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Kathedralbasilika St. Marien (Galveston)

2019-09-27T01:17:50Z

Tom.Bot: :Category:CS1 errors: deprecated parameters: migrate 3/3 |dead-url= to |url-status=; minor cleanup; WP:GenFixes on

{{About|the Cathedral Basilica of the Archdiocese of Galveston-Houston|other uses of St. Mary Basilica|Basilica of St. Mary (disambiguation)}}
{{Infobox church
| name = St. Mary Cathedral Basilica
| fullname =
| image = St_Mary's_Cathedral_Basilica,_Galveston.jpg
| imagesize = frameless
| imagelink =
| imagealt =
| caption = St. Mary Cathedral Basilica in 2006
| pushpin map = Texas#USA
| pushpin label position =
| pushpin map alt =
| pushpin mapsize =
| map caption =
| relief = yes
| coordinates = {{coord|29|18|15|N|94|47|25|W|display=inline,title}}
| location = 2011 Church St. [[Galveston, Texas]]
| country = United States

| denomination = [[Roman Catholic]]
| membership =
| attendance =
| website = {{Official website|url=https://www.archgh.org/about/our-cathedrals/st-mary-cathedral-basilica/|name=St. Mary Cathedral Basilica}}

| founded date =
| founder =
| dedication = [[Blessed Virgin Mary]]
| consecrated date = November 26, 1848
| status = [[Cathedral]] - [[Minor Basilica]]
| functional status =
| heritage designation =
| designated date =
| architect=[[Theodore Eugene Giraud]], with later addition by [[Nicholas J. Clayton]]
| architectural type = [[Gothic Revival architecture|Gothic]]
| style =
| groundbreaking = 1843
| completed date = 1847
| construction cost =
| closed date =
| demolished date =
| capacity = 400+<ref name="majorrepair">{{Cite news|last=Vara|first=Richard|title=The state's first cathedral in need of major repair|newspaper=Houston Chronicle|date=March 30, 2008|url=http://www.chron.com/CDA/archives/archive.mpl?id=2008_4541739|accessdate=June 4, 2010}}</ref>
| length = {{convert|40|m|ft|sp=us}}
| width = {{convert|23|m|ft|sp=us}}
| height max =
| other dimensions = {{convert|1|acre}} (grounds area)
| spire quantity = Three
| spire height = {{convert|24.3|m|ft|sp=us}}
| materials = Imported Belgian brick and mortar

| parish = Holy Family
| archdiocese = [[Archdiocese of Galveston-Houston|Galveston-Houston]]
| archbishop = Cardinal [[Daniel N. DiNardo]]
| rector =
| embedded = {{Infobox NRHP
|embed = yes
| name = St. Mary's Cathedral
| nrhp_type =
| added = June 4, 1973
| refnum = 73001964<ref name="nris">{{NRISref|version=2013a}}</ref>
| designated_other1=RTHL
| designated_other1_date=1967
| designated_other1_number=[https://atlas.thc.state.tx.us/Details/5167007172 7172]
| designated_other1_num_position=bottom
}}}}

'''St. Mary Cathedral Basilica''', also known as ''St. Mary's Cathedral Basilica'' is a Roman Catholic place of worship situated in [[Galveston, Texas]]. It is the primary [[cathedral]] of the [[Archdiocese of Galveston-Houston]] and the [[mother church]] of the Catholic Church in Texas, as well as a [[minor basilica]].<ref>[http://www.archgh.org/cocathedral/cathedral-history.htm Archdiocese Cathedral History] {{webarchive |url=https://web.archive.org/web/20070125042116/http://www.archgh.org/cocathedral/cathedral-history.htm |date=January 25, 2007 }}</ref> Along with the [[Co-Cathedral of the Sacred Heart in Houston]], St. Mary's serves more than 1.5 million Catholics living in the Archdiocese.<ref name="abtdio">{{cite web| title=About Our Diocese| publisher=Archdiocese of Galveston-Houston| year=2007| url=http://www.archgh.org/about.htm| archive-url=https://web.archive.org/web/20050216172245/http://www.archgh.org/about.htm| url-status=dead| archive-date=February 16, 2005| access-date=June 4, 2010}}</ref><ref name="shining">{{cite news| last=Dooley| first=Tara| title=A shining achievement| newspaper=Houston Chronicle| date=March 30, 2008| url=http://www.chron.com/CDA/archives/archive.mpl?id=2008_4540450| accessdate=June 4, 2010}}</ref>

==History==

In 1840, the Rev. John Timon, the newly appointed Apostolic Prefect of Texas, named fellow Vincentian priest Rev. [[Jean-Marie Odin|John Odin]], [[Congregation of the Mission|C.M.]], to be the resident Vice-Prefect of [[Texas]]. Fr. Odin embarked from [[New Orleans]] on a schooner bound for the Texas coast, arriving in Galveston early in 1841. There he found a community of Catholics eager to build a church for their small congregation.

In the months that followed, Father Odin procured enough money to begin construction of a wooden-frame church.<ref name="woodframe">{{cite news| last=Dooley| first=Tara| title=The Archdiocese of Galveston-Houston began in a wood-frame church during the Republic of Texas| newspaper=Houston Chronicle| date=March 30, 2008| url=http://www.chron.com/CDA/archives/archive.mpl?id=2008_4541743| accessdate=June 4, 2010}}</ref> He was assisted in this venture by Colonel Michael B. Menard and Dr. Nicholas Labadie, prominent Galvestonians. Colonel Menard is to be remembered as one of the founders of the City of Galveston.

