A General Hypothesis of Species Diversity

A new hypothesis, based on differences in the rates at which populations of competing species approach competitive equilibrium (reduction or exclusion of some species), is proposed to explain patterns of species diversity. The hypothesis assumes that most communities exist in a state of nonequilibrium where competitive equilibrium is prevented by periodic population reductions and environmental fluctuations. When competitive equilibrium is prevented, a dynamic balance may be established between the rate of competitive displacement and the frequency of population reduction, which results in a stable level of diversity. Under conditions of infrequent reductions, an increase in the population growth rates of competitors generally results in decreased diversity. This model clarifies an underlying pattern of variation in diversity and points out the common elements of previous hypotheses. Rather than arguing that either competition, predation, or productivity control diversity, it demonstrates that all of these may contribute to the same basic mechanism. In doing so, it not only explains the correlations of the other hypotheses with patterns of diversity, but also explains the exceptions that these hypotheses could not explain. This hypothesis may be applied to variations of diversity both on a latitudinal gradient and within specific regions.

[1]  John Bennet Lawes,et al.  XXVIII. Agricultural, botanical, and chemical results of experiments on the mixed herbage of permanent meadow, conducted for more than twenty years in succession on the same land. — Part. II. The botanical results , 1882, Philosophical Transactions of the Royal Society of London.

[2]  The effect of manuring, grazing and cutting on the yield, botanical and chemical composition of natural hill pastures. I. Yield and botanical section. , 1940 .

[3]  W. Milton,et al.  The Yield, Botanical and Chemical Composition of Natural Hill Herbage Under Manuring, Controlled Grazing and Hay Conditions , 1947 .

[4]  G. E. Hutchinson,et al.  CIRCULAR CAUSAL SYSTEMS IN ECOLOGY , 1948, Annals of the New York Academy of Sciences.

[5]  H. Swingle Experiments with Combinations of Largemouth Black Bass, Bluegills, and Minnows in Ponds , 1949 .

[6]  A. Tansley,et al.  The Tropical Rain Forest. , 1953 .

[7]  E. Nielsen On Organic Production in the Oceans , 1954 .

[8]  H. W. Harvey The chemistry and fertility of sea waters , 1955 .

[9]  J. Yount Factors That Control Species Numbers in Silver Springs, Florida1 , 1956 .

[10]  C. Elton Interspecific Competition , 1957, Nature.

[11]  R. Macarthur,et al.  Population Ecology of Some Warblers of Northeastern Coniferous Forests , 1958 .

[12]  J. T. Hack,et al.  Geomorphology and forest ecology of a mountain region in the central Appalachians , 1960 .

[13]  W. Murphy Ecological changes induced in moorland pastures by different fertilizer treatments. , 1960 .

[14]  J. P. Schulz The vegetation of Suriname: a series of papers on the plant communities of Suriname and their origin, distribution and relation to climate and habitat. Vol. II. Ecological studies on rain forest in northern Suriname. , 1960 .

[15]  J. Connell The Influence of Interspecific Competition and Other Factors on the Distribution of the Barnacle Chthamalus Stellatus , 1961 .

[16]  G. E. Hutchinson,et al.  The Balance of Nature and Human Impact: The paradox of the plankton , 2013 .

[17]  A. J. Willis Braunton Burrows: The Effects on the Vegetation of the Addition of Mineral Nutrients to the Dune Soils , 1963 .

[18]  THE STRUCTURE OF DIATOM COMMUNITIES UNDER VARYING ECOLOGICAL CONDITIONS , 1963, Annals of the New York Academy of Sciences.

[19]  P. Raven,et al.  BUTTERFLIES AND PLANTS: A STUDY IN COEVOLUTION , 1964 .

[20]  L. Williams Possible Relationships Between Plankton‐Diatom Species Numbers and Water‐Quality Estimates , 1964 .

[21]  J. Connell,et al.  The Ecological Regulation of Species Diversity , 1964, The American Naturalist.

[22]  A. Southward Limpet grazing and the control of vegetation on rocky shores , 1964 .

[23]  R. Macarthur PATTERNS OF SPECIES DIVERSITY , 1965 .

[24]  G. R. Hampson,et al.  An introduction to the study of deep-sea benthic faunal assemblages along the Gay Head-Bermuda transect* , 1965 .

[25]  Robert H. Whittaker,et al.  Vegetation of the Santa Catalina Mountains, Arizona: A Gradient Analysis of the South Slope , 1965 .

[26]  E. Pianka Latitudinal Gradients in Species Diversity: A Review of Concepts , 1966, The American Naturalist.

[27]  R. Paine Food Web Complexity and Species Diversity , 1966, The American Naturalist.

[28]  Harry F. Recher,et al.  On the Relation between Habitat Selection and Species Diversity , 1966, The American Naturalist.

