ADENOVIRUS TAXONOMY

ICTV Logo: link to ICTV Home Page

 

 

 

 

 

Official taxonomy of adenoviruses (updated after the electronic balloting of ICTV by February, 2013; changes in the species names since Report 9 are shown by red letters)

 

(Summary: short list)
 

Harrach B, Benkö M, Both GW, Brown M, Davison AJ, Echavarria M, Hess M, Jones MS, Kajon A, Lehmkuhl HD, Mautner V,
Mittal SK, Wadell G (2011) Family Adenoviridae. King AMQ, Adams MJ, Carstens EB, Lefkowitz EJ (eds) 
Virus Taxonomy: Classification and Nomenclature of Viruses. Ninth Report of the International Committee on Taxonomy of Viruses. 
Elsevier, San Diego pp 125-141 

 

Graphical representation of available adenovirus sequences.

 

  

 

Family           Adenoviridae

Genus                                Mastadenovirus

Genus                                Aviadenovirus

Genus                                Atadenovirus

Genus                                Siadenovirus

Genus                                Ichtadenovirus

 

 

 

Genome organization

 

 

Figure 1:  Schematic illustration of the various genome organizations found in members of four adenovirus genera. Black arrows depict genes conserved in every genus, grey arrows show genes present in more than one genus, and coloured arrows show genus-specific genes.

 

Genus           Mastadenovirus

Type Species               Human adenovirus C

Distinguishing features

 

Mastadenoviruses infect mammals only, and can be distinguished from members of other adenovirus genera traditionally by serology (genus members share complement-fixing antigen) and more recently (and preferably) by genome organization characteristics and phylogenetic distances. Virus infectivity is inactivated after heating at 56°C for more than 10 min. Mastadenovirus genomes fully sequenced to date range between 30,536 bp (CAdV-1) and 37,860 bp (simian adenovirus 31.2; SAdV-31.2). The G+C content of the DNA varies between 43.6% (bovine adenovirus 2; BAdV-2) and 63.9% (porcine adenovirus 3; PAdV-3). The ITRs of mastadenoviruses are considerable longer (93-371 bp) and more complex (containing a variety of cellular factor binding sites) than in members of the other genera. HAdV-2 comprises 35,937 bp and its ITR is 103 bp long.

 

Unique proteins of mastadenoviruses are proteins V and IX, and most of those coded by the E1A, E1B, E3 and E4 regions. As well as cementing the hexons on the outer surface of the capsid, protein IX also acts as a transcriptional activator and takes part in nuclear re-organization. Protein V is a core protein that, in association with cellular protein p32, seems to be involved in transport of viral DNA into the nucleus of the infected cell. The E3 and E4 proteins also often differ substantially between different mastadenovirus species.

 

Genome organization and replication have been most extensively studied for isolates of the species Human adenovirus C (Fig. 3), and the findings seem to be generally applicable to all mastadenoviruses, except in the E3 and E4 regions. These early regions are also different in the non-primate mastadenoviruses. In the E4 region, a single homologue of the HAdV-2 34K protein exists in all mastadenoviruses and is duplicated in bovine adenovirus 3 and porcine adenovirus 5. The E3 region is also considerably shorter and less complex in the non-primate mastadenoviruses. The simplest E3 region, comprising a single gene, occurs in murine adenovirus 1 (MAdV-1) and MAdV-3.

Species demarcation criteria in the genus

Species demarcation is based on evolutionary distance as reflected by phylogenetic distances and genome organizational differences. The species contain similar types (designated by Arabic numbers) that were traditionally distinguished serologically (by virus neutralization). The serological type demarcation criterion is currently being replaced by criteria similar to those used for species demarcation. Species designation depends on several of the following characteristics:

ˇ        Phylogenetic distance (>5-15%, based primarily on distance matrix analysis of the DNA polymerase amino acid sequence)

ˇ        Genome organization (characteristically in the E3 region)

ˇ        Nucleotide composition (G+C%)

ˇ        Oncogenicity in rodents

ˇ        Host range

ˇ        Cross-neutralization

ˇ        Ability to recombine

ˇ        Number of VA RNA genes

ˇ        Haemagglutination

 

For example, if virus neutralization data are available, lack of cross-neutralization combined with a phylogenetic distance of more than 15% separates two types into different species. If the phylogenetic distance is less than 5%, any additional common grouping criteria from the list above may classify separate types into the same species even if they were isolated from different hosts. As an example, the most numerous types from the same host, the human adenoviruses, can be clearly separated into 7 species supported by phylogenetic analysis, ability to recombine (e.g. between HAdV-1, 2, 5 and 6), growth characteristics (HAdV-40 and 41 show similar restricted capacity), oncogenicity and nucleotide composition (HAdV-12, 18 and 31, which are members of the species Human adenovirus A, share high oncogenicity in rodents and low G+C percentage in their genome). Adenoviruses isolated from chimpanzees resemble certain HAdVs to such an extent that they are classified into "human" adenovirus species. For example, simian adenoviruses (SAdVs) 22 to 25 belong to the species Human adenovirus E, and SAdV-21 belongs to the species Human adenovirus B.

