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Archives of Virology

, Volume 159, Issue 5, pp 1239–1247 | Cite as

The family Parvoviridae

  • Susan F. CotmoreEmail author
  • Mavis Agbandje-McKenna
  • John A. Chiorini
  • Dmitry V. Mukha
  • David J. Pintel
  • Jianming Qiu
  • Maria Soderlund-Venermo
  • Peter Tattersall
  • Peter Tijssen
  • Derek Gatherer
  • Andrew J. Davison
Virology Division News

Abstract

A set of proposals to rationalize and extend the taxonomy of the family Parvoviridae is currently under review by the International Committee on Taxonomy of Viruses (ICTV). Viruses in this family infect a wide range of hosts, as reflected by the longstanding division into two subfamilies: the Parvovirinae, which contains viruses that infect vertebrate hosts, and the Densovirinae, encompassing viruses that infect arthropod hosts. Using a modified definition for classification into the family that no longer demands isolation as long as the biological context is strong, but does require a near-complete DNA sequence, 134 new viruses and virus variants were identified. The proposals introduce new species and genera into both subfamilies, resolve one misclassified species, and improve taxonomic clarity by employing a series of systematic changes. These include identifying a precise level of sequence similarity required for viruses to belong to the same genus and decreasing the level of sequence similarity required for viruses to belong to the same species. These steps will facilitate recognition of the major phylogenetic branches within genera and eliminate the confusion caused by the near-identity of species and viruses. Changes to taxon nomenclature will establish numbered, non-Latinized binomial names for species, indicating genus affiliation and host range rather than recapitulating virus names. Also, affixes will be included in the names of genera to clarify subfamily affiliation and reduce the ambiguity that results from the vernacular use of “parvovirus” and “densovirus” to denote multiple taxon levels.

Keywords

Host Taxon Shrimp Virus Blatella Germanica Host Order Core Capsid Protein 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.

Introduction

A set of proposals to update the taxonomy of the family Parvoviridae has been submitted by a review group that includes all members of the International Committee on Taxonomy of Viruses (ICTV) Parvoviridae Study Group (SG), and is currently under review. Until a final ICTV decision is reached, the proposal can be downloaded at http://talk.ictvonline.org/files/proposals/taxonomy_proposals_vertebrate1/default.aspx. The taxonomy of this family was last modified in 2004, prior to publication of the 8th ICTV Report [13], and is now significantly dated. In the interim, many new candidate viruses and previously unsuspected viral hierarchies have been identified, often by the use of viral discovery approaches that rely on polymerase chain reaction DNA amplification. Unfortunately, this approach typically confounds characterization of complex secondary structures in the viral hairpin telomeres that are essential for viability [8, 10], making the recovery of viruses from DNA challenging. To accommodate these important new viruses, while avoiding inclusion of viral sequence fragments integrated into host genomes [1, 9] or metagenomic data that lack integrity or clear host attribution (for example, a full-length Blatella germanica densovirus-like virus sequence, GenBank JQ320376, with a probable cockroach host that was identified in bat faeces [7]), the SG developed a polythetic definition of a virus in the family Parvoviridae. This requires the complete DNA sequence of all viral protein-coding sequences but no longer absolutely requires isolation of a viable virus provided an infectious etiology is supported by the structure and arrangement of the genome, serology, or other biological data. The viral definition used throughout these proposals is: “In order for an agent to be classified in the family Parvoviridae, it must be judged to be an authentic parvovirus on the basis of having been isolated and sequenced or, failing this, on the basis of having been sequenced in tissues, secretions, or excretions of unambiguous host origin, supported by evidence of its distribution in multiple individual hosts in a pattern that is compatible with dissemination by infection. The sequence must be in one piece, contain all the non-structural (NS) and viral particle (VP) coding regions, and meet the size constraints and motif patterns characteristic of the family”.

