Epibiosis and parasitism of coquina clam Donax spp.: Location, location, location!

Cryptic species of coquina clams Donax fossor and D. variabilis carry the hydroid epibiont Lovenella gracilis and are infected with metacercariae of the monorchiid parasites Lasiotocus trachinoti and L. choanura. The associations among this host–epibiont–parasite system were investigated. Fifty clams were collected at low tide over 3 days in June 2020 in South Carolina from each of three groups: clams with no hydroid from the upper intertidal zone and clams with and without hydroids from the swash zone. Clams were measured, identified using a newly developed PCR-RFLP, and examined for infection by metacercariae. Parasites were identified based on cercarial morphology and on metacercariae habitat in the clams. D. fossor was most often found in the swash zone and D. variabilis in the upper intertidal zone. The hydroid was most often associated with D. fossor, which was more infected by both digeneans than D. variabilis. Mean abundance of metacercariae of L. choanura was higher than that of L. trachinoti in both clams and increased over time for both parasites, because higher infection was correlated with larger clams. Greater time spent in the water by individuals of D. fossor appears to best explain these results, with the presence of the hydroids also being associated with higher infection by metacercariae in this coquina. Integration of D. variabilis in both digenean life cycles appears to lead to a positive outcome for the parasites as prevalence and abundance of infection were high; however, because D. variabilis is most frequent in the upper intertidal, more emersed, zone, it is likely deleterious to the epibiont to establish on this clam.