Abstract
One of the basic questions in ecology is the relationship between species living at the same trophic level, and in particular the relationship between species that may compete for the available resources of food and places to live. One generalization relevant to this problem is known as the competitive exclusion principle, or Gause’s principle. According to this principle, two species competing for the same limited resources cannot coexist in the same locality; that is, one or the other species will sooner or later be eliminated. Considerable disagreement exists among ecologists concerning the validity and the importance of this principle. It is my purpose to show that the principle of competitive exclusion cannot be defended in its generality: two species may compete for limited resources and still coexist. Some of the conditions that allow competitive coexistence are discussed in the latter part of this paper. An investigation of the concepts of ecological niche and competition, which are essential for a precise formulation of the principle of competitive exclusion, precedes the consideration of the principle itself, and of the Volterra equations for species competition, which constitute the mathematical foundation of the principle.
Supported by PHS career development award K3 GM37265 from the National Institute of General Medical Sciences, and by contract AT-(30-1)-3096, U. S. Atomic Energy Commission.
Access this chapter
Tax calculation will be finalised at checkout
Purchases are for personal use only
Preview
Unable to display preview. Download preview PDF.
References
Anderson, W. W. 1969. Polymorphism resulting from the mating advantage of rare male genotypes (in press).
Andrewartha, H. G., and L. C. Birch. 1954. The Distribution and Abundance of Animals. Chicago, University of Chicago Press.
Anonymous. 1944. Symposium on “The ecology of closely allied species.” J. Anim. Ecol., 13:176–177.
Ayala, F. J. 1965a. Relative fitness of populations of Drosophila serrata and Drosophila birchii. Genetics, 51:527–544.
Ayala, F. J. 1965b. Sibling species of the Drosophila serrata group. Evolution, 19:538–545.
Ayala, F. J. 1966a. Reversal of dominance in competing species of Drosophila. Amer. Natural., 100:81–83.
Ayala, F. J. 1966b. Dynamics of populations. I. Factors controlling population growth and population and population size in Drosophila serrata. Amer. Natural. 100:333–344.
Ayala, F. J. 1967. Dynamics of populations. II. Factors controlling population growth and population size in Drosophila pseudoobscura and Drosophila melanogaster. Ecology, 48:67–75.
Ayala, F. J. 1968a. Evolution of fitness. II. Correlated effects of natural selection on the productivity and size of experimental populations of Drosophila serrata. Evolution, 22:55–65.
Ayala, F. J. 1968b. Environmental factors limiting the productivity and size of experimental populations of Drosophila serrata and D. birchii. Ecology, 49:562–565.
Ayala, F. J. 1968c. Genotype, environment, and population numbers. Science, 162:1453–1459.
Ayala, F. J. 1969a. Evolution of fitness. IV. Genetic evolution of interspecific competitive ability in Drosophila. Genetics (in press).
Ayala, F. J. 1969b. Genetic polymorphism and interspecific competitive ability in Drosophila. Genet. Res. (in press).
Bagenal, T. B. 1951. A note on the papers of Elton and Williams on the generic relations of species in small ecological communities. J. Anim. Ecol., 20:242–245.
Bakker, K. 1961. An analysis of factors which determine success in competition for food among larvae of Drosophila melanogaster. Arch. Neerl. Zool., 14:200–281.
Birch, L. C. 1955. Selection in Drosophila pseudoobscura in relation to crowding. Evolution, 9:389–399.
Birch, L. C. 1957. The meanings of competition. Amer. Natural., 91:5–18.
Brown, W. L., and E. O. Wilson. 1956. Character displacement. System. Zool., 5:49–64.
Chitty, D. 1960. Population processes in the vole and their relevance to general theory. Canad. J. Zool., 38:99–113.
Chitty, D. 1965. Predicting qualitative changes in insect populations. Proc. XIII Int. Congr. Entom., London (1964). pp. 384–386.
Clarke, B. 1962. Balanced polymorphism and the diversity of sympatric species. Systematics Assoc. Publ., 4:47–70.
