Indirect effects of an invasive annual grass on seed fates of two native perennial grass species

Invasive plants exhibit both direct and indirect negative effects on recruitment of natives following invasion. We examined indirect effects of the invader Bromus tectorum (cheatgrass) on seed fates of two native grass species, Elymus elymoides and Pseudoroegneria spicata, by removing B. tectorum and by adding inoculum of the shared seed pathogen Pyrenophora semeniperda in factorial experiments at xeric and mesic field sites. We also included a supplemental watering treatment to increase emergence and also the potential for pathogen escape. We recorded emergence and survival of native seedlings and also determined the fate of unemerged seeds. At the xeric site, Pyrenophora-caused mortality was high (34 %), and effects of other pathogens and failed emergence of germinants were smaller. Cheatgrass removal negatively affected both emergence (35 vs. 25 %) and spring survival (69 vs. 42 %). Pyrenophora-caused seed mortality increased with inoculum augmentation for both species (22 vs. 47 % overall), but emergence was negatively impacted only for P. spicata (20 vs. 34 %). At the mesic site, Pyrenophora-caused mortality was low (6 %). Cheatgrass removal doubled emergence (26 vs. 14 %). Seed mortality increased significantly with inoculum augmentation for P. spicata (12 vs. 5 %) but not E. elymoides, while emergence was not significantly affected in either species. A large fraction of seeds produced germinants that failed to emerge (37 %), while another large fraction (35 %) was killed by other pathogens. We conclude that facilitation by cheatgrass at the xeric site but interference at the mesic site was probably mediated through litter effects that could be ameliorative or suppressive. Apparent competition between cheatgrass and native grasses could occur through Pyrenophora, especially in a xeric environment, but effects were weak or absent at emergence. This was probably because Pyrenophora attacks the same slow-germinating fraction that is subject to pre-emergence mortality from other causes, including attack by other pathogens such as Fusarium.