List of species in the genus Mastadenovirus

Bat adenovirus A

 

 

     Bat adenovirus 3 (TJM)

[GU226970] = NC_016895]

(BtAdV-3)

Bat adenovirus B

 

 

     Bat adenovirus 2 (PPV1)

[FJ983127 = NC_015932]

(BtAdV-2)

Bovine adenovirus A

 

 

     Bovine adenovirus 1

[BD269513 = NC_006324]

(BAdV-1)

Bovine adenovirus B

 

 

     Bovine adenovirus 3

[AF030154 = AC_000002]

(BAdV-3)

Bovine adenovirus C

 

 

     Bovine adenovirus 10

[AF027599]

(BAdV-10)

Canine adenovirus A

 

 

     Canine adenovirus 1

[Y07760 = AC_000003]

(CAdV-1)

     Canine adenovirus 2

[U77082 = AC_000020]

(CAdV-2)

Equine adenovirus A

 

 

     Equine adenovirus 1

[JN418926]

(EAdV-1)

Equine adenovirus B

 

 

     Equine adenovirus 2

[L80007]

(EAdV-2)

Human adenovirus A

 

 

     Human adenovirus 12

[X73487 = NC_001460]

(HAdV-12)

     Human adenovirus 18

[GU191019]

(HAdV-18)

     Human adenovirus 31

[AM749299]

(HAdV-31)

Human adenovirus B

 

 

     Human adenovirus 3

[DQ086466 = NC_011203]

(HAdV-3)

     Human adenovirus 7

[AY495969 = AC_000018]

(HAdV-7)

     Human adenovirus 11

[AY163756 = NC_011202]

(HAdV-11)

     Human adenovirus 14

[AY803294]

(HAdV-14)

     Human adenovirus 16

[AY601636]

(HAdV-16)

     Human adenovirus 21

[AY601633]

(HAdV-21)

     Human adenovirus 34

[AY737797]

(HAdV-34)

     Human adenovirus 35

[AY271307]

(HAdV-35)

     Human adenovirus 50

[AJ272612]

(HAdV-50)

     Simian adenovirus 21

[AR101858 = AC_000010]

(SAdV-21)

     Simian adenovirus 27 [HC084988]

(SAdV-27)

     Simian adenovirus 28 [HC084950]

(SAdV-28)

     Simian adenovirus 29 [HC085020]

(SAdV-29)

     Simian adenovirus 32 [HC085052]

(SAdV-32)

     Simian adenovirus 33 [HC085083]

(SAdV-33)

     Simian adenovirus 35 [HC085115]

(SAdV-35)

     Simian adenovirus 41 [HI964271]

(SAdV-41)

     Simian adenovirus 46 [FJ025930]

(SAdV-46)

     Simian adenovirus 47 [FJ025929]

(SAdV-47)

Human adenovirus C

 

 

     Bovine adenovirus 9

 

(BAdV-9)

     Human adenovirus 1

[AF534906]

(HAdV-1)

     Human adenovirus 2

[J01917 = NC_001405]

(HAdV-2)

     Human adenovirus 5

[M73260 = AC_000008]

(HAdV-5)

     Human adenovirus 6

[HC492785]

(HAdV-6)

     Simian adenovirus 31

[FJ025904]

(SAdV-31)

     Simian adenovirus 34

[HC000847]

(SAdV-34)

     Simian adenovirus 40

[HC000785]

(SAdV-40)

     Simian adenovirus 42

[HC191035]

(SAdV-42)

     Simian adenovirus 43

[FJ025900]

(SAdV-43)

     Simian adenovirus 44

[HC191097]

(SAdV-44)

     Simian adenovirus 45

[HC191097]

(SAdV-45)

Human adenovirus D

 

 

     Human adenovirus 8

[AB448767]

(HAdV-8)

     Human adenovirus 9

[AJ854486 = NC_010956]

(HAdV-9)

     Human adenovirus 10

[DQ149615]

(HAdV-10)

     Human adenovirus 13

[DQ149616]

(HAdV-13)

     Human adenovirus 15

[AB562586]

(HAdV-15)

     Human adenovirus 17

[AF108105 = AC_000006]

(HAdV-17)

     Human adenovirus 19

[EF121005]

(HAdV-19)

     Human adenovirus 20

[DQ149619]

(HAdV-20)

