Abstract
The taxonomy of herpesviruses has been updated by the International Committee on Taxonomy of Viruses (ICTV). The former family Herpesviridae has been split into three families, which have been incorporated into the new order Herpesvirales. The revised family Herpesviridae retains the mammal, bird and reptile viruses, the new family Alloherpesviridae incorporates the fish and frog viruses, and the new family Malacoherpesviridae contains a bivalve virus. Three new genera have been created in the family Herpesviridae, namely Proboscivirus in the subfamily Betaherpesvirinae and Macavirus and Percavirus in the subfamily Gammaherpesvirinae. These genera have been formed by the transfer of species from established genera and the erection of new species, and other new species have been added to some of the established genera. In addition, the names of some nonhuman primate virus species have been changed. The family Alloherpesviridae has been populated by transfer of the genus Ictalurivirus and addition of the new species Cyprinid herpesvirus 3. The family Malacoherpesviridae incorporates the new genus Ostreavirus containing the new species Ostreid herpesvirus 1.
The classification of herpesviruses has been updated as a result of recommendations made to the International Committee on Taxonomy of Viruses (ICTV; www.ictvonline.org) by the Herpesviridae Study Group as it was constituted in the period 2002–2005. The purpose of this report is to convey the taxonomic changes.
Morphologically, herpesviruses are distinct from all other viruses. A linear, double-stranded DNA genome of 125–290 kbp is contained within a T=16 icosahedral capsid, which is surrounded by a proteinaceous matrix dubbed the tegument and then by a lipid envelope containing membrane-associated proteins [18]. Genetically, herpesviruses fall into three distinct groupings that are related only tenuously to each other [4–5, 7]. These groupings consist of viruses of mammals, birds and reptiles, viruses of fish and frogs, and a single virus of bivalves. Of the few proteins that are detectably conserved in sequence among the three groupings, none has homologues that are found only in herpesviruses. The protein that comes nearest to being herpesvirus-specific is the putative ATPase subunit of the terminase (a complex that is responsible for packaging virus DNA into nascent capsids), which is conserved in all herpesviruses and, to a lesser degree, T4-like bacteriophages in the family Myoviridae. The taxonomic outcome of the apparently large genetic distances among the groupings is that the former family Herpesviridae [6], which contained all herpesviruses, has now been divided into three families that comprise the new order Herpesvirales. The revised family Herpesviridae retains the viruses of mammals, birds and reptiles, the new family Alloherpesviridae (from ἅλλoς – other, different) incorporates the fish and frog viruses, and the new family Malacoherpesviridae (from μαλακός – soft; μαλάκιoν – mollusc) contains the bivalve virus. The revised classification is shown in Table 1.
Table 1.
The order Herpesvirales
| TAXON | NAMEa | ACRONYMb | FORMER NAMEc | COMMON NAMEd |
|---|---|---|---|---|
| Order | Herpesvirales | |||
|
| ||||
| Family | Herpesviridae | |||
|
| ||||
| Subfamily | Alphaherpesvirinae | |||
|
| ||||
| Genus | Simplexvirus | |||
