Novel and co-evolved associations between insects and microorganisms as drivers of forest pestilence

Some of the most devastating diseases of trees involve associations between forest insects and microorganisms. Although a small number of native insect-microorganism symbioses can cause tree mortality, the majority of associations with tree health implications involve one or more exotic organisms. Here, we divide damaging symbioses between forest insects and microorganisms into four categories based on the native/exotic status of the species involved: (1) insect and microorganism are native; (2) insect is native, microorganism is exotic; (3) insect is exotic, microorganism is native; and (4) insect and microorganism are both exotic. For each category, we describe several well-researched examples of forest insect symbioses and discuss some of the consequences of the types of interactions within each category. We then discuss priorities for research on forest insect symbioses that could help to further elucidate patterns in the complexity of such interactions in the context of invasion biology. We argue that a nuanced understanding of insect-pathogen relationships is lacking, even for the few well-studied examples. Because novel associations between insects, microorganisms, and trees are increasing with globalization, such symbioses and their potential to negatively impact forest ecosystems demand focused research in the future.