Positive interactions enhance biodiversity and ecosystem function, but can also exacerbate biological invasions. Facilitation of exotic invaders by exotic foundation species (invasional meltdown) has been studied extensively, but facilitation of exotic invaders by native foundation species has attracted less attention. Specifically, very few studies have examined the extent that native foundation species facilitate native and exotic competitors. Understanding the processes that mediate interactions between native and exotic species can help explain, predict, and improve management of biological invasions. Here, we examined the effects of native foundation shrubs on the relative abundance of the annual plant community – including native and exotic taxa – from 2015–2018 in a desert ecosystem at Carrizo Plain National Monument, California, USA (elevation: 723 m). Shrub effects varied by year and by the identity of annual species, but shrubs consistently enhanced the abundance of the annual plant community and facilitated both native (n=17 species) and exotic (n=4 species) taxa. However, at the provenance level, exotic annuals were facilitated 2.75 times stronger in abundance than native annuals, and exotic annuals were always more abundant than natives both near and away from shrubs. Our study reaffirms facilitation as an important process in the organisation of plant communities and confirms that both native and exotic species can form positive associations with native foundation species. However, facilitation by native foundation species can exacerbate biological invasions by increasing the local abundance of exotic invaders. Thus, the force of facilitation can have a dark side relevant to ecosystem function and management.

Invasive plants have tremendous potential to enrich native food webs by subsidizing net primary productivity. Here, we explored how a potential food subsidy, seeds produced by the aggressive invader cheatgrass (Bromus tectorum), is utilized by an important guild of native consumers – granivorous small mammals – in the Great Basin Desert, USA. In a series of field experiments we examined 1) how cheatgrass invasion affects the density and biomass of seed rain at the ecosystem-level; 2) how seed resources from cheatgrass numerically affect granivorous small mammals; and 3) how the food preferences of native granivores might mediate the trophic integration of cheatgrass seeds. Relative to native productivity, cheatgrass invasion increased the density and biomass of seed rain by over 2000% (P < 0.01) and 3500% (P < 0.01), respectively. However, granivorous small mammals in native communities showed no positive response in abundance, richness, or diversity to experimental additions of cheatgrass seeds over one year. This lack of response correlated with a distinct preference for seeds from native grasses over seeds from cheatgrass. Our experiments demonstrate that increased primary productivity associated with exotic plant invasions may not necessarily subsidize consumers at higher trophic levels. In this context, cheatgrass invasion could disrupt native food webs by providing less-preferred resources that fail to enrich higher trophic levels.

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The enemy release hypothesis (ERH) of plant invasion asserts that natural enemies limit populations of invasive plants more strongly in native ranges than in non‐native ranges. Despite considerable empirical attention, few studies have directly tested this idea, especially with respect to generalist herbivores. This knowledge gap is important because escaping the effects of generalists is a critical aspect of the ERH that may help explain successful plant invasions. Here, we used consumer exclosures and seed addition experiments to contrast the effects of granivorous rodents (an important guild of generalists) on the establishment of cheatgrass ( Bromus tectorum ) in western Asia, where cheatgrass is native, versus the Great Basin Desert, USA, where cheatgrass is exotic and highly invasive. Consistent with the ERH, rodent foraging reduced cheatgrass establishment by nearly 60% in western Asia but had no effect in the Great Basin. This main result corresponded with a region‐specific foraging pattern: rodents in the Great Basin but not western Asia generally avoided seeds from cheatgrass relative to seeds from native competitors. Our results suggest that enemy release from the effects of an important guild of generalists may contribute to the explosive success of cheatgrass in the Great Basin. These findings corroborate classic theory on enemy release and expand our understanding of how generalists can influence the trajectory of exotic plant invasions.

An important hypothesis for how plants respond to introduction to new ranges is the evolution of increased competitive ability (EICA). EICA predicts that biogeographical release from natural enemies initiates a trade‐off in which exotic species in non‐native ranges become larger and more competitive, but invest less in consumer defences, relative to populations in native ranges. This trade‐off is exceptionally complex because detecting concomitant biogeographical shifts in competitive ability and consumer defence depends upon which traits are targeted, how competition is measured, the defence chemicals quantified, whether defence chemicals do more than defend, whether ‘herbivory’ is artificial or natural, and where consumers fall on the generalist‐specialist spectrum. Previous meta‐analyses have successfully identified patterns but have yet to fully disentangle this complexity. We used meta‐analysis to reevaluate traditional metrics used to test EICA theory and then expanded on these metrics by partitioning competitive effect and competitive tolerance measures and testing Leaf‐Specific Mass in detail as a response trait. Unlike previous syntheses, our meta‐analyses detected evidence consistent with the classic trade‐off inherent to EICA. Plants from non‐native ranges imposed greater competitive effects than plants from native ranges and were less quantitatively defended than plants from native ranges. Our results for defence were not based on complex leaf chemistry, but instead were estimated from tannins, toughness traits and primarily Leaf‐Specific Mass. Species specificity occurred but did not influence the general patterns. As for all evidence for EICA‐like trade‐offs, we do not know if the biogeographical differences we found were caused by trade‐offs per se, but they are consistent with predictions derived from the overarching hypothesis. Underestimating physical leaf structure may have contributed to two decades of tepid perspectives on the trade‐offs fundamental to EICA.

Numerous pressures influence the ecological capacity and health of drylands globally. Shrubs are often a critical component of these systems and can function positively as foundation species through facilitation of other species. Nonetheless, limited attention has been paid to the potential negative and indirect effects of shrubs. Here, we tested the hypothesis that plant facilitation can both accelerate the invasion process and amplify the negative effects of an invader on the native community. The invasive species Bromus madritensis ssp. rubens capitalized on facilitation by resident native shrub species. This in turn further degraded California mixed grasslands by negatively impacting other annual protégé plant species in these specific microhabitats. Indirect shrub‐mediated interactions were thus a critical component of the ecological community assembly processes, and this suggests that we need to move beyond pairwise interactions to more rapidly advance grassland management and restoration theory.

This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.