The dark side of facilitation: native shrubs facilitate exotic annuals more strongly than native annuals

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.

Provenance of invaders has scale-dependent impacts in a changing wetland ecosystem

Exotic species are associated with a variety of impacts on biodiversity, but it is unclear whether impacts of exotic species differ from those of native species with similar growth forms or native species invading disturbed sites. We compared presence and abundance of native and exotic invaders with changes in wetland plant species diversity over a 28-year period by re-surveying 22 ponds to identify factors correlated with observed changes. We also compared communities found within dense patches of native and exotic emergent species with similar habits. Within patches, we found no categorical diversity differences between areas dominated by native or exotic emergent species. At the pond scale, the cover of the exotic grass Phragmitesaustralis best predicted change in diversity and evenness over time, likely owing to its significant increase in coverage over the study period. These changes in diversity and evenness were strongest in younger, less successionally-advanced ponds. Changes associated with cover of P.australis in these ponds were not consistent with expected diversity decreases, but instead with a dampening of diversity gains, such that the least-invaded ponds increased in diversity the most over the study period. There were more mixed effects on evenness, ranging from a reduction in evenness gains to actual losses of evenness in the ponds with highest invader cover. In this wetland complex, the habit, origin and invasiveness of species contribute to diversity responses in a scale- and context-dependent fashion. Future efforts to preserve diversity should focus on preventing the arrival and spread of invaders that have the potential to cover large areas at high densities, regardless of their origin. Future studies should also investigate more thoroughly how changes in diversity associated with species invasions are impacted by other ongoing ecosystem changes.

Exotic invaders gain foraging benefits by shoaling with native fish

Freshwater habitats are under increasing threat due to invasions of exotic fish. These invasions typically begin with the introduction of small numbers of individuals unfamiliar with the new habitat. One way in which the invaders might overcome this disadvantage is by associating with native taxa occupying a similar ecological niche. Here we used guppies ( Poecilia reticulata ) from a feral population in Mexico to test the prediction that exotic shoaling fish can associate with heterospecifics, and that they improve their foraging efficiency by doing so. Guppies have invaded the Mexican High Plateau and are implicated in the declines of many native topminnow (Goodeinae) species. We show that heterospecific associations between guppies and topminnows can deliver the same foraging benefits as conspecific shoals, and that variation in foraging gains is linked to differences in association tendency. These results uncover a mechanism enabling founding individuals to survive during the most vulnerable phase of an invasion and help explain why guppies have established viable populations in many parts of Mexico as well in every continent except Antarctica.