[29]  R. Whittaker Forest Dimensions and Production in the Great Smoky Mountains , 1966 .

[30]  R. Macarthur,et al.  The Theory of Island Biogeography , 1969 .

[31]  M. C. Whiteside,et al.  Species Diversity in Chydorid (Cladocera) Communities , 1967 .

[32]  Richard S. Miller Pattern and Process in Competition , 1967 .

[33]  A. Kohn Environmental Complexity and Species Diversity in the Gastropod Genus Conus on Indo-West Pacific Reef Platforms , 1967, The American Naturalist.

[34]  E. Pianka On Lizard Species Diversity: North American Flatland Deserts , 1967 .

[35]  H. L. Sanders,et al.  Faunal diversity in the deep-sea , 1967 .

[36]  J. Karr Habitat and Avian Diversity on Strip-Mined Land in East-Central Illinois , 1968 .

[37]  S. McNaughton,et al.  Structure and Function in California Grasslands , 1968 .

[38]  Martin L. Cody,et al.  On the Methods of Resource Division in Grassland Bird Communities , 1968, The American Naturalist.

[39]  R. Levins Evolution in Changing Environments , 1968 .

[40]  A. Kohn MICROHABITATS, ABUNDANCE AND FOOD OF CONUS ON ATOLL REEFS IN THE MALDIVE AND CHAGOS ISLANDS , 1968 .

[41]  F. Ayala,et al.  Experimental Invalidation of the Principle of Competitive Exclusion , 1969, Nature.

[42]  R. H. Lowe-McConell Speciation in Tropical Environments , 1969 .

[43]  R. Macarthur,et al.  Patterns of communities in the tropics , 1969 .

[44]  J. Rieley,et al.  Ecological Aspects of the Mineral Nutrition of Plants. , 1969 .

[45]  H. L. Sanders Benthic marine diversity and the stability-time hypothesis. , 1969, Brookhaven symposia in biology.

[46]  R. Margalef,et al.  Diversity and stability: a practical proposal and a model of interdependence. , 1969, Brookhaven symposia in biology.

[47]  Miller Rs Competition and species diversity. , 1969, Brookhaven symposia in biology.

[48]  H. L. Sanders,et al.  Ecology of the deep-sea benthos. , 1969, Science.

[49]  J. Harper The role of predation in vegetational diversity. , 1969, Brookhaven symposia in biology.

[50]  R. Paine,et al.  THE EFFECTS OF GRAZING BY SEA URCHINS, STRONGYLOCENTROTUS SPP., ON BENTHIC ALGAL POPULATIONS1 , 1969 .

[51]  H. Recher Bird Species Diversity and Habitat Diversity in Australia and North America , 1969, The American Naturalist.

[52]  Sanders Hl,et al.  On the contribution of environmental predictability to species diversity. , 1969 .

[53]  M. Rosenzweig,et al.  Population Ecology of Desert Rodent Communities: Habitats and Environmental Complexity , 1969 .

[54]  L. Slobodkin,et al.  On the contribution of environmental predictability to species diversity. , 1969, Brookhaven symposia in biology.

[55]  E. Duffey The scientific management of animal and plant communities for conservation , 1973 .

[56]  Joseph H. Connell,et al.  A Predator‐Prey System in the Marine Intertidal Region. I. Balanus glandula and Several Predatory Species of Thais , 1970 .

[57]  D. Janzen Herbivores and the Number of Tree Species in Tropical Forests , 1970, The American Naturalist.

[58]  O. Loucks,et al.  Evolution of diversity, efficiency, and community stability. , 1970, American zoologist.

[59]  J. Vandermeer The Community Matrix and the Number of Species in a Community , 1970, The American Naturalist.

[60]  H. G. Baker,et al.  Evolution in the Tropics , 1970 .

[61]  P. Richerson,et al.  Contemporaneous disequilibrium, a new hypothesis to explain the "paradox of the plankton". , 1970, Proceedings of the National Academy of Sciences of the United States of America.

[62]  T. Whitmore,et al.  On the influence of soil properties on species distribution in a Malayan lowland Dipterocarp rain forest. , 1970 .

[63]  M. Soulé,et al.  The "Niche-Variation" Hypothesis: A Test and Alternatives , 1970, The American Naturalist.

[64]  M. Rosenzweig Paradox of Enrichment: Destabilization of Exploitation Ecosystems in Ecological Time , 1971, Science.

[65]  F J Ayala,et al.  Competition between Species: Frequency Dependence , 1971, Science.

[66]  L. Holdridge,et al.  Forest environments in tropical life zones: a pilot study. , 1971 .

[67]  A. Auclair,et al.  Diversity Relations of Upland Forests in the Western Great Lakes Area , 1971, The American Naturalist.

