Multiple spatial scale analyses provide new clues on patterns and drivers of deep-sea nematode diversity

The deep sea is the largest biome of the biosphere. The knowledge of the spatial variability of deep-sea biodiversity is one of the main challenges of marine ecology and evolutionary biology. The choice of the observational spatial scale is assumed to play a key role for understanding processes structuring the deep-sea benthic communities and one of the most typical features of marine biodiversity distribution is the existence of bathymetric gradients. However, the analysis of biodiversity bathymetric gradients and the associated changes in species composition (beta diversity) typically compared large depth ranges (with intervals of 500 to 1000 or even 2000 m depth among sites). To test whether significant changes in alpha and beta diversity occur also at fine-scale bathymetric gradients (i.e., within few hundred-meter depth intervals) the variability of deep-sea nematode biodiversity and assemblage composition along a bathymetric transect (200–1200m depth) with intervals of 200m among sampling depths, was investigated. A hierarchical sampling strategy for the analysis of nematode species richness, beta diversity, functional (trophic) diversity, and related environmental variables, was used. The results indicate the lack of significant differences in taxonomic and functional diversity across sampling depths, but the presence of high beta diversity at all spatial scales investigated: between cores collected from the same box corer (on average 56%), among deployments at the same depth (58%), and between all sampling depths (62%). Such high beta diversity is influenced by the presence of small-scale patchiness in the deep sea and is also related to the large number of rare or very rare species (typically accounting for 480% of total species richness). Moreover, the number of ubiquitous nematode species across all sampling depths is quite low (ca. 15%). Multiple regression analyses provide evidence that such patterns could be related to the different availability, composition and size spectra of food particles in the sediments. Additionally, though to a lesser extent, our results indicate, that selective predation can influence the nematode trophic composition. These findings suggest that a multiple scale analysis based on a nested sampling design could significantly improve our knowledge of bathymetric patterns of deep- sea biodiversity and its drivers.