| Summary: | 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) ...
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