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Access Type

Open Access Thesis

Document Type


Degree Program

Organismic & Evolutionary Biology

Degree Type

Master of Science (M.S.)

Year Degree Awarded


Month Degree Awarded



1. The success of invasive species is often thought to be due to release from natural enemies. This hypothesis relies on the assumption that species are regulated by top-down forces in their native range and implies that species are likely to be regulated by bottom-up forces in the invasive range. Neither of these assumptions has been consistently supported with insects, a group which include many highly destructive invasive pest species.

2. Winter moth (Operophtera brumata) is an invasive defoliator in North America that appears to be regulated by mortality in the larval stage in its invasive range. To assess whether regulation in the invasive range is caused by top-down or bottom-up forces, we sought to identify the main causes of larval mortality.

3. To measure the importance of different sources of winter moth larval mortality, we used observational and manipulative field studies to measure dispersal, predation, parasitism, disease. We measured the response of larval dispersal in the field to multiple aspects of foliar quality, including total phenolics, pH 10 oxidized phenolics, trichome density, total nitrogen, total carbon, and carbon-nitrogen ration. We also used manipulative laboratory studies to measure the presence of cannibalism and dispersal.

4. Tree-level declines in density were driven by density-dependent larval dispersal of early instars with very little mortality caused by other factors. Later instar larvae dispersed at increased rates from previously damaged vs. undamaged foliage, and field larval dispersal rates were related to proportion of oxidative phenolics in 2015, suggesting that larval dispersal may have been mediated by an induced decline in foliar quality.

5. We conclude that winter moth population densities are regulated in New England by density-dependent larval dispersal possibly mediated by phenolic oxidative capacity. The suggested role of host plant quality in mediating dispersal means that winter moth population densities in New England appear to be regulated by bottom up forces, aligning with the assumptions of the natural enemy release hypothesis. This is the first study known to the authors presenting data showing a negative effect on insect herbivore performance from pH 10 oxidized phenolics.


First Advisor

Joseph S Elkinton