Abstract
The scour protection layer (SPL) is a layer of large stones placed around man-made structures in the marine environment, preventing sediment scouring while also providing new hard substrate and potentially increasing the structural complexity of the original environment. This fosters development of diverse benthic communities, supporting high abundance of organisms. Future SPLs are therefore a potential tool for the ecological enhancement of degrading marine habitats following the principles of nature-inclusive design. Yet, factors that shape the benthic communities on SPLs are poorly understood. Here, we analysed existing data from SPLs from offshore wind farms and a gas platform in the southern North Sea to determine how SPL characteristics affect the biofouling community structure. We combined this analysis with an in-situ experiment testing for the effects of habitat complexity on SPL communities. Our results demonstrate that abundant and diverse communities are present on all SPLs. On a regional scale, communities are mainly affected by depth and location. Increasing habitat complexity has significant and positive effects on species richness yet was non-significant for biomass and abundance of the biofouling community. If applied thoughtfully, nature-inclusive design of the SPL habitat, including manipulation of the physical complexity of the structure, can effectively promote biodiversity.