Archives of Virology

, Volume 154, Issue 1, pp 171–177

The order Herpesvirales

Authors

    • MRC Virology Unit, Institute of VirologyUniversity of Glasgow
  • Richard Eberle
    • Center for Veterinary Health SciencesOklahoma State University
  • Bernhard Ehlers
    • Robert Koch Institute
  • Gary S. Hayward
    • Johns Hopkins School of Medicine
  • Duncan J. McGeoch
    • MRC Virology Unit, Institute of VirologyUniversity of Glasgow
  • Anthony C. Minson
    • Department of PathologyUniversity of Cambridge
  • Philip E. Pellett
    • Wayne State University School of Medicine
  • Bernard Roizman
    • Marjorie B. Kovler Viral Oncology LaboratoriesUniversity of Chicago
  • Michael J. Studdert
    • School of Veterinary ScienceUniversity of Melbourne
  • Etienne Thiry
    • Faculty of Veterinary MedicineUniversity of Liège
Virology Division News

DOI: 10.1007/s00705-008-0278-4

Cite this article as:
Davison, A.J., Eberle, R., Ehlers, B. et al. Arch Virol (2009) 154: 171. doi:10.1007/s00705-008-0278-4

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; http://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 ἄλλος—other, different) incorporates the fish and frog viruses, and the new family Malacoherpesviridae (from μαλακός—soft; μαλάκιον—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 1e

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 3e

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 2e

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 1e

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 5e

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 1e

MuHV1

 

Mouse cytomegalovirus

Murid herpesvirus 2

MuHV2

 

Rat cytomegalovirus

Genus

Roseolovirus

   

 Species in the genus

Human herpesvirus 6e

HHV6

 

Human herpesvirus 6

Human herpesvirus 7

HHV7

 

Human herpesvirus 7

Genus

Proboscivirus

   

 Species in the genus

Elephantid herpesvirus 1e

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 4e

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 2e

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 1e

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 2e

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 1e

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 1e

OsHV1

None

Oyster herpesvirus

aFormal 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

bAcronyms apply to viruses, not species, and have no taxonomic standing. A hyphen may be included prior to the number

cWhere 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

dCommon 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

eType species in the genus

Three new genera have been created in the family Herpesviridae. One is the genus Proboscivirus (from προβoσκίς 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 ὄστρεον 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 (chaired by P.E. Pellett) 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.

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© Springer-Verlag 2008