Herbaceous invaders in temperate forests: a systematic review of their ecology and proposed mechanisms of invasion

We present a systematic literature review of exotic understory forest herbaceous invasions with a focus on the forests of East Asia (EAS) and Eastern North America (ENA), two dominant regions of the north temperate deciduous forest biome. We examined the biogeographic origins of herbaceous invaders in EAS and ENA forests, summarized their life histories and ecology, and compiled the relevant literature on the 10 leading mechanistic hypotheses proposed for these invasions. We asked whether invasions of EAS and ENA forests by herbs are shared between regions, and whether a common suite of ecological traits unite these invaders into a functionally distinct group. In a focused summary of the empirical literature, we investigated if leading hypothesized mechanisms for biological invasions at large are also invoked and supported for this ecologically important, but relatively understudied, group of species. In contrast to ENA, forest invaders in EAS are overwhelmingly herbaceous (78% of forest invasions vs. 34% for ENA) and originate from different regions. Plant families represented and species traits between regions differed. Within a single species, multiple invasion mechanisms were often supported, highlighting the need for future research that simultaneously investigates multiple mechanistic hypotheses. Further, because results for a single invader often differed across space and time, a shift in focus to incorporate the complex dynamics across temporal and spatial scales with the consideration of spatial heterogeneity and the interplays among natural and anthropogenic factors to study exotic invasions is needed.