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| image_caption =[[Transmission electron micrograph]] of ranaviruses (dark hexagons) gathering at the cell border and leaving the cell via a process called "[[budding]]".
| image_caption =[[Transmission electron micrograph]] of ranaviruses (dark hexagons) gathering at the cell border and leaving the cell via a process called "[[budding]]".
| taxon = Ranavirus
| taxon = Ranavirus
⚫
| type_species = ''[[Frog virus 3]]''
| subdivision_ranks = Species
| subdivision =
*''[[Epizootic haematopoietic necrosis virus]]''
⚫
*''[[Frog virus 3]]''
}}
}}
[[File:CSIRO ScienceImage 2315 Ranaviruses.jpg|thumb|Transmission electron micrograph of a cell infected with ranaviruses, which gather in the [[cytoplasm]] and in the assembly bodies next to the contorted nucleus.]]
[[File:CSIRO ScienceImage 2315 Ranaviruses.jpg|thumb|Transmission electron micrograph of a cell infected with ranaviruses, which gather in the [[cytoplasm]] and in the assembly bodies next to the contorted nucleus.]]


'''''Ranavirus''''' is a genus of [[viruses]], in the family ''[[Iridoviridae]]''.<ref name=ICTVReport>{{cite web|title=Iridoviridae|url=https://talk.ictvonline.org/ictv-reports/ictv_online_report/dsdna-viruses/w/iridoviridae|website=ICTV Online (10th) Report}}</ref> There are four other [[genus|genera]] of [[virus]]es within the family ''[[Iridoviridae]]'', but ''Ranavirus'' is the only one that includes viruses that are infectious to amphibians and reptiles. Additionally, it is one of the three genera within this family which infect [[teleostei|teleost]] [[fish]]es, along with ''[[Lymphocystivirus]]'' and ''[[Megalocytivirus]]''.<ref name=Whittington>{{cite journal | last1 = Whittington | first1 = RJ | last2 = Becker | first2 = JA | last3 = Dennis | first3 = MM | title = Iridovirus infections in finfish – critical review with emphasis on ranaviruses | journal = Journal of Fish Diseases | volume = 33 | issue = 2 | pages = 95–122 | year = 2010 | pmid = 20050967 | doi = 10.1111/j.1365-2761.2009.01110.x}}</ref> The family ''Iridoviridae'' is one of the five families of [[nucleocytoplasmic large DNA viruses]].
'''''Ranavirus''''' is a genus of [[viruses]], in the family ''[[Iridoviridae]]''.<ref name=ICTVReport>{{cite web|title=Iridoviridae|url=https://talk.ictvonline.org/ictv-reports/ictv_online_report/dsdna-viruses/w/iridoviridae|website=ICTV Online (10th) Report}}</ref> There are five other [[genus|genera]] of [[virus]]es within the family ''[[Iridoviridae]]'', but ''Ranavirus'' is the only one that includes viruses that are infectious to amphibians and reptiles. Additionally, it is one of the three genera within this family which infect [[teleostei|teleost]] [[fish]]es, along with ''[[Lymphocystivirus]]'' and ''[[Megalocytivirus]]''.<ref name=Whittington>{{cite journal | last1 = Whittington | first1 = RJ | last2 = Becker | first2 = JA | last3 = Dennis | first3 = MM | title = Iridovirus infections in finfish – critical review with emphasis on ranaviruses | journal = Journal of Fish Diseases | volume = 33 | issue = 2 | pages = 95–122 | year = 2010 | pmid = 20050967 | doi = 10.1111/j.1365-2761.2009.01110.x}}</ref> The family ''Iridoviridae'' is one of the five families of [[nucleocytoplasmic large DNA viruses]].