     Human adenovirus 22

[FJ404771]

(HAdV-22)

     Human adenovirus 23

[DQ149621]

(HAdV-23)

     Human adenovirus 24

[DQ149622]

(HAdV-24)

     Human adenovirus 25

[DQ149623]

(HAdV-25)

     Human adenovirus 26

[EF153474]

(HAdV-26)

     Human adenovirus 27

[DQ149625]

(HAdV-27)

     Human adenovirus 28

[FJ824826]

(HAdV-28)

     Human adenovirus 29

[AB562587]

(HAdV-29)

     Human adenovirus 30

[DQ149628]

(HAdV-30)

     Human adenovirus 32

[DQ149629]

(HAdV-32)

     Human adenovirus 33

[DQ149630]

(HAdV-33)

     Human adenovirus 36

[GQ384080]

(HAdV-36)

     Human adenovirus 37

[DQ900900]

(HAdV-37)

     Human adenovirus 38

[DQ149633]

(HAdV-38)

     Human adenovirus 39

[DQ149634]

(HAdV-39)

     Human adenovirus 42

[DQ149635]

(HAdV-42)

     Human adenovirus 43

[DQ149636]

(HAdV-43)

     Human adenovirus 44

[DQ149637]

(HAdV-44)

     Human adenovirus 45

[DQ149638]

(HAdV-45)

     Human adenovirus 46

[AY875648]

(HAdV-46)

     Human adenovirus 47

[DQ149640]

(HAdV-47)

     Human adenovirus 48

[EF153473]

(HAdV-48)

     Human adenovirus 49

[DQ393829]

(HAdV-49)

     Human adenovirus 51

[DQ149642]

(HAdV-51)

     Human adenovirus 53

[FJ169625]

(HAdV-53)

     Human adenovirus 54

[AB333801 = NC_012959]

(HAdV-54)

Human adenovirus E

 

 

     Human adenovirus 4

[AY487947 = NC_003266]

(HAdV-4)

     Simian adenovirus 22

[AY530876]

(SAdV-22)

     Simian adenovirus 23

[AY530877]

(SAdV-23)

     Simian adenovirus 24

[AY530878]

(SAdV-24)

     Simian adenovirus 25

[AR101859 = AC_000011]

(SAdV-25)

     Simian adenovirus 26

[HB426768]

(SAdV-26)

     Simian adenovirus 30

[HB426704]

(SAdV-30)

     Simian adenovirus 36

[HB426704]

(SAdV-36)

     Simian adenovirus 37

[HB426704]

(SAdV-37)

     Simian adenovirus 38

[HB426671]

(SAdV-38)

     Simian adenovirus 39

[HB426607]

(SAdV-39)

Human adenovirus F

 

 

     Human adenovirus 40

[L19443 = NC_001454]

(HAdV-40)

     Human adenovirus 41

[DQ315364]

(HAdV-41)

Human adenovirus G

 

 

     Human adenovirus 52

[DQ923122]

(HAdV-52)

     Simian adenovirus 1

[AY771780 = NC_006879]

(SAdV-1)

     Simian adenovirus 2

[SAU03008]

(SAdV-2)

     Simian adenovirus 7

[DQ792570]

(SAdV-7)

     Simian adenovirus 11

[SAU03014]

(SAdV-11)

     Simian adenovirus 12

 

(SAdV-12)

     Simian adenovirus 15

[SAU03016]

(SAdV-15)

Murine adenovirus A

 

 

     Murine adenovirus 1

[NC_000942]

(MAdV-1)

Murine adenovirus B

 

 

     Murine adenovirus 2

[HM049560 = NC_014899]

(MAdV-2)

Murine adenovirus C

 

 

     Murine adenovirus 3

[EU835513 = NC_012584]

(MAdV-3)

Ovine adenovirus A

 

 

     Bovine adenovirus 2

[AF252854 = AC_000001]

(BAdV-2)

     Ovine adenovirus 2

[DQ630755]

(OAdV-2)

     Ovine adenovirus 3

[DQ630756]

(OAdV-3)

     Ovine adenovirus 4

[DQ630757]

(OAdV-4)

     Ovine adenovirus 5

[DQ630758]

(OAdV-5)

Ovine adenovirus B

 

 

     Goat adenovirus 2

[DQ630760]

(GAdV-2)

 

     Ovine adenovirus 1

[DQ630754]

(OAdV-1)

Porcine adenovirus A

 

 

     Porcine adenovirus 1

[L43364]

(PAdV-1)

     Porcine adenovirus 2

[L43365]

(PAdV-2)

     Porcine adenovirus 3

[AF083132 = NC_005869]

(PAdV-3)

Porcine adenovirus B

 

 

     Porcine adenovirus 4

[U13893]

