From beginning to end: the synecology of tree-killing bark beetles, fungi, and trees.

Over a century of research has revealed an amazing complexity of behaviours and physiological adaptations that allow tiny bark beetles to overcome large trees, sometimes resulting in outbreaks that kill millions of trees. Turning a tree into a home and successfully raising offspring involves constant interactions among the beetles, the tree, its microbiome, and the beetles' associated microbes, all influenced by abiotic factors that can determine success or failure. While we have learned much about these systems, substantial knowledge gaps remain. This synthesis aims to clarify and integrate current understanding, identify gaps, challenge long-held assumptions, and address interpretative issues that impede progress toward a holistic understanding of these systems. We advocate for expanding perspectives using synecological approaches to understand these complex systems better. We encourage expanding research into how colonization by the bark beetle-fungi complex influences subsequent tree decay and forest carbon dynamics. An explicit goal is to provide a comprehensive resource for new researchers while encouraging them to question established hypotheses and to explore new avenues of enquiry.