Specialized myrmecophily at the ecological dawn of modern ants

Myrmecophiles--species that depend on ant societies--include some of the most morphologically and behaviorally specialized animals known. Remarkable adaptive characters enable these creatures to bypass fortress-like security, integrate into colony life, and exploit abundant resources and protection inside ant nests. Such innovations must result from intimate coevolution with hosts, but a scarcity of definitive fossil myrmecophiles obscures when and how this lifestyle arose. Here, we report the earliest known morphologically specialized and apparently obligate myrmecophile, in Early Eocene (∼ 52 million years old) Cambay amber from India. Protoclaviger trichodens gen. et sp. nov. is a stem-group member of Clavigeritae, a speciose supertribe of pselaphine rove beetles (Coleoptera: Staphylinidae) heavily modified for myrmecophily via reduced mouthparts for trophallaxis with worker ants, brush-like trichomes that exude appeasement compounds, and fusions of many body and antennal segments. Protoclaviger captures a transitional stage in the evolutionary development of this novel body plan, most evident in its still-distinct abdominal tergites. The Cambay paleobiota marks one of the first occurrences in the fossil record of a significant presence of modern ants. Protoclaviger reveals that sophisticated social parasites were nest intruders throughout, and probably before, the ascent of ants to ecological dominance, with ancient groups such as Clavigeritae primed to radiate as their hosts became increasingly ubiquitous.