(PAdV-4)

Porcine adenovirus C

 

 

     Porcine adenovirus 5

[AF289262 = NC_002702]

(PAdV-5)

Simian adenovirus A

 

 

     Simian adenovirus 3

[AY598782 = NC_006144]

(SAdV-3)

     Simian adenovirus 4

[SAU03010]

(SAdV-4)

     Simian adenovirus 6

[CQ982401]

(SAdV-6)

     Simian adenovirus 9

[SAU03012]

(SAdV-9)

     Simian adenovirus 10

[SAU03013]

(SAdV-10)

     Simian adenovirus 14

[SAU03016]

(SAdV-14)

     Simian adenovirus 48

[HQ241818]

(SAdV-48)

Tree shrew adenovirus A

 

 

     Tree shrew adenovirus 1

[AF258784 = NC_004453]

(TSAdV-1)

 

Species names are in italic script; names of types are in roman script. Sequence accessions [ ], and assigned abbreviations ( ) are also listed.

 

List of other related viruses which may be members of the genus Mastadenovirus but have not been approved as species

 

Alpaca adenovirus 1

[GQ499375]

(AlAdV-1)

Bat adenovirus 1 (FBV1)

[AB303301]

(BtAdV-1)

Guinea pig adenovirus 1

[X95630]

(GPAdV-1)

Ovine adenovirus 6

[DQ630759]

(OAdV-6)

Simian adenovirus 5, 8, 13, 16-20

[18: FJ025931]

(SAdV-5, 8, 13, 16-20)

Squirrel adenovirus 1

[GU735084]

(SqAdV-1)

 

  

Genus           Aviadenovirus

Type Species               Fowl adenovirus A

Distinguishing features

Aviadenoviruses are serologically distinct from members of the other adenovirus genera and they only infect birds. The virions contain two fibers per vertex. Fowl adenovirus 1 (FAdV-1), FAdV-4 and turkey adenovirus 1 (TAdV-1) have two fiber genes, and two projections (in case of FAdV-1, of considerably different lengths) on each penton base. Other FAdVs also have two fibers per vertex, but apparently only one fiber gene, and the fiber shafts are of similar lengths. The long fiber of FAdV-1 uses the coxsackievirus and adenovirus receptor (CAR) for attachment to the cell.

Aviadenovirus genomes are considerably larger (20-45%) than those of mastadenoviruses. Five aviadenovirus [FAdV-1, FAdV-4, "FAdV-8", FAdV-9 and TAdV-1] genomes have been fully sequenced, and range between 43,804 bp (FAdV-1) and 45,667 (FAdV-4). These are thought to represent the longest adenovirus DNA molecules after that of white sturgeon adenovirus. The G+C content of partial or complete sequences of aviadenovirus genomes varies between 53.8 and 66.9%. The size of sequenced aviadenovirus ITRs are between 54 and 95 bp long. The genomic organization of aviadenoviruses is also different from that of adenoviruses in other genera (Figure 1). The genes of proteins V and IX, as well as genes in mastadenovirus early regions E1 and E3, are missing. The E4 region may be translocated from its position in mastadenovirus genomes, resulting in the gene encoding dUTP pyrophosphatase (dUTPase, not present in every mastadenovirus) being on the left end of the genome, rather than the right end. (Alternatively, this gene may have been captured independently by ancestors of aviadenoviruses and mastadenoviruses.) The organization of the central part of the genome containing the late genes and the E2 region is similar to that of mastadenoviruses. The right end of the genome contains several transcription units, which are unique to aviadenoviruses. The majority of genes and proteins from this region have not yet been characterized in detail. A novel protein GAM-1 of FAdV-1 has been demonstrated to have an anti-apoptotic effect, and to activate the heat-shock response in the infected cell. GAM-1, in synergy with another novel protein encoded by ORF22, binds the retinoblastoma protein and can activate the E2F pathway. Additional, and as yet uncharacterized predicted gene products, exhibit sequence homology to proteins of other viruses, such as the non-structural protein NS1 (also known as Rep) of parvoviruses, and a triacylglycerol lipase, a homolog of which also occurs in an avian herpesvirus (Marek's disease virus). Aviadenoviruses possess no complement-fixing antigen in common with the members of the other genera. There are isolates where serum neutralization cannot differentiate clearly between the serotypes. The introduction of FAdV-8a and 8b was deemed necessary because of the inconsistency in the type-numbering scheme used in different countries and continents over the years. Avian adenoviruses have been associated with diverse disease patterns, including inclusion body hepatitis, bronchitis, pulmonary congestion and oedema in different bird species. Hydropericardium syndrome is caused by FAdV-4 in chickens, mainly in Asia. Falcon adenovirus 1 has caused fatalities in different falcon species. FAdV-1 (CELO virus), 9 and 10 are being studied for their feasibility as gene delivery vectors.

