A user's guide to functional diversity indices

Functional diversity is the diversity of species traits in ecosystems. This concept is increasingly used in ecological research, yet its formal definition and measurements are currently under discussion. As the overall behavior and consistency of functional diversity indices have not been described so far, the novice user risks choosing an inaccurate index or a set of redundant indices to represent functional diversity. In our study we closely examine functional diversity indices to clarify their accuracy, consistency, and independence. Following current theory, we categorize them into functional richness, evenness, or divergence indices. We considered existing indices as well as new indices developed in this study. The new indices aimed at remedying the weaknesses of currently used indices (e.g., by taking into account intraspecific variability). Using virtual data sets, we test (1) whether indices respond to community changes as expected from their category and (2) whether the indices within each category are consistent and independent of indices from other categories. We also test the accuracy of methods proposed for the use of categorical traits. Most classical functional richness indices either failed to describe functional richness or were correlated with functional divergence indices. We therefore recommend using the new functional richness indices that consider intraspecific variability and thus empty space in the functional niche space. In contrast, most functional evenness and divergence indices performed well with respect to all proposed tests. For categorical variables, we do not recommend blending discrete and real-valued traits (except for indices based on distance measures) since functional evenness and divergence have no transposable meaning for discrete traits. Nonetheless, species diversity indices can be applied to categorical traits (using trait levels instead of species) in order to describe functional richness and equitability.