|
| ||||
| Species in the genus | Ateline herpesvirus 1 | AtHV1 | Spider monkey herpesvirus | |
| Bovine herpesvirus 2 | BoHV2 | Bovine mammillitis virus | ||
| Cercopithecine herpesvirus 2 | CeHV2 | SA8 | ||
| Human herpesvirus 1 * | HHV1 | Herpes simplex virus type 1 | ||
| Human herpesvirus 2 | HHV2 | Herpes simplex virus type 2 | ||
| Macacine herpesvirus 1 | McHV1 | Cercopithecine herpesvirus 1 | B-virus | |
| Macropodid herpesvirus 1 | MaHV1 | Parma wallaby herpesvirus | ||
| Macropodid herpesvirus 2 | MaHV2 | Dorcopsis wallaby herpesvirus | ||
| Papiine herpesvirus 2 | PaHV2 | Cercopithecine herpesvirus 16 | Herpesvirus papio 2 | |
| Saimiriine herpesvirus 1 | SaHV1 | Marmoset herpesvirus | ||
|
| ||||
| Genus | Varicellovirus | |||
|
| ||||
| Species in the genus | Bovine herpesvirus 1 | BoHV1 | Infectious bovine rhinotracheitis virus | |
| Bovine herpesvirus 5 | BoHV5 | Bovine encephalitis herpesvirus | ||
| Bubaline herpesvirus 1 | BuHV1 | Water buffalo herpesvirus | ||
| Canid herpesvirus 1 | CaHV1 | Canine herpesvirus | ||
| Caprine herpesvirus 1 | CpHV1 | Goat herpesvirus | ||
| Cercopithecine herpesvirus 9 | CeHV9 | Simian varicella virus | ||
| Cervid herpesvirus 1 | CvHV1 | Red deer herpesvirus | ||
| Cervid herpesvirus 2 | CvHV2 | Reindeer herpesvirus | ||
| Equid herpesvirus 1 | EHV1 | Equine abortion virus | ||
| Equid herpesvirus 3 | EHV3 | Equine coital exanthema virus | ||
| Equid herpesvirus 4 | EHV4 | Equine rhinopneumonitis virus | ||
| Equid herpesvirus 8 | EHV8 | Asinine herpesvirus 3 | ||
| Equid herpesvirus 9 | EHV9 | Gazelle herpesvirus | ||
| Felid herpesvirus 1 | FeHV1 | Feline rhinotracheitis virus | ||
| Human herpesvirus 3 * | HHV3 | Varicella-zoster virus | ||
| Phocid herpesvirus 1 | PhoHV1 | Harbour seal herpesvirus | ||
| Suid herpesvirus 1 | SuHV1 | Pseudorabies virus | ||
|
| ||||
| Tentative species in the genus | Equid herpesvirus 6 | EHV6 | Asinine herpesvirus 1 | |
|
| ||||
| Genus | Mardivirus | |||
|
| ||||
| Species in the genus | Columbid herpesvirus 1 | CoHV1 | None | Pigeon herpesvirus |
| Gallid herpesvirus 2 * | GaHV2 | Marek’s disease virus type 1 | ||
| Gallid herpesvirus 3 | GaHV3 | Marek’s disease virus type 2 | ||
| Meleagrid herpesvirus 1 | MeHV1 | Turkey herpesvirus | ||
|
| ||||
| Genus | Iltovirus | |||
|
| ||||
| Species in the genus | Gallid herpesvirus 1 * | GaHV1 | Infectious laryngotracheitis virus | |
| Psittacid herpesvirus 1 | PsHV1 | None | Pacheco’s disease virus | |
|
| ||||
| Unassigned species in the subfamily | Chelonid herpesvirus 5 | ChHV5 | None | Chelonid fibropapilloma-associated herpesvirus |
| Chelonid herpesvirus 6 | ChHV6 | None | Lung-eye-trachea disease-associated virus | |
|
| ||||
| Subfamily | Betaherpesvirinae | |||
|
| ||||
| Genus | Cytomegalovirus | |||
|
| ||||
| Species in the genus | Cercopithecine herpesvirus 5 | CeHV5 | African green monkey cytomegalovirus | |
| Human herpesvirus 5 * | HHV5 | Human cytomegalovirus | ||
| Macacine herpesvirus 3 | McHV3 | Cercopithecine herpesvirus 8 | Rhesus cytomegalovirus | |
| Panine herpesvirus 2 | PnHV2 | Pongine herpesvirus 4 | Chimpanzee cytomegalovirus | |
|
| ||||
| Tentative species in the genus | Aotine herpesvirus 1 | AoHV1 | Herpesvirus aotus type 1 | |
| Aotine herpesvirus 3 | AoHV3 | Herpesvirus aotus type 3 | ||
|
| ||||
| Genus | Muromegalovirus | |||