Species demarcation criteria in the genus

Species designation depends on several of the following characteristics:

ˇ        Phylogenetic distance (>5-15%, based primarily on distance matrix analysis of the DNA polymerase amino acid sequence)

ˇ        Genome organization (characteristically in the region at the right end of the genome)

ˇ        RFLP analysis

ˇ        Host range

ˇ        Pathogenicity

ˇ        Cross-neutralization

ˇ        Ability to recombine

For example, the fowl adenovirus serotypes can be grouped into five species on the basis of phylogeny, genome organization, RFLP profiles and the lack of significant cross-neutralization.

List of species in the genus Aviadenovirus

Note:  A specific problem that has been addressed but not resolved is the lack of consensus in the numbering of the individual fowl adenovirus serotypes. Strains deposited in the American Type Culture Collection are numbered inconsistently in relation to the majority of newer publications. For this reason, one representative strain of each serotype is also listed (in parentheses).

 

 

Falcon adenovirus A

 

 

     Falcon adenovirus 1

[AY683541]

(FaAdV-1)

Fowl adenovirus A

 

 

     Fowl adenovirus 1 (CELO)

[U46933 = AC_000014]

(FAdV-1)

Fowl adenovirus B

 

 

     Fowl adenovirus 5 (340)

[AF508952]

(FAdV-5)

Fowl adenovirus C

 

 

     Fowl adenovirus 4 (ON1)

[GU188428 = NC_015323]

(FAdV-4)

     Fowl adenovirus 10 (CFA20)

[AF160185]

(FAdV-10)

Fowl adenovirus D

 

 

     Fowl adenovirus 2 (P7-A)

[AF339915]

(FAdV-2)

     Fowl adenovirus 3 (75)

[AF508949]

(FAdV-3)

     Fowl adenovirus 9 (A2-A)

[AF083975 = AC_000013

(FAdV-9)

     Fowl adenovirus 11 (380)

[AF339925]

(FAdV-11)

Fowl adenovirus E

 

 

     Fowl adenovirus 6 (CR119)

[AF508954]

(FAdV-6)

     Fowl adenovirus 7 (YR36)

[AF508955]

(FAdV-7)

     Fowl adenovirus 8a (CFA40)

[AF155911]

(FAdV-8a)

     Fowl adenovirus 8b (764)

[AF508958]

(FAdV-8b)

Goose adenovirus A

 

 

     Goose adenovirus 1

 

(GoAdV-1)

Turkey adenovirus B

 

 

     Turkey adenovirus 1

[GU936707 =  NC_014564]

(TAdV-1)

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Species names are in italic script; names of types and isolates ( ) are in roman script. Sequence accessions [ ], and assigned abbreviations ( ) are also listed.

 

List of other related viruses which may be members of the genus Aviadenovirus but have not been approved as species

Duck adenovirus 2

 

(DAdV-2)

Meyer's parrot adenovirus 1

[AY644731]

 

Pigeon adenovirus 1

[FN824512]

(PiAdV-1)

Psittacine adenovirus 1

[EF442329]

(PsAdV-1)

Turkey adenovirus 2

[GU936708]

(TAdV-2)

 

  

Genus           Atadenovirus

Type Species               Ovine adenovirus D

Distinguishing features

Atadenoviruses are serologically distinct from viruses in the other adenovirus genera, and their genomic organization and capsid protein complement also differ. Atadenoviruses have been detected in a broad range of hosts, including scaled reptiles (order Squamata) and species ranging from birds to ruminants and a marsupial. Virions are relatively heat stable and retain substantial infectivity after treatment for 30 min at 56 °C, conditions that inactivate mastadenovirions. The genome size of sequenced isolates ranges from 29,576 (ovine adenovirus 7, OAdV-7) to 33,213 bp (duck adenovirus 1, DAdV-1) with ITRs of 46 (OAdV-7) to 118 bp (snake adenovirus 1, SnAdV-1). For ruminant, marsupial and avian atadenoviruses, the G+C content of the DNA is low and varies between 33.6 (OAdV-7) and 43.0% (DAdV-1). The corresponding high A+T content was deemed sufficiently characteristic to be used as the basis of the name of the genus, though atadenoviruses originating from scaled reptiles have a non-biased nucleotide composition (50.0% G+C in SnAdV-1). The proteins encoded by an atadenovirus genome are summarized in Table 2. The central part of the atadenovirus genome is similar to that of mastadenoviruses (except that there are no protein V and IX genes), whereas the extremities differ markedly. Atadenoviruses have several unique proteins, including some that may be distant homologs of proteins in other adenovirus genera. The left end of the genome carries a gene for p32K, a unique structural protein. At this end, gene LH1 is also unique to the genus but not present in all members. The right end of the genome contains genes that are related to each other, suggestive of gene duplication. There are two E4 34K gene homologs (E4.2 and E4.3), and one to five gene RH homologs. At this end, genes E4.1 and RH1-6 are also unique to the genus but are not present in all members. The proteins encoded by genes LH3 and E4.3 (and its paralog, E4.2) show limited similarity to mastadenovirus proteins E1B 55K and E4 34K, respectively. However, LH3 is a structural protein that forms prominent "knobs" on the virion surface that distinguish atadenoviruses structurally from other known AdVs. LH3 is located in the same relative position among the hexon subunits as protein IX (present only in mastadenoviruses) but sits on top of the "central" (so called H3) hexon trimers, appearing to hold the outer capsid together. Particles that lack the LH3 or p32K proteins are viable although less stable. (Mastadenoviruses that lack proteins V or IX are also viable.) No immunomodulatory genes such as those found in the mastadenovirus E3 region have yet been identified. DAdV-1 has a unique genome region at the far right end with seven uncharacterized ORFs that may be host-specific in function. ORF5 and ORF6 are related to each other. This unique region of DAdV-1 also contains a VA RNA gene that is supposedly homologous to that of FAdV-1. 