This definition allows inclusion of 134 new viruses and virus strains in the family Parvoviridae, together with 47 of the 53 previously recognized isolates [14]. Six recognized viruses for which no sequence information is currently available have been withdrawn from the formal taxonomy pending further analysis but will remain listed in subsequent reports as unassigned in their current genus. To improve taxonomic clarity and to facilitate the ready assimilation of present and future candidate viruses, a root-and-branch re-evaluation of the taxonomic structure and nomenclature of the family was also instituted, leading to the development of new systematic guidelines. Proposed changes to the taxonomy are summarized in Table 1. In the subfamily Parvovirinae, these changes include the introduction of three new genera and the expansion of five existing genus names with the affixes “parvo” or “proto”. In the subfamily Densovirinae, proposed changes include the introduction of two new genera for shrimp viruses and the expansion of the existing genus names Iteravirus and Densovirus to Iteradensovirus and Ambidensovirus, respectively. In both subfamilies, species identity levels will be lowered, numbered, binomial species names adopted, and new species introduced.
Table 1

Summary of changes between the current and proposed taxonomy

Current taxonomy

Proposed taxonomy

Genus

# Species

# viruses or strains

Genus

# Species

# viruses or variants

Subfamily Parvovirinae - vertebrate hosts

Amdovirus

1

1

Amdo parvo virus

2

2

-

-

-

Aveparvovirus a

1

2

Bocavirus

2

6

Boca parvo virus

12

22

-

-

-

Copiparvovirus a

2

2

Dependovirus

12

13

Dependo parvo virus

7

23

Erythrovirus

4

9

Erythro parvo virus

6

12

Parvovirus

12

18

Proto parvovirus

5

25

-

-

-

Tetraparvovirus a

6

10

Subfamily Densovirinae - arthropod hosts

Densovirus

2

2

Ambi densovirus

6

11

Brevidensovirus

2

2

Brevidensovirus

2

8

-

-

-

Hepandensovirus a

1

7

Iteravirus

1

1

Itera denso virus

5

6

-

-

-

Penstyldensovirus a

1

4

Pefudensovirus b

1

1

-

-

-

Bold type in genus names denotes new affixes

aDenotes new genera

bDenotes a retired genus

Changes in taxonomic structure and nomenclature

Taxon demarcation criteria

Parvoviruses encode two gene cassettes: an NS gene essential for replication and a VP gene encoding various forms of the structural protein [reviewed in 2, 14]. The amino acid sequence of the NS1 protein is used for phylogenetic analysis in the current proposals. NS1 is a multidomain ~70-80 kDa nuclear phosphoprotein that encodes highly conserved enzymatic activities, including a site-specific DNA-binding and single-strand endonuclease function and an AAA+ helicase [reviewed in 2, 3]. These well-conserved domains facilitate amino acid sequence alignment, allowing insights from structural biology and the derivation of a reliable sequence-based phylogeny (see the summary tree in Fig. 1). However, the core capsid protein sequence (defined as the smallest VP protein that contains all residues comprising the virion shell, as determined by X-ray crystallography) was analyzed in parallel, with conspicuously similar overall results although the data appeared less reliable at the greater distances apparent between the two subfamilies. Thus, the proposed taxonomic changes are supported by protein alignments of both major viral proteins. Previously, genera were defined largely by non-quantifiable criteria, including helper-virus requirements and genome characteristics, which provided little taxonomic structure [13, 14]. The current proposals add the requirement that all viruses in a genus should be monophyletic and encode NS1 proteins that are generally >30 % identical to each other at the amino acid sequence level but <30 % identical to those of other genera as determined by pairwise sequence alignments. Within the subfamily Parvovirinae, these criteria work well to separate all current and proposed genera, with the minor exception of the proposed genus Erythroparvovirus, where marginally greater divergence is evident between some virus pairs. The resulting eight genera in this subfamily are well supported by phylogenic analysis, as illustrated in Fig. 1 and detailed in the proposals.
Fig. 1