|
| ||||
| Species in the genus | Murid herpesvirus 1 * | MuHV1 | Mouse cytomegalovirus | |
| Murid herpesvirus 2 | MuHV2 | Rat cytomegalovirus | ||
|
| ||||
| Genus | Roseolovirus | |||
|
| ||||
| Species in the genus | Human herpesvirus 6 * | HHV6 | Human herpesvirus 6 | |
| Human herpesvirus 7 | HHV7 | Human herpesvirus 7 | ||
|
| ||||
| Genus | Proboscivirus | |||
|
| ||||
| Species in the genus | Elephantid herpesvirus 1 * | ElHV1 | None | Elephant endotheliotropic herpesvirus |
|
| ||||
| Unassigned species in the subfamily | Caviid herpesvirus 2 | CavHV2 | Guinea pig cytomegalovirus | |
| Suid herpesvirus 2 | SuHV2 | None | Pig cytomegalovirus | |
| Tupaiid herpesvirus 1 | TuHV1 | Tree shrew herpesvirus | ||
|
| ||||
| Subfamily | Gammaherpesvirinae | |||
|
| ||||
| Genus | Lymphocryptovirus | |||
|
| ||||
| Species in the genus | Callitrichine herpesvirus 3 | CalHV3 | Marmoset lymphocryptovirus | |
| Cercopithecine herpesvirus 14 | CeHV14 | African green monkey EBV-like virus | ||
| Gorilline herpesvirus 1 | GoHV1 | Pongine herpesvirus 3 | Gorilla herpesvirus | |
| Human herpesvirus 4 * | HHV4 | Epstein-Barr virus | ||
| Macacine herpesvirus 4 | McHV4 | Cercopithecine herpesvirus 15 | Rhesus lymphocryptovirus | |
| Panine herpesvirus 1 | PnHV1 | Pongine herpesvirus 1 | Herpesvirus pan | |
| Papiine herpesvirus 1 | PaHV1 | Cercopithecine herpesvirus 12 | Herpesvirus papio | |
| Pongine herpesvirus 2 | PoHV2 | Orangutan herpesvirus | ||
|
| ||||
| Genus | Rhadinovirus | |||
|
| ||||
| Species in the genus | Ateline herpesvirus 2 | AtHV2 | Herpesvirus ateles strain 810 | |
| Ateline herpesvirus 3 | AtHV3 | Herpesvirus ateles strain 73 | ||
| Bovine herpesvirus 4 | BoHV4 | Movar virus | ||
| Human herpesvirus 8 | HHV8 | Kaposi’s sarcoma-associated herpesvirus | ||
| Macacine herpesvirus 5 | McHV5 | Cercopithecine herpesvirus 17 | Rhesus rhadinovirus | |
| Murid herpesvirus 4 | MuHV4 | Murine gammaherpesvirus 68 | ||
| Saimiriine herpesvirus 2 * | SaHV2 | Herpesvirus saimiri | ||
|
| ||||
| Tentative species in the genus | Leporid herpesvirus 1 | LeHV1 | Cottontail rabbit herpesvirus | |
| Leporid herpesvirus 2 | LeHV2 | Herpesvirus cuniculi | ||
| Leporid herpesvirus 3 | LeHV3 | Herpesvirus sylvilagus | ||
| Marmodid herpesvirus 1 | MarHV1 | Woodchuck herpesvirus | ||
|
| ||||
| Genus | Macavirus | |||
|
| ||||
| Species in the genus | Alcelaphine herpesvirus 1 * | AlHV1 | Malignant catarrhal fever virus | |
| Alcelaphine herpesvirus 2 | AlHV2 | Hartebeest malignant catarrhal fever virus | ||
| Bovine herpesvirus 6 | BoHV6 | None | Bovine lymphotropic herpesvirus | |
| Caprine herpesvirus 2 | CpHV2 | None | Caprine herpesvirus 2 | |
| Hippotragine herpesvirus 1 | HiHV1 | Roan antelope herpesvirus | ||
| Ovine herpesvirus 2 | OvHV2 | Sheep-associated malignant catarrhal fever virus | ||
| Suid herpesvirus 3 | SuHV3 | None | Porcine lymphotropic herpesvirus 1 | |
| Suid herpesvirus 4 | SuHV4 | None | Porcine lymphotropic herpesvirus 2 | |
| Suid herpesvirus 5 | SuHV5 | None | Porcine lymphotropic herpesvirus 3 | |
|
| ||||
| Genus | Percavirus | |||
|
| ||||
| Species in the genus | Equid herpesvirus 2 * | EHV2 | Equine herpesvirus 2 | |
| Equid herpesvirus 5 | EHV5 | Equine herpesvirus 5 | ||
| Mustelid herpesvirus 1 | MusHV1 | Badger herpesvirus | ||
|
| ||||
