Long-term survey discloses a shift in the dynamics pattern of an emerging disease of cockles Cerastoderma edule, marteiliosis, and raises hypotheses to explain it.

Drivers of marine disease outbreaks are poorly understood in spite of their growing impact. We present here results from a unique case study examining how cockles Cerastoderma edule have responded to the introduction of the novel protistan Marteilia cochillia, which led in 2012 to cockle fishery collapse in Galician rias. Based on intensive survey for eight years (2011-2019) of two affected shellfish beds, inner and outer in the Ría de Arousa, involving monthly evaluation of cockle health status and estimation of mortality, detailed information is provided of the declining impact of marteiliosis over a wild cockle population with evidence suggesting its increasing resistance. Disease dynamics involved an annual "breaking wave" of prevalence and subsequent cockle mass mortality, causing the near extinction of every recruited cohort. A shift in this pattern, from a severe epidemic towards an endemic profile, was observed in the inner shellfish bed since the cohort that was recruited in 2016, suggesting the hypothesis of increasing marteiliosis resistance through natural selection. Risk factors that may contribute to trigger marteiliosis outbreaks were analysed. Host age and sex did not influence susceptibility to marteiliosis. No clear relationships between environmental conditions (temperature, salinity and upwelling index) or cockle density and disease dynamics were found. Spatial differences in disease dynamics could be due to differences in the abundance of infective stages hypothetically linked to spatial differences in the population dynamics of a putative planktonic intermediate host. All these findings have potential implications for the management of diseased populations.

Insights into Environmental Drivers on the Reproductive Cycle of Diopatra neapolitana (Polychaeta: Onuphidae).

The reproductive cycle of Diopatra neapolitana was studied over two years in Redondela (NW Spain), considering both the monthly evolution of the percentage of individuals with oocytes and the variation of the mean oocyte size. Moreover, since three other species of Diopatra have recently been documented in regions geographically close to NW Spain, we first confirmed the identity of this species by carrying out a morphological identification of a representative number of the specimens collected. Our data showed a discontinuous reproductive season, with a period of proliferation and maturation of oocytes from January to September and asynchronous spawning events between July and October. Then, a resting period in the months of November and December was observed. We also explored the influence of some relevant environmental factors (namely, water temperature, photoperiod, salinity, primary production and upwelling index) on the observed reproductive cycle. The results suggest that water temperature is one of the most important drivers of the reproductive cycle for this species. Particularly, in Ría de Vigo, the reproductive cycle seems to be influenced by upwelling events of deep cold water that occur frequently between March and September, and that could act as a spawning-inducing switch.

Parasites, pathological conditions and resistance to Marteilia cochillia in lagoon cockle Cerastoderma glaucum from Galicia (NW Spain).

A histopathological survey revealed parasites and pathological conditions affecting lagoon cockles Cerastoderma glaucum along the Galician coast; serious pathological threats were not detected because the potentially pathogenic conditions (infections with a Marteilia-like parasite and bucephalid sporocysts, disseminated neoplasia and a condition involving large foci of heavy haemocytic reaction) were rare, while more prevalent parasites had negligible or limited pathogeny. Considering that C. edule and C. glaucum are sympatric in some Galician rias, it is remarkable that C. glaucum was not seriously affected by Marteilia cochillia while C. edule suffered an intense outbreak of this parasite associated with massive mortality. Comparison of the digestive gland between cockle species showed co-occurrence of digestive tubules in different phases, with abundant disintegrated tubules, in the case of C. glaucum, while C. edule showed synchronicity and absence of fully disintegrated tubules; these differences could influence their susceptibility to M. cochillia because the main location of this parasite in common cockles is the epithelia of the digestive gland. Moreover, the observation of histological sections through the digestive gland easily allows differentiating the 2 cockle species.

Cockle Cerastoderma edule fishery collapse in the Ría de Arousa (Galicia, NW Spain) associated with the protistan parasite Marteilia cochillia.

The highest shellfishery catch in Galicia (NW Spain) has traditionally been cockle Cerastoderma edule. The shellfish bed located in Lombos do Ulla (Ría de Arousa) used to be among those with the highest cockle production; however, cockle mortality rate increased sharply in this bed in April 2012, reaching 100% in May 2012. Salinity and temperature were discounted as potential causes of the mortality. Marteiliosis, which was first detected in February 2012 and reached 100% prevalence in April 2012, was identified as the most probable cause. Marteiliosis had never been detected in Galician cockles, but extensive surveillance of the Galician coast in May to July 2012 detected marteiliosis in most cockle beds of the Ría de Arousa, whereas it was not found in other rías; 2 mo later, the cockle catch in the Ría de Arousa became negligible. Examination of the aetiological agent of marteiliosis with light and transmission electron microscopy supported its assignation to the genus Marteilia; morphological features showed similarity, but not complete identity, with the recently described species M. cochillia Carrasco et al., 2013. Regarding its molecular characterisation, a consensus sequence of 4433 bp containing a partial sequence of the intergenic spacer region, the complete 18S rRNA gene and a partial sequence of the first internal transcribed spacer region was obtained. The obtained sequences were compared with those available for Marteilia spp. and other Paramyxida. Molecular data support that this parasite corresponds to the species M. cochillia, and a PCR assay was designed for its specific diagnosis. The association of huge cockle mortality with M. cochillia infection urges extreme caution to avoid spreading this disease.