Eukaryotic single-stranded DNA (ssDNA) viruses are classified into 10 families (Table 1), but many remain unclassified (1, 2). Viruses from seven families (Bacilladnaviridae, Circoviridae, Geminiviridae, Genomoviridae, Nanoviridae, Redondoviridae, and Smacoviridae) have small circular genomes, with some encompassing only two genes, encoding the replication initiation protein (Rep) and the capsid protein (3–5). The genomes are replicated by the rolling-circle mechanism initiated by the virus-encoded Rep proteins of the HUH endonuclease superfamily (6–10). In eukaryotic ssDNA viruses, the HUH endonuclease domain is followed by a superfamily 3 helicase (S3H) domain, which is not found in prokaryotic ssDNA virus Reps, although homologous two-domain Reps are encoded by several groups of bacterial plasmids (5). Informally, this assemblage of ssDNA viruses has been referred to as “circular, Rep-encoding single-stranded (CRESS) DNA viruses” (4, 8). Viruses from the seven families infect hosts across the eukaryotic domain, including animals (Circoviridae), diatoms (Bacilladnaviridae), fungi (Genomoviridae), and plants (Nanoviridae and Geminiviridae), whereas members of the Smacoviridae and the newly created Redondoviridae have been discovered by metagenomics and are suspected to infect animals (3, 11), although an association of smacovirids with methanogenic archaea has also been suggested (12).