==Ecological impact==
==Ecological impact==
The Ranaviruses, like the Megalocytiviruses, are an [[emerging infectious disease|emerging]] group of closely related [[DNA|dsDNA]] viruses which cause [[systemic disease|systemic infections]] in a wide variety of wild and cultured fresh and saltwater fishes. As with Megalocytiviruses, ''Ranavirus'' outbreaks are therefore of considerable economic importance in [[aquaculture]], as [[epizootic]]s can result in moderate fish loss or mass mortality events of cultured fishes. Unlike Megalocytiviruses, however, ''Ranavirus'' infections in amphibians have been implicated as a contributing factor in the global decline of amphibian populations.<ref>{{Cite journal|last=Teacher|first=A. G. F.|last2=Cunningham|first2=A. A.|last3=Garner|first3=T. W. J.|date=2010-06-10|title=Assessing the long-term impact of Ranavirus infection in wild common frog populations: Impact of Ranavirus on wild frog populations|journal=Animal Conservation|volume=13|issue=5|pages=514–522|doi=10.1111/j.1469-1795.2010.00373.x}}</ref><ref>{{Cite journal|last=Price|first=Stephen J.|last2=Garner|first2=Trenton W.J.|last3=Nichols|first3=Richard A.|authorlink4=Francois Balloux|last4=Balloux|first4=François|last5=Ayres|first5=César|last6=Mora-Cabello de Alba|first6=Amparo|last7=Bosch|first7=Jaime|date=November 2014|title=Collapse of Amphibian Communities Due to an Introduced Ranavirus|journal=Current Biology|volume=24|issue=21|pages=2586–2591|doi=10.1016/j.cub.2014.09.028|pmid=25438946|doi-access=free}}</ref> The impact of Ranaviruses on amphibian populations has been compared to the [[chytrid]] [[fungus]] ''[[Batrachochytrium dendrobatidis]]'', the causative agent of [[chytridiomycosis]].<ref>{{cite journal | doi=10.1016/j.virol.2003.08.001 | title=Genomic sequence of a ranavirus (family Iridoviridae) associated with salamander mortalities in North America | year=2003 | journal=Virology | volume=316 | pages=90–103 | pmid=14599794 |first1=James K |last1=Jancovich |first2=Jinghe |last2=Mao |first3=V.Gregory |last3=Chinchar |first4=Christopher |last4=Wyatt |first5=Steven T |last5=Case |first6=Sudhir |last6=Kumar |first7=Graziela |last7=Valente |first8=Sankar |last8=Subramanian |first9=Elizabeth W |last9=Davidson | first10=James P |last10=Collins |first11=Bertram L |last11=Jacobs | issue=1 }}</ref><ref>{{cite journal|doi=10.1890/02-0374|title=Intraspecific Reservoirs: Complex Life History and the Persistence of a Lethal Ranavirus|journal=Ecology|volume=85|issue=2|pages=560|year=2004|last1=Brunner|first1=Jesse L.|last2=Schock|first2=Danna M.|last3=Davidson|first3=Elizabeth W.|last4=Collins|first4=James P.}}</ref><ref>{{cite journal|doi = 10.1111/j.1461-0248.2005.00735.x|title = Susceptibility of Italian agile frog populations to an emerging strain of Ranavirus parallels population genetic diversity|year = 2005|author = Pearman, Peter B.|journal = Ecology Letters|volume = 8|issue = 4|pages = 401|last2 = Garner|first2 = Trenton W. J.}}</ref> In the UK, the severity of disease outbreaks is thought to have increased due to climate change.<ref>{{Cite journal|last=Price|first=Stephen J.|last2=Leung|first2=William T. M.|last3=Owen|first3=Christopher J.|last4=Puschendorf|first4=Robert|last5=Sergeant|first5=Chris|last6=Cunningham|first6=Andrew A.|last7=Balloux|first7=Francois|last8=Garner|first8=Trenton W. J.|last9=Nichols|first9=Richard A.|date=2019-05-09|title=Effects of historic and projected climate change on the range and impacts of an emerging wildlife disease|journal=Global Change Biology|volume=25|issue=8|pages=2648–2660|doi=10.1111/gcb.14651|issn=1354-1013|hdl=10026.1/13802|hdl-access=free}}</ref>
The Ranaviruses, like the Megalocytiviruses, are an [[emerging infectious disease|emerging]] group of closely related [[DNA|dsDNA]] viruses which cause [[systemic disease|systemic infections]] in a wide variety of wild and cultured fresh and saltwater fishes. As with Megalocytiviruses, ''Ranavirus'' outbreaks are therefore of considerable economic importance in [[aquaculture]], as [[epizootic]]s can result in moderate fish loss or mass mortality events of cultured fishes. Unlike Megalocytiviruses, however, ''Ranavirus'' infections in amphibians have been implicated as a contributing factor in the global decline of amphibian populations.<ref>{{Cite journal|last=Teacher|first=A. G. F.|last2=Cunningham|first2=A. A.|last3=Garner|first3=T. W. J.|date=2010-06-10|title=Assessing the long-term impact of Ranavirus infection in wild common frog populations: Impact of Ranavirus on wild frog populations|journal=Animal Conservation|volume=13|issue=5|pages=514–522|doi=10.1111/j.1469-1795.2010.00373.x}}</ref><ref>{{Cite journal|last=Price|first=Stephen J.|last2=Garner|first2=Trenton W.J.|last3=Nichols|first3=Richard A.|authorlink4=Francois Balloux|last4=Balloux|first4=François|last5=Ayres|first5=César|last6=Mora-Cabello de Alba|first6=Amparo|last7=Bosch|first7=Jaime|date=November 2014|title=Collapse of Amphibian Communities Due to an Introduced Ranavirus|journal=Current Biology|volume=24|issue=21|pages=2586–2591|doi=10.1016/j.cub.2014.09.028|pmid=25438946|doi-access=free}}</ref> The impact of Ranaviruses on amphibian populations has been compared to the [[chytrid]] [[fungus]] ''[[Batrachochytrium dendrobatidis]]'', the causative agent of [[chytridiomycosis]].<ref>{{cite journal | doi=10.1016/j.virol.2003.08.001 | title=Genomic sequence of a ranavirus (family Iridoviridae) associated with salamander mortalities in North America | year=2003 | journal=Virology | volume=316 | pages=90–103 | pmid=14599794 |first1=James K |last1=Jancovich |first2=Jinghe |last2=Mao |first3=V.Gregory |last3=Chinchar |first4=Christopher |last4=Wyatt |first5=Steven T |last5=Case |first6=Sudhir |last6=Kumar |first7=Graziela |last7=Valente |first8=Sankar |last8=Subramanian |first9=Elizabeth W |last9=Davidson | first10=James P |last10=Collins |first11=Bertram L |last11=Jacobs | issue=1 }}</ref><ref>{{cite journal|doi=10.1890/02-0374|title=Intraspecific Reservoirs: Complex Life History and the Persistence of a Lethal Ranavirus|journal=Ecology|volume=85|issue=2|pages=560|year=2004|last1=Brunner|first1=Jesse L.|last2=Schock|first2=Danna M.|last3=Davidson|first3=Elizabeth W.|last4=Collins|first4=James P.}}</ref><ref>{{cite journal|doi = 10.1111/j.1461-0248.2005.00735.x|title = Susceptibility of Italian agile frog populations to an emerging strain of Ranavirus parallels population genetic diversity|year = 2005|author = Pearman, Peter B.|journal = Ecology Letters|volume = 8|issue = 4|pages = 401|last2 = Garner|first2 = Trenton W. J.}}</ref> In the UK, the severity of disease outbreaks is thought to have increased due to climate change.<ref>{{Cite journal|last=Price|first=Stephen J.|last2=Leung|first2=William T. M.|last3=Owen|first3=Christopher J.|last4=Puschendorf|first4=Robert|last5=Sergeant|first5=Chris|last6=Cunningham|first6=Andrew A.|last7=Balloux|first7=Francois|last8=Garner|first8=Trenton W. J.|last9=Nichols|first9=Richard A.|date=2019-05-09|title=Effects of historic and projected climate change on the range and impacts of an emerging wildlife disease|journal=Global Change Biology|volume=25|issue=8|pages=2648–2660|doi=10.1111/gcb.14651|issn=1354-1013|hdl=10026.1/13802|hdl-access=free}}</ref>


