Variability in annual recruitment success as a determinant of long-term and large-scale variation in annual production of intertidal Wadden Sea mussels (Mytilus edulis)

To understand the background of the strong variation and recent decline of stocks and production of mussels ( Mytilus edulis) on tidal flats of the Wadden Sea, we analysed long-term ( twice- annual for 26 years) and multi-station ( 15 sites) estimates of numbers, mean individual weights, biomass, and annual production on Balgzand, a 50- km(2) tidal-flat area in the westernmost part of the Wadden Sea ( The Netherlands). Somatic production was estimated from summed growth increments of soft tissues per half-year period and expressed in ash-free dry mass ( AFDM). In adults, positive values in spring/ summer regularly alternated with negative values in autumn/ winter, when up to similar to 25% ( mean: 14%) of individual weight gains in the preceding season were lost. No weight losses were observed during the Wrst winter of the life of mussels. The 26-year mean of net somatic tissue production P amounted to 5.5 g AFDM m(-2) a(-1) at a mean biomass B of 3.2 g AFDM m(-2); the ratio P/ B varied strongly with age composition of the mussel population and ranged between 0.5 and 3.0 a(-1) ( mean: 1.7). Within the restricted areas of mussel beds, mean biomass and annual production values were two orders of magnitude higher. In the Wadden Sea, mussel beds cover a typical 1% of extensive tidal flat areas. Numerical densities of recruits showed straight- line relationships with subsequent life- time year- class production. Once recruits had reached an age of similar to 10 months, their numbers predicted subsequent production within narrow limits. Production per recruit averaged 0.21 g AFDM for 10-mo recruits and was not related to recruit density. Local variation in annual production varied strongly, with maximal values between mid- tide and low- tide level, where recruitment was also maximal. Production per recruit was higher at low than at high intertidal levels. Frequently failing recruitment is indicated as the main cause of declining mussel stocks in the Wadden Sea. As in other bivalve species, a declining frequency of the occurrence of cold winters appears to govern declining recruitment success and consequently declining production.