Controls of biological soil crust cover and composition shift with succession in sagebrush shrub-steppe

Successional stage may determine strength and causal direction of interactions among abiotic and biotic factors; e.g., species that facilitate the establishment of other species may later compete with them. We evaluated multivariate hypotheses about abiotic and biotic factors shaping biological soil crusts (BSCs) in early and late successional stages. We surveyed vegetation and BSC in the shrub-steppe ecosystem of the Columbia Basin. We analyzed the relationships with bryophyte and lichen covers using structural equation models, and analyzed the relationships with BSC composition using Indicator Species Analysis and distance-based linear models. Cover, indicator species, and composition varied with successional stage. Increasing elevation and bryophyte cover had higher lichen cover early in succession; these relationships were negative in the later successional stage. Lichen cover did not appear to impede B. tectorum cover, but B. tectorum appeared to strongly negatively affect lichen cover in both stages. Biological soil crust composition varied with bunchgrass cover in the early successional stage, but with elevation and B. tectorum cover later in succession. Our findings support the hypotheses that as succession progresses, the strength and direction of certain community interactions shift, and B. tectorum leads to reductions in biological soil crust cover regardless of successional stage.