== Etymology ==
==Etymology==
''Rana'' is derived from the [[Latin]] for "frog",<ref>{{OEtymD|frog}}</ref> reflecting the first isolation of a ''Ranavirus'' in 1960s from the Northern leopard frog (''[[Lithobates pipiens]]'').<ref name=Granoff>{{cite journal | last1 = Granoff | first1 = A | last2 = Came | first2 = PE | last3 = Rafferty | first3 = KA | title = The isolation and properties of viruses from Rana pipiens: their possible relationship to the renal adenocarcinoma of the leopard frog | journal = Annals of the New York Academy of Sciences | volume = 126 | issue = 1 | pages = 237–255 | year = 1965 | pmid = 5220161 | doi = 10.1111/j.1749-6632.1965.tb14278.x| bibcode = 1965NYASA.126..237G }}</ref><ref name=Gray/><ref name=Rafferty>{{cite journal | last1 = Rafferty | first1 = KA | title = The cultivation of inclusion-associated viruses from Lucke tumor frogs | journal = Annals of the New York Academy of Sciences | volume = 126 | issue = 1 | pages = 3–21 | year = 1965 | pmid = 5220167 | doi = 10.1111/j.1749-6632.1965.tb14266.x| bibcode = 1965NYASA.126....3R }}</ref>
''Rana'' is derived from the [[Latin]] for "frog",<ref>{{OEtymD|frog}}</ref> reflecting the first isolation of a ''Ranavirus'' in 1960s from the Northern leopard frog (''[[Lithobates pipiens]]'').<ref name=Granoff>{{cite journal | last1 = Granoff | first1 = A | last2 = Came | first2 = PE | last3 = Rafferty | first3 = KA | title = The isolation and properties of viruses from Rana pipiens: their possible relationship to the renal adenocarcinoma of the leopard frog | journal = Annals of the New York Academy of Sciences | volume = 126 | issue = 1 | pages = 237–255 | year = 1965 | pmid = 5220161 | doi = 10.1111/j.1749-6632.1965.tb14278.x| bibcode = 1965NYASA.126..237G }}</ref><ref name=Gray/><ref name=Rafferty>{{cite journal | last1 = Rafferty | first1 = KA | title = The cultivation of inclusion-associated viruses from Lucke tumor frogs | journal = Annals of the New York Academy of Sciences | volume = 126 | issue = 1 | pages = 3–21 | year = 1965 | pmid = 5220167 | doi = 10.1111/j.1749-6632.1965.tb14266.x| bibcode = 1965NYASA.126....3R }}</ref>


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=Hosts=
=Hosts=
⚫
===Anuran Hosts===

⚫
==Anuran Hosts==
*Wood frogs (''Lithobates sylvaticus'')
*Wood frogs (''Lithobates sylvaticus'')
*American Bullfrog (''Lithobates catesbieanus'')
*American Bullfrog (''Lithobates catesbieanus'')


==Urodelan Hosts==
===Urodelan Hosts===


==Reptilian Hosts==
===Reptilian Hosts===


*Green pythons (''[[Chondropython viridis]]'')<ref name=Hyatt2002>{{cite journal|author=First identification of a ranavirus from green pythons (''Chondropython viridis'')|title=First identification of a ranavirus from green pythons (Chondropython viridis)|journal=Journal of Wildlife Diseases|volume=38|issue=2|pages=239–52|pmid=12038121|year=2002|last2=Williamson|last3=Coupar|last4=Middleton|last5=Hengstberger|last6=Gould|last7=Selleck|last8=Wise|last9=Kattenbelt|last10=Cunningham|last11=Lee|doi=10.7589/0090-3558-38.2.239}}</ref>
*Green pythons (''[[Chondropython viridis]]'')<ref name=Hyatt2002>{{cite journal|author=First identification of a ranavirus from green pythons (''Chondropython viridis'')|title=First identification of a ranavirus from green pythons (Chondropython viridis)|journal=Journal of Wildlife Diseases|volume=38|issue=2|pages=239–52|pmid=12038121|year=2002|last2=Williamson|last3=Coupar|last4=Middleton|last5=Hengstberger|last6=Gould|last7=Selleck|last8=Wise|last9=Kattenbelt|last10=Cunningham|last11=Lee|doi=10.7589/0090-3558-38.2.239}}</ref>
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*Eastern Fence Lizard (''[[Sceloporus undulatus]]'') <ref name="Goodman2018">{{cite journal|title=Detection of Ranavirus in Eastern Fence Lizards and Eastern Box Turtles in Central Virginia|journal=Northeastern naturalist|volume=25|issue=3|pages=391–398|year=2018|last1=Goodman|first1=R.|last2=Hargadon|first2=K|last3=Carter|first3=E. |doi=10.1656/045.025.0306}}</ref>
*Eastern Fence Lizard (''[[Sceloporus undulatus]]'') <ref name="Goodman2018">{{cite journal|title=Detection of Ranavirus in Eastern Fence Lizards and Eastern Box Turtles in Central Virginia|journal=Northeastern naturalist|volume=25|issue=3|pages=391–398|year=2018|last1=Goodman|first1=R.|last2=Hargadon|first2=K|last3=Carter|first3=E. |doi=10.1656/045.025.0306}}</ref>


=Taxonomy=
==Taxonomy==
<big>'''Group: dsDNA'''</big>
<big>'''Group: dsDNA'''</big>
{{Collapsible list|title= <big>Order: Unassigned</big>
{{Collapsible list|title= <big>Order: Pimascovirales</big>
|1={{Collapsible list| framestyle=border:none; padding:1.0em;|title=Family: [[Iridoviridae]]
|1={{Collapsible list| framestyle=border:none; padding:1.0em;|title=Family: [[Iridoviridae]]
|1={{hidden begin|title=<small>Genus: Ranavirus</small>}}
|1={{hidden begin|title=<small>Genus: Ranavirus</small>}}
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}}<ref name=ICTVReport/>
}}<ref name=ICTVReport/>