Clarke, B. 1964. Frequency-dependent selection for the dominance of rare polymorphic genes. Evolution, 18:364–369.
Cole, L. C. 1960. Competitive exclusion. Science, 132:348–349.
Cooper, D. M., and Th. Dobzhansky. 1956. Studies on the ecology of Drosophila in the Yosemite region of California. I. The occurrence of species of Drosophila in different life zones and at different seasons. Ecology, 37:526–533.
Crombie, A. C. 1945. On competition between different species of graminivorous insects. Proc. Roy. Soc. [Biol.], 132:362–395.
Crombie, A. C. 1946. Further experiments on insect competition. Proc. Roy. Soc. [Biol.], 133: 76–109.
Crombie, A. C. 1947. Interspecific competition. J. Anim. Ecol., 16:44–73.
Da Cunha, A. B., Th. Dobzhansky, and A. Sokoloff. 1951. On food preferences of sympatric species of Drosophila. Evolution, 5:97–101.
Dawood, M. M., and M. W. Strickberger. 1969. The effect of larval interaction on viability in Drosophila melanogaster. III. Effects of biotic residues (in press).
Dawson, P. 1968. Mutation as a source of indeterminism in Tribolium “competition” experiments. Genetics, 60:172.
DeBach, P. 1966. The competitive displacement and coexistence principles. Ann. Rev. Entom., 11:183–212.
Del Solar, E. 1968. Selection for and against gregariousness in the choice of oviposition sites by Drosophila pseudoobscura. Genetics, 58:275–282.
Del Solar, E. and H. Palomino. 1966. Choice of oviposition in Drosophila melanogaster. Amer. Natural., 100:127–133.
Dobzhansky, Th. 1944. Chromosomal races in Drosophila pseudoobscura and Drosophila persimilis. Carnegie Instit. Washington Publ. 554, pp. 47–144.
Dobzhansky, Th. and B. Spassky. 1953. Genetics of natural poulations. XXI. Concealed variability in two sympatric species of Drosophila. Genetics, 38:471–484.
Elton, C. S. 1927. Animal Ecology. London, Sidgwick and Jackson.
Elton, C. S. 1946. Competition and the structure of animal communities. J. Anim. Ecol., 15:54–68.
Elton, C. S. and R. S. Miller. 1954. The ecological survey of animal communities: with a practical system of classifying habitats by structural characters. J. Ecol., 42: 460–496.
Feller, W. 1940. On the logistic law of growth and its empirical verifications in Biology. Acta Biotheor. [A] (Leiden), 5:51–66.
Ford, E. B. 1964. Ecological Genetics. London, Methuen.
Gause, G. F. 1934. The Struggle for Existence. Baltimore, The Williams & Wilkins Co. (Repr., 1964. New York, Hafner Publishing Co., Inc.)
Gause, G. F. and A. A. Witt. 1935. Behavior of mixed populations and the problem of natural selection. Amer. Natural., 69:596–609.
Grinnell, J. 1904. The origin and distribution of the chestnut-backed chickadee. Auk, 21:364–382.
Grinnell, J. 1917. The niche-relationships of the California thrasher. Auk, 34:427–433.
Grinnell, J. 1924. Geography and evolution. Ecology 5:225–229.
Haldane, J. B. S. 1924. A mathematical theory of natural and artificial selection. Trans. Cambridge Philos. Soc, 23:19–41.
Haldane, J. B. S. and S. D. Jayakar. 1963. Polymorphism due to selection of varying direction. J. Genetics, 58:237–242.
Hardin, G. 1960. The competitive exclusion principle. Science, 131:1292–1298.
Harris, P., D. F. Riordan, and D. Cooke. 1969. Mosquitoes feeding on insect larvae. Science, 164:184–185.
Hutchinson, G. E. 1957. Concluding remarks. Sympos. Quant. Biol., 22:415–427.
