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A 40-y series of consistently collected samples (15 fixed sampling sites, constant sampled area of 15 × 0.95 m2, annual sampling only in late-winter/early-spring seasons, and consistent sieving and sorting procedures; restriction to 50 easily recognizable species) of macrozoobenthos on Balgzand, a tidal flat area in the westernmost part of the Wadden Sea (The Netherlands), revealed significantly increasing trends of species richness. Total numbers of species annually encountered increased from ~28 to ~38. Mean species density (number of species found per sampling site) increased from ~13 to ~18 per 0.95 m2. During the 40 years of the 1970–2009 period of observation, 4 exotic species invaded the area: (in order of first appearance) Ensis directus, Marenzelleria viridis, Crassostrea gigas, and Hemigrapsus takanoi. Another 5 species recently moved to Balgzand from nearby (subtidal) locations. Together, these 9 new species on the tidal flats explained by far most of the increase in total species numbers, but accounted for only one-third of the observed increase in species density (as a consequence of the restricted distribution of most of them). Species density increased particularly by a substantial number of species that showed increasing trends in the numbers of tidal flat sites they occupied. Most of these wider-spreading species were found to suffer from cold winters. During the 40-y period of observation, winter temperatures rose by about 2°C and cold winters became less frequent. The mean number of cold-sensitive species found per site significantly increased by almost 2 per 0.95 m2. Among the other species (not sensitive to low winter temperatures), 6 showed a rising and 2 a declining trend in number of occupied sites, resulting in a net long-term increase in species density amounting to another gain of 1.6 per 0.95 m2. Half of the 50 studied species did not show such long-term trend, nor were invaders. Thus, each of 3 groups (local or alien invaders/winter-sensitive species/other increasing species) contributed to a roughly similar extent to the overall increase in species density.

Increasing species richness of the macrozoobenthic fauna on tidal flats of the Wadden Sea by local range expansion and invasion of exotic species