On February 6, 1842, one month before his consecration as a bishop, Odin dedicated the completed structure to the [[Blessed Virgin Mary]]. The small, rectangular building measured {{convert|22|ft|m}}. Odin, now the Apostolic Vicar of Texas, purchased a five-room cottage as the episcopal residence. He made an addition to the church structure of a small [[sacristy]], and bought thirty benches for the convenience of his parishioners.
[[File:St. Mary Cathedral, Galveston, exterior, by A. V. Latourette.jpg|thumb|left|[[Stereoscopic]] view of the Cathedral, circa 1865.]]
[[Image:Interior 1847 St Mary Cathedral Basilica, Galveston.jpg|thumb|left|200px|Sanctuary of St. Mary Cathedral Basilica]]
[[File:St Mary's Cathedral, Galveston.jpg|thumb|St Mary's Cathedral, Galveston]]

In 1845, Bishop Odin purchased 500,000 bricks from [[Belgium]], which were shipped to Galveston as ballast. He would use the bricks in the construction of his dream: a larger, permanent church.<ref>{{cite news| last=Long| first=Steve| title=Floods and storms, and now pestilence| newspaper=Houston Chronicle| date=January 2, 1989| url=http://www.chron.com/CDA/archives/archive.mpl?id=1989_593982| accessdate=June 4, 2010}}</ref> The little frame church was moved out into the street, and work on the new St. Mary's began in 1847. The ceremony of laying the cornerstone took place on Sunday, March 14. Father Timon came to Galveston for the event and preached the sermon before a large crowd. On May 4, 1847 [[Pope Pius IX]] approved the establishment of the Diocese of Galveston and named Odin as its first bishop.<ref name="woodframe"/><ref name="diocese">{{cite web| title=History| url=http://www.archgh.org/About/History/| archive-url=https://web.archive.org/web/20120415135013/http://www.archgh.org/About/History/| url-status=dead| archive-date=2012-04-15| publisher=Archdiocese of Galveston-Houston| access-date=2011-12-29}}</ref><ref name="storms">{{cite news| last=Dooley| first=Tara| title=At 155, Galveston's St. Mary's still battles the storms| newspaper=Houston Chronicle| date=January 4, 2003| url=http://www.chron.com/CDA/archives/archive.mpl?id=2003_3615026| accessdate=June 4, 2010}}</ref>

On November 26, 1848, the Cathedral was ready for dedication.<ref name="woodframe"/> Once more Father John Timon was chosen as the principal speaker because of his close association with, and his pioneer work in the diocese.

The Cathedral Basilica is notable as being one of the few buildings in Galveston that survived the devastating [[1900 Galveston Hurricane]] with only minimal damage.<ref name="storms"/>

Due to the tremendous growth in the City of [[Houston]], in 1959 the Most Reverend [[Wendelin J. Nold]], fifth bishop of the diocese, asked that the Diocese be re-designated the Diocese of Galveston-Houston. This created a co-capital or "see" city in Houston, and Sacred Heart Church in Houston was named the "Co-Cathedral" of the Diocese. This did not change the status of Galveston as a see city nor St. Mary Cathedral's place in the Diocese.<ref>{{cite news| last=Duin| first=Julia| title=Bishops celebrate 200th birthday| newspaper=Houston Chronicle| date=November 4, 1989| url=http://www.chron.com/CDA/archives/archive.mpl?id=1989_661494| accessdate=June 4, 2010}}</ref> Since St. Mary Cathedral was the first Catholic cathedral in the State of Texas, and the original Diocese of Galveston encompassed the entire state, it has the distinction of being the mother church of all the Catholic dioceses in Texas.<ref name="majorrepair"/>
[[File:St. Mary's Cathedral, Galveston, Texas.jpg|thumb|left|St. Mary's Cathedral, Galveston, Texas (postcard, circa 1890-1924)]]
St. Mary Cathedral was named a Texas state historic landmark in 1968 and a national historic landmark in 1973. In 1979, in recognition of the Cathedral's importance to the community and the State of Texas, as well as the historical impact it had on Catholicism in the state of Texas, [[Pope John Paul II]] elevated St. Mary Cathedral to the status of a minor basilica.<ref>{{cite web| title=Histories of the Cathedral| url=http://www.marycath.org/index.htm| archive-date=20 March 2008| access-date=2016-03-23| archive-url=https://web.archive.org/web/20080320073844/http://www.marycath.org/index.htm| publisher=St. Mary's Cathedral Basilica| url-status=dead| df=}}</ref>
{{Clear}}

==The basilica today==

The Cathedral Basilica sustained significant water damage during Hurricane Ike in 2008 and was closed for repairs until Easter 2014.<ref>{{cite web| url=http://www.galvestondailynews.com/story/142412/| work=[[The Daily News (Texas)|Galveston Daily News]]| title=Parish consolidates Galveston, Bolivar Catholics| last=Cousins| first=Rick| accessdate=2010-06-04}}</ref>

In 2009, the Archdiocese appointed a director of special projects to oversee the Cathedral Basilica's restoration. As of July 2012, the roof has been replaced, the pews have been rebuilt and refinished, steel armature reinforcements have been added to the two front [[spire]]s, the confessionals and [[Stations of the Cross]] have been refinished, and exterior masonry repairs, coating and chemical remediation have all been completed. A new concrete substructure is being built to support the floor, which is currently supported by the original wooden beams that were installed when the Cathedral Basilica was constructed in 1847.<ref name=ResHist>{{cite news| url=http://www.archgh.org/default.asp?id=500&hid=1040| work=Texas Catholic Herald News| title=Resurrecting history: Repairs continue at St. Mary Basilica| last=Torrellas| first=Rebecca| date=2012-07-17| accessdate=2012-08-02}}</ref>

==See also==
{{Portal|National Register of Historic Places|Catholicism|Texas}}
*[[National Register of Historic Places listings in Galveston County, Texas]]
*[[List of Recorded Texas Historic Landmarks (Eastland-Gray)#Galveston County|Recorded Texas Historic Landmarks in Galveston County]]
*[[List of Catholic cathedrals in the United States]]
*[[List of cathedrals in the United States]]

