Seeds of 52 species of herbaceous plants typical from European grassland ecosystems were obtained from a commercial supplier (Planta naturalis). When species germinated in Petri dishes the seedlings were then transplanted to plastic pots (11 x 11 x 12 cm height, 1L volume). Pots were filled with a mixture of a potting substrate (Biolan Murumuld) and sand. Pots were randomly placed in the greenhouse of the University of Tartu, Estonia. Then, we established monocultures with seven individuals of a single species per pot which were grown under well-watered conditions. One month after transplanting the seedlings to the pots, a drought treatment was applied to half of the pots (five pots per species). The experiment was harvested in late July 2020, when the first individuals started flowering, after month-long drought treatment. Plant traits related to drought responses and resource use strategies were selected and measured for each species following established protocols. These included seven above- and belowground traits: Vegetative plant height (H, cm), Leaf Area (LA, mm2), Specific Leaf Area (SLA, mm2 mg-1), Leaf Dry Matter Content (LDMC, mg g-1), Specific Root Length (SRL, cm g-1), Average root Diameter (AvgD, mm), Root Dry Matter Content (RDMC, mg g-1). Before harvesting, we measured the plant height and collected one leaf per individual for three individuals per pot. Afterward, we collected the aboveground biomass and belowground biomass of all the individuals in each pot. Due to the difficulty in untangling the roots of the different individuals in a pot, root traits were estimated at the pot level. Roots were washed and a sample of finest roots (10-50mg) was collected. Leaves and fine roots were scanned at 300dpi and 600dpi, respectively, using an Epson perfection 3200 Photo scanner for leaves and Epson V700 Photo scanner for fine roots. After scanning, leaves and roots were oven-dried at 60°C for 72h. AvgD and root length were determined using WinRHIZO Pro 2015 (Regent Instruments Inc., Canada), and leaf area with ImageJ software. We averaged all traits values at the species level, attaining a single value for each trait in each treatment. The total aboveground biomass and total belowground biomass of each pot were oven-dried at 60°C for 72h and weighed.

Drought is expected to increase in future climate scenarios. Although responses to drought of individual functional traits are relatively well-known, simultaneous changes across multiple traits in response to water scarcity remain poorly understood despite its importance to understand alternative strategies to resist drought. We grew 52 herbaceous species in monocultures under drought and control treatments and characterized the functional space using seven measured above- and belowground traits: plant height, leaf area, specific leaf area, leaf dry matter content, specific root length, average root diameter, and root dry matter content. Then, we estimated how each species occupied this space and the amount of functional space occupied in both treatments using trait probability density functions. We also estimated intraspecific trait variability (ITV) for each species as the dissimilarity in trait values between the individuals of each treatment. We then mapped drought resistance and ITV in the functional space using generalized additive models. The response of species to drought strongly depended on their traits, with species that invested more in root tissues and conserved small size being both more resistant to drought and having higher ITV. We also observed a significant trend of trait displacement towards less conservative strategies. However, these changes depended strongly on the trait values of species in the control treatment, with species with different traits having opposing responses to drought. These contrasting responses resulted in lower trait variability in the species pool in drought compared to control conditions. Our results suggest strong trait filtering acting on conservative species as well as the existence of an optimal part in the functional space to which species converge under drought. Our results show that changes in species trait-space occupancy are key to understand plant strategies to withstand drought, highlighting the importance of individual variation in response to environmental changes, and suggest that community-wide functional diversity and biomass productivity could decrease in a drier future. Knowing these shifts will help to anticipate changes in ecosystem functioning facing climate change.

The complete dataset is in the file.