Dynamique des systèmes parasites - hôte, entre trematodes digènes et coque Cerastoderma edule : comparaison de la lagune de Merja Zerga avec le bassin d'Arcachon

La dynamique globale des systèmes parasites-hôte (SPH) ‘trématode digènes - coque Cerastoderma edule’ a été étudiée simultanément à Merja Zerga (Maroc) et dans le Bassin d’Arcachon (France). La dynamique de populations de la coque a montré une variabilité intra et inter-site et a été expliquée essentiellement par la température, la prédation, la compétition et la dynamique sédimentaire. Dans les deux lagunes, le recrutement s’opère à une température de 19°C avec un décalage temporel en relation avec les positions latitudinales des sites (avril-mai à Merja Zerga et juin-juillet à Arcachon). Les taux de croissance restent comparables pour les deux populations de coque (K=1,5 an-1) mais les performances de croissance sont plus élevées à Arcachon (Φ’=3,3) qu’à Merja Zerga (Φ’=3,1). La production était plus élevée à Merja Zerga (36-65 g poids sec.m-2.an-1). P/B est faible sur les deux sites et légèrement plus élevée à Arguin (1,1-1,5 contre 1,0-1,1 an-1). Les deux sites présentent deux riches communautés parasitaires regroupant à Merja Zerga (11 espèces) et à Arguin (13 espèces), respectivement, 69% et 81% de la richesse parasitaire (digènes) connue des coques. Les communautés parasitaires restent comparables et l’éloignement géographique et climatique n’a eu qu’un faible retentissement sur la structure des peuplements parasitaires. La comparaison latitudinale de la structure et de la dynamique des SPH a permis de relativiser l’importance de la température dans la dynamique de l’hôte et des parasites, du moins dans cet intervalle climatique. A Merja Zerga, la température ne semble pas être un facteur limitant de l’infestation contrairement aux sites plus au nord. Mais finalement, les niveaux d’infestation des coques adultes sont similaires. Les échinostomatidés et les gymnophalidés ont été les plus contributives dans la mortalité parasito- dépendante observée. L’impact des parasites digènes en situation normale se traduit par une augmentation de E/B, cependant plus marqué à Merja Zerga. Cette différence d’effet dénote d’un gradient positif de virulence en allant du nord vers le sud. La présence d’herbiers, la situation ‘intertidal versus subtidal’ et la densité des coques ont un effet modéré sur l’infestation parasitaire.The global dynamics of the parasites-host systems (PHS) ‘digenean trematodes – cockle Cerastoderma edule’ was simultaneously studied in Merja Zerga (Morocco) and Arcachon Bay (France). Cockle population dynamics exhibited an intra- and inter-site variability mostly explained by temperature, predation, competition and sedimentary dynamics. In both lagoons, recruitment occurred at 19° C, with a temporal delay due to latitudinal position (April-May at Merja Zerga, and June-July at Arcachon). Growth rates were similar in both cockle populations (K=1.5 an-1) but growth performances were higher at Arcachon (Φ’=3.3) than at Merja Zerga (Φ’=3.1). Production was higher at Merja Zerga (36-65 g dry weight.m-2.yr-1). P/B was low in both sites and slighly higher at Arguin (1.1-1.5 contre 1.0-1.1 yr-1). Both sites exhibited rich parasite communities with 11 species at Merja Zerga and 13 species at Arguin, corresponding to 69 and 81% of the identified cockle parasite species richness (digeneans). Great distances and climate differences did not alter the structure of parasite communities. Structure and dynamics of PHS related to latitude showed the moderate effect of temperature, within this climate range. At Merja Zerga, temperature was not a limiting factor for parasite infestation, in contrast with what happens in northern countries. But eventually, parasite loads in adult cockles were similar in both sites. Echinostomatids and gymnophallids were the most contributive families explaining parasite- dependent mortality. Digenean impact on cockle resulted in an increasing of E/B, with a higher effect at Merja Zerga. This difference could be linked to a higher virulence in southern sites. The presence of seagrass beds, the position between subtidal and intertidal and the cockle density remained secondary factors in controlling parasite infestation

Dynamique des systèmes parasites - hôte, entre trematodes digènes et coque Cerastoderma edule : comparaison de la lagune de Merja Zerga avec le bassin d'Arcachon

