Phylogenetic Evidence Supporting the Nonenveloped Nature of Hepadnavirus Ancestors

Reverse-transcribing animal DNA viruses include the hepadnaviruses, a well-characterized family of small enveloped viruses that infect vertebrates but also a sister group of nonenveloped viruses more recently discovered in fish and termed the nackednaviruses. Here, we describe the complete sequence of a virus found in the feces of an insectivorous bat, which encodes a core protein and a reverse transcriptase but no envelope protein. A database search identified a viral sequence from a permafrost sample as its closest relative. The two viruses form a cluster that occupies a basal phylogenetic position relative to hepadnaviruses and nackednaviruses, with an estimated divergence time of 500 My. These findings may lead to the definition of a "proto-nackednavirus" family and support the hypothesis that the ancestors of hepadnaviruses were nonenveloped.