Niche restriction in parasites: proximate and ultimate causes.

Hutchinson's (1957) definition of an ecological niche as a multidimensional hypervolume determined by a number of physical and biotic variables is adopted. The number of niche dimensions is very great, but as a working hypothesis it is assumed that a few are sufficient to characterize the niche of a parasite species to a high degree of accuracy. They are host species, microhabitat(s), macrohabitat(s), geographical range, sex and age of host, season, food and hyperparasites. Methods to measure niche width, in particular specificity indices, are discussed, and some examples of niche restriction are described. Proximate and ultimate causes of niche restriction are discussed, mainly using marine parasites as examples. Among proximate causes of one niche dimension, host specificity, are ecological factors restricting exposure to infection to certain host species; host-specific chemical factors that induce hatching, direct infective stages to a host and bring about settlement of a parasite; factors that lead to mortality in or on the wrong host; morphological adaptations that guarantee survival in or on the 'correct' host; and availability of suitable hosts. Many factors are likely to be responsible for microhabitat specificity, but have been little studied, except for some physiological and morphological adaptations to particular microhabitats. Macrohabitats and geographical range may be determined by the distribution of intermediate hosts and certain food items, and by a variety of chemical and physical factors. Hosts of different sexes may differ in feeding habits and the composition of the skin, and thus acquire parasites differentially. Hosts of different age may be differentially infected due to accumulation of parasites with age, loss of parasites due to developing resistance (or immunity), and different size and feeding habits. Among ultimate causes of niche restriction and segregation are avoidance of competition, predation and hyperparasites; facilitation of mating; reinforcement of reproductive barriers; and adaptations to environmental complexity. Few studies permit a decision on which factor or factors are responsible in particular cases. Interspecific competition may play a greater role in helminth communities of some host groups than of others, but it seems that, overall, its role has been exaggerated at least for marine parasites. Some 'classical' examples of microhabitat segregation explained by interspecific competition can also be explained by reinforcement of reproductive barriers. There is evidence for the importance of facilitation of mating in microhabitat restriction, and the availability of many vacant niches indicates that competition, overall, is not of great importance.