The family ''Iridoviridae'' is divided into five genera which include ''[[Chloriridovirus]]'', ''[[Iridovirus]]'', ''[[Lymphocystivirus]]'', ''[[Megalocytivirus]]'', and ''Ranavirus''.<ref name=ICTVReport/> The genus ''Ranavirus '' is composed of 6 recognized [[viral species]], 3 of which are known to infect amphibians ([[Ambystoma tigrinum virus]] (ATV), [[Bohle iridovirus]] (BIV), and [[frog virus 3]]).<ref name = Chinchar/>
The family ''Iridoviridae'' is divided into six genera which include ''[[Chloriridovirus]]'', ''[[Iridovirus]]'', ''[[Lymphocystivirus]]'', ''[[Megalocytivirus]]'', and ''Ranavirus''.<ref name=ICTVReport/> The genus ''Ranavirus '' is composed of 6 recognized [[viral species]], 3 of which are known to infect amphibians ([[Ambystoma tigrinum virus]] (ATV), [[Bohle iridovirus]] (BIV), and [[frog virus 3]]).<ref name = Chinchar/>


=Structure=
==Structure==
Ranaviruses are large [[icosahedral]] DNA viruses measuring approximately 150&nbsp;nm in diameter with a large single linear dsDNA [[genome]] of roughly 105 kbp<ref name=Williams>Williams T, Barbosa-Solomieu V, Chinchar GD (2005). "A decade of advances in iridovirus research" 173-148. ''In'' Maramorosch K, Shatkin A (eds). ''Advances in virus research, Vol. 65'' Academic Press, New York, USA.</ref> which codes for around 100 gene products.<ref name=Chinchar2002>{{cite journal | last1 = Chinchar | first1 = VG | title = Ranaviruses (family ''Iridoviridae'') emerging cold-blooded killers | journal = Archives of Virology | volume = 147 | issue = 3 | pages = 447–470 | year = 2002 | pmid = 11958449 | doi = 10.1007/s007050200000 }}</ref> The main structural component of the [[protein]] [[capsid]] is the [[major capsid protein]] (MCP).
Ranaviruses are large [[icosahedral]] DNA viruses measuring approximately 150&nbsp;nm in diameter with a large single linear dsDNA [[genome]] of roughly 105 kbp<ref name=Williams>Williams T, Barbosa-Solomieu V, Chinchar GD (2005). "A decade of advances in iridovirus research" 173-148. ''In'' Maramorosch K, Shatkin A (eds). ''Advances in virus research, Vol. 65'' Academic Press, New York, USA.</ref> which codes for around 100 gene products.<ref name=Chinchar2002>{{cite journal | last1 = Chinchar | first1 = VG | title = Ranaviruses (family ''Iridoviridae'') emerging cold-blooded killers | journal = Archives of Virology | volume = 147 | issue = 3 | pages = 447–470 | year = 2002 | pmid = 11958449 | doi = 10.1007/s007050200000 }}</ref> The main structural component of the [[protein]] [[capsid]] is the [[major capsid protein]] (MCP).
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=Replication=
==Replication==
Ranaviral replication is well-studied using the [[type species]] for the genus, frog virus 3 (FV3).<ref name=Chinchar/><ref name = Williams/> Replication of FV3 occurs between 12 and 32 degrees Celsius.<ref name=Chinchar2002/> Ranaviruses enter the host cell by [[receptor-mediated endocytosis]].<ref>{{cite journal|title=The genomic diversity and phylogenetic relationship in the family ''Iridoviridae''|journal=Viruses|volume=2|issue=7|pages=1458–75|doi=10.3390/v2071458|pmid=21994690|year=2010|last1=Eaton|first1=Heather E.|last2=Ring|first2=Brooke A.|last3=Brunetti|first3=Craig R.|pmc=3185713}}</ref> Viral particles are uncoated and subsequently move into the [[cell nucleus]], where viral [[DNA replication]] begins via a virally encoded [[DNA polymerase]].<ref name=Goorha>{{cite journal | last1 = Goorha | first1 = R | title = Frog virus 3 DNA replication occurs in two stages | journal = Journal of Virology | volume = 43 | issue = 2 | pages = 519–28 | year = 1982 | pmid = 7109033| pmc = 256155}}</ref> Viral DNA then abandons the cell nucleus and begins the second stage of DNA replication in the cytoplasm, ultimately forming DNA [[concatemers]].<ref name=Goorha/> The viral DNA is then packaged via a [[headful mechanism]] into infectious virions.<ref name=Chinchar>Chinchar VG, Essbauer S, He JG, Hyatt A, Miyazaki T, Seligy V, Williams T (2005). "Family ''Iridoviridae''" pp. 145–162 in Fauquet CM, Mayo MA, Maniloff J, Desselburger U, Ball LA (eds). ''Virus Taxonomy, Eighth report of the International Committee on Taxonomy of Viruses.'' Academic Press, San Diego, USA.</ref> The ''ranavirus'' genome, like other iridoviral genomes is [[Circular permutation in proteins|circularly permuted]] and exhibits [[terminally redundant DNA]].<ref name=Goorha/>
Ranaviral replication is well-studied using the [[type species]] for the genus, frog virus 3 (FV3).<ref name=Chinchar/><ref name = Williams/> Replication of FV3 occurs between 12 and 32 degrees Celsius.<ref name=Chinchar2002/> Ranaviruses enter the host cell by [[receptor-mediated endocytosis]].<ref>{{cite journal|title=The genomic diversity and phylogenetic relationship in the family ''Iridoviridae''|journal=Viruses|volume=2|issue=7|pages=1458–75|doi=10.3390/v2071458|pmid=21994690|year=2010|last1=Eaton|first1=Heather E.|last2=Ring|first2=Brooke A.|last3=Brunetti|first3=Craig R.|pmc=3185713}}</ref> Viral particles are uncoated and subsequently move into the [[cell nucleus]], where viral [[DNA replication]] begins via a virally encoded [[DNA polymerase]].<ref name=Goorha>{{cite journal | last1 = Goorha | first1 = R | title = Frog virus 3 DNA replication occurs in two stages | journal = Journal of Virology | volume = 43 | issue = 2 | pages = 519–28 | year = 1982 | pmid = 7109033| pmc = 256155}}</ref> Viral DNA then abandons the cell nucleus and begins the second stage of DNA replication in the cytoplasm, ultimately forming DNA [[concatemers]].<ref name=Goorha/> The viral DNA is then packaged via a [[headful mechanism]] into infectious virions.<ref name=Chinchar>Chinchar VG, Essbauer S, He JG, Hyatt A, Miyazaki T, Seligy V, Williams T (2005). "Family ''Iridoviridae''" pp. 145–162 in Fauquet CM, Mayo MA, Maniloff J, Desselburger U, Ball LA (eds). ''Virus Taxonomy, Eighth report of the International Committee on Taxonomy of Viruses.'' Academic Press, San Diego, USA.</ref> The ''ranavirus'' genome, like other iridoviral genomes is [[Circular permutation in proteins|circularly permuted]] and exhibits [[terminally redundant DNA]].<ref name=Goorha/>
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=Transmission=
==Transmission==
Transmission of ranaviruses is thought to occur by multiple routes, including contaminated soil, direct contact, waterborne exposure, and ingestion of infected tissues during [[predation]], [[necrophagy]] or [[cannibalism]].<ref name=Gray/> Ranaviruses are relatively stable in aquatic environments, persisting several weeks or longer outside a host organism.<ref name=Gray>{{cite journal | last1 = Gray | first1 = MJ | last2 = Miller | first2 = DL | last3 = Hoverman | first3 = JT | title = Ecology and pathology of amphibian ranaviruses | journal = Diseases of Aquatic Organisms | volume = 87 | issue = 3 | pages = 243–266 | year = 2009 | pmid = 20099417 | doi = 10.3354/dao02138| doi-access = free }}</ref>
Transmission of ranaviruses is thought to occur by multiple routes, including contaminated soil, direct contact, waterborne exposure, and ingestion of infected tissues during [[predation]], [[necrophagy]] or [[cannibalism]].<ref name=Gray/> Ranaviruses are relatively stable in aquatic environments, persisting several weeks or longer outside a host organism.<ref name=Gray>{{cite journal | last1 = Gray | first1 = MJ | last2 = Miller | first2 = DL | last3 = Hoverman | first3 = JT | title = Ecology and pathology of amphibian ranaviruses | journal = Diseases of Aquatic Organisms | volume = 87 | issue = 3 | pages = 243–266 | year = 2009 | pmid = 20099417 | doi = 10.3354/dao02138| doi-access = free }}</ref>