| Unassigned species in the subfamily | Equid herpesvirus 7 | EHV7 | Asinine herpesvirus 2 | |
| Phocid herpesvirus 2 | PhoHV2 | None | Phocid herpesvirus 2 | |
| Saguinine herpesvirus 1 | SgHV1 | Callitrichine herpesvirus 1 | Herpesvirus saguinus | |
|
| ||||
| Unassigned species in the family | Iguanid herpesvirus 2 | IgHV2 | None | Iguana herpesvirus |
|
| ||||
| Unassigned viruses in the family | Acciptrid herpesvirus 1 | AcHV1 | Bald eagle herpesvirus | |
| Anatid herpesvirus 1 | AnHV1 | Duck plague herpesvirus | ||
| Boid herpesvirus 1 | BoiHV1 | Boa herpesvirus | ||
| Callitrichine herpesvirus 2 | CalHV2 | Marmoset cytomegalovirus | ||
| Caviid herpesvirus 1 | CavHV1 | Guinea pig herpesvirus | ||
| Caviid herpesvirus 3 | CavHV3 | Guinea pig herpesvirus 3 | ||
| Cebine herpesvirus 1 | CbHV1 | Capuchin herpesvirus AL-5 | ||
| Cebine herpesvirus 2 | CbHV2 | Capuchin herpesvirus AP-18 | ||
| Cercopithecine herpesvirus 3 | CeHV3 | SA6 | ||
| Cercopithecine herpesvirus 4 | CeHV4 | SA15 | ||
| Chelonid herpesvirus 1 | ChHV1 | Grey patch disease-associated virus | ||
| Chelonid herpesvirus 2 | ChHV2 | Pacific pond turtle herpesvirus | ||
| Chelonid herpesvirus 3 | ChHV3 | Painted turtle herpesvirus | ||
| Chelonid herpesvirus 4 | ChHV4 | Argentine turtle herpesvirus | ||
| Ciconiid herpesvirus 1 | CiHV1 | Black stork herpesvirus | ||
| Cricetid herpesvirus | CrHV1 | Hamster herpesvirus | ||
| Elapid herpesvirus 1 | EpHV1 | Indian cobra herpesvirus | ||
| Erinaceid herpesvirus 1 | ErHV1 | European hedgehog herpesvirus | ||
| Falconid herpesvirus 1 | FaHV1 | Falcon inclusion body disease virus | ||
| Gruid herpesvirus 1 | GrHV1 | Crane herpesvirus | ||
| Iguanid herpesvirus 1 | IgHV1 | Green iguana herpesvirus | ||
| Lacertid herpesvirus | LaHV1 | Green lizard herpesvirus | ||
| Macacine herpesvirus 6 | McHV6 | Cercopithecine herpesvirus 10 | Rhesus leukocyte-associated herpesvirus strain 1 | |
| Macacine herpesvirus 7 | McHV7 | Cercopithecine herpesvirus 13 | Herpesvirus cyclopis | |
| Murid herpesvirus 3 | MuHV3 | Mouse thymic herpesvirus | ||
| Murid herpesvirus 5 | MuHV5 | Field mouse herpesvirus | ||
| Murid herpesvirus 6 | MuHV6 | Sand rat nuclear inclusion agent | ||
| Ovine herpesvirus 1 | OvHV1 | Sheep pulmonary adenomatosis-associated herpesvirus | ||
| Perdicid herpesvirus 1 | PdHV1 | Bobwhite quail herpesvirus | ||
| Phalacrocoracid herpesvirus 1 | PhHV1 | Cormorant herpesvirus | ||
| Procyonid herpesvirus 1 | PrHV1 | Lorisine herpesvirus 1 | Kinkajou herpesvirus | |
| Sciurid herpesvirus 1 | ScHV1 | Ground squirrel cytomegalovirus | ||
| Sciurid herpesvirus 2 | ScHV2 | Ground squirrel herpesvirus | ||
| Sphenicid herpesvirus 1 | SpHV1 | Black footed penguin herpesvirus | ||
| Strigid herpesvirus 1 | StHV1 | Owl hepatosplenitis virus | ||
|
| ||||
| Family | Alloherpesviridae | |||
|
| ||||
| Genus | Ictalurivirus | |||
|
| ||||
| Species in the genus | Ictalurid herpesvirus 1 * | IcHV1 | Channel catfish virus | |
|
| ||||
| Unassigned species in the family | Cyprinid herpesvirus 3 | CyHV3 | None | Koi herpesvirus |
|
| ||||
| Unassigned viruses in the family | Acipenserid herpesvirus 1 | AciHV1 | White sturgeon herpesvirus 1 | |
| Acipenserid herpesvirus 2 | AciHV2 | White sturgeon herpesvirus 2 | ||
| Anguillid herpesvirus 1 | AngHV1 | Japanese eel herpesvirus | ||
| Cyprinid herpesvirus 1 | CyHV1 | Carp pox herpesvirus | ||
