Abstract
Forest understories play a vital role in ecosystem functioning and the provision of ecosystem services. However, the extent to which environmental conditions drive dominant ecological strategies in forest understories at the continental scale remains understudied. Here, we used ~29 500 forest vegetation plots sampled across Europe and classified into 25 forest types to explore the relative role of macroclimate, soil pH and tree canopy cover in driving abundance-weighted patterns in the leaf economic spectrum (LES) and plant size spectrum (PSS) of forest understories (shrub and herb layers). We calculated LES using specific leaf area (SLA) and leaf dry matter content (LDMC) and PSS using plant height and seed mass of vascular plant species found in the understories. We found that forest understories had more conservative leaf economics in areas with more extreme mean annual temperatures (mainly Fennoscandia and the Mediterranean Basin), more extreme soil pH and under more open canopies. Warm and summer-dry regions around the Mediterranean Basin and areas of Atlantic Europe also had taller understories with heavier seeds than continental temperate or boreal areas. Understories of broadleaved deciduous forests, such as Fagus forests on non-acid soils, or ravine forests, more commonly hosted species with acquisitive leaf economics. In contrast, some coniferous forests, such as Pinus, Larix and Picea mire forests, or Pinus sylvestris light taiga and sclerophyllous forests, more commonly hosted species with conservative leaf economics. Our findings highlight the importance of macroclimate and soil factors in driving trait variation of understory communities at the continental scale and the mediator effect of canopy cover on these relationships. We also provide the first maps and analyses of LES and PSS of forest understories across Europe and give evidence that the understories of European forest types are differently positioned along major axes of trait variation.