=Epizoology=
==Epizoology==
Amphibian mass mortality events due to ''Ranavirus'' have been reported in Asia, Europe, North America, and South America.<ref name=Gray/> Ranaviruses have been isolated from wild populations of amphibians in Australia, but have not been associated with mass mortality on that continent.<ref name=Gray/><ref name=Speare>{{cite journal | last1 = Speare | first1 = R | last2 = Smith | first2 = JR | title = An iridovirus-like agent isolated from the ornate burrowing frog ''Limnodynastes ornatus'' in northern Australia | journal = Diseases of Aquatic Organisms | volume = 14 | pages = 51–57 | year = 1992 | doi = 10.3354/dao014051| doi-access = free }}</ref><ref name=Cullen>{{cite journal | last1 = Cullen | first1 = BR | last2 = Owens | first2 = L | title = Experimental challenge and clinical cases of Bohle iridovirus (BIV) in native Australian anurans | journal = Diseases of Aquatic Organisms | volume = 49 | issue = 2 | pages = 83–92 | year = 2002 | pmid = 12078986 | doi=10.3354/dao049083| doi-access = free }}</ref>
Amphibian mass mortality events due to ''Ranavirus'' have been reported in Asia, Europe, North America, and South America.<ref name=Gray/> Ranaviruses have been isolated from wild populations of amphibians in Australia, but have not been associated with mass mortality on that continent.<ref name=Gray/><ref name=Speare>{{cite journal | last1 = Speare | first1 = R | last2 = Smith | first2 = JR | title = An iridovirus-like agent isolated from the ornate burrowing frog ''Limnodynastes ornatus'' in northern Australia | journal = Diseases of Aquatic Organisms | volume = 14 | pages = 51–57 | year = 1992 | doi = 10.3354/dao014051| doi-access = free }}</ref><ref name=Cullen>{{cite journal | last1 = Cullen | first1 = BR | last2 = Owens | first2 = L | title = Experimental challenge and clinical cases of Bohle iridovirus (BIV) in native Australian anurans | journal = Diseases of Aquatic Organisms | volume = 49 | issue = 2 | pages = 83–92 | year = 2002 | pmid = 12078986 | doi=10.3354/dao049083| doi-access = free }}</ref>


