Termites are a clade of eusocial wood-feeding roaches with > 3000 described species. Eusociality emerged ~ 150 million years ago in the ancestor of modern termites, which, since then, have acquired and sometimes lost a series of adaptive traits defining of their evolution. Termites primarily feed on wood, and digest cellulose in association with their obligatory nutritional mutualistic gut microbes. Recent advances in our understanding of termite phylogenetic relationships have served to provide a tentative timeline for the emergence of innovative traits and their consequences on the ecological success of termites. While all "lower" termites rely on cellulolytic protists to digest wood, "higher" termites (Termitidae), which comprise ~ 70% of termite species, do not rely on protists for digestion. The loss of protists in Termitidae was a critical evolutionary step that fostered the emergence of novel traits, resulting in a diversification of morphology, diets, and niches to an extent unattained by "lower" termites. However, the mechanisms that led to the initial loss of protists and the succession of events that took place in the termite gut remain speculative. In this review, we provide an overview of the key innovative traits acquired by termites during their evolution, which ultimately set the stage for the emergence of "higher" termites. We then discuss two hypotheses concerning the loss of protists in Termitidae, either through an externalization of the digestion or a dietary transition. Finally, we argue that many aspects of termite evolution remain speculative, as most termite biological diversity and evolutionary trajectories have yet to be explored.

Division of Entomology Natural History Museum and Department of Ecology and Evolutionary Biology University of Kansas 1501 Crestline Drive Suite 140 Lawrence KS 66045 USA

Entomology and Nematology Department Institute of Food and Agricultural Science Ft Lauderdale Research and Education Center University of Florida Davie FL USA

Faculty of Tropical AgriSciences Czech University of Life Sciences Prague Czech Republic

Okinawa Institute of Science and Technology Graduate University Onna son Okinawa Japan

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Termites and subsocial roaches inherited many bacterial-borne carbohydrate-active enzymes (CAZymes) from their common ancestor

Genomic data provide insights into the classification of extant termites

Impact of Wood Age on Termite Microbial Assemblages

Mycophagy: A Global Review of Interactions between Invertebrates and Fungi

Alarm communication predates eusociality in termites

The functional evolution of termite gut microbiota

Termite dispersal is influenced by their diet

Termite nest evolution fostered social parasitism by termitophilous rove beetles