Phylogenetic tree showing genera in the family Parvoviridae. Phylogenetic analysis based on the amino acid sequence of the viral replication initiator protein, NS1, which contains a conserved AAA+ helicase domain corresponding to the Parvo_NS1 Pfam domain: http://pfam.sanger.ac.uk/family/Parvo_NS1. This region was aligned by incorporating insights from structural biology using the ehmmalign application in EMBASSY [6], and sequences flanking the Pfam domain were aligned using the modification of the Needleman-Wunsch local alignment method [11] as implented in MOE-Align (http://www.chemcomp.com). Pairwise p-distance matrices were constructed from this alignment using MEGA version 5.10 [12]. Bayesian trees were calculated over one billion iterations using BEAST [5], using a Yule model of speciation and an exponential relaxed molecular clock [4]. Trees were viewed in FigTree (part of BEAST) in ultrametric format on an arbitrary scale, midpoint-rooted, and with posterior probability scores indicated at statistically significant nodes. Bold type in genus names indicates affixes used to expand existing names. Asterisks denote the names of new genera

In the subfamily Densovirinae, attributable sequences are available only for a small number of economically significant viruses, which is likely to reflect poorly the diverse nature of viruses infecting hosts from the immense phylum Arthropoda. Accordingly, the >30 % identity requirement is applied less rigorously within this subfamily, in order to allow clustering of monophyletic viruses with conspicuously similar characteristics from host orders separated by large evolutionary distances. Constituent viruses in most of these genera are closely related and infect arthropods from the same host order. However, in the proposed genus Ambidensovirus, certain pairs of viruses that infect different orders of hosts fall short of the proposed identity requirements. Nevertheless, all of these viruses exhibit a complex genomic rearrangement that allows them to co-ordinate bidirectional transcription, which is not seen in viruses from any other parvovirus taxon. In the existing taxonomy, most of these “ambisense” viruses cluster in the genus Densovirus, with a single outlier (Periplaneta fuliginosa densovirus) that infects a blattodean host and is the sole member of genus Pefudensovirus. Recently, this organization was challenged by the identification of four new isolates with ambisense organization, which infect insects from different host orders and are closely related to, albeit not monophyletic with, viruses from both of the existing ambisense genera. To resolve this situation, it is proposed that the six groups should be combined as distinct species in a single monophyletic Ambidensovirus genus, which will have slightly relaxed demarcation criteria that likely reflect the host diaspora. Members of the proposed genus Ambidensovirus thus appear to illustrate how host divergence may mask and complicate underlying sequence-based phylogeny: using current approaches, it would prove challenging to track viral lineages with less conspicuous genomic rearrangements against this background of host-related genetic drift.

Previously, species in the family were generally required to be >95 % related in the NS1 DNA sequence, which is so high a level that many current species consist of single isolates. This has fuelled confusion between taxa and viruses in the literature, and it allows the species level to contribute little to taxonomic structure. The proposals will decrease species identity criteria significantly, requiring viruses in a species to encode NS1 proteins that show >85 % amino acid sequence identity while diverging by >15 % from viruses in other species. This adjustment permits a species to contain a greater diversity of viruses than is currently the case, so that it typically designates a distinct phylogenetic branch and thus adds useful structure within the genus. Other existing criteria, such as host, antigenic properties, and genome characteristics, are still considered.

The two subfamilies, Parvovirinae and Densovirinae, are distinguished primarily by their respective ability to infect vertebrate and arthropod hosts, and this remains the case in the proposals. This separation is supported by Bayesian phylogeny, although it is not immediately apparent under the rooting procedure used in Fig. 1.