Certain atadenoviruses can cause hemorrhagic epizooty in free-living ruminants. DAdV-1 is also associated with a specific disease of hens that is characterized globally by sharp decreases in egg production (egg drop syndrome). Due to the lack of pre-existing immunity in humans and its bio-safety profile, OAdV-7 has been developed as a gene delivery vector intended for human vaccine and gene therapy applications.

Species demarcation criteria in the genus

Species designation depends on several of the following characteristics:

ˇ        Phylogenetic distance (>5-15%, based primarily on distance matrix analysis of the DNA polymerase amino acid sequence)

ˇ        Host range

ˇ        DNA hybridization

ˇ        Nucleotide composition

ˇ        Cross-neutralization

ˇ        Gene organization of the right end of the genome

List of species in the genus Atadenovirus

Bovine adenovirus D

 

 

     Bovine adenovirus 4

[AF036092 = NC_002685]

(BAdV-4)

     Bovine adenovirus 5

[AF207658]

(BAdV-5)

     Bovine adenovirus 8

[AF238233]

(BAdV-8)

     Bovine adenovirus strain Rus

[AF238880]

(BAdV-Rus)

Duck adenovirus A

 

 

     Duck adenovirus 1

[Y09598 = AC_000004]

(DAdV-1)

Ovine adenovirus D

 

 

     Goat adenovirus 1

[AF207660]

(GAdV-1)

     Ovine adenovirus 7

[U40839 = NC_004037]

(OAdV-7)

Possum adenovirus A

 

 

     Possum adenovirus 1

[AF338822]

(PoAdV-1)

Snake adenovirus A

 

 

     Snake adenovirus 1

[DQ106414 = NC_009989]

(SnAdV-1)

 

Species names are in italic script; names of types and isolates are in roman script. Sequence accessions [ ], and assigned abbreviations ( ) are also listed.

List of other related viruses which may be members of the genus Atadenovirus but have not been approved as species

Asp viper adenovirus

[EU914209]

 

Bearded dragon adenovirus 1

(agamid adenovirus 1)

[AY576678]

(BDAdV-1)

Blue-tongued skink adenovirus

[AY576682]

 

Bovine adenovirus 6

[AF207659]

(BAdV-6)

Bovine adenovirus 7

[AF238232]

(BAdV-7)

Chameleon adenovirus 1

[AY576679]

(ChAdV-1)

Emerald monitor adenovirus

[EU914208]

 

Gecko adenovirus 1

[AY576677]

(GeAdV-1)

Gila adenovirus 1

[AY576680]

 

Hagen's Pit-viper adenovirus

[EU851415]

 

Mexican beaded lizard adenovirus 1

[EU914207]

 

Odocoileus adenovirus 1

[AF198354]

(OdAdV-1)

Snake adenovirus 2

[FJ012163]

(SnAdV-2)

Snake adenovirus 3

[FJ012164]

(SnAdV-3)

Tokay gecko adenovirus

[AY576681]

 

 

  

