Legume effects in a native community invaded by alien Asteraceae in a multi-species comparison

Facilitation has been a long-neglected type of interaction but received more attention recently. Legumes are commonly involved in facilitative interactions due to their nitrogen fixation. Facilitative interactions are so far underappreciated yet potentially important for biological invasions, especially given increasing numbers of alien species. In a common garden experiment using 30 annual Asteraceae species (neophytes, archaeophytes, plus some natives), grown in communities with or without legume presence, we measured functional traits and fitness in focal Asteraceae, as well as nitrogen characteristics of Asteraceae and two native community phytometer species. We investigated how legume presence affects relationships between trait and nitrogen concentration and Asteraceae fitness; and whether mechanisms of facilitation in legume presence and its effects on aboveground performance differ among native phytometer, neophyte, and archaeophyte Asteraceae using the delta N-15 natural abundance method. Lower specific leaf area was associated with higher aboveground biomass and seed production, with a stronger effect in legume absence. Nitrogen concentration had a positive relationship with biomass, but did not generally increase seed production. Our results hint at N facilitation for the native grass phytometer Festuca rupicola when growing in legume presence, whereas the forb Potentilla argentea and 27 alien Asteraceae species did not indicate facilitative effects. Intriguingly, direct legume facilitation in native phytometer species was only detected when growing with archaeophytes neighbors, not with neophytes. This hints at varied mechanisms of competition for nitrogen between natives and alien species of different residence time and deepens the understanding of altered facilitative leguminous effects in alien species presence.