==References==
{{Reflist}}

==External links==
{{Commons category|St. Mary's Cathedral Basilica, Galveston|{{PAGENAMEBASE}}}}
{{Wikisource1913CatholicEnc|Galveston|The Diocese of Galveston}}
*[https://web.archive.org/web/20060905130456/http://www.marycath.org/ Official Cathedral Site]
*[http://www.archgh.org/ Roman Catholic Archdiocese of Galveston-Houston Official Site]

{{Roman Catholic Archdiocese of Galveston–Houston}}
{{National Register of Historic Places}}
{{Galveston, Texas}}

{{DEFAULTSORT:Saint Mary Cathedral Basilica, Galveston, Texas}}
[[Category:Basilica churches in Texas|Mary's Cathedral, Galveston]]
[[Category:Roman Catholic cathedrals in Texas|Mary Galveston]]
[[Category:Roman Catholic Archdiocese of Galveston–Houston|Cathedral St. Mary's]]
[[Category:Churches on the National Register of Historic Places in Texas]]
[[Category:National Register of Historic Places in Galveston County, Texas]]
[[Category:1847 establishments in Texas]]
[[Category:Churches in Galveston, Texas]]
[[Category:Recorded Texas Historic Landmarks]]

Marriages

2018-05-20T22:14:13Z

Tom.Bot: +{{Authority control}} (1 source from Wikidata), WP:GenFixes on, using AWB

{{Infobox musical artist 
| name = Marriages
| image =
| caption =
| background = group_or_band
| origin = Los Angeles, California, US
| genre = [[Post-rock]], [[alternative rock]], [[experimental rock]]
| years_active = 2011–present
| label = [[Sargent House]]
| associated_acts = The Nocturnes, Halifax Pier, [[Red Sparowes]], the Headless Prince of Zolpidem
| website = [http://marriagesband.com marriagesband.com]
| current_members = [[Emma Ruth Rundle]] Greg Burns Andrew Clinco
| past_members = Dave Clifford
}}

'''Marriages''' is an American [[rock music|rock]] band from [[Los Angeles]], [[California]] that formed in 2012.<ref name="AllMusic">{{cite web|url=http://www.allmusic.com/artist/marriages-mn0002897552|title=Marriages Music Biography, Credits and Discography|publisher=[[AllMusic]]|accessdate=August 4, 2014}}</ref> The band is composed of vocalist and guitarist [[Emma Ruth Rundle]], bassist Greg Burns and drummer Andrew Clinco.<ref name="AllMusic"/>

==History==
Following the hiatus of [[Red Sparowes]], two members of that band, Rundle (also formerly of the Nocturnes) and Burns (ex-Halifax Pier), formed Marriages.<ref>{{cite web|url=http://ghettoblastermagazine.com/2013/interview-greg-burns-of-marriages/|title=Interview: Greg Burns of Marriages|publisher=Ghettoblaster Magazine|author=Timothy Anderl|date=February 22, 2013|accessdate= June 5, 2015}}</ref> The group recorded ''[[Kitsune (Marriages EP)|Kitsune]]'' EP with drummer Dave Clifford. It was released in 2012 on [[Sargent House]],<ref name="AllMusic"/> followed by tours with [[Russian Circles]] and [[Chelsea Wolfe]] in 2012, and [[Deafheaven]] in 2013.<ref>{{cite web|url=http://www.stereogum.com/1730692/marriages-skin-stereogum-premiere/mp3s/|title=Marriages – "Skin" (Stereogum Premiere)|publisher=[[Stereogum]]|author=Michael Nelson|date=January 22, 2015|accessdate=June 5, 2015}}</ref><ref>{{cite web|url=http://www.brooklynvegan.com/archives/2013/05/deafheaven_anno.html|title=Deafheaven announce tour with Marriages, including show at Saint Vitus with NOTHING (dates)|publisher=[[BrooklynVegan]]|date=May 14, 2013|accessdate=June 5, 2015}}</ref> Clinco was brought in as a permanent member of the band, replacing Clifford.

In 2014, the group completed recording for their debut full-length album, ''[[Salome (Marriages album)|Salome]]'', while Rundle simultaneously recorded and released her solo debut ''[[Some Heavy Ocean]]'' (she has also recorded under the name the Headless Prince of Zolpidem).<ref>{{cite web|url=http://noisey.vice.com/en_ca/blog/marriages-interview-2015|title=Marriages Breathes New Life Into Post-Rock On Their New Album, 'Salome'|publisher=[[Noisey]]|author=Jamie Ludwig|date=March 10, 2015|accessdate=June 5, 2015}}</ref> ''Salome'' was released on April 7, 2015 by Sargent House,<ref>{{cite web|url=http://www.avclub.com/article/settle-down-marriages-new-record-217325|title=Settle down with Marriages’ new record|publisher=[[The A.V. Club]]|author=David Anthony|date=April 1, 2015|accessdate=June 3, 2015}}</ref> preceded by a U.S. tour with [[Helms Alee]] and followed by a European tour with [[Wovenhand]].<ref>{{cite web|url=http://exclaim.ca/Music/article/marriages_reveal_salome_for_sargent_house|title=Marriages Reveal 'Salome' for Sargent House|publisher=[[Exclaim!]]|author=Gregory Adams|date=January 22, 2015|accessdate=June 5, 2015}}</ref>

==Band members==
'''Current members'''
*[[Emma Ruth Rundle]] - [[Electric guitar|guitar]], [[Singing|vocals]] (2011–present)
*Greg Burns - [[Bass guitar|bass]], keyboards, programming (2011–present)
*Andrew Clinco - [[Drum kit|drums]] (2013–present)