| Cyprinid herpesvirus 2 | CyHV2 | Haematopoietic necrosis virus | ||
| Esocid herpesvirus 1 | EsHV1 | Northern pike herpesvirus | ||
| Percid herpesvirus 1 | PeHV1 | Walleye epidermal hyperplasia herpesvirus | ||
| Pleuronectid herpesvirus 1 | PlHV1 | Turbot herpesvirus | ||
| Ranid herpesvirus 1 | RaHV1 | Lucké tumor herpesvirus | ||
| Ranid herpesvirus 2 | RaHV2 | Frog virus 4 | ||
| Salmonid herpesvirus 1 | SalHV1 | Herpesvirus salmonis | ||
| Salmonid herpesvirus 2 | SalHV2 | Oncorhynchus masou herpesvirus | ||
|
| ||||
| Family | Malacoherpesviridae | |||
|
| ||||
| Genus | Ostreavirus | |||
|
| ||||
| Species in the genus | Ostreid herpesvirus 1 * | OsHV1 | None | Oyster herpesvirus |
Type species in the genus.
Formal taxonomic names are in italicized font. The names of tentative species and unassigned viruses are in non-italicized font, since these ranks have no taxonomic standing.
Acronyms apply to viruses, not species, and have no taxonomic standing. A hyphen may be included prior to the number.
Where the name has changed, the former name is given. Where the name is new, the word “None” is given. Where the name has not changed, no information is given.
Common names apply to viruses, not species, and have no taxonomic standing. They may correspond to the formal name (e.g. human herpesvirus 7 for Human herpesvirus 7), or they may not (e.g. Epstein-Barr virus for Human herpesvirus 4), or they may take multiple forms (e.g. human herpesvirus 8 and Kaposi’s sarcoma-associated herpesvirus for Human herpesvirus 8; pseudorabies virus and Aujeszky’s disease virus for Suid herpesvirus 1) and variants thereof. This list is limited to a single, common, English name for each virus.
Three new genera have been created in the family Herpesviridae. One is the genus Proboscivirus (from Πρoβoσκίς or proboscis: elephant’s trunk), which recognizes a distinct lineage of the subfamily Betaherpesvirinae containing the new species Elephantid herpesvirus 1 [9, 16, 19]. The others are the genera Macavirus (sigla from malignant catarrhal fever) and Percavirus (sigla from perissodactyl and carnivore), which form two lineages in the subfamily Gammaherpesvirinae that are separable from the established genera [14, 15]. The genus Macavirus contains the established species Alcelaphine herpesvirus 1, Alcelaphine herpesvirus 2, Hippotragine herpesvirus 1 and Ovine herpesvirus 2, which were transferred from the genus Rhadinovirus, plus the new species Bovine herpesvirus 6, Caprine herpesvirus 2, Suid herpesvirus 3, Suid herpesvirus 4 and Suid herpesvirus 5 [1–2, 23]. The genus Percavirus contains the established species Equid herpesvirus 2, Equid herpesvirus 5 and Mustelid herpesvirus 1, which were also transferred from the genus Rhadinovirus. Other new species include Columbid herpesvirus 1 and Psittacid herpesvirus 1 in the genera Mardivirus and Iltovirus, respectively, of the subfamily Alphaherpesvirinae [8, 22]. In addition, several new species were classified at the level of subfamily or family, with assignment to genera awaiting further data. These include the reptilian species Chelonid herpesvirus 5 and Chelonid herpesvirus 6 in the subfamily Alphaherpesvirinae [3, 11, 13, 17, 26], Phocid herpesvirus 2 in the subfamily Gammaherpesvirinae [10, 12], and Iguanid herpesvirus 2 in the family Herpesviridae [13, 25].
