Parasite-modified behaviour in non-trophic transmission: trematode parasitism increases the attraction between snail intermediate hosts

Many parasites with complex life cycles cause host behavioural changes that increase the likelihood of transmission to the next host. Parasite modification is often found in trophic transmission, but its influence on non-trophic transmission is unclear. In trematodes, transmission from the first to second intermediate host is non-trophic, suggesting that free-swimming larvae (cercariae) emerging in closer proximity to the next host would have higher transmission success. We performed a series of behavioural experiments with echinostome trematodes and their snail hosts to determine if potential second hosts (ramshorn snail, genus Planorbella Haldeman, 1842) were more attracted to parasitized first hosts (marsh pondsnail, Lymnaea elodes Say, 1821). In a Y maze, a responding snail (Planorbella sp.) was placed in the base and its response to five treatments was assessed: no stimulus, turion duckweed (Lemna turionifera Landolt; a food item), non-parasitized L. elodes, parasitized L. elodes, and finally parasitized versus non-parasitized L. elodes. Snails showed some attraction to uninfected snails, but had a stronger response to infected first host snails. These results indicate that potential second host snails were more attracted to parasitized, heterospecific first host snails over non-parasitized heterospecific snails. This study demonstrates that echinostome trematodes alter snail behaviour by changing navigational choices in uninfected potential hosts through a chemical communication mechanism.The accepted manuscript in pdf format is listed with the files at the bottom of this page. The presentation of the authors' names and (or) special characters in the title of the manuscript may differ slightly between what is listed on this page and what is listed in the pdf file of the accepted manuscript; that in the pdf file of the accepted manuscript is what was submitted by the author