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Coexistence of Competitors

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image of Netherlands Journal of Zoology
For more content, see Archives Néerlandaises de Zoologie (Vol 1-17) and Animal Biology (Vol 53 and onwards).

Some critical remarks have been made concerning Darwin's statement that "two animal species with similar needs and habits cannot inhabit the same habitat". Until now this "exclusion principle" has been studied quantitatively using the model of logistic growth. In this paper the principle is tested by some variants of the Lotka-Volterra model of interaction between predator and prey and some analyses and simulations are carried out. The shrews Sorex araneus L. and S. minutus L. were chosen as examples of predators with earthworms and spiders as examples of prey species. The impact of territoriality, density dependent predation and handling time on the coexistence both of competitors and predator sharing prey species is studied. The model is continuous, implies a constant, homogeneous environment, and takes no account of delayed reactions. If the densities of two predator species are regulated by one common prey species only, one of the competitors will exclude the other. This also applies when two prey species share one predator species. The latter type of exclusion has been called "klompetition". In accordance with Levin's formulation that "no stable equilibrium is possible if some r species are limited by less than r factors", it appears that in a system with two common predators and two common prey species there is no trend towards extermination in any of the species. The introduction of intraspecific territoriality of the predators stabilizes the system in such a way that permanent coexistence of competitors is possible. A sensitivity study for all parameters of the model is carried out. The introduction of density dependent predation counteracts "klompetition" in such a way that it enables two prey species which share one predator to coexist permanently. The introduction of handling time has only a destabilizing effect on the system and does not affect coexistence or exclusion. Summaring we can formulate the following "exclusion" or "coexistence" rule: Permantent coexistence of competitors is possible. A sufficient condition for stability in an ecosystem in a constant, homogeneous environment is that for each arbitrarily chosen pair of species there must be two different combinations of density limiting factors, provided that each species itself can be conceived as such a factor.

Affiliations: 1: (Department of Ecology, Zoological Laboratory, University of Leiden, The Netherlands


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