Genus           Siadenovirus

Type Species               Frog adenovirus

Distinguishing features

Siadenoviruses are serologically and phylogenetically distinct from members of the other adenovirus genera. This genus comprises only three accepted members, frog adenovirus 1 (FrAdV-1), turkey adenovirus 3 (TAdV-3) and raptor adenovirus 1 (RAdV-1). FrAdV-1 was isolated from an amphibian (frog) and TAdV-3 from birds (turkey, pheasant and chicken). RAdV-1 was detected in captive raptors by PCR. The RAdV-1 genome was the first adenovirus genome to be fully sequenced solely by PCR-based methods without virus isolation. The genomes of the three siadenovirus types represent the shortest adenovirus genomes known to date. Their lengths are between 26,163 and 26,282 bp, with G+C contents of 34.9 to 38.5%, and ITRs of 29-39 bp. The genomic organization of siadenoviruses is also characteristic and different from that of other genera. The genes of proteins V and IX, and homologues in mastadenovirus early regions E1, E3 and E4 are absent. Apart from the proteins conserved in all adenoviruses, there are only five ORFs potentially encoding novel proteins. At the left end of the genome, the first putative gene encodes a protein that is related to sialidases. Adjacent to it is a novel ORF predicted to code for a highly hydrophobic protein. The gene named "E3" solely because of its position between the pVIII and fiber genes is not homologous to any of the mastadenovirus E3 genes (or to any other known genes). The right end of the genome harbours ORF7 and ORF8. TAdV-3 has no common complement-fixing antigen with other adenoviruses isolated from birds and classified in the aviadenovirus or atadenovirus genera. FrAdV-1 is supposedly non-pathogenic. TAdV-3 is associated with specific diseases in different hosts (haemorrhagic enteritis in turkey, marble spleen disease in pheasants and splenomegaly in chickens).

 

List of species in the genus Siadenovirus

 

Frog adenovirus

 

 

     Frog adenovirus 1

[AF224336 = NC_002501]

(FrAdV-1)

Great tit adenovirus A

 

 

     Great tit adenovirus 1

[FJ849795]

(GTAdV-1)

Raptor adenovirus A

 

 

     Raptor adenovirus 1

[EU715130 = NC_015455]

(RAdV-1)

Skua adenovirus A

 

 

     South Polar skua adenovirus 1 [HM585353 = NC_016437]

(SPAdV-1)

Turkey adenovirus A

 

 

     Turkey adenovirus 3

[AF074946 = AC_000016]

(TAdV-3)

 

Species names are in italic script; names of types are in roman script. Sequence accessions [ ], and assigned abbreviations ( ) are also listed.

List of other related viruses which may be members of the genus Siadenovirus but have not been approved as species

Budgerigar adenovirus 1

[AB485763]

 

Psittacine adenovirus 2

[EU056825]

(PsAdV-2)

Sulawesi tortoise adenovirus 1

[EU056826]

(STAdV-1)

 

Genus           iCHTadenovirus

Type Species               Sturgeon adenovirus A

Distinguishing features

The single known member of this genus, white sturgeon adenovirus 1 (WSAdV-1), is the only confirmed fish adenovirus. While the host is very divergent from those of other AdVs, phylogenetic calculations and a unique genome organization further distinguish this virus from all other adenoviruses. The WSAdV-1 genome is 48,395 bp, and thus it is longer than the genome of any known adenovirus. The G+C content is 42.7%. The fiber gene was not found in its usual position towards the right end of the genome, but fiber gene homologues were discovered at the left end. The virus seems to be non-pathogenic for fish.

 List of species in the genus Ichtadenovirus

 

Sturgeon adenovirus A

 

 

     White sturgeon adenovirus 1

[AJ495768]

(WSAdV-1)

 

Other related viruses which may be members of the genus but have not been approved as species

None

 

 

Figure 2 Phylogenetic tree of adenoviruses based on a distance matrix analysis of hexon amino acid sequences. Primate adenovirus hexons known to have resulted from homologous recombination were excluded from the analysis. Only selected members of primate adenovirus species with more than three members are shown (boxed in grey). The Seqboot (bootstrap), Protdist (categories matrix), Fitch (global rearrangements), and Consense programs of the PHYLIP 3.68 package were used. The tree was gained by unrooted calculation, and is shown with white sturgeon adenovirus 1 chosen as outgroup. Species names are indicated, but for reasons of presentation the word "adenovirus" is abbreviated to AdV followed by a hyphen. Abbreviations after the type numbers of simian adenoviruses: bo, bonobo; ch, chimpanzee; go, gorilla; cy, cynomolgus macaque; rh, rhesus macaque. Bootstrap values higher than 50 (from 100 re-samplings) are shown for every confirmed branching.

 

Derivation erivation of names

Adeno: from Greek aden, adenos, "gland"; in recognition of the fact that adenoviruses were first isolated from human adenoid tissue.

At: from English adenine and thymine, in recognition that the genome of the first recognized members of the genus (from ruminant, avian and marsupial hosts) has a remarkably high AT content.

Avi: from Latin avis, "bird".

Icht:truncated from Greek ichthys, "fish".