Taxon nomenclature

Systematic changes are proposed at the level of species, in part because decreasing the mandatory level of sequence identity for this taxon effectively does away with current species divisions. The call for new names that encompass broader groups of viruses thus provides an opportunity for the field to adopt a non-Latinized, binomial system that has been discussed extensively in the literature [15, 16] and is commonly in use in other viral families. In the proposed nomenclature, species names are emphatically different from virus names and typically consist of a host taxon, a genus affiliation, and a distinguishing numerical or letter suffix, for example, Rodent protoparvovirus 1 (type species of the genus Protoparvovirus, which includes both the existing type species, Minute virus of mice, and a group of closely related rodent viruses, as detailed in Table 2). Since these names indicate the range of viruses included and their branch within the family, they provide useful information about the likely properties of the virus, and allow for facile addition of new species by simply advancing the numerical suffix, as in Rodent protoparvovirus 2, which currently contains a single virus, rat parvovirus 1. In addition, because the proposed species distinguish major branches within each genus, they will provide taxonomic names for groups of viruses that are now commonly discussed together in the literature. The only exceptions to this standard naming pattern involve two species from the genus Dependoparvovirus, which contain viruses that show excellent potential for clinical use as gene therapy delivery vectors. Viruses in one species are named “adeno-associated virus” plus a hyphenated numeral between 1-4 or 6-13, with individual isolates showing important differences in receptor-binding and tissue-specific transduction efficiency. Because these virus names are so well recognized both inside and outside the field, and because their specific numerals have such important implications, the SG considered it unwise to introduce host taxa or additional numerals into the species name. Accordingly, the proposed name for this species is Adeno-associated dependoparvovirus A (instead of the more systematic Primate dependoparvovirus 1). A second species in this genus is named Adeno-associated dependoparvovirus B and includes one virus, adeno-associated virus-5, which is also of current interest for gene therapy applications.
Table 2

Proposed taxonomy for the subfamily Parvovirinae

Genus

Species

Virus or virus variants

Abbreviation

Accession #

Amdoparvovirus

Carnivore amdoparvovirus 1

Aleutian mink disease virus

AMDV

JN040434

 

Carnivore amdoparvovirus 2

gray fox amdovirus

GFAV

JN202450

Aveparvovirus

Galliform aveparvovirus 1

chicken parvovirus

ChPV

GU214704

  

turkey parvovirus

TuPV

GU214706

Bocaparvovirus

Carnivore bocaparvovirus 1

canine minute virus

CnMV

FJ214110

 

Carnivore bocaparvovirus 2

canine bocavirus 1

CBoV

JN648103

 

Carnivore bocaparvovirus 3

feline bocavirus

FBoV

JQ692585

 

Pinniped bocaparvovirus 1

California sea lion bocavirus 1

CslBoV1

JN420361

  

California sea lion bocavirus 2

CslBoV2

JN420366

 

Pinniped bocaparvovirus 2

California sea lion bocavirus 3

CslBoV3

JN420365

 

Primate bocaparvovirus 1

human bocavirus 1

HBoV1

JQ923422

  

human bocavirus 3

HBoV3

EU918736

  

gorilla bocavirus

GBoV

HM145750

 

Primate bocaparvovirus 2

human bocavirus 2a

HBoV2a

FJ973558

  

human bocavirus 2b

HBoV2b

FJ973560

  

human bocavirus 2c

HBoV2c

FJ170278

  

human bocavirus 4

HBoV4

FJ973561

 

Ungulate bocaparvovirus 1

bovine parvovirus

BPV

DQ335247

 

Ungulate bocaparvovirus 2

porcine bocavirus 1

PBoV1

HM053693

  

porcine bocavirus 2

PBoV2

HM053694

  

porcine bocavirus 6

PBoV6

HQ291309

 

Ungulate bocaparvovirus 3

porcine bocavirus 5

PBoV5

HQ223038

 

Ungulate bocaparvovirus 4

porcine bocavirus 7

PBoV7

HQ291308

 

Ungulate bocaparvovirus 5

porcine bocavirus 3

PBoV3

JF429834

  

porcine bocavirus 4-1

PBoV4-1

JF429835

  

porcine bocavirus 4-2

PBoV4-2

JF429836

Copiparvovirus

Ungulate copiparvovirus 1

bovine parvovirus 2

BPV2

AF406966

 

Ungulate copiparvovirus 2

porcine parvovirus 4

PPV4

GQ387499

Dependoparvovirus

Adeno-associated dependoparvovirus A

adeno-associated virus-1

AAV1

AF063497

  

adeno-associated virus-2

AAV2

AF043303

  

adeno-associated virus-3

AAV3

AF028705

  

adeno-associated virus-4

AAV4

U89790

  

adeno-associated virus-6

AAV6

AF028704

  

adeno-associated virus-7

AAV7

AF513851

  

adeno-associated virus-8

AAV8

AF513852

  

adeno-associated virus-9

AAV9

AX753250

  

adeno-associated virus-10

AAV10

AY631965

  

adeno-associated virus-11

AAV11

AY631966

  

adeno-associated virus-12

AAV12

DQ813647

  

adeno-associated virus-13

AAV13

EU285562

  

adeno-associated virus-S17

AAVS17

AY695376

 

