A barrier island perspective on species-area relationships

Predictions of species richness by island area are a classical cornerstone in ecology, while the specific features of barrier islands have been little appreciated. Many shorelines are occupied by barrier islands, which are shaped by offshore sedimentation processes and annual storm tide events. Hence, the appearance of these islands may vary between years if they are not protected by dykes. Here, we analyzed more than 2,990 species across 36 taxonomic groups (including vertebrates, invertebrates, and land plants) on German barrier islands, the East Frisian Islands. We tested for relationships between species richness or species incidence and island area (SAR), island habitat diversity and further island parameters using a range of generalized linear and mixed-effects models. Overall species richness was explained best by habitat diversity (Shannon index of habitat types). Analyses on the occurrence probability of individual species showed that changes of barrier island area by sedimentation and erosion, that is, barrier island-specific dynamics, explained the occurrence of 17 of 34 taxa, including most beetles, plants, and birds. Only six taxa such as spiders (249 species) and mammals (27 species) were primarily related to area. The diversity of habitat types was a key predictor for the incidence of twenty-five taxa, including ground beetles, true bugs and grasshoppers, amphibians, and reptiles. Overall, richness and incidence of taxa differed greatly in their responses, with area (although varying from 0.1 to 38.9 km²) playing a minor and island heterogeneity a major role, while barrier island-specific sedimentation and erosion turned out to additionally explain species richness and occurrence.