The role of the invasive bivalve Ensis directus as food source for fish and birds in the Dutch coastal zone.

The razor clam Ensis directus was introduced to Europe presumably as larvae in ballast water around 1978. Starting in the German Bight it spread northward and southward along the continental coastline. Currently it is the most common shellfish species in the Dutch coastal zone, where it mainly occurs in the Voordelta and off the Wadden Sea islands. The mean density of E. directus in the Dutch coastal zone increased from around 2–5 individuals m-2 in the late ‘90’s to around 12–19 individuals m-2 from 2002 onwards. Diet studies show that E. directus makes up a significant proportion in the current diet of plaice, sole, dab, flounder and dragonet and in the diet of eider and common scoter. In recent years E. directus contributed 20–100% of the total wet weight in fish stomachs. The proportion E. directus in the diet increases with fish length. Based on stomach contents of oiled and beached birds and of faeces samples the recent frequency of occurrence is 85–90% in eider and 26% in common scoter. Also waders, gulls and corvids prey on E. directus but the contribution to the diet is still unquantified. Because of its great burying depth the species is not easily accessible. Fish either profit from massive die-offs that regularly occur, or they extract (probably only the smaller) individuals from the sediment. Sea ducks can extract E. directus from the sediment, while shorebirds and gulls feed on dying E. directus washing up on the shore. E. directus is possibly an important food item for fish and seabirds when they occur in high densities and in the right size classes. Since the availability depends greatly on massive die-offs, shell size, burying depth and water depth, it is probably not a very reliable food source. Judging from the role E. directus currently plays for the higher trophic levels, its introduction must have caused a major change in the food relations in its distribution area.