The current rate of invasive species introductions is unprecedented, and the dramatic impacts of exotic invasive plants on community and ecosystem properties have been well documented. Despite the pressing management implications, the mechanisms that control exotic plant invasion remain poorly understood. Several factors, such as disturbance, propagule pressure, species diversity, and herbivory, are widely believed to play a critical role in exotic plant invasions. However, few studies have examined the relative importance of these factors, and little is known about how propagule pressure interacts with various mechanisms of ecological resistance to determine invasion success.We quantified the relative importance of canopy disturbance, propagule pressure, species diversity, and herbivory in determining exotic plant invasion in 10 eastern hemlock forests in Pennsylvania and New Jersey (USA). Use of a maximum-likelihood estimation framework and information theoretics allowed us to quantify the strength of evidence for alternative models of the influence of these factors on changes in exotic plant abundance. In addition, we developed models to determine the importance of interactions between ecosystem properties and propagule pressure. These analyses were conducted for three abundant, aggressive exotic species that represent a range of life histories: Alliaria petiolata, Berberis thunbergii, and Microstegium vimineum.Of the four hypothesized determinants of exotic plant invasion considered in this study, canopy disturbance and propagule pressure appear to be the most important predictors of A. petiolata, B. thunbergii, and M. vimineum invasion. Herbivory was also found to be important in contributing to the invasion of some species. In addition, we found compelling evidence of an important interaction between propagule pressure and canopy disturbance. This is the first study to demonstrate the dominant role of the interaction between canopy disturbance and propagule pressure in determining forest invasibility relative to other potential controlling factors. The importance of the disturbance-propagule supply interaction, and its nonlinear functional form, has profound implications for the management of exotic plant species populations. Improving our ability to predict exotic plant invasions will require enhanced understanding of the interaction between propagule pressure and ecological resistance mechanisms.

Hemlock woolly adelgid (HWA; Adelges tsugae Annand) infestations have resulted in the continuing decline of eastern hemlock (Tsuga canadensis (L.) Carrière) throughout much of the eastern United States. In 1994 and 2003, we quantified the vegetation composition and structure of two hemlock ravines in the Delaware Water Gap National Recreation Area. This is the first study to use pre-adelgid disturbance data, annual monitoring of infestation severity, and annual records of hemlock health to assess forest response to HWA infestation. In 2003, 25% of monitored hemlock trees were either dead or in severe decline. Measures of hemlock decline (crown vigor, transparency, density, and dieback) were correlated with HWA infestation severity and changes in light availability over the study period. Average percent total transmitted radiation more than doubled at these sites from 5.0% in 1994 to 11.7% in 2003. The total percent cover of vascular plants increased from 3.1% in 1994 to 11.3% in 2003. Species richness increased significantly, and more species were gained (53) than lost (19) from both ravine floras over the 9-year study period. Though exotic invasive plants were absent from these ravines in 1994, our 2003 resurvey found invasive plants in 35% of the permanent vegetation plots.

The successful invasion of exotic plants is often attributed to the absence of coevolved enemies in the introduced range (i.e., the enemy release hypothesis). Nevertheless, several components of this hypothesis, including the role of generalist herbivores, remain relatively unexplored. We used repeated censuses of exclosures and paired controls to investigate the role of a generalist herbivore, white-tailed deer (Odocoileus virginianus), in the invasion of 3 exotic plant species (Microstegium vimineum, Alliaria petiolata, and Berberis thunbergii) in eastern hemlock (Tsuga canadensis) forests in New Jersey and Pennsylvania (U.S.A.). This work was conducted in 10 eastern hemlock (T. canadensis) forests that spanned gradients in deer density and in the severity of canopy disturbance caused by an introduced insect pest, the hemlock woolly adelgid (Adelges tsugae). We used maximum likelihood estimation and information theoretics to quantify the strength of evidence for alternative models of the influence of deer density and its interaction with the severity of canopy disturbance on exotic plant abundance. Our results were consistent with the enemy release hypothesis in that exotic plants gained a competitive advantage in the presence of generalist herbivores in the introduced range. The abundance of all 3 exotic plants increased significantly more in the control plots than in the paired exclosures. For all species, the inclusion of canopy disturbance parameters resulted in models with substantially greater support than the deer density only models. Our results suggest that white-tailed deer herbivory can accelerate the invasion of exotic plants and that canopy disturbance can interact with herbivory to magnify the impact. In addition, our results provide compelling evidence of nonlinear relationships between deer density and the impact of herbivory on exotic species abundance. These findings highlight the important role of herbivore density in determining impacts on plant abundance and provide evidence of the operation of multiple mechanisms in exotic plant invasion.

Determining the ubiquity of top-down control effects of predators on their prey and ecosystem processes is important for understanding community and ecosystem-level consequences that may result from predator loss. We conducted experiments at two spatial scales to investigate the effects of terrestrial frogs (Eleutherodactylus coqui) on aerial and litter invertebrates, plant growth and herbivory, and litter decomposition. At both scales, frogs reduced aerial invertebrates and leaf herbivory, but had no effect on litter invertebrates. At the smaller scale, frogs increased foliage production rates, measured as the number of new leaves and new leaf area produced, by 80% and decomposition rates by 20%. The influence of E. coqui on increasing primary productivity and decomposition rates at the smaller scale appeared to be a result of elimination and excretion rather than of controlling prey. While the results provide evidence for frogs controlling herbivorous prey at both scales, species effects on ecosystem processes were only detectable at the smaller scale. The results highlight the difficulties in conducting experiments at large spatial scales. The findings from this study imply that the loss of amphibians and other species of higher trophic levels may affect nutrient cycling rates in tropical forests.

The widely held belief that riparian communities are highly invasible to exotic plants is based primarily on comparisons of the extent of invasion in riparian and upland communities. However, because differences in the extent of invasion may simply result from variation in propagule supply among recipient environments, true comparisons of invasibility require that both invasion success and propagule pressure are quantified. In this study, we quantified propagule pressure in order to compare the invasibility of riparian and upland forests and assess the accuracy of using a community's level of invasion as a surrogate for its invasibility. We found the extent of invasion to be a poor proxy for invasibility. The higher level of invasion in the studied riparian forests resulted from greater propagule availability rather than higher invasibility. Furthermore, failure to account for propagule pressure may confound our understanding of general invasion theories. Ecological theory suggests that species-rich communities should be less invasible. However, we found significant relationships between species diversity and invasion extent, but no diversity-invasibility relationship was detected for any species. Our results demonstrate that using a community's level of invasion as a surrogate for its invasibility can confound our understanding of invasibility and its determinants.