'''Former members'''
*Dave Clifford - [[Drum kit|drums]] (2011-2013)

==Discography==
===Studio albums===
*''[[Salome (Marriages album)|Salome]]'' (2015, [[Sargent House]])

===EPs===
*''[[Kitsune (Marriages EP)|Kitsune]]'' (2012, Sargent House)

==References==
{{Reflist}}

==External links==
*{{Official website|http://marriagesband.com}}

{{Authority control}}

[[Category:Musical groups from Los Angeles]]
[[Category:Musical groups established in 2011]]

Antonio Albergati

2018-05-20T21:59:37Z

Tom.Bot: +{{Authority control}} (1 source from Wikidata), WP:GenFixes on, using AWB

{{Multiple issues|
{{more citations needed|date=February 2017}}
{{cleanup biography|date=February 2017}}
}}
{{Infobox Christian leader
| type = Bishop
| honorific-prefix = Most Reverend
| name = Antonio Albergati
| title = Bishop of Bisceglie
| image =
| alt =
| caption =
| church = [[Catholic Church]]
| archdiocese =
| diocese =
| see =
| term = 1609–1627
| predecessor = [[Alessandro Cospi]]
| successor = [[Nicola Bellolatto]]

| ordination = 2 August 1609
| ordained_by =
| consecration = 23 August 1609
| consecrated_by = [[Giovanni Garzia Mellini]]
| cardinal =
| rank =

| birth_date = 16 September 1566
| birth_place = [[Bologna]], [[Italy]]
| death_date = 13 January 1634 (age 60)
| death_place = [[Bisceglie]], [[Italy]]
| previous_post = [[Apostolic Nuncio to Germany]] (1610–1621) [[Apostolic Nuncio to Portugal|Apostolic Collector to Portugal]] (1621–1624)
| nationality =
}}

'''Antonio Albergati''' (16 September 1566 – 13 January 1634) was a Roman Catholic prelate who served as [[Bishop of Bisceglie]] (1609–1627), [[Apostolic Nuncio to Germany]] (1610–1621), and [[Apostolic Nuncio to Portugal|Apostolic Collector to Portugal]] (1621–1624).<ref name=CathHierAntAlber>[http://www.catholic-hierarchy.org/bishop/balberga.html "Archbishop Antonio Albergati"] ''[[Catholic-Hierarchy.org]]''. David M. Cheney. Retrieved November 24, 2016</ref><ref name=CathHierBisceglie>[http://www.catholic-hierarchy.org/diocese/db251.html "Diocese of Bisceglie"][[Catholic-Hierarchy.org]]''. David M. Cheney. Retrieved October 7, 2016</ref><ref name=GCathBisceglie>[http://www.gcatholic.org/dioceses/former/bisc0.htm "Diocese of Bisceglie"] ''GCatholic.org''. Gabriel Chow. Retrieved October 7, 2016</ref><ref name=CathHierNuncioGermany>[http://www.catholic-hierarchy.org/diocese/dxxde.html "Nunciature to Germany"] ''[[Catholic-Hierarchy.org]]''. David M. Cheney. Retrieved July 18, 2017</ref>

==Biography==
Antonio Albergati was born in [[Bologna]], [[Italy]] on 16 September 1566 and ordained a priest on 2 August 1609.<ref name=CathHierAntAlber />
On 3 August 1609, he was appointed during the papacy of [[Pope Paul V]] as [[Bishop of Bisceglie]].<ref name=CathHierAntAlber />
On 23 August 1609, he was consecrated bishop by [[Giovanni Garzia Mellini]], [[Bishop of Imola]], with [[Domenico Rivarola]], [[Titular Archbishop]] of ''Nazareth'', and [[Antonio d'Aquino]], [[Bishop of Sarno]], serving as co-consecrators.<ref name=CathHierAntAlber />
On 26 April 1610, he was appointed during the papacy of Pope Paul V as [[Apostolic Nuncio to Germany]]<ref name=CathHierAntAlber />
On 15 September 1621, he was appointed during the papacy of Pope Paul V as [[Apostolic Nuncio to Portugal|Apostolic Collector to Portugal]] where he served until his resignation in 1624.<ref name=CathHierAntAlber />
He served as Bishop of Bisceglie until his resignation in 1627.<ref name=CathHierAntAlber /> He died on 13 Jan 1634.<ref name=CathHierAntAlber />

While bishop, he was the principal consecrator of [[Stephen Strecheus]], [[Bishop of Liège|Auxiliary Bishop of Liège]] (1615) and [[Gereon Otto von Gutmann zu Sobernheim]], [[Bishop of Cologne|Auxiliary Bishop of Cologne]] (1616); and the principal co-consecrator of [[Vincenzo Napoli]], [[Bishop of Patti]] (1609).<ref name=CathHierAntAlber />

== References ==
{{Reflist}}

{{s-start}}
{{s-rel|ca}}
{{succession box
| title = [[Apostolic Nuncio to Germany]]
| years = 1610–1621
| before = [[Attilio Amalteo]]
| after = [[Pietro Francesco Montorio]]}}
{{succession box
| title = [[Apostolic Nuncio to Portugal|Apostolic Collector to Portugal]]
| years = 1621–1624
| before = [[Vincenzo Landinelli]]
| after = [[Giovanni Battista Maria Pallotta]]}}
{{succession box
| title = [[Bishop of Bisceglie]]
| years = 1609–1627
| before = [[Alessandro Cospi]]
| after = [[Nicola Bellolatto]]}}
{{s-end}}
{{Authority control}}

{{DEFAULTSORT:Albergati, Antonio}}
[[Category:17th-century Roman Catholic bishops]]
[[Category:Bishops appointed by Pope Paul V]]
[[Category:1566 births]]
[[Category:1634 deaths]]