=Pathogenesis=
==Pathogenesis==
Synthesis of viral proteins begins within hours of viral entry<ref name=Chinchar2002/> with [[necrosis]] or [[apoptosis]] occurring as early as a few hours post-infection.<ref name=Williams/><ref name= Chinchar2003>{{cite journal | last1 = Chinchar | first1 = VG | last2 = Bryan | first2 = L | last3 = Wang | first3 = J | last4 = Long | first4 = S | last5 = Chinchar | first5 = GD |title = Induction of apoptosis in frog virus 3-infected cells | journal = Virology | volume = 306 | pages = 303–312 | year = 2003 | pmid = 12642103 | doi = 10.1016/S0042-6822(02)00039-9 | issue = 2}}</ref>
Synthesis of viral proteins begins within hours of viral entry<ref name=Chinchar2002/> with [[necrosis]] or [[apoptosis]] occurring as early as a few hours post-infection.<ref name=Williams/><ref name= Chinchar2003>{{cite journal | last1 = Chinchar | first1 = VG | last2 = Bryan | first2 = L | last3 = Wang | first3 = J | last4 = Long | first4 = S | last5 = Chinchar | first5 = GD |title = Induction of apoptosis in frog virus 3-infected cells | journal = Virology | volume = 306 | pages = 303–312 | year = 2003 | pmid = 12642103 | doi = 10.1016/S0042-6822(02)00039-9 | issue = 2}}</ref>


=Gross pathology=
==Gross pathology==
Gross lesions associated with ''Ranavirus'' infection include erythema, generalized swelling, hemorrhage, limb swelling, and swollen and friable livers.<ref name=Gray/>
Gross lesions associated with ''Ranavirus'' infection include erythema, generalized swelling, hemorrhage, limb swelling, and swollen and friable livers.<ref name=Gray/>


⚫
==See also==
= Information Sources =
⚫
More information on ''Ranavirus'' and other pathogens impacting amphibian populations, including ''[[Batrachochytrium dendrobatidis]]'' and ''[[Batrachochytrium salamandrivorans]]'' can be found at the Southeast Partners in Amphibian and Reptile Conservation disease task team web-page. [http://separc.org/diseases-herpetofauna]

⚫
=See also=
*[[Decline in amphibian populations]]
*[[Decline in amphibian populations]]


= References =
==References==
{{Reflist|35em}}
{{Reflist|35em}}


=External links=
==External links==
{{Commons category|Ranavirus}}
{{Commons category|Ranavirus}}
{{Wikispecies-inline|List of viruses}}
{{Wikispecies-inline|List of viruses}}
Line 118: Line 111:
* [http://www.ranavirus.net Ranavirus Research Project]
* [http://www.ranavirus.net Ranavirus Research Project]
* [https://web.archive.org/web/20090411062041/http://www.jcu.edu.au/school/phtm/PHTM/frogs/otherdiseases-viruses.htm Viral Diseases of Amphibians]
* [https://web.archive.org/web/20090411062041/http://www.jcu.edu.au/school/phtm/PHTM/frogs/otherdiseases-viruses.htm Viral Diseases of Amphibians]
⚫
* More information on ''Ranavirus'' and other pathogens impacting amphibian populations, including ''[[Batrachochytrium dendrobatidis]]'' and ''[[Batrachochytrium salamandrivorans]]'' can be found at the Southeast Partners in Amphibian and Reptile Conservation disease task team web-page. [http://separc.org/diseases-herpetofauna]

{{Baltimore classification}}
{{Baltimore classification}}


Revision as of 22:02, 1 May 2020

Ranavirus
Transmission electron micrograph of ranaviruses (dark hexagons) gathering at the cell border and leaving the cell via a process called "budding".
Virus classification Edit this classification
(unranked): Virus
Realm: Varidnaviria
Kingdom: Bamfordvirae
Phylum: Nucleocytoviricota
Class: Megaviricetes
Order: Pimascovirales
Family: Iridoviridae
Subfamily: Alphairidovirinae
Genus: Ranavirus
Transmission electron micrograph of a cell infected with ranaviruses, which gather in the cytoplasm and in the assembly bodies next to the contorted nucleus.

Ranavirus is a genus of viruses, in the family Iridoviridae.[1] There are five other genera of viruses within the family Iridoviridae, but Ranavirus is the only one that includes viruses that are infectious to amphibians and reptiles. Additionally, it is one of the three genera within this family which infect teleost fishes, along with Lymphocystivirus and Megalocytivirus.[2] The family Iridoviridae is one of the five families of nucleocytoplasmic large DNA viruses.

Ecological impact

The Ranaviruses, like the Megalocytiviruses, are an emerging group of closely related dsDNA viruses which cause systemic infections in a wide variety of wild and cultured fresh and saltwater fishes. As with Megalocytiviruses, Ranavirus outbreaks are therefore of considerable economic importance in aquaculture, as epizootics can result in moderate fish loss or mass mortality events of cultured fishes. Unlike Megalocytiviruses, however, Ranavirus infections in amphibians have been implicated as a contributing factor in the global decline of amphibian populations.[3][4] The impact of Ranaviruses on amphibian populations has been compared to the chytrid fungus Batrachochytrium dendrobatidis, the causative agent of chytridiomycosis.[5][6][7] In the UK, the severity of disease outbreaks is thought to have increased due to climate change.[8]

Etymology

Rana is derived from the Latin for "frog",[9] reflecting the first isolation of a Ranavirus in 1960s from the Northern leopard frog (Lithobates pipiens).[10][11][12]

Evolution

VOA report about Ranavirus

The ranaviruses appear to have evolved from a fish virus which subsequently infected amphibians and reptiles.[13]

Hosts

Anuran Hosts

  • Wood frogs (Lithobates sylvaticus)
  • American Bullfrog (Lithobates catesbieanus)

Urodelan Hosts

Reptilian Hosts

Taxonomy

Group: dsDNA

Order: Pimascovirales

[1]

The family Iridoviridae is divided into six genera which include Chloriridovirus, Iridovirus, Lymphocystivirus, Megalocytivirus, and Ranavirus.[1] The genus Ranavirus is composed of 6 recognized viral species, 3 of which are known to infect amphibians (Ambystoma tigrinum virus (ATV), Bohle iridovirus (BIV), and frog virus 3).[24]

Structure

Ranaviruses are large icosahedral DNA viruses measuring approximately 150 nm in diameter with a large single linear dsDNA genome of roughly 105 kbp[25] which codes for around 100 gene products.[26] The main structural component of the protein capsid is the major capsid protein (MCP).