La dynamique globale des systèmes parasites-hôte (SPH) ‘trématode digènes - coque Cerastoderma edule’ a été étudiée simultanément à Merja Zerga (Maroc) et dans le Bassin d’Arcachon (France). La dynamique de populations de la coque a montré une variabilité intra et inter-site et a été expliquée essentiellement par la température, la prédation, la compétition et la dynamique sédimentaire. Dans les deux lagunes, le recrutement s’opère à une température de 19°C avec un décalage temporel en relation avec les positions latitudinales des sites (avril-mai à Merja Zerga et juin-juillet à Arcachon). Les taux de croissance restent comparables pour les deux populations de coque (K=1,5 an-1) mais les performances de croissance sont plus élevées à Arcachon (F’=3,3) qu’à Merja Zerga (F’=3,1). La production était plus élevée à Merja Zerga (36-65 g poids sec.m-2.an-1). P/B est faible sur les deux sites et légèrement plus élevée à Arguin (1,1-1,5 contre 1,0-1,1 an-1). Les deux sites présentent deux riches communautés parasitaires regroupant à Merja Zerga (11 espèces) et à Arguin (13 espèces), respectivement, 69% et 81% de la richesse parasitaire (digènes) connue des coques. Les communautés parasitaires restent comparables et l’éloignement géographique et climatique n’a eu qu’un faible retentissement sur la structure des peuplements parasitaires. La comparaison latitudinale de la structure et de la dynamique des SPH a permis de relativiser l’importance de la température dans la dynamique de l’hôte et des parasites, du moins dans cet intervalle climatique. A Merja Zerga, la température ne semble pas être un facteur limitant de l’infestation contrairement aux sites plus au nord. Mais finalement, les niveaux d’infestation des coques adultes sont similaires. Les échinostomatidés et les gymnophalidés ont été les plus contributives dans la mortalité parasito- dépendante observée. L’impact des parasites digènes en situation normale se traduit par une augmentation de E/B, cependant plus marqué à Merja Zerga. Cette différence d’effet dénote d’un gradient positif de virulence en allant du nord vers le sud. La présence d’herbiers, la situation ‘intertidal versus subtidal’ et la densité des coques ont un effet modéré sur l’infestation parasitaire.The global dynamics of the parasites-host systems (PHS) ‘digenean trematodes – cockle Cerastoderma edule’ was simultaneously studied in Merja Zerga (Morocco) and Arcachon Bay (France). Cockle population dynamics exhibited an intra- and inter-site variability mostly explained by temperature, predation, competition and sedimentary dynamics. In both lagoons, recruitment occurred at 19° C, with a temporal delay due to latitudinal position (April-May at Merja Zerga, and June-July at Arcachon). Growth rates were similar in both cockle populations (K=1.5 an-1) but growth performances were higher at Arcachon (F’=3.3) than at Merja Zerga (F’=3.1). Production was higher at Merja Zerga (36-65 g dry weight.m-2.yr-1). P/B was low in both sites and slighly higher at Arguin (1.1-1.5 contre 1.0-1.1 yr-1). Both sites exhibited rich parasite communities with 11 species at Merja Zerga and 13 species at Arguin, corresponding to 69 and 81% of the identified cockle parasite species richness (digeneans). Great distances and climate differences did not alter the structure of parasite communities. Structure and dynamics of PHS related to latitude showed the moderate effect of temperature, within this climate range. At Merja Zerga, temperature was not a limiting factor for parasite infestation, in contrast with what happens in northern countries. But eventually, parasite loads in adult cockles were similar in both sites. Echinostomatids and gymnophallids were the most contributive families explaining parasite- dependent mortality. Digenean impact on cockle resulted in an increasing of E/B, with a higher effect at Merja Zerga. This difference could be linked to a higher virulence in southern sites. The presence of seagrass beds, the position between subtidal and intertidal and the cockle density remained secondary factors in controlling parasite infestation

Population dynamics and secondary production of the cockle Cerastoderma edule: A comparison between Merja Zerga (Moroccan Atlantic Coast) and Arcachon Bay (French Atlantic Coast)