{{RC-bishop-stub}}

Fort Regent

2018-04-06T01:29:57Z

Tom.Bot: Task 5: fix :Category:Pages using deprecated image syntax; WP:GenFixes on, using AWB

{{Infobox Military Structure
|name=Fort Regent
|partof=
|location=[[Saint Helier]], [[Jersey]]
|map_type=Channel Islands
|coordinates = {{coord|49.1813|-2.1059|type:landmark|display=inline}}
|image=Fort Regent View.JPG
|image_size=300px
|caption=Fort Regent in 2008.
|type=[[Fortification]]
|code=
|built=1806 to 1814
|builder=
|materials=[[Granite]] [[Carboniferous limestone]]
|height=
|used=
|demolished=
|condition=Intact
|ownership=People of Jersey
|open_to_public=Yes
|controlledby=[[States of Jersey]]
|garrison = [[British Army]] [[Royal Militia of the Island of Jersey|Jersey Militia]] [[Wehrmacht]] (1940–1945)
|current_commander =
|commanders =
|occupants =
|battles =
|events =
|image2=Historical and Topographical Description of the Channel Islands 1840 Robert Mudie 10.jpg
|image2_size=300px
|caption2=Fort Regent in the 19th century.
}}

'''Fort Regent''' is a 19th-century fortification, and [[leisure centre]], on Mont de la Ville (Town Hill), in [[Saint Helier, Jersey|St. Helier]], Jersey.<ref>{{cite web|url=http://www.gov.je/leisure/sport/facilities/indoor/pages/fortregent.aspx|title=Fort Regent facilities, bookings and opening hours|accessdate=8 August 2015|work=gov.je}}</ref><ref name="conservation">{{cite web|url=http://www.statesassembly.gov.je/ScrutinyReviewResearches/2009/S-1921-7210-27102009.pdf|title=A Conservation Statement for Fort Regent|author=Education, Sport and Culture|date=September 2006|accessdate=8 August 2015}}</ref> The fort is in close proximity to the fortified South Hill, Engineers Barracks at La Collette, and overlooks the 16th-century [[Elizabeth Castle]] and harbour to the west.

The fort's main features are substantial [[Curtain wall (fortification)|curtain walls]], [[Ditch (fortification)|ditches]], a [[glacis]], [[redoubt]]s, [[bastion]]s, and [[redan]]s (or demi-bastions). A [[Military parade|parade ground]] was in the centre, which is now built upon, and covered with a roof.

==Pre-history of Le Mont de la Ville==
A [[Jersey Dolmens|dolmen]] was located on the hill, prior to the construction of Fort Regent. In 1785, workmen, who were leveling the area for use as a parade ground, uncovered the dolmen.<ref name="conservation"/> The dolmen was gifted to [[Henry Seymour Conway]], who removed it to his home, [[Park Place, Berkshire|Park Place]], near [[Henley-on-Thames]], in 1788.

==Middle Ages and 16th century==
During the Middle Ages the Town Hill, and near-by ''Petit Mont de la Ville'' were used as common land. The ''Chapel of Notre Dame des Pas'' was situated at the foot of the hill during this period, but was demolished by the [[Board of Ordnance]] in 1814.<ref name="conservation"/>

In 1550, [[Edward VI]] ordered the town to be relocated onto the hill, because it would be easier to defend in that position. However, the town was never moved.<ref name="conservation"/>

In 1591, an intention to fortify the hill arose in the form of consent, from the Procureurs of the [[Vingtaine de la Ville|Vingtaine]], to acquire the common land from the people, with their consent, so that fortifications can be constructed. The document referred to letters from [[Elizabeth I]] promising to fortify the top of the hill to provide defence for the town.<ref name="conservation"/> Despite this, no evidence exists of any work being carried out during the 16th century.

==17th century==
In October 1651, during the [[Third English Civil War]], the [[Roundhead|Parliamentarian]] Colonel James Heane, [[Siege|besieged]] Elizabeth Castle by firing [[Shell (projectile)#History|explosive shells]] with [[mortar (weapon)|mortars]] located on high ground between Town Hill and South Hill.<ref>{{cite web|url=http://www.british-civil-wars.co.uk/military/1651-jersey.htm|title=1651: Jersey and the Channel Isles|first=David|last=Plant|date=17 January 2010|accessdate=8 August 2015}}</ref> This bombardment forced the eventual surrender of [[George Carteret]], at Elizabeth Castle, in December 1651.

==18th century==
An illustration by J. Heath, dated 1757, shows the first signs of fortification on the Town Hill, in form of lines, possibly [[Earthworks (engineering)|earthworks]] rather than stone walls.<ref name="conservation"/> A map based on a survey carried out in 1787, under the order of the [[Charles Lennox, 3rd Duke of Richmond|Duke of Richmond]], supports this possibility.<ref name="conservation"/>

A later map produced by James Stead, known as the ''Bouillon Map of 1799'', indicates that the main citadel, at this time, was located on South Hill, rather than the Town Hill.<ref name="conservation"/>

The hill was used in 1781, during the [[Battle of Jersey]], by the [[72nd Regiment, Duke of Albany's Own Highlanders|78th Regiment of Foot]] as a suitable position to prevent the retreat of the invading [[French Army]].

==19th century==
[[File:Fort Regent rampart 1.jpg|thumb|left|Most walls are built from large granite blocks and [[Quoin (architecture)|quoin]]s.]]
[[File:Map Fort Regent Jersey.png|thumb|19th century map: 1: North [[Outwork]]s; 2: East Ditch; 3: East Outworks; 4: Parade ground; 5: [[Counterguard]]; 6: [[Glacis]]; 7: South Hill]]
The construction of the fortress we see today on Town Hill began on 7 November 1806, during the [[Napoleonic Wars]], with the laying of a foundation stone by [[George Don (British Army officer)|George Don]] the [[Lieutenant Governor of Jersey]]. The fort was built using local workers and men from the [[Royal Engineers]], with an average of 800 men working at any given time. This enabled the substantial amount of work to be completed 8 years later, in 1814.