Adeno-associated dependoparvovirus B

adeno-associated virus-5

AAV5

AF085716

  

bovine adeno-associated virus

BAAV

AY388617

  

caprine adeno-associated virus

CapAAV

DQ335246

 

Anseriform dependoparvovirus 1

duck parvovirus

DPV

U22967

  

goose parvovirus-PT

GPV2

JF926695

  

goose parvovirus

GPV

U25749

 

Avian dependoparvovirus 1

avian adeno-associated virus

AAAV

AY186198

 

Chiropteran dependoparvovirus 1

bat adeno-associated virus

BtAAV

GU226971

 

Pinniped dependoparvovirus 1

California sea lion adeno-associated virus

CslAAV

JN420372

 

Squamate dependoparvovirus 1

snake adeno-associated virus

SAAV

AY349010

Erythroparvovirus

Primate erythroparvovirus 1

human parvovirus B19-Au

B19V-Au

M13178

  

human parvovirus B19-J35

B19V-J35

AY386330

  

human parvovirus B19-Wi

B19V-Wi

M24682

  

human parvovirus B19-A6

B19V-A6

AY064475

  

human parvovirus B19-Lali

B19V-Lali

AY044266

  

human parvovirus B19-V9

B19V-V9

AJ249437

  

human parvovirus B19-D91

B19-D91

AY083234

 

Primate erythroparvovirus 2

simian parvovirus

SPV

U26342

 

Primate erythroparvovirus 3

rhesus macaque parvovirus

RhMPV

AF221122

 

Primate erythroparvovirus 4

pig-tailed macaque parvovirus

PtMPV

AF221123

 

Rodent erythroparvovirus 1

chipmunk parvovirus

ChpPV

GQ200736

 

Ungulate erythroparvovirus 1

bovine parvovirus 3

BPV3

AF406967

Protoparvovirus

Carnivore protoparvovirus 1

feline parvovirus

FPV

EU659111

  

canine parvovirus

CPV

M19296

  

mink enteritis virus

MEV

D00765

  

racoon parvovirus

RaPV

JN867610

 

Primate protoparvovirus 1

bufavirus 1a

BuPV1a

JX027296

  

bufavirus 1b

BuPV1b

JX027295

  

bufavirus 2

BuPV2

JX027297

 

Rodent protoparvovirus 1

H-1 parvovirus

H1

X01457

  

Kilham rat virus

KRV

AF321230

  

LuIII virus

LuIII

M81888

  

minute virus of mice (prototype)

MVMp

J02275

  

minute virus of mice (immunosuppressive)

MVMi

M12032

  

minute virus of mice (Missouri)

MVMm

DQ196317

  

minute virus of mice (Cutter)

MVMc

U34256

  

mouse parvovirus 1

MPV1

U12469

  

mouse parvovirus 2

MPV2

DQ196319

  

mouse parvovirus 3

MPV3

DQ199631

  

mouse parvovirus 4

MPV4

FJ440683

  

mouse parvovirus 5

MPV5

FJ441297

  

hamster parvovirus

HaPV

U34255

  

tumor virus X

TVX

In preparation

  

rat minute virus 1

RMV1

AF332882

 

Rodent protoparvovirus 2

rat parvovirus 1

RPV1

AF036710

 

Ungulate protoparvovirus 1

porcine parvovirus Kresse

PPV-Kr

U44978

  

porcine parvovirus NADL-2

PPV-NADL2

L23427

Tetraparvovirus

Chiropteran tetraparvovirus 1

Eidolon helvum (bat) parvovirus

Ba-PARV4

JQ037753

 

Primate tetraparvovirus 1

human parvovirus 4 G1

PARV4G1

AY622943

  

human parv4 G2

PARV4G2

DQ873391

  

human parv4 G3

PARV4G3

EU874248

  

chimpanzee parv4

Ch-PARV4

HQ113143

 

