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Knowledge of how diversity changes across spatial scales is important for conservation of biodiversity. Alpha (α), beta (β) and gamma (γ) species richness and Shannon-Wiener diversity index (H') of benthic macroinvertebrates were analysed by additive partitioning across 3 different nested spatial scales (cm, m and km scales) in the estuarine Baltic Sea-North Sea transition area. The data set consisted of abundance of 324 species from a total of 638 samples taken at 64 sites distrib- uted over 6 salinity-defined regions. Results were compared to a null model, randomly assigning individuals of species among samples. β-richness among regions was significantly high and the major contributor to γ-richness, while α- and β-richness were less than expected at finer scales, suggesting that salinity-defined regions largely determined γ-richness. Salinity effects on α-richness were posi- tive and most evident at a regional scale, likely reflecting differential evolutionary adaptation to salinity among species. For H', the greatest contribution to γ-diversity was from α and β at the finest scale, but significantly high contributions at larger scales likely indicated that different species dom- inated abundance in different sites and regions. Effects of rare species on partitioning of total rich- ness was less different from random, compared to common species, for β among regions, suggesting that the occurrence of rare species was more affected by chance than for common species. Results suggest that additive partitioning is a simple and effective tool to unravel changes and sources in diversity over spatial scales in marine and/or estuarine benthic environments. This method may be used to assess effects of habitat homogenisation and as a basis for the design of conservation protocols.

Additive partitioning of estuarine benthic macroinvertebrate diversity across multiple spatial scales