Biological crusts as a model system for examining the biodiversity-ecosystem function relationship in soils

Despite that soils may be the greatest repository of biodiversity on Earth, and that most terrestrial ecosystem functions occur in the soil, research on the role of soil biodiversity in ecosystem function has lagged behind corresponding research on aboveground organisms. Soil organisms pose special problems to biodiversity-function research, including the fact that we usually do not know their identity nor what they do in soil ecosystems, cannot easily estimate their biodiversity, and cannot culture the majority of the organisms for use in manipulative experiments. We propose here that biological soil crusts (BSCs) of deserts and many other ecosystems may serve as a useful model system for diversity-function research because the species concept is relatively well-defined within BSC organisms, their functional attributes are relatively well-known, and estimation and manipulation of biodiversity in experiments are feasible, at least within some groups of BSC biota. In spite of these features, there is a pronounced lack of research on biodiversity-function using these organisms. At least two complementary approaches are possible: experiments using artificially-constructed BSCs, and observational studies which statistically control for the effects of other factors which are likely to covary with biodiversity. We applied the latter to four observational datasets collected at multiple spatial scales in Spain and the United States using structural equation models or path analysis using ecosystem function indicators relating to hydrology, trapping and retention of soil resources, and nutrient cycling. We found that, even when total BSC abundance and key environmental gradients are controlled for, direct and approximately linear relationships between species richness and/or evenness and indicators of ecosystem functioning were common. Such relationships appear to vary independently of region or spatial scale, but their strength seems to differ in every dataset. Functional group richness did not seem to adequately capture biodiversity-function relationships, suggesting that bryophyte and lichen components of BSC may exhibit low redundancy. More research employing the multi-trophic, multi-functional, and manipulable BSC system may enable more rapid understanding of the consequences of biodiversity loss in soils, and help enable a biodiversity-function theory that is pertinent to the numerous ecosystem services provided by soil organisms.