In gastropods, variation in shell morphology can be caused by the action of several biotic and abiotic factors. While much of this variation is seen in comparisons between different sites or populations, there is also substantial variation in shell morphology among individuals living side by side. We investigate the effect of trematode parasitism on both intra- as well as inter-site variation in the morphology of the New Zealand whelk Cominella glandiformis. We found that both infection by the trematode Curtuteria australis and site of origin had significant effects on several morphometric dimensions of the snail shell, with some interactions between the two factors. On its own, infection by C. australis accounted for 20 to 60% of the variance in shell morphology, depending on the dimension measured. Infected snails also had smoother shells, with less prominent ridges, than their uninfected conspecifics. Other trematode species, infecting whelks at much lower prevalence, also had impacts on shell morphology, but not necessarily in the same direction as C. australis. Overall, parasitism may be an important factor in explaining intra- and inter-site variation in snail phenotype, with potential repercussions for snail populations and their interactions with other community members.

The New Zealand cockle Austrovenus stutchburyi, whose foot is commonly infected by the digenean trematode Curtuteria australis (Echinostomatidae), is often found heavily infected and unable to burrow on the sediment surface of tidal flats. This has been interpreted as a Curtuteria-manipulation with the purpose of increasing the transmission of the parasite to shorebirds acting as final hosts. Using a field-experimental approach the alternative hypothesis was tested, that surface-dwelling cockles, caught on the surface for other reasons than parasites, accumulate larval C. australis at a higher rate than buried cockles. During the 3-month experiment, larval trematodes accumulated with a rate of approximately 0·5 metacercariae/day in both surface and buried cockles. The result strengthens the manipulation hypothesis indirectly by rejecting the alternative hypothesis. The metacercariae were unevenly distributed along the cockle-foot, with about 4 times as many cysts being found in the tip than in either the mid or hind part of the foot. In light of existing knowledge of the burrowing behaviour and apparatus in bivalves, and a negative relationship between foot mobility and infection intensity, it is suggested that C. australis manipulates its host through a mechanical obstruction of foot muscles and the dynamic hydrostatic skeleton, both necessary for successful burrowing.

Different parasite species sharing the same intermediate host species may have similar or conflicting interests, depending on whether they are at the same stage in their life-cycle or whether they share the same definitive host. In the New Zealand cockle, Austrovenus stutchburyi, metacercariae of the digenean Meiogymnophallus sp. are positively associated with metacercariae of Curtuteria australis. This relationship is found in different cockle samples, and is independent of cockle shell size, which suggests that it is not merely the product of metacercariae accumulation over time. Both digenean species have the same definitive host, oystercatchers. Metacercariae of C. australis manipulate the phenotype of cockles, impairing the cockle's ability to burrow in the sediments. This makes the host more susceptible to oystercatcher predation. Thus Meiogymnophallus sp. can benefit by associating with C. australis and may hitch a ride with the manipulator parasite. This is supported by the finding that cockles impaired by C. australis and lying at the sediment surface harbour greater numbers of Meiogymnophallus than buried cockles. A third digenean species, whose sporocysts are found in cockles and which is not transmitted by predation, occurred only in surface cockles. Finally, a parasitic copepod with a direct life-cycle was found evenly distributed among buried and surface cockles, independently of their metacercarial loads. These results show that different parasite species do not use cockles in a random fashion, and that not all patterns of host use are consistent with shared or conflicting interests among parasites.

We investigated the influence of infection by the trematode Curtuteria australis on the burrowing behaviour of its intermediate host, the bivalve Austrovenus stutchburyi. Laboratory experiments and field observations revealed that cockles, unable to bury completely or even partially under the sediment, have a reduced foot length compared with buried individuals. The ability to bury proved to be highly repeatable in field experiments: cockles found at the surface and transplanted to an experimental area did not bury themselves, and cockles found buried stayed buried when relocated. All metacercariae of C. australis were found strictly in the foot and for each of 3 samples collected in different sites, there was a negative and significant relationship between the relative length of the foot and the parasite load. A predation test conducted under natural conditions indicated that cockles with the stunted foot and the altered behaviour are significantly more susceptible to predation by aquatic birds than other cockles. Given that the definitive host of C. australis is an oystercatcher, we first discuss our results in the context of transmission strategy. Comparisons with other studies on more or less related trematode species parasitic in bivalves and evolving under similar constraints for their transmission, shed light on the origin of this adaptation in C. australis.