Hidden structural heterogeneity enhances marine hotspots' biodiversity

Studies in terrestrial and shallow-water ecosystems have unravelled the key role of interspecific interactions in enhancing biodiversity, but important knowledge gaps persist for the deep sea. Cold-water coral reefs are hotspots of biodiversity, but the role of interspecific interactions and "habitat cascades" (i.e. positive effects on focal organisms mediated by biogenic habitat formation) in shaping their biodiversity is unknown. Associations between macrofaunal hosts and epifauna were examined in 47 stations at the Mingulay Reef Complex (northeast Atlantic). In total, 101 (group level) and 340 (species level) unique types of facultative associations formed by 43 hosts and 39 epifaunal species were found. Molluscs and empty polychaete tubes had higher values for the type and number of host-epifaunal associations, the Shannon-Wiener (H) and Margalef (d) indices of the epifauna than the rest of the taxonomic groups (p < 0.05). Hosts' body size, orientation, surface smoothness, and growth form explained a significant amount of variability (32.96%) in epifauna community composition. Epifaunal species richness (S), H and d were 27.4 (+/- 2.2%), 56.2 (+/- 2.8%) and 39.9 (+/- 2.3%) of the respective values for the total sessile communities living on coral framework. This is intriguing as coral framework is orders of magnitude larger than the size of macrofaunal hosts. It is suggested that bivalves, tunicates and empty polychaete tubes increase habitat heterogeneity and enhance biodiversity through "habitat cascades", in a similar way that epiphytes do in tropical rainforests. Most macrofaunal habitat suppliers in the studied cold-water coral reef are calcified species and likely susceptible to ocean acidification. This indicates that the impacts of climate change on the total biodiversity, structure and health of cold-water coral reefs may potentially be more severe than previously thought.