Mast: from Greek mastos, "breast".

Si: from English sialidase, in recognition that members of the genus have a putative sialidase homolog.

 

Further reading

 

Benkő, M. (2008). Adenoviruses. Pathogenesis. In: Mahy, B.W.J., van Regenmortel, M.H.V. (Eds.), Encyclopedia of Virology. Third edition. Elsevier, Oxford, vol. 1, pp. 24-29.

Corredor, J.C., Garceac, A., Krell, P.J., Nagy, É. (2008). Sequence comparison of the right end of fowl adenovirus genomes. Virus Genes, 36, 331-344.

Davison, A.J., Benkő, M., Harrach, B. (2003). Genetic content and evolution of adenoviruses. J. Gen. Virol., 84, 2895-2908.

Farkas, S.L., Harrach, B., Benkő, M. (2008). Completion of the genome analysis of snake adenovirus type 1, a representative of the reptilian lineage within the novel genus Atadenovirus. Virus Res., 132, 132-139.

Harrach, B. (2008). Adenoviruses. General features. In: Mahy, B.W.J., van Regenmortel, M.H.V. (Eds.), Encyclopedia of Virology. Third Edition. Elsevier, Oxford vol. 1, pp. 1-9.

Hemmi, S., Vidovszky, M.Z., Ruminska, J., Ramelli, S., Decurtins, W., Greber, U.F., Harrach, B. (2011) Virus Res. 160, 128-135.

Jones, M.S. 2nd, Harrach, B., Ganac, R.D., Gozum, M.M., Dela Cruz, W.P., Riedel, B., Pan, C., Delwart, E.L., Schnurr, D.P. (2007). New adenovirus species found in a patient presenting with gastroenteritis. J. Virol., 81, 5978-5984.

Kaján, G.L., Davison, A.J., Palya, V., Harrach, B., Benkő, M. (2012) Genome sequence of a waterfowl aviadenovirus, goose adenovirus 4. J. Gen. Virol. 93, (11) 2457-2465

Kaján, G.L., Stefancsik, R., Ursu, K., Palya, V., Benkő, M. (2010) The first complete genome sequence of a non-chicken aviadenovirus, proposed to be turkey adenovirus 1. Virus Res. 153, 226-233.

Kohl, C., Vidovszky, M.Z., Mühldorfer, K., Dabrowski, P.W., Radonic, A., Nitsche, A., Wibbelt, G., Kurth, A., Harrach, B. (2012) Genome analysis of bat adenovirus 2: indications of interspecies transmission. J. Virol. 86, 1888-1892.

Kovács, E.R., Benkő, M. (2011) Complete sequence of raptor adenovirus 1 confirms the characteristic genome organization of siadenoviruses. Infect. Genet. Evol. 11, 1058-1065.

Kovács, E.R., Jánoska, M., Dán, Á., Harrach, B., Benkő, M. (2010). Recognition and partial genome characterization by non-specific DNA amplification and PCR of a new siadenovirus species in a sample originating from Parus major, a great tit. J. Virol. Methods, 163, 262-268.

Marek, A., Nolte, V., Schachner, A., Berger, E., Schlötterer, C., Hess, M. (2012) Two fiber genes of nearly equal lengths are a common and distinctive feature of Fowl adenovirus C members. Vet. Microbiol. 156, 411-417.

Pantelic, R.S., Lockett, L.J., Rothnagel, R., Hankamer, B., Both, G.W. (2008). Cryoelectron microscopy map of Atadenovirus reveals cross-genus structural differences from human adenovirus. J. Virol., 82, 7346-7356.

Park, Y.M., Kim, J.H., Gu, S.H., Lee, S.Y., Lee, M.G., Kang,Y.K., Kang, S.H., Kim, H.J., Song, J.W. (2012) Full genome analysis of a novel adenovirus from the South Polar skua (Catharacta maccormicki) in Antarctica. Virology 422, 144-150.

Roy, S., Vandenberghe, L.H., Kryazhimskiy, S., Grant, R., Calcedo, R., Yuan, X., Keough, M., Sandhu, A., Wang, Q., Medina-Jaszek, C.A., Plotkin, J.B., Wilson, J.M. (2009). Isolation and characterization of adenoviruses persistently shed from the gastrointestinal tract of non-human primates. PLoS Pathog., 5, e1000503.

Wellehan, J.F.X., Johnson, A.J., Harrach, B., Benkő, M., Pessier, A.P., Johnson, C.M., Garner, M.M., Childress, A., Jacobson, E.R. (2004). Detection and analysis of six lizard adenoviruses by consensus primer PCR provides further evidence of a reptilian origin for the atadenoviruses. J. Virol., 78, 13366-13369.

 

Last update by Balazs Harrach on March 2, 2013.