We describe the digenean fauna of one of the dominant intertidal hosts, the common cockle Cerastoderma edule, in terms of biomass, off north-eastern Atlantic shores. Using published and unpublished literature we have prepared an identification key and provide an up-date of the large-scale distributional patterns of digenean species of the common cockle. At least sixteen digenean species, belonging to seven families, use cockles as intermediate host. Among these species two utilize cockles as first intermediate host only, whereas two species utilize cockles as both first and second intermediate host. The remaining eleven species have cockles as their second intermediate host. Water birds and fish are the definitive hosts to twelve and four species, respectively.Cockles are infected with digeneans along the latitudinal gradient from southern Morocco to the western region of the Barents Sea often with high infection levels. Whereas some of these digenean species occur along most of the latitudinal gradient others show a more restricted northern or southern distribution mostly caused by an underlying latitudinal gradient of host species.Knowledge of digenean species and their large-scale distribution pattern may serve as a baseline for future studies dealing with the effects of climate change on parasite-host systems. For such studies the cockle and its digenean community could be an ideal model system.

Cercaria cerastodermae I, a digenean parasite of Cerastoderma edule, was recorded for the first time in the Atlantic Ocean off the Iberian peninsula. Sporocysts were present in the hemolymph of the digestive gland, gonad, gills, and foot of the mollusc. Most of the cercariae present within sporocysts were encysted as metacercariae. The corresponding adult stages were obtained after experimental infection of several Diplodus sargus artificially reared in fish farms and that had previously been protected against natural infections. Numerous adult specimens of Monorchis parvus were collected in Diplodus annularis along the French Mediterranean coast. Comparison of wild and experimental adults allowed the adult stage of Cercaria cerastodermae I to be identified as M. parvus. Another monorchid, Monorchis monorchis, a parasite of Spondyliosoma cantharus, was found in the same Mediterranean area and compared with M. parvus. Additionally, ITS1 nuclear ribosomal DNA sequences of C. cerastodermae I and of the adults collected in naturally infected D. annularis and S. cantharus were obtained. Sequence data indicate that C. cerastodermae I corresponds to the adult of M. parvus found in D. annularis and is clearly distinct from M. monorchis found in S. cantharus.

This work reviews the significant community of digenean trematodes observed infecting the edible cockle, Cerastoderma edule, and the snail Nassarius reticulatus in the channel of S. Jacinto, Ria de Aveiro Estuary, Portugal. The Digenea observed comprise 15 species, belonging to the families Bucephallidae (1), Echinostomatidae (4), Gymnophallidae (3), Lepocreadiidae (1), Microphallidae (1), Monorchiidae (1), Renicolidae (1), Strigeidae, (1), and Zoogonidae (2). The species were studied by light microscopy and scanning electron microscopy. Information is included for hosts, sites of infection, prevalence, and a brief description of parasite surface morphology. The significance of these observations is compared with the presence of these digeneans from other regions, mainly the Atlantic Ocean and Mediterranean Sea. The Ria de Aveiro is a geographical location conducive to extensive species co-occurrence resulting from the overlap between southern (Mediterranean) and northern definitive hosts of the parasites. Differences in life cycle strategies were observed and discussed.

Diphtherostomum brusinae was first recorded by the present study in the north of Portugal. Sporocysts, containing cercariae and encysted metacercariae, were observed in the gonads and digestive gland of the gastropod Nassarius reticulatus. Metacercariae were also found infecting the foot, mantle border and gills of the cockle Cerastoderma edule. The adult form was lodged in the rectum of the definitive host Diplodus sargus. The morphology of the three parasitic stages was studied by light (LM) and scanning electron microscopy (SEM). Despite the close similarity between cercaria and metacercaria, SEM data provided information that allowed their differentiation, namely the presence of a dense crown of microvilli around the oral cavity of the cercariae, which was absent in the metacercariae. In addition, the metacercariae presented a specific pre-acetabular rectangular band with conspicuous triangular spines. The adult showed characteristics of D. brusinae species, in particular the presence of acetabular lips, compact vitellaria and large elliptical eggs. Sequenced ITS1 data clearly demonstrated that the cercariae and metacercarial cysts from N. reticulatus, the cysts from C. edule and the adult isolated from D. sargus were life cycle stages that belonged to the same species, i.e. D. brusinae. Two transmission strategies in the life cycle of this species were observed: (1) cercariae encyst within the sporocysts of N. reticulatus and await ingestion by the definitive host; and (2) N. reticulatus naturally emits cercariae; they encyst in C. edule or the environment and are ingested by the definitive host.

The morphology and ecology are described for four sibling species of digenean metacercariae: Meiogymnophallus minutus (Cobbold 1859), M. strigatus (Lebour 1908) from the estuarine cockle Cerastoderma edule L. and M. fossarum (Bartoli, 1965) and M. rebecqui (Bartoli, 1983) comb. nov. from the brackish water lagoon cockle C. glaucum (Poiret). Although occupying different sites within the two species of cockle, both M. minutus and M. fossarum infections result in pathological effects that may cause heavy host mortality. Remarkably, they also result in an inversion of the cockles in situ in the sand such that predation by the final host species is enhanced.