Biotic resistance describes the ability of resident species in a community to reduce the success of exotic invasions. Although resistance is a well-accepted phenomenon, less clear are the processes that contribute most to it, and whether those processes are strong enough to completely repel invaders. Current perceptions of strong, competition-driven biotic resistance stem from classic ecological theory, Elton's formulation of ecological resistance, and the general acceptance of the enemies-release hypothesis. We conducted a meta-analysis of the plant invasions literature to quantify the contribution of resident competitors, diversity, herbivores and soil fungal communities to biotic resistance. Results indicated large negative effects of all factors except fungal communities on invader establishment and performance. Contrary to predictions derived from the natural enemies hypothesis, resident herbivores reduced invasion success as effectively as resident competitors. Although biotic resistance significantly reduced the establishment of individual invaders, we found little evidence that species interactions completely repelled invasions. We conclude that ecological interactions rarely enable communities to resist invasion, but instead constrain the abundance of invasive species once they have successfully established.

Recent efforts to clear invasive plants from the fynbos of South Africa forces managers to think about how N 2 -fixing invasives have altered ecosystem processes and the implications of these changes for community development. This study investigated the changes in nitrogen (N) cycling regimes in fynbos with the invasion of Acacia saligna, the effects of clear-cutting acacia stands on soil microclimate and N cycling, and how altered N resources affected the growth of a weedy grass species. Litterfall, litter quality, soil nutrient pools, and ion exchange resin (IER)-available soil N were measured in uninvaded fynbos, intact acacia, and cleared acacia stands. In addition, a bioassay experiment was used to ascertain whether the changes in soil nutrient availability associated with acacia would enhance the success of a weedy grass species. Acacia plots had greater amounts of litterfall, which had higher concentrations of N. This led to larger quantities of organic matter, total N, and IER-available N in the soil. Clearing acacia stands caused changes in soil moisture and temperature, but did not result in differences in IER-available N. The alteration of N availability by acacias was shown to increase growth rates of the weedy grass Ehrharta calycina, suggesting that secondary invasions by nitrophilous weedy species may occur after clearing N 2 -fixing alien species in the fynbos. It is suggested that managers use controlled burns, the addition of mulch, and the addition of fynbos seed after clearing to lower the levels of available N in the soil and initiate the return of native vegetation.

Plant invaders have been suggested to change soil microbial communities and biogeochemical cycling in ways that can feedback to benefit themselves. In this paper, we ask when do these feedbacks influence the spread of exotic plants. Because answering this question is empirically challenging, we show how ecological theory on 'pushed' and 'pulled' invasions can be used to examine the problem. We incorporate soil feedbacks into annual plant invasion models, derive the conditions under which such feedbacks affect spread, and support our approach with simulations. We show that in homogeneous landscapes, strong positive feedbacks can influence spreading velocity for annual invaders, but that empirically documented feedbacks are not strong enough to do so. Moreover, to influence spread, invaders must modify the soil environment over a spatial scale larger than is biologically realistic. Though unimportant for annual invader spread in our models, feedbacks do affect invader density and potential impact. We discuss how future research might consider the way landscape structure, dispersal patterns, and the time scales over which plant-soil feedbacks develop regulate the effects of such feedbacks on invader spread.

Returning native species to habitats degraded by biological invasions is a critical conservation goal. A leading hypothesis poses that exotic plant dominance is self-reinforced by impacts on ecosystem processes, leading to persistent stable states. Invaders have been documented to modify fire regimes, alter soil nutrients or shift microbial communities in ways that feed back to benefit themselves over competitors. However, few studies have followed invasions through time to ask whether ecosystem impacts and feedbacks persist. Here we return to woodland sites in Hawai'i Volcanoes National Park that were invaded by exotic C4 grasses in the 1960s, the ecosystem impacts of which were studied intensively in the 1990s. We show that positive feedbacks between exotic grasses and soil nitrogen cycling have broken down, but rather than facilitating native vegetation, the weakening feedbacks facilitate new exotic species. Data from the 1990s showed that exotic grasses increased nitrogen-mineralization rates by two- to fourfold, but were nitrogen-limited. Thus, the impacts of the invader created a positive feedback early in the invasion. We now show that annual net soil nitrogen mineralization has since dropped to pre-invasion levels. In addition, a seedling outplanting experiment that varied soil nitrogen and grass competition demonstrates that the changing impacts of grasses do not favour native species re-establishment. Instead, decreased nitrogen availability most benefits another aggressive invader, the nitrogen-fixing tree Morella faya. Long-term studies of invasions may reveal that ecosystem impacts and feedbacks shift over time, but that this may not benefit native species recovery.