Ungulate tetraparvovirus 1

bovine hokovirus 1

B-PARV4-1

EU200669

  

bovine hokovirus 2

B-PARV4-2

JF504697

 

Ungulate tetraparvovirus 2

porcine hokovirus

P-PARV4

EU200677

 

Ungulate tetraparvovirus 3

porcine Cn virus

CnP-PARV4

GU938300

 

Ungulate tetraparvovirus 4

ovine hokovirus

O-PARV4

JF504699

The type species for each genus is indicated in bold type. Viruses in the current species Canine adeno-associated virus, Equine adeno-associated virus, Ovine adeno-associated virus, HB parvovirus, Lapine parvovirus and RT parvovirus do not meet the new criteria for inclusion in the family. Viruses in the current species Chicken parvovirus are transferred from genus Protoparvovirus to the new genus Aveparvovirus in the new species Galliform aveparvovirus 1

In general, host taxon descriptors at the level of order, rather than family, are selected for species names to accommodate potential host-range disparity among viruses. However, where such host names were considered confusing or awkward to pronounce, less rigid terms were preferred, as in the use of “pinniped” (fin-footed mammals, including walrus, seals, and sea lions) instead of “carnivore” for viruses of the Californian sea lion, and “ungulate” (hoofed animals) instead of “artiodactyl” for viruses of cows, pigs, and sheep.

The proposals also expand the names of most existing genera by introducing an affix into each name. Two distinct problems are addressed in this way. First, it requires specialist knowledge to recognize that Amdovirus, Bocavirus, Dependovirus, and Erythrovirus are genera within one subfamily of the family Parvoviridae, and that Iteravirus is a genus in the subfamily Densovirinae. This dislocation will be addressed by adding the infixes “parvo” or “denso” to indicate subfamily affiliation, as in the genera Amdoparvovirus, Bocaparvovirus, Dependoparvovirus, Erythroparvovirus, and Iteradensovirus. One remaining genus in the subfamily Densovirinae, Brevidensovirus, already contains the infix, and proposed names for all new genera will include the appropriate notation. It is hoped that this modification will improve family recognition, thus providing information about the general properties of a virus in any given genus to people outside the field, and will obviate the need to explain the taxonomy whenever viruses in different parvovirus genera are compared. Practically, it was becoming challenging to invent names for new genera, since these commonly appeared to suggest affiliation to a different virus family. For example, a previously proposed genus name Partetravirus, which is widely in use in the field to encompass viruses related to human parvovirus 4 (PARV4, GenBank AY622943), was not welcomed by the ICTV because it arguably suggested that these viruses were members of the family Alphatetraviridae. In the current proposals, we again seek recognition for this group of viruses, but under the genus name Tetraparvovirus, since the infix should substantially limit ambiguity.

Although subfamily affiliations of viruses in the existing genera Parvovirus and Densovirus are explicit, vernacular use of “parvovirus” and “densovirus” is ambiguous because the terms indicate multiple taxa. Thus, “parvovirus” can refer to members of the genus Parvovirus, the subfamily Parvovirinae, or the family Parvoviridae, while “densovirus” can indicate genus or subfamily affiliation. To provide greater taxonomic precision, the proposals also insert the prefix “Proto” before Parvovirus, creating the genus name Protoparvovirus (from Greek, “proto” meaning “first”, in this case the first viruses identified), and “Ambi” before Densovirus, creating the genus name Ambidensovirus (from Latin or Celtic, meaning “both”, referring to ambisense transcription). Overall, these changes should provide the field with a more self-explanatory framework and greater precision when using taxonomically derived terms.