It was given the name Fort Regent in honour of [[George III of the United Kingdom|Prince Regent]], who was King of the United Kingdom of Great Britain and Ireland at this time.<ref name="conservation"/>

The design of the fort is credited to Lieutenant-General John Humfrey, and it is thought that Lieutenant-Colonel John Evelegh would have also worked on the final plans.<ref name="conservation"/>

===Western flank and rampart===
Along the top of the cliffs on the west side, between the West Bastion and Northwest Redan, is a 5.5-m- thick (18 ft) curtain wall that provided protection from attacks on that side.<ref name="conservation"/> Quarrying also steepened the cliffs.

===East Rampart===
A curtain wall, similar to the one on the west, provides protection to bombardment from the east. The East Bastion and south [[redan]]s are positioned behind this wall. The height of this rampart allows for a view to the south-east coast of Jersey, including a view of Icho Tower, a [[Martello tower]] built around 1811.

===Bastions, redans, ditches and glacis===
Two [[bastion]]s are at the fort, one facing west and the other facing east, and four redans, two to the south, and two at the north end. Cannons placed in these areas would have been able to target forces attacking Fort Regent from any direction. The redans are not typical examples because they have more than two sides, are closer to demi-bastions, as seen in [[hornwork]].

The fort has one 210-m-long (689 ft) glacis at the south end, which is a flat, sloping open area of grass, known as the Glacis Field.<ref name="conservation"/> The only road and foot access to the fort is in this area; all other sides are very steep or vertical cliffs.

The East Ditch has a [[masonry]]-faced [[counterscarp]] and scarp, with a further outer ditch in the form of a large cutting. The [[Jersey Eastern Railway]] enlarged the cutting in 1807 to use it as a train station.

===Cannons===
[[File:Cannon at Fort Regent.JPG|thumb|A [[carronade]] in 2008, on the west rampart]]
Positions and embrasures for 100 cannons are in place within the fortress.<ref name="conservation"/> However, a report dated 8 March 1810, counts only 55 cannons and six mortars at Fort Regent.<ref name="conservation"/>

===Water well===
No [[Water well|well]] existed on the hill, so between December 1806 and October 1808, a deep well-shaft was blasted into the notoriously hard Jersey granite to a depth of about 71 m (235 ft)<ref name="conservation"/> by blasting a shaft of an average diameter of 8 ft with gunpowder charges. The Fort Regent well is believed to be the deepest well-shaft in the island [excluding artesian bores], and is a stunning tribute to the persistence and ingenuity of Georgian military engineers. When the sappers and miners blasted through to the spring, at a depth of 221 ft below the well-curb in the underground well-head chamber, they did so unexpectedly and water rose rapidly in the shaft. The miners were hanging in a basket halfway up the shaft. The major of engineers in charge of the works recorded that "great difficulty was experienced in recovering the men to the surface before they were drowned by the inrush from the Spring". The construction records, including the commanding officer's daily diary, are now in the National Archives at Kew, and were researched by the architect and the main contractor – C.G. Dumond (Builders) Ltd. – for useful information during the conversion of the fort into a leisure centre in the 1970s. The granite was drilled with "jumper drill" irons hit with sledgehammers. As originally commissioned, water was raised to the surface by an above-ground "horse whim" at parade-ground level, but this proved to be both time-consuming and unpopular with the soldiery, and lasted for only a year after the official opening in 1814. The first rocking-beam pumps, operated in series all the way down the well-shaft by one long pump-rod, and operated by a man-powered capstan through a line shaft and gearing, were delivered and installed by Henry Maudeslay and Son in 1815, in a double circular underground chamber excavated just below the level of the parade ground. One chamber contained the capstan, and connected to the adjacent chamber which contained the well-head machinery, and the mouth of the shaft. At some later time, the man-powered capstan was converted to operate by donkey power. This method, too, proved less efficient than was thought desirable; it proved difficult to get the donkeys into and out of the chambers through the long, sloping tunnel which originally began at ground level in the East Bastion, The motive power was changed again to be provided by a small steam engine. The steam engine, in turn, gave way to a gas engine, which is still ''in-situ'', a rare survival of that late-Victorian technology, along with Henry Maudeslay's original well-head machinery from 1815. During the 1970s reconstruction, the local government's client organisation – the Fort Regent Development Committee (FRDC) ordered that all of the well-shaft access-ladders, cast-iron floor-gratings (supplied from Ironbridge by order of the War Office), and pump rods and valves – should be removed from the well-shaft and scrapped, "because the Fort requires the well-water to be used for commercial and domestic supplies within the new leisure complex, and contamination from this old machinery cannot be risked". This act of "cultural vandalism" was carried out under the supervision of the clerk of the works, Mr Greenwell, which was protested by a few knowledgeable persons at the time, but the reconstruction contract was politically sensitive; knowledge of the machinery's destruction and scrapping was limited to members of the managing FRDC, and to the project architect and main contractors' staff. So, the general public of Jersey, who might have expressed their concerns, never became aware of the destruction until long after the fact: even now it is not common knowledge amongst those who care about preserving as much of Jersey's historic past as possible. Though the main contractor salvaged all the recovered machinery parts, and handed them over to the client, as required under the construction contract for "all antiquities recovered from the Site", the Fort Regent authorities scrapped them 2 years after contract completion as being "of no historic interest and not worthy of preservation". Access to the underground well-head and machinery chambers down a steep set of concrete steps and a steeply sloping tunnel is deemed too dangerous under modern health and safety concerns, this historic 19th-century well-head machinery seems doomed to be forever hidden away from members of the public. During the clearing and cleaning of the well-shaft in 1979, the main contractor discovered a 40-ft-tall flat sheet of granite caused by a natural fracture-plane which formed one face of the shaft about 100 ft below the surface. It was so hot, water dripping onto it from further up the shaft turned to steam, hot enough to blister a bare hand accidentally brought into contact with the granite surface. This may be the only example in the island of geothermal heat.

