Developing the multitrait concept for functional diversity: lessons from a system rich in functions but poor in species

Studies focusing on the linkage between numerical and functional trait diversity frequently consider functional diversity indices but rarely evaluate them empirically or evaluate the use of other than continuous traits such as body size. Here, we present an extensive compilation on functional knowledge of benthic macrofauna using the categorical trait approach and scores of both common and rare species for 25 biological traits, including 102 modalities. We empirically quantify functional trait richness, within-trait species richness (redundancy), and trait variability on a large regional scale (.1000 km), in three environmentally different areas (basins of a sea), over a long time-span (10 years). To develop further the usage of multiple categorical traits as an analysis tool, we examine the effect of sampling effort for the understanding of the functional properties of the benthic meta-assemblages. We also evaluate the relationship between species richness and trait richness in order to understand co-variation between trait modalities and how traits are packaged within species. Results show that the biological diversity in terms of traits could be distinguished between areas of higher and lower salinity, higher and lower anthropogenic stress, and higher and lower species richness. A considerably lower number of samples are needed to portray the functional structure of an area in relation to the taxonomic structure, thereby demonstrating the advantage of using traits when considering management and conservation issues. Using categorical traits empirically requires, as shown within this study, an understanding of the relationship between species richness and expression of traits, co- variation of traits at different species richness and composition levels, acknowledgment of differences in trait expressions between common and rare species, and variability in abundance of species. Empirical trait-based analysis can reveal large-scale differences and insights into complexities between assemblage structure and function, and simultaneously be a valid tool for finding generalities.