Genus Structure Symmetry Capsid Genomic arrangement Genomic segmentation
Ranavirus Polyhedral T=133 or 147 Linear Monopartite

Replication

Ranaviral replication is well-studied using the type species for the genus, frog virus 3 (FV3).[24][25] Replication of FV3 occurs between 12 and 32 degrees Celsius.[26] Ranaviruses enter the host cell by receptor-mediated endocytosis.[27] Viral particles are uncoated and subsequently move into the cell nucleus, where viral DNA replication begins via a virally encoded DNA polymerase.[28] Viral DNA then abandons the cell nucleus and begins the second stage of DNA replication in the cytoplasm, ultimately forming DNA concatemers.[28] The viral DNA is then packaged via a headful mechanism into infectious virions.[24] The ranavirus genome, like other iridoviral genomes is circularly permuted and exhibits terminally redundant DNA.[28]

Genus Host details Tissue tropism Entry details Release details Replication site Assembly site Transmission
Ranavirus Frogs; snakes None Cell receptor endocytosis Lysis; budding Nucleus Cytoplasm Contact

Transmission

Transmission of ranaviruses is thought to occur by multiple routes, including contaminated soil, direct contact, waterborne exposure, and ingestion of infected tissues during predation, necrophagy or cannibalism.[11] Ranaviruses are relatively stable in aquatic environments, persisting several weeks or longer outside a host organism.[11]

Epizoology

Amphibian mass mortality events due to Ranavirus have been reported in Asia, Europe, North America, and South America.[11] Ranaviruses have been isolated from wild populations of amphibians in Australia, but have not been associated with mass mortality on that continent.[11][29][30]

Pathogenesis

Synthesis of viral proteins begins within hours of viral entry[26] with necrosis or apoptosis occurring as early as a few hours post-infection.[25][31]

Gross pathology

Gross lesions associated with Ranavirus infection include erythema, generalized swelling, hemorrhage, limb swelling, and swollen and friable livers.[11]

