The understorey of gorgonian forests in mesophotic temperate reefs

1. In the Mediterranean Sea, dense populations of the gorgonian Paramuricea clavata shape marine animal forests, characterizing the seascapes of coralligenous habitats. Despite concerns for its health, with several anthropogenic threats and recent mass mortality events, mainly triggered by thermal anomalies, the understorey of its forests and the ecological processes that they promote are still little known. Here, the abundance and composition of epibenthic assemblages inside and outside P. clavata forests were investigated across the central and western Mediterranean Sea, by applying a multifactorial sampling design. 2. In spite of the large variability in the structures of epibenthic assemblages at local and regional scales, the gorgonian understoreys share some common features, such as higher abundances of calcareous builder organisms and reduced invasion by the non-indigenous alga Caulerpa cylindracea, compared with the adjacent unforested rocky bottoms. Paramuricea clavata showed non-linear density-dependent relationships with algal turfs and non-encrusting algae belonging to the genus Peyssonnelia. Moreover, by entrapping benthic mucilaginous aggregates with their branches, these gorgonians risk topical necrotic lesions, but may reduce the suffocation risks for understorey organisms. 3. Overall, P. clavata forests may enhance bioconstruction processes and increase resistance and resilience of the benthic assemblages in the Mediterranean coralligenous habitats. This species and its forests, together with their understoreys, should be considered as essential elements of the ecology of the Mediterranean Sea, and therefore worthy of specific and effective protection measures. 4. Conservation strategies should reduce the risk of mechanical damage by regulating fishing activities, anchorages, and scuba diving where gorgonian forests are present. Moreover, when evident alterations are documented, restoration actions should be implemented to recover the integrity of gorgonian forests.