LAUSR

We explored the most influential stand-scaled drivers of ectomycorrhizal, terricolous saprotrophic, and wood-inhabiting (main functional groups) macrofungal species richness in mixed forests by applying regression models. We tested 67 potential explanatory variables representing tree species composition, stand structure, soil and litter conditions, microclimate, landscape structure, and management history. Within the main functional groups, we formed and modeled guilds and used their drivers to more objectively interpret the drivers of the main functional groups. Terricolous saprotrophic fungi were supported by air humidity and litter mass. Ectomycorrhizal fungi were suppressed by high soil nitrogen content and high air temperature. Wood saprotrophs were enhanced by litter pH (deciduous habitats), deadwood cover, and beech proportion. Wood saprotrophic guilds were determined often by drivers with hidden effects on all wood saprotrophs: non-parasites: total deadwood cover; parasites: beech proportion; white rotters: litter pH; brown rotters: air temperature (negatively); endophytes: beech proportion; early ruderals: deciduous stands that were formerly meadows; combative invaders: deciduous tree taxa; heart rotters: coarse woody debris; late stage specialists: deciduous deadwood. Terricolous saprotrophic cord formers positively responded to litter mass. Studying the drivers of guilds simultaneously, beech was a keystone species to maintain fungal diversity in the region, and coniferous stands would be more diverse by introducing deciduous tree species. Guilds were determined by drivers different from each other underlining their different functional roles and segregated substrate preferences. Modeling guilds of fungal species with concordant response to the environment would be powerful to explore and understand the functioning of fungal communities.