Understanding the importance of biotic interactions in driving the distribution and abundance of species is a central goal of plant ecology. Early vascular plants likely colonized land occupied by biocrusts — photoautotrophic, surface-dwelling soil communities comprised of cyanobacteria, bryophytes, lichens, and fungi — suggesting biotic interactions between biocrusts and plants may have been at play for some 2,000 million years. Today, biocrusts coexist with plants in dryland ecosystems worldwide, and have been shown to both facilitate or inhibit plant species performance depending on ecological context. Yet, the factors that drive the direction and magnitude of these effects remain largely unknown. We conducted a meta-analysis of plant responses to biocrusts using a global dataset encompassing 1,004 studies from six continents. Our meta-analysis revealed there is no simple positive or negative effect of biocrusts on plants. Rather, plant responses differ by biocrust composition and plant species traits and vary across plant ontogeny. Moss-dominated biocrusts facilitated, while lichen-dominated biocrusts inhibited overall plant performance. Plant responses also varied among plant functional groups: C4 grasses received greater benefits from biocrusts compared to C3 grasses, and plants without N-fixing symbionts responded more positively to biocrusts than plants with N-fixing symbionts. Biocrusts decreased germination but facilitated growth of non-native plant species. Our results suggest that interspecific variation in plant responses to biocrusts, contingent on biocrust type, plant traits, and ontogeny can have strong impacts on plant species performance. These findings have important implications for understanding plant community assembly processes and ecosystem responses to global change.

BSC-Plant DatabaseDataset used in multi-factor meta-analyses (i.e., overall plant performance, germination, survival, growth, cover) and univariate meta-regression models in Havrilla et al. (2019). Columns G-K indicate whether or not a particular study (row) was utilized in a particular analysis group. Columns L-U contain effect sizes (log response ratios) and variance estimates (ESTVAR3) for the five analyses. Full citations for each publication can be found in Appendix S2. Columns Z-AG contain the eight candidate explanatory predictor variables. See text for detailed description of each variable (Table 1). Columns AH-AK contain additional plant species information. Columns AL-AP contain geographic and climatic information.BSC_PLANT_Database.xlsx