Google Scholar 

  • Pontin A.J. (1982): Competition and coexistence of species. — Pitman, London.

    Google Scholar 

  • Popper K. (1935): Logik der Forschung: Zur Erkenntnistheorie der modernen Natuwissenschaft. — Springer, Wien.

    Google Scholar 

  • Quinn J.F. &Harrison S.P. (1988): Effects of habitat fragmentation and isolation on species richness: evidence from biogeographic patterns. — Oecologia 75: 132–140.

    Article  Google Scholar 

  • Real R., Vargas J.M. &Antnez A. (1993): Environmental influences on local amphibian diversity: the role of floods on river basins. — Biodivers. Conserv. 2: 376–399.

    Article  Google Scholar 

  • Rebele F. (1992): Colonization and early succession on anthropogenic soils. — J. Veg. Sci. 3: 201–208.

    Article  Google Scholar 

  • Rex M.A. (1981): Community structure in the deep sea benthos. — Ann. Rev. Ecol. Syst. 12: 331–353.

    Article  Google Scholar 

  • Ricklefs R.E. (1976): The economy of nature. — Chiron Press, Portland.

    Google Scholar 

  • Ricklefs R.E. (1979): Ecology. — Chiron Press, New York.

    Google Scholar 

  • Ricklefs R.E. (1987): Community diversity: relative roles of local and regional processes. — Science 235: 167–171.

    Article  PubMed  Google Scholar 

  • Robinson G.R. &Quinn J.F. (1988): Extinction, turnover and species diversity in an experimentally fragmented California annual grassland. — Oecologia 76: 71–82.

    Google Scholar 

  • Samson D.A., Philippi T.E. &Davidson D.W. (1992): Granivory and competition as determinants of annual plant diversity in the Chihuahuan desert. — Oikos 65: 61–80.

    Article  Google Scholar 

  • Sanders H.L. (1968): Marine benthic diversity: a comparative study. — Am. Nat. 102: 243–282.

    Article  Google Scholar 

  • Schoener A. (1988): Experimental island biogeography. — In:Myers A.A. &Giller P.S. [eds.]: Analytical Biogeography, Chapman and Hall, New York, pp. 483–512.

    Google Scholar 

  • Schupp E.W. (1992): The Janzen-Connell model for tropical tree diversity: population implications and the importance of spatial scale. — Am. Nat. 140: 526–530.

    Article  PubMed  CAS  Google Scholar 

  • Schwartz M.W. (1988): Species diversity patterns in woody flora on three North American peninsulas. — J. Biogeogr. 15: 759–774.

    Article  Google Scholar 

  • Shmida A. &Ellner S. (1984): Coexistence of plant species with similar niches. — Vegetatio 58: 29–55.

    Google Scholar 

  • Shmida A. &Wilson M.V. (1985): Biological determinants of species diversity. — J. Biogeogr. 12: 1–21.

    Article  Google Scholar 

  • Silvertown J. (1980): The dynamics of a grassland ecosystem: botanical equilibrium in the park grass experiment. — J. Appl. Ecol. 17: 491–504.

    Article  Google Scholar 

  • Simpson G.G. (1964): Species density of North American recent mammals. — Syst. Zool. 13: 57–73.

    Article  Google Scholar 

  • Smith R.L. (1990): Ecology and field biology. — Harper & Row, New York.

    Google Scholar 

  • Stevens G.C. (1992): The elevational gradient in altitudinal range: an extension of Rapoport’s latitudinal rule to altitude. — Am. Nat. 140: 893–911.

    Article  PubMed  CAS  Google Scholar 

  • Stone L. &Roberts A. (1991): Conditions for a species to gain advantage from the presence of competitors. — Ecology 72: 1964–1972.

    Article  Google Scholar 

  • Szentkiralyi F. &Kozar F. (1991): How many species are there in apple insect communities? Testing the resource diversity and intermediate disturbance hypotheses. — Ecol. Ent. 16: 491–504.

    Article  Google Scholar 

  • Tangney R.S., Wilson J.B. &Mark A.F. (1990): Bryophyte island biogeography: a study in Lake Manapouri, New Zealand. — Oikos 59: 21–26.

    Article  Google Scholar 

  • Taylor D.R., Aarssen L.W. &Loehle C. (1990): On the relationship between r/K selection and environmental carrying capacity: a new habitat templet for plant life history strategies. — Oikos 58: 239–250.

    Article  Google Scholar 

  • Terborgh J. (1977): Bird species diversity on an Andean elevational gradient. — Ecology 58: 1007–1019.

    Article  Google Scholar 

  • Tilman D. (1982): Resource competition and community structure. — Princeton University Press, Princeton, New Jersey.

    Google Scholar 

  • Tilman D. (1985): The resource ratio hypothesis of succession. — Am. Nat. 125: 827–852.

    Article  Google Scholar 

  • Tscharntke T. (1992): Cascade effects among four trophic levels: bird predation on galls affect density-dependent parasitism. — Ecology 73: 1689–1698.

    Article  Google Scholar 

  • Warner R.R. &Chesson P.L. (1985): Coexistence mediated by recruitment fluctuations: a field guide to the storage effect. — Am. Nat. 125: 769–787.

    Article  Google Scholar 

  • Whittaker R.H. (1965): Dominance and diversity in land plant communities. — Science 147: 250–260.

    Article  PubMed  Google Scholar 

  • Whittaker R.H., Levin S.A. &Root R.B. (1973): Niche, habitat, and biotope. — Am. Nat. 107: 321–338.

    Article  Google Scholar 

  • Williamson M.H. &Lawton J.H. (1990): Fractal geometry of ecological habitats. 73- In:Bell S.S. [ed.]: Habitat structure, Chapman and Hall, London, pp. 69–86.

    Google Scholar 

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

    Article  Google Scholar 

  • Wilson E.O. &Bossert W.H. (1971): A primer of population biology. — Sinauer Associates, Sunderland.

    Google Scholar 

  • Wilson J.B. (1990): Mechanisms of species coexistence: twelve explanations for Hutchinson’s paradox of the plankton: evidence from New Zealand plant communities. — N.Z. J. Ecol. 13: 17–42.

    Google Scholar 

  • Wilson J.B., Gitay H. &Agnew A.D.Q. (1987): Does niche limitation exist? — Funct. Ecol. 1: 391–397.

    Article  Google Scholar 

  • Woods K.D. (1979): Reciprocal replacement and the maintenance of codominance in a beech-maple forest. — Oikos 33: 31–39.

    Article  Google Scholar 

  • Zimmerman J.L. (1992): Density-independent factors affecting the avian diversity of the tallgrass prairie community. — Wilson Bull. 104: 85–94.

    Google Scholar 

  • Zobel M. (1992): Plant species coexistence — the role of historical, evolutionary and ecological factors. — Oikos 65: 314–320.

    Article  Google Scholar 

Download references

Author information

Authors and Affiliations

  1. Botany Department, Oklahoma State University, 74078, Stillwater, OK, USA

    Michael W. Palmer

  2. Department of Ecological Botany, Uppsala University, Uppsala, Sweden

    Michael W. Palmer

Authors
  1. Michael W. Palmer
    View author publications

    You can also search for this author in PubMed Google Scholar

Corresponding author

Correspondence to Michael W. Palmer.

Rights and permissions

Reprints and permissions

About this article

Cite this article

Palmer, M.W. Variation in species richness: Towards a unification of hypotheses. Folia geobot. phytotax. 29, 511–530 (1994). https://doi.org/10.1007/BF02883148

Download citation

  • Issue Date:

  • DOI: https://doi.org/10.1007/BF02883148

Keywords

Search

Navigation