Richness and diversity of helminth communities in tropical freshwater fishes: Empirical evidence

Aim: Published information on the richness and diversity of helminth parasite communities in tropical freshwater fishes is reviewed in response to expectations of species-rich parasite communities in tropical regions. Location: Areas covered include the tropics and some subtropical areas. In addition, the north temperate area of the nearctic zone is included for comparison. Methods: Data from 159 communities in 118 species of tropical freshwater fish, summarized from 46 published studies, were used for this review. Parasite community descriptors used in the analyses included component community richness and calculated mean species richness. Data from 130 communities in 47 species of nearctic north temperate freshwater fish were summarized from 31 studies and used for comparison. Results: The component helminth communities of many tropical freshwater fish are species-poor, and considerable proportions of fish from certain parts of the tropics, e.g. West African drainages, are uninfected or lightly infected. Mean helminth species richness was low and equaled or exceeded 2.0 in only 22 of 114 communities. No single group of helminths was identified as a dominant component of the fauna and species composition was variable among and within broader geographical areas. The richest enteric helminth assemblages were found in mochokid and clariid catfish with a mixed carnivorous diet, whereas algal feeders, herbivores and detritivores generally had species-poor gut helminth communities. Comparisons indicated that certain areas in the north temperate region had higher helminth species richness in fishes than areas in the tropics. Main conclusions: Expectations of high species richness in helminth communities of tropical freshwater fishes are not fulfilled by the data. Direct comparisons of infracommunities and component communities in host species across widely separated phylogenetic and geographical lines are inappropriate. Examination of latitudinal differences in richness of monophyletic parasite groups or of compound communities may uncover patterns different from those found in this study. Richness of helminth communities may be ultimately determined not by the number of host species present but by the degree of divergence of host lineages and by their diversification modes. A phylogenetic framework for hosts and parasites will reveal if increased host species richness within host clades, when host speciation is accompanied by habitat or diet specialization, or both, leads to lower helminth diversity in host species by fragmentation of a core helminth fauna characteristic or specific of the larger host clade. This pattern may be analysed in the context of cospeciation and acquisition from other unrelated hosts (host-sharing or host-switching).