Plant–soil feedbacks depend on drought stress, functional group, and evolutionary relatedness in a semiarid grassland

Plant-soil feedback (PSF) occurs when plants change the biota and physicochemical properties of the soil, and these changes affect future survival or growth of plants. PSF depends on several factors such as plant functional attributes (e.g., life cycle or photosynthetic metabolism) and the environment. PSF often turn positive under dry conditions because soil biota confers drought tolerance. Conspecifics and close relatives share pathogens and consume similar resources, exerting negative PSF on each other. These ideas have mostly been tested under controlled conditions, while field studies remain scarce. To reevaluate these findings in nature, we analyzed plant-soil feedbacks over a drought-stress gradient in a phosphorus-limited semiarid grassland. We planted seedlings of 17 species in plots where community composition had been monitored for six years. To determine PSF intensity, we measured how seedling longevity was affected by previous occupancy of conspecifics and heterospecifics. The previous occupancy-survival relationship (OSR) was used as a proxy for PSF. Evidence for OSRs was found in one-third of the species pairs, with inconclusive evidence for the rest suggesting weak feedbacks. This is in line with the expectation that PSFs in the field are weaker than under controlled conditions. As expected, positive PSFs were more frequent as drought stress increased. The strongest OSRs were caused in dry plots by C4 perennial grasses, which had very positive OSRs on several C3 annual forbs, but negative effects on each other. Well-documented differences between these two functional groups may explain this result: C3 plants are more sensitive to drought, and thus may be favored by tolerance-conferring microbiota; in contrast, water-efficient C4 perennial grasses compete for phosphorus strongly, perhaps driving strong negative PSFs between them. Finally, close relatives had more negative OSRs on each other than on distant relatives as expected, although only in dry plots. This pattern was mostly due to the negative effects of closely related C4 grasses under dry conditions, and their positive effects on distantly related dicots. Our results highlight the importance of plant traits and of the environmental context in determining the direction and strength of PSFs under field conditions.