Although a range of studies have suggested that competition plays a critical role in determining herbivore assemblages, there has been little work addressing the nature of interactions between competing invasive herbivores. We report the results of research on the hemlock woolly adelgid Adelges tsugae ('HWA') and elongate hemlock scale Fiorinia externa ('EHS'), invasive herbivores that both feed on eastern hemlock (Tsuga canadensis). HWA has been linked to hemlock mortality throughout the East Coast of the US; the loss of hemlock threatens to permanently alter surrounding ecosystems. We assessed the spread and impact of both species by resurveying 142 hemlock stands across a 7,500 km 2 latitudinal transect, running from coastal CT to northern MA, for HWA and EHS density as well as hemlock mortality. These stands had been previously surveyed in either 1997-1998 (CT) or 2002. While the number of HWAinfested stands has increased, per-stand HWA density has substantially decreased. In contrast, EHS distribution and density has increased dramatically since 1997-1998. Hemlock mortality was much more strongly related to HWA density than to EHS density, and many stands remain relatively healthy despite an overall increase in hemlock mortality. There was a positive correlation between HWA and EHS densities in stands with low mean HWA densities, suggesting the potential for host-plant-mediated facilitation of EHS by HWA. Our findings underline the importance of research explicitly addressing interactions between competing invasive species, and of determining the potential consequences of these interactions for the invaded ecosystem.
Common buckthorn (Rhamnus cathartica L.) is one of the most abundant and ecologically harmful non-native plants in forests of the Upper Midwest United States. At the same time, European earthworms are invading previously glaciated areas in this region, with largely anecdotal evidence suggesting they compound the negative effects of buckthorn and influence the invasibility of these forests. Germination and seedling establishment are important control points for colonization by any species, and manipulation of the conditions influencing these life history stages may provide insight into why invasive species are successful in some environments and not others. Using a greenhouse microcosm experiment, we examined the effects of important biotic and abiotic factors on the germination and seedling establishment of common buckthorn. We manipulated light levels, leaf litter depth and earthworm presence to investigate the independent and interactive effects of these treatments on buckthorn establishment. We found that light and leaf litter depth were significant predictors of buckthorn germination but that the presence of earthworms was the most important factor; earthworms interacted with light and leaf litter to increase the number and biomass of buckthorn across all treatments. Path analysis suggested both direct and moisture-mediated indirect mechanisms controlled these processes. The results suggest that the action of earthworms may provide a pathway through which buckthorn invades forests of the Upper Midwest United States. Hence, researchers and managers should consider co-invasion of plants and earthworms when investigating invasibility and creating preemptive or post-invasion management plans.
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