A long-term study of the recovery of the macrozoobenthos on large defaunated plots on a tidal flat in the Wadden Sea

The process of recovery of the macrozoobenthos community on tidal flats after disturbance by anoxia was studied by monitoring changes in the macrozoobenthos over periods of up to 4.5 year within and around eleven defaunated squares of about 120 m2 each. Regardless of the season, nearly all species that were present in the surrounding area had settled again in the central parts of the plots within 6 months, though their numbers were generally low if only a winter half-year had elapsed. The total numbers of macrobenthic animals reached values similar to those of the surrounding areas as soon as the recovery period had included at least one summer. Numerical recovery thus proceeded much faster if it started in spring than in autumn. As opposed to species richness and total number of animals, biomass recovery needed several years, as did the recovery of the size and age structure of the long-lived species. Most species settled primarily as early postlarvae in summer, a few species (Macoma balthica, Arenicola marina) also in high numbers as juveniles in winter. Moreover, several species showed slow colonisation in all seasons by adults transported by tidal currents or moving across the sediment. Seasonal migration abilities of the constituent species and their life stages appear to be the most important mechanism explaining the course of the recovery of the infaunal community on tidal flats. The extraordinarily successful settlement of larvae of several species within still sparsely populated plots led to (sometimes substantially) higher densities of these species within than outside the plots. Such abundance overshoots were not only frequently observed in short-lived opportunistic species (such as Pygospio elegans and Capitella capitata) but also in the juveniles of several long-lived species (such as Macoma balthica and Mya arenaria). It is concluded that inhibition outside rather than facilitation within the plots was an important cause of these abundance overshoots.