Taxon and virus lists for the proposed classification are shown in Tables 2 and 3. In the subfamily Parvovirinae, there are three new genera, to be called Aveparvovirus, to indicate the bird (Aves) hosts of the founding members, Copiparvovirus, a siglum for cow and pig, which were the hosts of the first two species identified, and Tetraparvovirus, from the name of the founder virus, human parvovirus 4 (PARV4), using Latin “tetra” in place of the numeral 4. In the subfamily Densovirinae, two new genera are proposed, in order to accommodate shrimp viruses. These will be called Hepandensovirus, to reflect the original name of these viruses, “hepatopancreatic parvovirus”, and Penstyldensovirus, a siglum for Penaeus stylirostris, the host, and name, of the founding member of this species.
Table 3

Proposed taxonomy for the subfamily Densovirinae

Genus

Species

Virus or virus variants

Abbreviation

Accession #

Ambidensovirus

Blattodean ambidensovirus 1

Periplaneta fuliginosa densovirus

PfDV

AF192260

 

Blattodean ambidensovirus 2

Blattella germanica densovirus 1

BgDV1

AY189948

 

Dipteran ambidensovirus 1

Culex pipens densovirus

CpDV

FJ810126

 

Hemipteran ambidensovirus 1

Planococcus citri densovirus

PcDV

AY032882

 

Lepidopteran ambidensovirus 1

Diatraea saccharalis densovirus

DsDV

AF036333

  

Galleria mellonella densovirus

GmDV

L32896

  

Helicoverpa armigera densovirus

HaDV1

JQ894784

  

Junonia coenia densovirus

JcDV

S47266

  

Mythimna loreyi densovirus

MlDV

AY461507

  

Pseudoplusia includens densovirus

PiDV

JX645046

 

Orthopteran ambidensovirus 1

Acheta domesticus densovirus

AdDV

HQ827781

Brevidensovirus

Dipteran brevidensovirus 1

Aedes aegypti densovirus 1

AaeDV1

M37899

  

Aedes albopictus densovirus 1

AalDV1

AY095351

  

Culex pipiens pallens densovirus

CppDV

EF579756

  

Anopheles gambiae densovirus

AgDV

EU233812

  

Aedes aegypti densovirus 2

AaeDV2

FJ360744

 

Dipteran brevidensovirus 2

Aedes albopictus densovirus 2

AalDV2

X74945

  

Aedes albopictus densovirus 3

AalDV3

AY310877

  

Haemagogus equinus densovirus

HeDV

AY605055

Hepandensovirus a

Decapod hepandensovirus 1

Penaeus monodon hepandensovirus 1

PmoHDV1

DQ002873

  

Penaeus chinensis hepandensovirus

PchDV

AY008257

  

Penaeus monodon hepandensovirus 2

PmoHDV2

EU247528

  

Penaeus monodon hepandensovirus 3

PmoHDV3

EU588991

  

Penaeus merguiensis hepandensovirus

PmeDV

DQ458781

  

Penaeus monodon hepandensovirus 4

PmoHDV4

FJ410797

  

Fenneropenaeus chinensis hepandensovirus

FchDV

JN082231

Iteradensovirus

Lepidopteran iteradensovirus 1

Bombyx mori densovirus

BmDV

AY033435

 

Lepidopteran iteradensovirus 2

Casphalia extranea densovirus

CeDV

AF375296

  

Sibine fusca densovirus

SfDV

JX020762

 

Lepidopteran iteradensovirus 3

Dendrolimus punctatus densovirus

DpDV

AY665654

 

Lepidopteran iteradensovirus 4

Papilio polyxenes densovirus

PpDV

JX110122

 

Lepidopteran iteradensovirus 5

Helicoverpa armigera densovirus

HaDV2

HQ613271

Penstyldensovirus b

Decapod penstyldensovirus 1

Penaeus stylirostris penstyldensovirus 1

PstDV1

AF273215

  

Penaeus monodon penstyldensovirus 1

PmoPDV1

GQ411199

  

Penaeus monodon penstyldensovirus 2

PmoPDV2

AY124937

  

Penaeus stylirostris penstyldensovirus 2

PstDV2

GQ475529

The type species for each genus is indicated in bold type

aIndicates genus of viruses formerly known as hepatopancreatic parvovirus [HPV] of shrimp

bIndicates genus of viruses formerly known as infectious hypodermal and hematopoietic necrosis virus (IHHNV) of shrimp