Herpesvirus species are named after a taxon of the host that in its natural setting harbours the virus [20, 21]. The default host taxon employed is that of family, and, except for the species of humans, the name ends in ‘-id’. Owing to their larger numbers, species from nonhuman primates and the family Bovidae have been designated by host subfamily, with the name ending in ‘-ine’. In the revised taxonomy, species from nonhuman primates have been renamed by host genus, with the name again ending in ‘-ine’. This has resulted in name changes for several Old World and one New World nonhuman primate herpesvirus species, and has reduced the very large number of species that previously incorporated the Cercopithecine prefix. The new names are correlated to the former names in Table 1.
The established genus Ictalurivirus has been transferred to the family Alloherpesviridae, and the new species Cyprinid herpesvirus 3 been added to the family [24]. The new genus Ostreavirus (from ὄστρεoν or ostrea – oyster) has been created in the family Malacoherpesviridae, containing the single known species of an invertebrate host, Ostreid herpesvirus 1 [7].
The need to balance logic and utility makes taxonomy a challenging process, and matters are often not straightforward. In addition to keeping abreast of new viruses as they are discovered, current discussions in the Herpesviridae (now Herpesvirales) Study Group include updating the herpesvirus species definition and the methods used for taxonomic assignment, in order to accommodate the well-established value of sequence-based phylogenetic relationships, and defining a pathway to species status for the large and increasing number of herpesviruses that have been detected only by PCR.
Contributor Information
Andrew J. Davison, Email: a.davison@mrcvu.gla.ac.uk, MRC Virology Unit, Institute of Virology, University of Glasgow, Church Street, Glasgow G11 5JR, UK, Chair of the ICTV Herpesviridae Study Group (1999–2005)
Richard Eberle, Center for Veterinary Health Sciences, Oklahoma State University, Stillwater, OK, USA.
Bernhard Ehlers, Robert Koch Institute, Berlin, Germany.
Gary S. Hayward, Johns Hopkins School of Medicine, Baltimore, MD, USA
Duncan J. McGeoch, MRC Virology Unit, Glasgow, UK
Anthony C. Minson, Department of Pathology, University of Cambridge, Cambridge, UK
Philip E. Pellett, Email: ppellett@med.wayne.edu, Wayne State University School of Medicine, Detroit, MI, USA, Chair of the ICTV Herpesvirales Study Group (2005–2011)
Bernard Roizman, Marjorie B. Kovler Viral Oncology Laboratories, University of Chicago, Chicago, IL, USA.
Michael J. Studdert, School of Veterinary Science, University of Melbourne, Melbourne, Victoria, Australia
Etienne Thiry, Faculty of Veterinary Medicine, University of Liège, Liège, Belgium.
