Host specificity is a fundamental component of a parasite's life history. However, accurate assessments of host specificity, and the factors influencing it, can be obscured by parasite cryptic species complexes. We surveyed two congeneric species of intertidal snail intermediate hosts, Zeacumantus subcarinatus and Zeacumantus lutulentus, throughout New Zealand to identify the number of genetically distinct echinostome trematodes infecting them and determine the levels of snail host specificity among echinostomes. Two major echinostome clades were identified: a clade consisting of an unidentified species of the subfamily Himasthlinae and a clade consisting of five species of the genus Acanthoparyphium. All five Acanthoparyphium species were only found in a single snail species, four in Z. subcarinatus and one in Z. lutulentus. In contrast, the Himasthlinae gen. sp. was found in both hosts, but was more prevalent in Z. lutulentus (97 infections) than Z. subcarinatus (10 infections). At least two of the Acanthoparyphium spp. and the Himasthlinae gen. sp. are widespread throughout New Zealand, and can therefore encounter both snail species. Our results suggest that host specificity is determined by host–parasite incompatibilities, not geographic separation, and that it can evolve in different ways in closely related parasite lineages.

The evolution of host specificity and the potential trade-off between being a generalist and a specialist are central issues in the evolutionary ecology of parasites. Different species of parasites or even different populations of the same species often show different degrees of host specificity. However, less is known about intraspecific variation in host specificity within a population. We investigated intraspecific variation by experimentally exposing cercariae from different clones of the trematode Curtuteria australis to two species of second intermediate hosts, the New Zealand cockle Austrovenus stutchburyi and the wedge shell Macomona liliana. We found an overall difference in infection success between the two bivalve species, with A. stutchburyi being the more heavily infected host. However, the cercariae showed a consistent preference for encysting at the tip of the bivalve's foot, regardless of host species. Importantly, there were no significant differences among parasite clones in either relative infection success in the two hosts or preference for the host foot tip. This lack of intraspecific variation may be due to the life-history traits of both parasite and hosts in our system, which may limit opportunities for variation in performance and exploitation strategies in different hosts to evolve within the population.

Evidence for the existence of 2 first intermediate host-specific sibling species of the 45 collar-spined echinostome Echinoparyphium recurvatum is presented. Experimental studies on their life-cycles were carried out under controlled laboratory conditions. The two entities were found to be morphologically indistinguishable in all major respects, yet they exhibited distinct biological characteristics. One of the sibling species utilizes the freshwater lymnaeid pulmonate snail Lymnaea peregra as first intermediate host, and the adults occur in the anterior small intestine of the wildfowl experimental definitive host Anas platyrhynchos. The other utilizes the freshwater mesogastropod prosobranch snail Valvata piscinalis as first intermediate host, and the adults occur in the posterior small intestine and rectum of A. platyrhynchos. The existence of the two sibling species in sympatry may be explained by the fact that the life-cycles of the two forms represent two distinct cycles of transmission serving to reduce excessive competition between them.

Transmission of cercariae of the echinostome digenean Echinoparyphium recurvatum was observed in experimental populations of the second intermediate host snail Lymnaea peregra. Populations of 100 snails were distributed among plastic mesh cages in an infection arena to produce different patterns of host dispersion. Cercarial transmission was investigated in host populations showing three separate patterns of host dispersion; Ideal Regular, Random and Contagious (‘Clumped’). Levels of cercarial transmission in populations showing each type of dispersion pattern were found to be significantly different. The highest rate of cercarial transmission occurred in the contagiously dispersed host population, and the lowest in the randomly dispersed population. Analysis of results from the randomly dispersed population also showed that both the mean percentage prevalence of infection, and also the mean number of cysts recovered per snail, increased significantly with the number of snails per cage. The positive relationship between the mean number of cysts per snail and the number of snails per cage was found to be best described by an exponential function. Chemosensory location of L. peregra snail hosts by E. recurvatum cercariae may provide an explanation for the patterns of cercarial transmission observed.

Infectivity of Pseudechinoparyphium echinatum cercariae to 11 species of gastropod was examined experimentally. Broad specificity and differential host—parasite compatibility were exhibited. Nine gastropod species functioned as second intermediate hosts. Planorbarius corneus, Physa fontinalis, Lymnaea peregra and Biomphalaria alexandrina showed high levels of compatibility with the parasite. In single-species exposures over 90% of cercariae encysted in each of these hosts. Low compatibility with the first intermediate host species Lymnaea stagnalis may be a mechanism preventing super-infection of emitting snails. Cercariae did not infect the prosobranchs Bithynia tentaculata and Viviparus viviparus. Experimental infection of a host community comprised of 8 European gastropod species revealed an order of host utilization similar to that shown in single-species exposures. However, cercarial transmission success in P. fontinalis and L. peregra (compared to that in P. corneus) was significantly reduced. This may have been due to the marked preference to cercariae for P. corneus compared to the other two highly suitable hosts for whom cercariae showed equal preference.

The patterns of association between parasites within a particular host are determined by a number of factors. One of these factors is whether or not infection by one parasite influences the probability of acquiring other parasite species. This study investigates the pattern of association between various parasites of the New Zealand cockle Austrovenus stutchburyi. Hundreds of cockles were collected from one locality within Otago Harbour, New Zealand and examined for trematode metacercariae and other symbionts. Two interspecific associations emerged from the study. First, the presence of the myicolid copepod Pseudomyicola spinosus was positively associated with higher infection intensity by echinostomes. The side-effect of the copepod's activities within the cockle is suggested as the proximate mechanism that facilitates infection by echinostome cercariae, leading to a greater rate of accumulation of metacercariae in cockles harbouring the copepod. Second, a positive association was also found between infection intensity of the metacercariae of foot-encysting echinostomes and that of gymnophallid metacercariae. This supports earlier findings and suggests that the gymnophallid is a hitch-hiker parasite because, in addition to the pattern of positive association, it (a) shares the same transmission route as the echinostomes, and (b) unlike the echinostomes, it is not capable of increasing the host's susceptibility to avian predation. Thus, both active hitch-hiking and incidental facilitation lead to non-random infection patterns in this parasite community.

Survival and infectivity characteristics are described for cercariae of the echinostome Echinoparyphium recurvatum at 10, 15, 20, 25 and 30°C. Cercarial survival was markedly age- and temperature-dependent, maximum survival time being reduced from 68 h at 10°C to 12 h at 30°C, 50% survival times falling from 47·9 h at 10°C to 8·4 h at 30°C. The mean instantaneous per capita death rate of cercariae increased exponentially with temperature. Cercarial infectivity was also temperature dependent, the main effect of temperature being upon the rate at which infectivity diminished with increasing age of cercariae. Infectivity increased from zero at 10°C to a maximum at 25°C after which it declined to a low level at 30°C. Overall transmission efficiency of cercariae at each experimental temperature was calculated as the ratio of the mean instantaneous per capita death rate (μ) to the mean instantaneous per capita rate of infection (β). Transmission efficiency of cercariae was found to be maximal at 20°C and at a relatively high level between 10 and 25°C. Transmission efficiency was zero at 10°C and at a very low level at 30°C. These results suggest that the cercariae of E. recurvatum show transmission optima at water temperatures likely to be encountered in natural habitats in Britain and Europe.