As a general rule, the proposals do not tamper with existing viral names, which remain written in Roman script, for example, canine minute virus and Galleria mellonella densovirus (in this case capitalized because “Galleria” is derived from a formal name), whereas all formal taxonomic names, for family, subfamily, genus, and species, are capitalized and written in italics. Although abbreviations of viral names also have no formal standing, those listed in Tables 2 and 3 are recommended by the SG, in order to encourage uniformity. For viruses from the subfamily Densovirinae, viral names have typically been assembled from binomial host names plus the word “densovirus”, for example, Jujonia coenia densovirus, originally abbreviated to JcDNV (where the capitalized “N” harks back to a time when these viruses were called “densonucleosis viruses”). However, many host species share the same initials, and viruses from multiple densovirus genera can infect a single host species. Therefore, as new viruses were identified, their abbreviations were distinguished from pre-existing isolates by the insertion of additional letters, causing them to become progressively longer. In part to offset this continued expansion, the SG suggest eliminating the vestigial N from all abbreviations, as implemented in Table 3. Finally, the proposed establishment of two new genera for shrimp viruses, each encompassing viruses that are responsible for an economically significant disease but which all infect an overlapping group of host species, was deemed to require an unusual approach. As discussed above, one of these viral clusters, formerly known as “hepatopancreatic parvovirus” of shrimp (HPV), now constitutes the genus Hepandensovirus, whereas the other, formerly known as “infectious hypodermal and hematopoietic virus“ of shrimp (IHHNV), is classified in the genus Penstyldensovirus. Because these names and abbreviations do not meet standard densovirus conventions, in this particular instance the SG voted to rename the viruses. However, rather than use “densovirus”, the new genus name was included, in order to improve clarity. Accordingly, in Table 3 these viruses are called, for example, Penaeus monodon hepandensovirus 1-4, or Penaeus monodon penstyldensovirus 1-2, and are abbreviated to PmoHDV (1-4) and PmoPDV (1-2), respectively.

Notes

Acknowledgments

This work was supported in part by Public Health Service grants from the National Institutes of Health (CA029303 and AI026109), SFC and PTa; (AI046458 and AI091588), DJP; (AI070723), JQ; (GM082946), MAM; the National Science Foundation (MCB 0718948), MAM; the UK Medical Research Council, AD and DG; the Helsinki University Research Funds and Jusélius Foundation, Finland, MSV; and the Natural Sciences and Engineering Research Council of Canada, PTi.

Conflict of interest

The authors declare that they have no conflicts of interest.

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Copyright information

© Springer-Verlag Wien 2013

Authors and Affiliations

  • Susan F. Cotmore
    • 1
    Email author
  • Mavis Agbandje-McKenna
    • 2
  • John A. Chiorini
    • 3
  • Dmitry V. Mukha
    • 4
  • David J. Pintel
    • 5
  • Jianming Qiu
    • 6
  • Maria Soderlund-Venermo
    • 7
  • Peter Tattersall
    • 8
  • Peter Tijssen
    • 9
  • Derek Gatherer
    • 10
    • 11
  • Andrew J. Davison
    • 10
  1. 1.Department of Laboratory MedicineYale University School of MedicineNew HavenUSA
  2. 2.Department of Biochemistry and Molecular Biology, Center for Structural Biology, McKnight Brain Institute, College of MedicineUniversity of FloridaGainesvilleUSA
  3. 3.Molecular Physiology and Therapeutics Branch, National Institute of Dental and Craniofacial ResearchNational Institutes of HealthBethesdaUSA
  4. 4.Vavilov Institute of General Genetics, Russian Academy of SciencesMoscowRussia
  5. 5.Department of Molecular Microbiology and Immunology, Life Sciences Center, School of MedicineUniversity of Missouri-ColumbiaColumbiaUSA
  6. 6.Department of Microbiology, Molecular Genetics and ImmunologyUniversity of Kansas Medical CenterKansas CityUSA
  7. 7.Department of Virology, Haartman InstituteUniversity of HelsinkiHelsinkiFinland
  8. 8.Departments of Laboratory Medicine and GeneticsYale University School of MedicineNew HavenUSA
  9. 9.INRS-Institut Armand-FrappierLavalCanada
  10. 10.MRC, University of Glasgow Centre for Virus ResearchGlasgowUK
  11. 11.Division of Biomedical and Life SciencesLancaster UniversityLancasterUK

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