The bark beetle holobiont: why microbes matter.

All higher organisms are involved in symbioses with microbes. The importance of these partnerships has led to the concept of the holobiont, defined as the animal or plant with all its associated microbes. Indeed, the interactions between insects and symbionts form much of the basis for the success and diversity of this group of arthropods. Insects rely on microbes to perform basic life functions and to exploit resources and habitats. By "partnering" with microbes, insects access new genomic variation instantaneously allowing the exploitation of new adaptive zones, influencing not only outcomes in ecological time, but the degree of innovation and change that occurs over evolutionary time. In this review, I present a brief overview of the importance of insect-microbe holobionts to illustrate how critical an understanding of the holobiont is to understanding the insect host and it interactions with its environment. I then review what is known about the most influential insect holobionts in many forest ecosystems-bark beetles and their microbes-and how new approaches and technologies are allowing us to illuminate how these symbioses function. Finally, I discuss why it will be critical to study bark beetles as a holobiont to understand the ramifications and extent of anthropogenic change in forest ecosystems.