Hutchinson, G. E. 1965. The Ecological Theater and the Evolutionary Play. New Haven, Conn., Yale University Press.
Kettlewell, H. B. D. 1955. Selection experiments on industrial melanism in the Lepidoptera. Heredity (London), 9:323–342.
Kojima, K., and K. Yarbrough. 1967. Frequency dependent selection at the Esterase 6 locus in Drosophila melanogaster. Proc. Nat. Acad. Sci. (U.S.A.), 57:645–649.
Lack, D. 1944. Ecological aspects of species formation in passerine birds. Ibis, 86: 260–286.
Lerner, I. M., and F. K. Ho. 1961. Genotype and competitive ability in Tribolium species. Amer. Natural., 95:329–343.
Levene, H. 1953. Genetic equilibrium when more than one ecological niche is available. Amer. Natural., 87:331–333.
Levene, H. O. Pavlovsky, and Th. Dobzhansky. 1954. Interaction of the adaptive values in polymorphic experimental populations of Drosophila pseudoobscura. Evolution, 8:335–349.
Levin, B. R. 1967. Genetic variability and selection in a system of species competition. Ph.D. thesis. Ann Arbor, University of Michigan.
Lewontin, R. C. 1955. The effects of population density and composition on viability in Drosophila melanogaster. Evolution, 9:27–41.
Lewontin, R. C. and Y. Matsuo. 1963. Interaction of genotypes determining viability in Drosophila busckii. Proc. Nat. Acad. Sci. (U.S.A.), 49:270–278.
Lewontin, R. C. and M. J. D. White. 1960. Interaction between inversion polymorphisms of two chromosomal pairs in the grasshopper, Moraba scurra. Evolution, 14:116–129.
L’Heritier, P., and G. Teissier. 1937. Elimination des forms mutants dans les populations des Drosophiles. Cas des Drosophiles Bar. C. R. Soc. Biol. (Paris), 124:880–882.
Lotka, A. J. 1925. Elements of Physical Biology. Baltimore, The Williams & Wilkins Co.
Lotka, A. J. 1932. The growth of mixed populations: Two species competing for a common food supply. J. Washington Acad. Sci., 22:461–469.
Merrell, D. J. 1951. Interspecific competition between Drosophila funebris and Drosophila melanogaster. Amer. Natural., 85:159–169.
Miller, R. S. 1964. Larval competition in Drosophila melanogaster and D. simulans. Ecology, 45:132–148.
Miller, R. S. 1967. Pattern and process in competition. Advances Ecol. Res., 4:1–74.
Moore, J. A. 1952a. Competition between Drosophila melanogaster and Drosophila simulans. II. The improvement of competitive ability through selection. Proc. Nat. Acad. Sci. (U.S.A.), 38:813–817.
Moore, J. A. 1952b. Competition between Drosophila melanogaster and Drosophila simulans. I. Population cage experiments. Evolution, 6:407–420.
Nicholson, A. J. 1954. An outline of the dynamics of animal populations. Aust. J. Zool., 2:9–65.
Nicholson, A. J. 1957. The self-adjustment of populations to change. Sympos. Quant. Biol., 22:153–172.
Narise, T. 1965. The effect of relative frequency of species in competition. Evolution, 19:350–354.
Park, T. 1948. Experimental studies of interspecies competition. I. Competition between populations of the flour beetles, Tribolium confusum Duval and Tribolium castaneum Herbst. Ecol. Monographs, 18:265–307.
Park, T. 1954. Experimental studies of interspecies competition. II. Temperature, humidity, and competition in two species of Tribolium. Physiol. Zool., 27:177–238.
Patten, B. C. 1961. Competitive exclusion. Science, 134:1599–1601.
Pearl, R. 1926. The Biology of Population Growth. New York, Alfred A. Knopf, Inc.
Pearl, R. J. R. Miner, and S. L. Parker. 1927. Experimental studies on the duration of life. XI. Density of population and life duration in Drosophila. Amer. Natural., 61:289–318.