Cockle (Cerastoderma edule) population dynamics were studied at the southern limit of the distribution of this marine bivalve in Merja Zerga, Morocco. Parameters such as growth, mortality, and production were compared with those of a population at Arcachon Bay (France) a site in the center of the cockle's range. At each sampling period between two and three cohorts were simultaneously observed at each site and the average total abundance was usually higher at Merja Zerga. Recruitment occurred at both sites in spring when temperature rose above 19 °C, independently of the month. In Merja Zerga, winter recruitment was also observed at one occasion, following high sediment disturbance. The first year (2005–06) at Merja Zerga, the mortality rate was close to nil for juveniles and was Z = 1.5 yr− 1 for adults, providing a high production (64 g dry weight m− 2 yr− 1). At Arcachon during the same period, the juvenile mortality rate was Z = 10.9 yr− 1, the adult mortality rate was 3.4 yr− 1 and production was 26 gDW m−2 yr− 1. The second year (2006–07), mortality after recruitment was much higher (Z = 8.6 yr− 1, for juveniles) and similar to what was observed at Arcachon (Z = 8.4 yr− 1). Mortality rate of adults was higher at Merja Zerga (Z = 3.0 yr− 1) than at Arcachon (Z = 1.5 yr− 1). Production was lower at Arcachon than at Merja Zerga although growth performances were higher at Arcachon. The higher growth performance at Arcachon (Φ′ = 3.3) was mainly due to high asymptotic length (L∞ = 38 mm) and was related to low intraspecific competition compared to Merja Zerga where cockle abundance was higher (Φ′ = 3.1, L∞ = 31 mm). P/B was low in both sites and slightly higher at Arcachon (1.1–1.5 against 1.0–1.1 yr− 1). At Arcachon, recruitment was correlated with temperature, a peak occurring when temperature rose above 19 °C (June–July). At Merja Zerga, recruitment was already 2–3 months earlier but was not significantly correlated to temperature. This study showed that population dynamics of cockles at the southern limit of this distribution fell in the range of what was observed elsewhere in the North-Eastern Atlantic coast. Most factors that were involved in population regulation (intraspecific competition, predation and sediment dynamics) were not strictly dependent on latitude. The direct role of temperature (latitude dependent factor) was not obvious. Variation in temperature could explain the recruitment delay between Arcachon and Merja Zerga and the low maximum shell length at Merja Zerga

Do trematode parasites affect cockle ( Cerastoderma edule) secondary production and elimination?

Digenean trematodes are omnipresent in the cockle Cerastoderma edule, a common coastal bivalve of (semi-)sheltered north-eastern Atlantic coasts. They can use their host as a second intermediate host where they remain in a relatively latent stage as metacercariae. Cockle population dynamics and trematode parasite load were monitored for two years in two sites, Arcachon (France) and Merja Zerga (Morocco) for the cockle cohort of 2005. Individual growth was slightly higher at Arcachon than at Merja Zerga (Von Bertalanffy parameters: K = 1.5 yr−1 in both sites but L∞ = 31.2 mm at Merja Zerga against 38.3 mm at Arcachon). Production during cockle life was twice as high at Merja Zerga (48.5 compared to 22.2 gDW.m−2 at Arcachon). Elimination compensated production at Merja Zerga while elimination was low at Arcachon (8.1 gDW.m−2) due to sediment dynamics which concentrated cockles within the sampling area. P/B was similar in both sites (2.4–2.6 yr−1) while E/B was higher at Merja Zerga (2.2 yr−1 versus 1.0 yr−1). Trematode communities were similar in both sites in terms of species and abundance. During the cockle cohort lifespan, there were 3 to 5 events in the parasite abundance survey that suggested parasite-dependent mortality. At Merja Zerga, trematodes metacercariae did not affect P/B (2.4 yr−1) but increased E/B (+14%). At Arcachon, they also had no effect on P/B (2.6 yr−1) but on the contrary increased E/B (+20%). The threshold of metacercariae abundance beyond which cockles are affected was lower for Echinostomatid trematodes at Merja Zerga, suggesting an interaction with other factors such as temperature which is higher at the southern limit of cockle distribution

Metazoan parasites in an intermediate host population near its southern border:The common cockle (Cerastoderma edule) and its trematodes in a Moroccan coastal lagoon (Merja Zerga)

International audienceThe metazoan parasite community of Cerastoderma edule was studied in the southern geographical range of the host (the coastal lagoon Merja Zerga, Morocco). A total of 11 metazoan species was found in cockles. Nine of these were trematodes using cockles as either first intermediate host (three species) or second intermediate host (six species). In addition, two other endo-metazoan species (Pinnotheres pisum and Paravortex cardii) were recorded from cockles in the studied lagoon. All the observed metazoans in cockles from Merja Zerga have previously been recorded at sites north of Africa.Up to 10% of the cockles in the studied size-groups were first intermediate hosts to castrating parasites (Gymnophallus choledochus, Labratrema minimus and Monorchis parvus). Among trematodes having metacercariae in cockles (second intermediate host) Meiogymnophallus minutus was the most widespread as it was observed in all cockles from all the examined habitats in the lagoon and it occurred in record high intensities. Different sub-communities of the trematode fauna using cockles as second intermediate host could be identified (subtidal vs intertidal associations).The richness and species composition of the macroparasite community in cockles from Morocco are discussed in relation to patterns seen in cockles from other sites along their geographical range. Migratory fish and waterbirds (final hosts) are generally responsible for the large scale spread (latitudinal spread) of trematodes. However, the distributional patterns of involved intermediate hosts in the life-cycles of the different trematode species in cockles are determining the richness and species composition patterns seen in cockles at shallow water sites along the east Atlantic shoreline

Digenean trematode species in the cockle cerastoderma edule: Identification key and distribution along the North-Eastern Atlantic Shoreline

International audienceWe 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