The practice of classifying organisms into hierarchical groups originated with Aristotle and was codified into nearly immutable biological law by Linnaeus. The heart of taxonomy is the biological species, which forms the foundation for higher levels of classification. Whereas species have long been established among sexual eukaryotes, achieving a meaningful species concept for prokaryotes has been an onerous task and has proven exceedingly difficult for describing viruses and bacteriophages. Moreover, the assembly of viral "species" into higher-order taxonomic groupings has been even more tenuous, since these groupings were based initially on limited numbers of morphological features and more recently on overall genomic similarities. The wealth of nucleotide sequence information that catalyzed a revolution in the taxonomy of free-living organisms necessitates a reevaluation of the concept of viral species, genera, families, and higher levels of classification. Just as microbiologists discarded dubious morphological traits in favor of more accurate molecular yardsticks of evolutionary change, virologists can gain new insight into viral evolution through the rigorous analyses afforded by the molecular phylogenetics of viral genes. For bacteriophages, such dissections of genomic sequences reveal fundamental flaws in the Linnaean paradigm that necessitate a new view of viral evolution, classification, and taxonomy.Biological taxonomy is rooted in the Linnaean "boxes within boxes" hierarchical paradigm (80). Here, groups of organisms are defined by their shared characteristics. These groups are subdivided (boxes formed within boxes) based on greater numbers of characters shared within subgroups and on the presence of characters that distinguish between subgroups. This framework is strictly hierarchical; that is, a group at any one taxonomic level can belong to only one parental group (e.g., a species can be a member of only one genus). When devised, the Linnaean paradigm provided an orderly classification of living things, allowing natural historians to place newly found creatures into its hierarchical framework with relative ease, merely by navigating the ever more detailed sets of characteristics that defined groups within groups.Although this concept was devised in the absence of evolutionary theory, it readily accommodated evolution as a driving force leading to such hierarchical classification of organisms. Soon after the publication of Darwin's Origin of Species (25), Haeckel (55) proposed that the more closely related forms in Linnaean classification shared more recent common ancestors than did more distantly related forms. In this way, taxonomy based on shared characteristics could be used as a framework for understanding the evolution of organisms, since it is fundamentally based on the vertical inheritance of genetic information from parent to offspring. As a caveat, if genetic information were to be transferred between distantly related forms (those residing in different "boxes"), a purely hierarchical f...