References
- 1.Chmielewicz B, Goltz M, Ehlers B. Detection and multigenic characterization of a novel gammaherpesvirus in goats. Virus Res. 2001;75:87–94. doi: 10.1016/s0168-1702(00)00252-5. [DOI] [PubMed] [Google Scholar]
- 2.Chmielewicz B, Goltz M, Franz T, Bauer C, Brema S, Ellerbrok H, Beckmann S, Rziha HJ, Lahrmann KH, Romero C, Ehlers B. A novel porcine gammaherpesvirus. Virology. 2003;308:317–329. doi: 10.1016/s0042-6822(03)00006-0. [DOI] [PubMed] [Google Scholar]
- 3.Coberley SS, Condit RC, Herbst LH, Klein PA. Identification and expression of immunogenic proteins of a disease-associated marine turtle herpesvirus. J Virol. 2002;76:10553–10558. doi: 10.1128/JVI.76.20.10553-10558.2002. [DOI] [PMC free article] [PubMed] [Google Scholar]
- 4.Davison AJ. Channel catfish virus: a new type of herpesvirus. Virology. 1992;186:9–14. doi: 10.1016/0042-6822(92)90056-u. [DOI] [PubMed] [Google Scholar]
- 5.Davison AJ. Evolution of the herpesviruses. Vet Microbiol. 2002;86:69–88. doi: 10.1016/s0378-1135(01)00492-8. [DOI] [PubMed] [Google Scholar]
- 6.Davison A, Eberle R, Hayward GS, McGeoch DJ, Minson AC, Pellett PE, Roizman B, Studdert MJ, Thiry E. Herpesviridae. In: Fauquet CM, Mayo MA, Maniloff J, Desselberger U, Ball LA, editors. Virus Taxonomy, Classification and Nomenclature of Viruses, Eighth Report of the International Committee on the Taxonomy of Viruses. Elsevier Academic Press; London: 2005a. pp. 193–212. [Google Scholar]
- 7.Davison AJ, Trus B, Cheng N, Steven AC, Watson MS, Cunningham C, Le Deuff RM, Renault T. A novel class of herpesvirus with bivalve hosts. J Gen Virol. 2005b;86:41–53. doi: 10.1099/vir.0.80382-0. [DOI] [PubMed] [Google Scholar]
- 8.Ehlers B, Borchers K, Grund C, Frölich K, Ludwig H, Buhk HJ. Detection of new DNA polymerase genes of known and potentially novel herpesviruses by PCR with degenerate and deoxyinosine-substituted primers. Virus Genes. 1999;18:211–220. doi: 10.1023/a:1008064118057. [DOI] [PubMed] [Google Scholar]
- 9.Ehlers B, Burkhardt S, Goltz M, Bergmann V, Ochs A, Weiler H, Hentschke J. Genetic and ultrastructural characterization of a European isolate of the fatal endotheliotropic elephant herpesvirus. J Gen Virol. 2001;82:475–482. doi: 10.1099/0022-1317-82-3-475. [DOI] [PubMed] [Google Scholar]
- 10.Harder TC, Harder M, Vos H, Kulonen K, Kennedy-Stoskopf S, Liess B, Appel MJ, Osterhaus AD. Characterization of phocid herpesvirus-1 and -2 as putative alpha- and gammaherpesviruses of North American and European pinnipeds. J Gen Virol. 1996;77:27–35. doi: 10.1099/0022-1317-77-1-27. [DOI] [PubMed] [Google Scholar]
- 11.Herbst L, Ene A, Su M, Desalle R, Lenz J. Tumor outbreaks in marine turtles are not due to recent herpesvirus mutations. Curr Biol. 2004;14:R697–699. doi: 10.1016/j.cub.2004.08.040. [DOI] [PubMed] [Google Scholar]
- 12.Martina BE, Verjans GM, Harder TC, Kuiken T, van de Bildt MW, van Bergen PA, Osterhaus AD. Seal gammaherpesviruses: identification, characterisation and epidemiology. Virus Res. 2003;94:25–31. doi: 10.1016/s0168-1702(03)00120-5. [DOI] [PubMed] [Google Scholar]
- 13.McGeoch DJ, Gatherer D. Integrating reptilian herpesviruses into the family Herpesviridae. J Virol. 2005;79:725–731. doi: 10.1128/JVI.79.2.725-731.2005. [DOI] [PMC free article] [PubMed] [Google Scholar]
