Document Type


Date of Original Version



Biological Sciences


Although biological invasions are of considerable concern to ecologists, relatively little attention has been paid to the potential for and consequences of indirect interactions between invasive species. Such interactions are generally thought to enhance invasives' spread and impact (i.e., the “invasional meltdown” hypothesis); however, exotic species might also act indirectly to slow the spread or blunt the impact of other invasives. On the east coast of the United States, the invasive hemlock woolly adelgid (Adelges tsugae, HWA) and elongate hemlock scale (Fiorinia externa, EHS) both feed on eastern hemlock (Tsuga canadensis). Of the two insects, HWA is considered far more damaging and disproportionately responsible for hemlock mortality. We describe research assessing the interaction between HWA and EHS, and the consequences of this interaction for eastern hemlock. We conducted an experiment in which uninfested hemlock branches were experimentally infested with herbivores in a 2 × 2 factorial design (either, both, or neither herbivore species). Over the 2.5‐year course of the experiment, each herbivore's density was ∼30% lower in mixed‐ vs. single‐species treatments. Intriguingly, however, interspecific competition weakened rather than enhanced plant damage: growth was lower in the HWA‐only treatment than in the HWA + EHS, EHS‐only, or control treatments. Our results suggest that, for HWA‐infested hemlocks, the benefit of co‐occurring EHS infestations (reduced HWA density) may outweigh the cost (increased resource depletion).