[68]  S. Hurlbert The Nonconcept of Species Diversity: A Critique and Alternative Parameters. , 1971, Ecology.

[69]  P. Dayton Competition, Disturbance, and Community Organization: The Provision and Subsequent Utilization of Space in a Rocky Intertidal Community , 1971 .

[70]  J. Connell On the role of the natural enemies in preventing competitive exclusion in some marine animals and in rain forest trees , 1971 .

[71]  J. Karr,et al.  Vegetation Structure and Avian Diversity in Several New World Areas , 1971, The American Naturalist.

[72]  R. Macarthur Mathematical Ecology and Its Place among the Sciences. (Book Reviews: Geographical Ecology. Patterns in the Distribution of Species) , 1974 .

[73]  J. Porter Predation by Acanthaster and Its Effect on Coral Species Diversity , 1972, The American Naturalist.

[74]  W. T. Williams,et al.  Regeneration and pattern in the subtropical rain forest , 1972 .

[75]  Reinterpretation of the Invalidation of the Principle of Competitive Exclusion , 1972, Nature.

[76]  P. Dayton,et al.  Role of biological disturbance in maintaining diversity in the deep sea , 1972 .

[77]  W. Murdoch,et al.  Diversity and Pattern in Plants and Insects , 1972 .

[78]  J. P. Grime Control of species density in herbaceous vegetation , 1973 .

[79]  S. Fretwell Populations in a seasonal environment. , 1973, Monographs in population biology.

[80]  C. Strobeck,et al.  N Species Competition , 1973 .

[81]  M. Rex Deep-Sea Species Diversity: Decreased Gastropod Diversity at Abyssal Depths , 1973, Science.

[82]  A. L. Koch,et al.  Coexistence resulting from an alternation of density dependent and density independent growth. , 1974, Journal of theoretical biology.

[83]  S. Levin Dispersion and Population Interactions , 1974, The American Naturalist.

[84]  John F. Riebesell PARADOX OF ENRICHMENT IN COMPETITIVE SYSTEMS , 1974 .

[85]  James P. Collins,et al.  Environmental Certainty, Trophic Level, and Resource Availability in Life History Evolution , 1974, The American Naturalist.

[86]  Enzyme variability in natural populations of Daphnia magna , 1976, Genetics.

[87]  P. Hebert Enzyme variability in natural populations of Daphnia magna. 3. Genotypic frequencies in intermittent populations. , 1974, Genetics.

[88]  Robert K. Peet,et al.  The Measurement of Species Diversity , 1974 .

[89]  A. L. Koch,et al.  Competitive coexistence of two predators utilizing the same prey under constant environmental conditions. , 1974, Journal of theoretical biology.

[90]  J. Karr Production, Energy Pathways, and Community Diversity in Forest Birds , 1975 .

[91]  J. Bunt Primary Productivity of Marine Ecosystems , 1975 .

[92]  P. Feeny Biochemical coevolution between plants and their insect herbivores , 1975, Coevolution of Animals and Plants.

[93]  J. Vandermeer,et al.  Interspecific competition: a new approach to the classical theory , 1975, Science.

[94]  J. Harper,et al.  Relationship Between Plant Weight and Numbers in Mixed Populations of Sinapsis alba (L.) Rabenh. and Lepidium sativum L. , 1976 .

[95]  B. Menge,et al.  Species Diversity Gradients: Synthesis of the Roles of Predation, Competition, and Temporal Heterogeneity , 1976, The American Naturalist.

[96]  J. Diamond,et al.  Ecology and Evolution of Communities , 1976, Nature.

[97]  R. McGehee,et al.  Coexistence of two competitors on one resource. , 1976, Journal of theoretical biology.

[98]  DAVID TITMAN,et al.  Ecological Competition Between Algae: Experimental Confirmation of Resource-Based Competition Theory , 1976, Science.

[99]  M. Rex Biological accomodation in the deep-sea benthos: comparative evidence on the importance of predation and productivity , 1976 .

[100]  R. McGehee,et al.  Coexistence of species competing for shared resources. , 1976, Theoretical population biology.

[101]  P. Grubb THE MAINTENANCE OF SPECIES‐RICHNESS IN PLANT COMMUNITIES: THE IMPORTANCE OF THE REGENERATION NICHE , 1977 .

[102]  P. Sale Maintenance of High Diversity in Coral Reef Fish Communities , 1977, The American Naturalist.

[103]  R. McGehee,et al.  Some mathematical problems concerning the ecological principle of competitive exclusion , 1977 .

[104]  Mark H Gromko,et al.  WHAT IS FREQUENCY‐DEPENDENT SELECTION? , 1977, Evolution; international journal of organic evolution.