==Signal station==
The Town Hill has been used as a commercial signal station since the late 18th century, prior to the construction of Fort Regent.<ref>{{cite web|url=http://www.maritime.je/the-signal-station/4555760496|title=The Signal Station|accessdate=8 August 2015}}</ref>

==20th century==
The last British force to garrison Fort Regent was the [[Royal Militia of the Island of Jersey]], who left the fort on 20 June 1940, and served in the UK as part of [[The Hampshire Regiment]]. The militia reformed as the Jersey Field Squadron in 1987, and are located in the Engineers Barracks at La Collette.

===German occupation===
[[File:Fort Regent German Occupation 4.jpg|thumb|One of the gun emplacements dating from the occupation]]
During the [[German occupation of the Channel Islands]], the German forces made some additions to the fort, including [[Anti-aircraft warfare|flak cannons]]. Some of these concrete structures remain today.

===Storage===
After the German occupation, the fort was used as a storage area for potatoes, wine, and coal.<ref>{{cite web|url=https://www.mygov.je//Planning/Pages/HistoricEnvironmentDetail.aspx?s=3&r=HE1195 |title=Historic Environment Detail – Historic Document Reference : HE1195|publisher=Mygov.je |date=2 September 1939|accessdate=16 October 2013}}</ref> The nearby [[La Collette Power Station|power station]] was coal-fired at the time.

===Leisure centre===
[[File:Fort regent pool.JPG|thumb|The swimming pool building can be seen in the upper area of the photo]]
In December 1967, the States of Jersey made a decision to adapt the site into a leisure centre.<ref>{{cite web|url=http://www.statesassembly.gov.je/ScrutinyReviewSubmissions/Submission%20-%20Fort%20Regent%20-%20Member%20of%20the%20Public%20Submission%202.23%20Appendix%201%20-%2031%20March%202009.pdf |title=The Future Use of Fort Regent – the Jersey Sports Village and Community Centre – a basis for change|first=Roy|last=Travert|date=September 2003|publisher=The Fort Users Association |accessdate=1 April 2013}}</ref> The swimming pool located on the glacis field, which opened in 1971 and closed in 2009,<ref>{{cite news|newspaper=[[Jersey Evening Post]]|url=http://jerseyeveningpost.com/news/2014/09/02/should-fort-regent-swimming-pool-be-refurbished/temps-passe-fort-pool-1971/|date=2 September 2014|accessdate=8 August 2015|title=Fort Regent swimming pool might finally return}}</ref> was the first modern addition to the fort.

==See also==
*[[Mont Orgueil]]

==References==
{{reflist}}

==External links==
{{Commons category}}

[[Category:Forts in Jersey]]
[[Category:Sports venues in Jersey]]
[[Category:19th century in Jersey]]
[[Category:Buildings and structures in Saint Helier]]
[[Category:Tourist attractions in Jersey]]
[[Category:Archaeological sites in Jersey]]
[[Category:World War II sites in the Channel Islands]]
[[Category:German occupation of Jersey during World War II]]
[[Category:1806 establishments in the British Empire]]

A Famosa

2018-04-05T22:28:29Z

Tom.Bot: Task 4: fix :Category:Pages using deprecated image syntax; WP:GenFixes on, using AWB

{{EngvarB|date=January 2015}}
{{Use dmy dates|date=January 2015}}
{{Infobox Military Structure
|name = A Famosa
|native_name = Kota A Famosa {{ms icon}}
|partof =
|location = [[Malacca]], [[Malaysia]]
|image = A Famosa Fortress.JPG
|caption = The surviving gate of the A Famosa Portuguese fort in Malacca.
|image2 = Melaka-Porta-de-Santiago-2Cimg2194.jpg
|image2_size = 300px
|caption2 = A view of the gate of the A Famosa fort from the rear.
|map_type =
|map_size =
|map_caption =
|type =
|coordinates = {{coord|2|11|29.82|N|102|15|1.10|E|type:landmark|display=inline,title}}
|code =
|built = 1511
|builder = [[Portuguese Empire]]
|materials =
|height =
|used = 1511–1807
|demolished = 10 August 1807 (except for a small gate house)
|condition = Largely destroyed except for a few remaining structures
|ownership =
|open_to_public = Yes
|controlledby = [[Kingdom of Portugal|Portugal]] (1511–1641) [[Dutch Republic|Netherlands]] (1641–1795) [[United Kingdom of Great Britain and Ireland|Britain]] (1795–1807)
|garrison =
|current_commander =
|past_commanders = [[Afonso de Albuquerque]]
|occupants =
|battles =
|events =
}}
[[File:Stad en Kasteel Malacca.JPG|thumb|right|The town and fortress of Malacca (1780)|280px]]
[[File:A Famosa - Reconstruction of watchtower.jpg|thumb|left|Reconstruction of watchtower on top of the unearthed remains|250px]]
[[File:A Famosa anitary sewer line remain.JPG|thumb|right|A Famosa [[sanitary sewer]] line ruin|250px]]

'''A Famosa''' ({{lang-ms|Kota A Famosa}}; "The Famous" in [[Portuguese language|Portuguese]]) was a Portuguese [[fortress]] located in [[Malacca]], [[Malaysia]]. It is among the oldest surviving [[Europe]]an architectural remains in [[Southeast Asia]] and the [[Far East]]. The ''Porta de Santiago'', a small gate house, is the only part of the fortress which still remains today.

