Viruses were discovered at the end of the nineteenth century as filterable agents causing infectious diseases of plants and animals
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5. Subsequently, their pathogenicity and ability to undergo rapid evolutionary change
6 has sparked a large body of research, often connected to the so-called ‘microevolution’ of relatively closely related viruses
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8. However, over the last decade, our appreciation of the importance and distribution of viruses has expanded beyond the original parasitic–pathogen model, and now virologists recognize the role of viruses in host regulation and the maintenance of natural ecosystems
9. Shotgun metagenomic sequencing has also revealed the presence of a vast variety of viruses in diverse environmental samples and in apparently healthy organisms from all divisions of life
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To understand the true extent of virus genomic diversity—which may be significantly broader than that of their hosts—and the origins and forces that shape this diversity, virologists will have to systematically rationalize the more distant relationships between viruses, ideally reflecting their ‘macroevolution’, and virus taxonomy should provide an inclusive yet dynamic classification framework to reflect these relationships. In contrast to the taxonomies of cellular organisms, this new virus taxonomic framework will have to accommodate the current view that viruses have multiple origins (polyphyly) and that their diversity cannot be represented by a single virosphere-wide tree
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