- 14.McGeoch DJ, Dolan A, Ralph AC. Toward a comprehensive phylogeny for mammalian and avian herpesviruses. J Virol. 2000;74:10401–10406. doi: 10.1128/jvi.74.22.10401-10406.2000. [DOI] [PMC free article] [PubMed] [Google Scholar]
- 15.McGeoch DJ, Gatherer D, Dolan A. On phylogenetic relationships among major lineages of the Gammaherpesvirinae. J Gen Virol. 2005;86:307–316. doi: 10.1099/vir.0.80588-0. [DOI] [PubMed] [Google Scholar]
- 16.McGeoch DJ, Rixon FJ, Davison AJ. Topics in herpesvirus genomics and evolution. Virus Res. 2006;117:90–104. doi: 10.1016/j.virusres.2006.01.002. [DOI] [PubMed] [Google Scholar]
- 17.Nigro O, Yu G, Aguirre AA, Lu Y. Sequencing and characterization of the full-length gene encoding the single-stranded DNA binding protein of a novel Chelonian herpesvirus. Arch Virol. 2004;149:337–347. doi: 10.1007/s00705-003-0204-8. [DOI] [PubMed] [Google Scholar]
- 18.Pellett PE, Roizman B. The Herpesviridae: a brief introduction. In: Knipe DM, Howley PM, Griffin DE, Lamb RA, Martin MA, Roizman B, Straus SE, editors. Fields Virology. 5. Lippincott, Williams & Wilkins; Philadelphia: 2006. pp. 2479–2499. [Google Scholar]
- 19.Richman LK, Montali RJ, Garber RL, Kennedy MA, Lehnhardt J, Hildebrandt T, Schmitt D, Hardy D, Alcendor DJ, Hayward GS. Novel endotheliotropic herpesviruses fatal for Asian and African elephants. Science. 1999;283:1171–1176. doi: 10.1126/science.283.5405.1171. [DOI] [PubMed] [Google Scholar]
- 20.Roizman B, Carmichael LE, Deinhardt F, de-The G, Nahmias AJ, Plowright W, Rapp F, Sheldrick P, Takahashi M, Wolf K. Herpesviridae. Definition, provisional nomenclature, and taxonomy. Intervirology. 1981;16:201–217. doi: 10.1159/000149269. [DOI] [PubMed] [Google Scholar]
- 21.Roizman B, Desrosiers RC, Fleckenstein B, Lopez C, Minson AC, Studdert MJ. The family Herpesviridae: an update. Arch Virol. 1992;123:425–449. doi: 10.1007/BF01317276. [DOI] [PubMed] [Google Scholar]
- 22.Thureen DR, Keeler CL., Jr Psittacid herpesvirus 1 and infectious laryngotracheitis virus: Comparative genome sequence analysis of two avian alphaherpesviruses. J Virol. 2006;80:7863–7872. doi: 10.1128/JVI.00134-06. [DOI] [PMC free article] [PubMed] [Google Scholar]
- 23.Ulrich S, Goltz M, Ehlers B. Characterization of the DNA polymerase loci of the novel porcine lymphotropic herpesviruses 1 and 2 in domestic and feral pigs. J Gen Virol. 1999;80:3199–3205. doi: 10.1099/0022-1317-80-12-3199. [DOI] [PubMed] [Google Scholar]
- 24.Waltzek TB, Kelley GO, Stone DM, Way K, Hanson L, Fukuda H, Hirono I, Aoki T, Davison AJ, Hedrick RP. Koi herpesvirus represents a third cyprinid herpesvirus (CyHV-3) in the family Herpesviridae. J Gen Virol. 2005;86:1659–1667. doi: 10.1099/vir.0.80982-0. [DOI] [PubMed] [Google Scholar]
- 25.Wellehan JFX, Jarchow JL, Reggiardo C, Jacobson ER. A novel herpesvirus associated with hepatic necrosis in a San Esteban chuckwalla, Sauromalus varius. J Herpetol Med Surg. 2003;13:15–19. [Google Scholar]
- 26.Yu Q, Hu N, Lu Y, Nerurkar VR, Yanagihara R. Rapid acquisition of entire DNA polymerase gene of a novel herpesvirus from green turtle fibropapilloma by a genomic walking technique. J Virol Methods. 2001;91:183–195. doi: 10.1016/s0166-0934(00)00267-6. [DOI] [PubMed] [Google Scholar]