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lead to a further doubling of the estimates of global parasite richness, suggesting that there could be >300,000 parasitic helminth species using vertebrates as hosts.

HOW MANY PARASITE SPECIES PER HOST SPECIES?

In the best-studied taxa, an average mammalian host species appears to harbor two cestodes, two trematodes, and four nematodes, and an acanthocephalan is found in every fourth mammalian species examined. Each bird species harbors on average three cestodes, two trematodes, three nematodes, and one acanthocephalan (Poulin, 1999; Poulin and Morand, 2000, 2004). None of these estimates take possible unrecognized cryptic species into account, but, in general, helminths that parasitize avian species seem to be less host-specific than those that parasitize mammals. Ultimately, the parasitic fauna of any host species reflects its interaction with the host’s feeding niche, latitudinal range, and social system.

The survey of parasite diversity provided by Poulin and Morand raises many unanswered questions. Do host species from monospecific genera harbor more specialized parasites than do species from more diverse genera or families? What is the status of parasite diversity in the tropics? Nearly all parasite data for nonhuman hosts have been collected from the commonest species of the temperate zone.

Studies of helminth parasites of fishes suggest that latitudinal gradients of diversity are more complex than are those of their hosts. There are many more fish species in the tropics, so we might initially expect there to be more parasite species as well. But, if high host diversity in the tropics leads to low densities of each host species, then some host-specific parasites might be unable to maintain viable populations in their low-density tropical hosts, in which case host-specific parasites and their hosts could exhibit reverse gradients of species diversity. Empirically, the two best studied parasite taxa show opposite trends: tropical fish species have more monogenean parasites per host species than do those in temperate zones (Rohde, 1982, 1999, 2002), whereas tropical fish species have less diverse gut parasites than do their temperate counterparts (Choudary and Dick, 2000; Poulin, 2001). The monogeneans predominantly live on the skin and gills of fish and are either transmitted directly by physical contact between hosts (in the case of the Gyrodactyloidea, the most speciose monogenean group) or via short-lived infectious stages known as oncomiracidia. Thus, monogeneans may be more host-specific, assuming that transmission occurs primarily between individuals living in conspecific social groups. In contrast, the gut parasites may tend to be host-generalists because they characteristically enter a host via predation on infected prey species that may be a component of the diet of many host species. More research is



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