See also

References

  1. ^ a b c "Iridoviridae". ICTV Online (10th) Report.
  2. ^ Whittington, RJ; Becker, JA; Dennis, MM (2010). "Iridovirus infections in finfish – critical review with emphasis on ranaviruses". Journal of Fish Diseases. 33 (2): 95–122. doi:10.1111/j.1365-2761.2009.01110.x. PMID 20050967.
  3. ^ Teacher, A. G. F.; Cunningham, A. A.; Garner, T. W. J. (10 June 2010). "Assessing the long-term impact of Ranavirus infection in wild common frog populations: Impact of Ranavirus on wild frog populations". Animal Conservation. 13 (5): 514–522. doi:10.1111/j.1469-1795.2010.00373.x.
  4. ^ Price, Stephen J.; Garner, Trenton W.J.; Nichols, Richard A.; Balloux, François; Ayres, César; Mora-Cabello de Alba, Amparo; Bosch, Jaime (November 2014). "Collapse of Amphibian Communities Due to an Introduced Ranavirus". Current Biology. 24 (21): 2586–2591. doi:10.1016/j.cub.2014.09.028. PMID 25438946. {{cite journal}}: no-break space character in |first2= at position 8 (help); no-break space character in |first3= at position 8 (help); no-break space character in |first= at position 8 (help); no-break space character in |last6= at position 13 (help)
  5. ^ Jancovich, James K; Mao, Jinghe; Chinchar, V.Gregory; Wyatt, Christopher; Case, Steven T; Kumar, Sudhir; Valente, Graziela; Subramanian, Sankar; Davidson, Elizabeth W; Collins, James P; Jacobs, Bertram L (2003). "Genomic sequence of a ranavirus (family Iridoviridae) associated with salamander mortalities in North America". Virology. 316 (1): 90–103. doi:10.1016/j.virol.2003.08.001. PMID 14599794.
  6. ^ Brunner, Jesse L.; Schock, Danna M.; Davidson, Elizabeth W.; Collins, James P. (2004). "Intraspecific Reservoirs: Complex Life History and the Persistence of a Lethal Ranavirus". Ecology. 85 (2): 560. doi:10.1890/02-0374.
  7. ^ Pearman, Peter B.; Garner, Trenton W. J. (2005). "Susceptibility of Italian agile frog populations to an emerging strain of Ranavirus parallels population genetic diversity". Ecology Letters. 8 (4): 401. doi:10.1111/j.1461-0248.2005.00735.x.
  8. ^ Price, Stephen J.; Leung, William T. M.; Owen, Christopher J.; Puschendorf, Robert; Sergeant, Chris; Cunningham, Andrew A.; Balloux, Francois; Garner, Trenton W. J.; Nichols, Richard A. (9 May 2019). "Effects of historic and projected climate change on the range and impacts of an emerging wildlife disease". Global Change Biology. 25 (8): 2648–2660. doi:10.1111/gcb.14651. hdl:10026.1/13802. ISSN 1354-1013.
  9. ^ Harper, Douglas. "frog". Online Etymology Dictionary.
  10. ^ Granoff, A; Came, PE; Rafferty, KA (1965). "The isolation and properties of viruses from Rana pipiens: their possible relationship to the renal adenocarcinoma of the leopard frog". Annals of the New York Academy of Sciences. 126 (1): 237–255. Bibcode:1965NYASA.126..237G. doi:10.1111/j.1749-6632.1965.tb14278.x. PMID 5220161.
  11. ^ a b c d e f Gray, MJ; Miller, DL; Hoverman, JT (2009). "Ecology and pathology of amphibian ranaviruses". Diseases of Aquatic Organisms. 87 (3): 243–266. doi:10.3354/dao02138. PMID 20099417.
  12. ^ Rafferty, KA (1965). "The cultivation of inclusion-associated viruses from Lucke tumor frogs". Annals of the New York Academy of Sciences. 126 (1): 3–21. Bibcode:1965NYASA.126....3R. doi:10.1111/j.1749-6632.1965.tb14266.x. PMID 5220167.
  13. ^ Jancovich, JK; Bremont, M; Touchman, JW; Jacobs, BL (2010). "Evidence for multiple recent host species shifts among the Ranaviruses (family Iridoviridae)". J Virol. 84 (6): 2636–2647. doi:10.1128/JVI.01991-09. PMC 2826071. PMID 20042506.
  14. ^ First identification of a ranavirus from green pythons (Chondropython viridis); Williamson; Coupar; Middleton; Hengstberger; Gould; Selleck; Wise; Kattenbelt; Cunningham; Lee (2002). "First identification of a ranavirus from green pythons (Chondropython viridis)". Journal of Wildlife Diseases. 38 (2): 239–52. doi:10.7589/0090-3558-38.2.239. PMID 12038121.
  15. ^ Benetka V. (2007). "First report of an iridovirus (genus Ranavirus) infection in a leopard tortoise (Geochelone pardalis pardalis)" (PDF). Vet Med Austria. 94: 243–248.
  16. ^ De Matos, A. P.; Caeiro, M. F.; Papp, T; Matos, B. A.; Correia, A. C.; Marschang, R. E. (2011). "New viruses from Lacerta monticola (Serra da Estrela, Portugal): Further evidence for a new group of nucleo-cytoplasmic large deoxyriboviruses (NCLDVs)". Microscopy and Microanalysis. 17 (1): 101–8. Bibcode:2011MiMic..17..101A. doi:10.1017/S143192761009433X. PMID 21138619.
  17. ^ Mao, J; Hedrick, RP; Chinchar, VG (1997). "Molecular characterization, sequence analysis, and taxonomic position of newly isolated fish iridoviruses". Virology. 229 (1): 212–220. doi:10.1006/viro.1996.8435. PMID 9123863.
  18. ^ a b Johnson, A. J.; Pessier, A. P.; Jacobson, E. R. (2007). "Experimental transmission and induction of ranaviral disease in Western Ornate box turtles (Terrapene ornata ornata) and red-eared sliders (Trachemys scripta elegans)". Veterinary Pathology. 44 (3): 285–97. doi:10.1354/vp.44-3-285. PMID 17491069.
  19. ^ Blahak S., Uhlenbrok C. "Ranavirus infections in European terrestrial tortoises in Germany". Proceedings of the 1st International Conference on Reptile and Amphibian Medicine; Munich, Germany. 4–7 March 2010; pp. 17–23
  20. ^ McKenzie, C. M.; Piczak, M.L.; Snyman, H. N.; Joseph, T.; Theijin, C.; Chow-Fraser, P.; Jardine, C. M. (2019). "First report of ranavirus mortality in a common snapping turtle Chelydra serpentina" (PDF). Diseases of Aquatic Organisms (132): 221–227. doi:10.3354/dao03324.
  21. ^ Chen, Z. X.; Zheng, J. C.; Jiang, Y. L. (1999). "A new iridovirus isolated from soft-shelled turtle". Virus Research. 63 (1–2): 147–51. doi:10.1016/S0168-1702(99)00069-6. PMID 10509727.
  22. ^ Marschang, R. E.; Braun, S; Becher, P (2005). "Isolation of a ranavirus from a gecko (Uroplatus fimbriatus)". Journal of Zoo and Wildlife Medicine. 36 (2): 295–300. doi:10.1638/04-008.1. JSTOR 20096453. PMID 17323572.
  23. ^ Goodman, R.; Hargadon, K; Carter, E. (2018). "Detection of Ranavirus in Eastern Fence Lizards and Eastern Box Turtles in Central Virginia". Northeastern naturalist. 25 (3): 391–398. doi:10.1656/045.025.0306.
  24. ^ a b c Chinchar VG, Essbauer S, He JG, Hyatt A, Miyazaki T, Seligy V, Williams T (2005). "Family Iridoviridae" pp. 145–162 in Fauquet CM, Mayo MA, Maniloff J, Desselburger U, Ball LA (eds). Virus Taxonomy, Eighth report of the International Committee on Taxonomy of Viruses. Academic Press, San Diego, USA.
  25. ^ a b c Williams T, Barbosa-Solomieu V, Chinchar GD (2005). "A decade of advances in iridovirus research" 173-148. In Maramorosch K, Shatkin A (eds). Advances in virus research, Vol. 65 Academic Press, New York, USA.
  26. ^ a b c Chinchar, VG (2002). "Ranaviruses (family Iridoviridae) emerging cold-blooded killers". Archives of Virology. 147 (3): 447–470. doi:10.1007/s007050200000. PMID 11958449.
  27. ^ Eaton, Heather E.; Ring, Brooke A.; Brunetti, Craig R. (2010). "The genomic diversity and phylogenetic relationship in the family Iridoviridae". Viruses. 2 (7): 1458–75. doi:10.3390/v2071458. PMC 3185713. PMID 21994690.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  28. ^ a b c Goorha, R (1982). "Frog virus 3 DNA replication occurs in two stages". Journal of Virology. 43 (2): 519–28. PMC 256155. PMID 7109033.
  29. ^ Speare, R; Smith, JR (1992). "An iridovirus-like agent isolated from the ornate burrowing frog Limnodynastes ornatus in northern Australia". Diseases of Aquatic Organisms. 14: 51–57. doi:10.3354/dao014051.
  30. ^ Cullen, BR; Owens, L (2002). "Experimental challenge and clinical cases of Bohle iridovirus (BIV) in native Australian anurans". Diseases of Aquatic Organisms. 49 (2): 83–92. doi:10.3354/dao049083. PMID 12078986.
  31. ^ Chinchar, VG; Bryan, L; Wang, J; Long, S; Chinchar, GD (2003). "Induction of apoptosis in frog virus 3-infected cells". Virology. 306 (2): 303–312. doi:10.1016/S0042-6822(02)00039-9. PMID 12642103.

External links

Wikimedia Commons has media related to Ranavirus.

Data related to List of viruses at Wikispecies

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