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 Ria 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.