Pearl, R. and L. J. Reed. 1920. On the rate of growth of the population of the United States since 1790 and its mathematical representation. Proc. Nat. Acad. Sci. (U.S.A.), 6:275–288.
Petit, C. 1966. La concurrence larvaire et le maintien du polymorphism. C. R. Acad. Sci. (Paris), 263:1262–1265.
Petit, C. and L. Ehrman. 1969. Sexual selection in Drosophila. In Dobzhansky, Th., Hecht, M. K., and Steere, W. C., eds. Evolutionary Biology, Vol. 3. New York, Appleton-Century-Crofts (in press).
Pimentel, D. 1961. Animal population regulation by the genetic feed-back mechanism. Amer. Natural., 95:65–79.
Pimentel, D. 1968. Population regulation and genetic feedback. Science, 159:1432–1437.
Pimentel, D. E. H. Feinberg, P. W. Wood, and J. T. Hayes. 1965. Selection, spatial distribution, and the coexistence of competing fly species. Amer. Natural., 99:97–109.
Prout, T. 1968. Sufficient conditions for multiple niche polymorphism. Amer. Natural., 102:493–496.
Ross, H. 1957. Principles of natural coexistence indicated by leafhopper populations. Evolution, 11:113–129.
Sang, J. H. 1950. Population growth in Drosophila cultures. Biol. Rev., 25:188–219.
Slobodkin, L. B. 1954. Population dynamics in Daphnia obtusa Kurz. Ecological Monographs, 24:69–88.
Slobodkin, L. B. 1957. A laboratory study of the effect of removal of newborn animals from a population. Proc. Nat. Acad. Sci. (U.S.A.), 43:780–782.
Slobodkin, L. B. 1961. Growth and Regulation of Animal Populations. New York, Holt, Rinehart & Winston, Inc.
Spassky, B. 1957. Morphological differences between sibling species of Drosophila. Univ. Tex. Publ. No. 5721:48–61.
Ullyett, G. C. 1950. Competition for food and allied phenomena in sheep-blowfly populations. Philos. Trans. Roy. Soc. London [B]., 234:77–174.
Utida, S. 1957. Population fluctuation, an experimental and theoretical approach. Sympos. Quant. Biol., 22:139–150.
Van Valen, L. 1960. Further competitive exclusion. Science, 132:1674–1675.
Verhulst, P. F. 1839. Notice sur la loi que la population suit dans son accroissement. Corr. Math. Phys., 10:113–121.
Volterra, V. 1926. Variazioni e fluttuazioni del numero d’individui in specie animali conviventi. Mem. Accad. Lincei, ser. 6, 2:31–113. (English transl., Variations and fluctuations of the number of individuals in animal species living together. In Chapman, R. N., Animal Ecology, 1931. New York, McGraw-Hill Book Company. pp. 409–448.
Volterra, V. 1931. Leçons sur la theorie mathematique de la lutte pour la vie. Paris, Gauthiers-Vallar s.
Weisbrot, D. R. 1966. Genotypic interactions among competing strains and species of Drosophila. Genetics, 53:427–435.
Williams, C. B. 1947. The generic relations of species in small ecological communities. J. Anim. Ecol., 16:11–18.
Author information
Authors and Affiliations
Editor information
Editors and Affiliations
Rights and permissions
Copyright information
© 1970 Meredith Corporation
About this chapter
Cite this chapter
Ayala, F.J. (1970). Competition, Coexistence, and Evolution. In: Hecht, M.K., Steere, W.C. (eds) Essays in Evolution and Genetics in Honor of Theodosius Dobzhansky. Springer, Boston, MA. https://doi.org/10.1007/978-1-4615-9585-4_4
Download citation
DOI: https://doi.org/10.1007/978-1-4615-9585-4_4
Publisher Name: Springer, Boston, MA
Print ISBN: 978-1-4615-9587-8
Online ISBN: 978-1-4615-9585-4
eBook Packages: Springer Book Archive