SUMMARYRecently, some authors (Kennedy, 1981; Price & Clancy, 1983) have argued that there are fundamental differences between the communities of helminths in fish and bird hosts. Such differences are foreshadowed by the work of Dogiel (1964) and are apparent from survey data (e.g. Threlfall, 1967; Bakke, 1972; Hair & Holmes, 1975 on birds, and compare Chubb, 1963; Mishra & Chubb, 1969; Wootten, 1973; Ingham & Dronen, 1980 on fish). Questions still remain, however, as to whether the distinctions are truly justified and whether the differences are really fundamental. In this paper, we address these questions by examining helminth diversity in a series of hosts. We then discuss and provide explanations for the observed differences.

SummaryWe examine patterns of community richness among intestinal parasitic helminth communities in fishes, herptiles, birds and mammals with respect to the comparative number of component species in a host population. We show that terrestrial hosts have, on average, fewer component species than aquatic hosts. We also show that the mean number of component species in aquatic hosts increases from fishes through herptiles to birds before declining slightly in mammals. For terrestrial hosts, the mean number of component species increases from herptiles, through birds, reaching a maximum in mammals. We conclude that: (i) habitat of the host is more important in determining community richness than is host phylogeny; (ii) the phenomenon of 'host capture' may be largely responsible for increased species richness in some host groups; (iii) aquatic birds harbour the richest intestinal helminth communities; and (iv) as we interpret them, our data refute the time hypothesis, which would predict that fishes as the oldest lifestyle should have the richest helminth communities.

In late 1987, immature Anguillicola crossus were reported for the first time in Britain from eels from two river systems. By late 1988, gravid adults were present in a number of rivers in the east of England in two discrete centres ofdistribution: one in East Anglia correlated with the route taken by lorries exporting eels to the continent, and one in the R. Thames correlated with the import of eels to London. The parasite was firmly established in the R. Trent, where prevalence levels reached 100% in some places.Laboratory investigations showed that adult parasites and their eggs remained viable even after infected eels had been maintained for 4 weeks in 100% sea water. Hatching of eggs declined with increasing salinity, but was not totally inhibited by sea water. Survival and infectivity of freeliving second stage larvae were maximal in natural fresh water (95% survival for 4 months, and 50% still infective to copepod intermediate hosts after 70 days), but declined in alkaline water and with increased salinity. Nevertheless, in 100% sea water, 50% of larvae were still infective after 8 days. Specificity to the intermediate host was low, and eels of all sizes could be infected.These characteristics, plus a high reproductive potential, give the parasite exceptional colonization potential and ability, enabling it to survive natural movements of eels from catchment to catchment and toincreaserapidlywithinanewlocality. Theabilityoffree-livinglarvae toadhere to the substratum and survive in sea water enables them to survive in eel-transport lorries from which they will not readily be removed by flushing, the normal cleansing procedure. It is concluded that there were two separate introductions of the parasite to Britain; via the eel import trade through London, and, totally unexpectedly, via the eel export trade in lorries traversing East Anglia. The parasite is now firm1 y established in Britain and willcontinue to spread by natural movementsofeels but especially by human-assisted movements of infected eels for stocking and market. This latter practice is recognized as a major factor in introducing and disseminating fish parasites.

Examples of the apparently stochastic nature of freshwater fish helminth communities illustrating the erratic and unpredictable occurrence and distribution of many species are provided for six species of fish from several localities throughout Britain. By focussing on parasite colonization strategies two categories of helminths are recognized: autogenic species which mature in fish and allogenic species which mature in vertebrates other than fish and have a greater colonization potential and ability. Three groups of fish are distinguished: salmonids, in which helminth communities are generally dominated by autogenic species which are also responsible for most of the similarity within and between localities; cyprinids, in which they are dominated by allogenic species which are also responsible for most of the similarity within and between localities; and anguillids, whose helminth communities exhibit intermediate features with neither category consistently dominating nor providing a clear pattern of similarity. Recognition and appreciation of the different colonization strategies of autogenic and allogenic helminths in respect of host vagility and ability to cross land or sea barriers and break down habitat isolation, and their period of residence in a locality, whether transient or permanent, provides an understanding of, and explanation for, the observed patchy spatial distribution of many helminths. Comparison with other parts of the world indicates that colonization is a major determinant of helminth community structure.

No abstract

One hundred and sixty eight species of parasite are listed from 36 species of freshwater fish from Britain and Ireland, and notes on their distribution are provided. A list of hosts together with the parasites recorded from them is also given. Although almost twice as many species are listed as in the most recent published reference list, the parasite fauna of Britain appears to be considerably poorer in number of species than that of mainland Europe, and that of Ireland poorer still. The distribution of most species, apart from two endemic species and a very few which have clearly been introduced into Britain in recent years from America or the mainland of Europe, appears to show no peculiarities and is probably contiguous with that of their hosts.

SummaryThe prediction that species richness and diversity of intestinal helminth communities in eels would change over time in response to habitat changes was tested over a period of 13 years in a small stream subjected to extensive human management. Nearly all measures of helminth community structure adopted indicated a decline in richness and diversity over the first 6 years followed by a recovery over the last few years to levels unexpectedly close to those at the start of the investigation. Changes in total number of species suggested that the component community was far richer (from 3–9 species) at the end of the period. By contrast, changes in diversity and dominance measures revealed less variation than expected and suggested that there was an underlying stability of community structure characterized by high dominance by a single species, although the identity of this changed, low diversity and a large proportion of the eel population harbouring 0 or only 1 species. A similar pattern of changes was recorded in the infra-communities, where values of species richness and diversity were very similar at the commencement and termination of the study. It appeared that those helminths that colonized in the recovery period contributed to community richness but had little impact on community structure. The helminth communities clearly did change in response to habitat changes, and the evidence for a fixed number of niches and an underlying constancy in helminth community structure in eels is evaluated.

Problems of pattern and scale are considered in relation to helminth communities of freshwater fish by examining them at different hierarchical taxonomic and spatial scales, with a view to seeking generalizations of heuristic value, assessing the importance of phylogenetic and ecological determinants of community structure and improving understanding of unpredictable communities. Initially, communities were analysed at the level of salmonid genera, focusing on Oncorhynchus, in its heartland in Canada: then in O. mykiss throughout its global range and finally in individual localities to which it has been introduced in Britain. In the heartland, communities are dominated by salmonid specialist helminths, forming a phylogenetic element: the minority ecological element comprises broad generalists and non-salmonid specialists. Most species except generic specialists are shared between host genera. As the distance to which O. mykiss was translocated from its heartland increases, so generic specialists disappear first and then salmonid specialists decline. The community is thus increasingly composed of generalists and it also becomes increasingly poor. Helminths may be acquired from native salmonids and/or unrelated hosts, depending on availability. This same pattern is paralleled in individual localities in a restricted region: the phylogenetic element reflects the native salmonid species present and the ecological element the presence of other genera of fish; i.e., a supply-side situation. The change of scale in analysis has thus enabled the recognition of generalizations and patterns of heuristic value and improved the understanding of unpredictable communities by interpreting local variation as ecological 'noise' that often obscures fundamental patterns. In this and other taxa of fish, phylogenetic elements dominate helminth communities in the heartlands, but ecological elements dominate as the host increasingly becomes a stranger in a strange land.