Breeding, planktonic and settlement factors shape recruitment patterns of one of the last remaining major population of Pinna nobilis within Spanish waters

The pen shell, Pinna nobilis L., is a critically endangered bivalve threatened by mass mortality events throughout the Mediterranean, but the Alfacs Bay (Ebro Delta) still hosts many healthy individuals. Herein, we study the main factors controlling recruitment patterns in this locality, including gonadal development and abundance of critical life-stages, as well as the effect of environmental factors. Growth records from empty shells suggested a single major peak of recruitment during a period of 11 years, although many juveniles were found in two very shallow sand bars possibly acting as a barrier for water circulation and as a trap for larvae. Collectors deployed outside these sand bar areas showed zero settlers, and the availability of planktonic larvae was very low. Gonadal examination evidenced breeding throughout the summer period with successive hermaphroditism, but 20% of individuals were simultaneous hermaphrodites, a condition that has been associated with environmental stress and that could lead to in-breeding depression and potentially reduced fertility. Yet, given the large size of the population and the wide breeding period observed, planktonic processes causing larval mortality such as freshwater discharges from rice locally important rice agriculture are also proposed as possible impacts accounting for patterns of low larval availability.