Biological determinants of species diversity

We consider four categories of biological mechanisms of deter- minants which cause and maintain species diversity: niche relations, habitat diver- sity, mass effects and ecological equivalency. Two of these determinants are origi- nal to this paper: mass effect, the establishment of species in sites where they cannot be self-maintaining; and ecological equivalency, the coexistence of species with effectively identical niche and habitat requirements. The mode of action and ecological implications of each biological determinant are discussed using a schematic method for measuring alpha (community), beta (differentiation), and gamma (regional) diversities. The importance of mass effects and ecological equivalency to species richness is documented with several types of field data from Israel and California, U.S.A. Floristic richness and, in particular, the richness of floristic transitions, are discussed and interpreted by use of the biological determinants of diversity. Contact transitions between distinct floras are rich predominantly because of mass effects. Transitions induced by marked environmental changes are rich because of the combined influences of habitat diversity and mass effects. The rate at which species richness increases with sample area is related to the combined effects of all four biological determinants. This complexity explains the failures of simple species-area models. The relative intensity of each determinant is related to area: niche relations are most important at within-community scales, habitat diversity most important at both within-community and land- scape scales, and ecological equivalency most important at regional scales. We suggest that understanding patterns of species diversity will be enhanced by the partitioning of total species richness into the richness caused by each of the four ecologically distinct determinants of diversity.

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