The name is often mispronounced {{IPAc-en|eɪ}} ''Famosa'', even among Malaysians, as though the Portuguese [[article (linguistics)|definite article]] ''a'' were the English letter ''[[A]]''. A more authentic pronunciation would be {{IPAc-en|ɑː}} ''Famosa''.

==History==

In 1511, a [[Kingdom of Portugal|Portuguese]] fleet arrived under the command of [[Afonso de Albuquerque]]. His forces [[Capture of Malacca (1511)|attacked and defeated]] the armies of the [[Sultanate of Malacca|Malacca Sultanate]]. Moving quickly to consolidate his gains, Albuquerque had the fortress built around a natural hill near the sea. [[Afonso de Albuquerque|Albuquerque]] believed that [[Malacca]] would become an important port linking [[Portugal]] to the [[Spice Route]] in China. At this time other [[Portuguese people|Portuguese]] were establishing outposts in such places as [[Portuguese Macau|Macau]], [[Ming China|China]] and [[Portuguese India|Goa]], India to create a string of friendly ports for ships heading to [[Ming China]] and returning home to [[Kingdom of Portugal|Portugal]].

The [[fortress]] once consisted of long [[defensive wall|ramparts]] and four major [[towers]]. One was a four-story [[keep]], while the others held an [[ammunition]] storage room, the residence of the [[Captain (land and air)|captain]], and an officers' quarters. Most of the village clustered in town houses inside the [[fortress]] walls. As Malacca's population expanded it outgrew the original fort and extensions were added around 1586.

The fort [[Battle of Malacca (1641)|changed hands in 1641]] when the [[Dutch Republic|Dutch]] drove the [[Kingdom of Portugal|Portuguese]] out of Malacca.<ref>Borschberg, Peter., “Ethnicity, Language and Culture in Melaka during the Transition from Portuguese to Dutch Rule”, Journal of the Malaysian Branch of the Royal Asiatic Society, 83.2, (2010): 93-117; Borschberg, P., The Singapore and Melaka Straits: Violence, Security and Diplomacy in the 17th Century (Singapore: NUS Press, 2010).</ref> The Dutch renovated the gate in 1670, which explains the logo "ANNO 1670" inscribed on the gate's [[arch]]. Above the arch is a bas-relief logo of the [[Dutch East India Company]].

The [[fortress]] changed hands again in the late 18th century when the [[Dutch Republic|Dutch]] handed it over to the [[Kingdom of Great Britain|British]] to prevent it from falling into the hands of [[Napoleon]]'s expansionist [[First French Empire|France]]. The [[England|English]] were wary of maintaining the [[fortification]] and ordered its destruction in 1806. The fort was almost totally demolished but for the timely intervention of [[Sir Stamford Raffles]], the founder of modern [[Singapore]], who was sent on sick leave from [[Penang]] to Malacca in 1807. It was Captain William Farquhar, tasked with the destruction of the fort and town, who decided to save two of the gateways to the fort, including the Santiago Gate, as well as the stadthuys, church and jail.

==Archaeological finding==
In late November 2006, a structure part of the fort, believed to be the [[Bastion Middleburg]] was accidentally uncovered during the construction of 110 meter revolving tower in Malacca Town.<ref name="nst.com.my">Lee, Cynthia. [[New Straits Times]]. [http://www.nst.com.my/Current_News/nst/Friday/NewsBreak/20061201192448/Article/index_html Excavation for Malacca tower project unearths ruins of Dutch fort]{{dead link|date=October 2016 |bot=InternetArchiveBot |fix-attempted=yes }}. 1 December 2006</ref> The construction of the tower was ceased and its site was subsequently shifted to the popular district of Bandar Hilir on Jalan Merdeka where it was officially opened to the public on 18 April 2008.
Malacca Museums Corporation suspects the structure was built by the Dutch during the Dutch occupation of Malacca from 1641 to 1824.

Earlier in June 2004, a watchtower named '''Santiago Bastion''' was discovered during the construction of Dataran Pahlawan.<ref>The Star. [https://web.archive.org/web/20061206033124/http://thestar.com.my/news/story.asp?file=%2F2006%2F12%2F4%2Fnation%2F16216405&sec=nation Old watchtower may be under site]. 4 December 2006.</ref>

==References==
{{reflist}}

===Notations===
* {{cite book|last=De Witt|first=Dennis|title=Melaka from the Top|location=Malaysia|publisher=Nutmeg Publishing|year=2010|isbn=978-983-43519-2-2}}

==External links==
{{commons category|A Famosa}}
* [http://www.tourism.gov.my/en/my/web-page/places/states-of-malaysia/melaka/st-pauls-hill-afamosa Tourism Malaysia - St.Paul's Hill (A'Famosa) ]
* [http://allmalaysia.info/news/attraction.asp?id=590&pt=7 A Famosa at All Malaysia.info]
* [https://web.archive.org/web/20081208111442/http://courses.nus.edu.sg/course/ellhpj/resources/abdullah3.HTM Description of the fort during English rules by ''Hikayat Abdullah''.]

{{Malaysian colonial architecture}}
{{Malaysian historical architectures and sites}}

[[Category:Buildings and structures in Melaka City]]
[[Category:Forts in Malaysia]]
[[Category:Portuguese forts]]
[[Category:Portuguese Malacca]]
[[Category:Tourist attractions in Melaka]]
[[Category:1511 establishments in Portuguese Malacca]]
[[Category:1641 disestablishments in Portuguese Malacca]]

Knappogue Castle

2018-04-05T22:26:49Z

Tom.Bot: Task 4: fix :Category:Pages using deprecated image syntax; WP:GenFixes on, using AWB

{{Infobox military installation
| name = Knappogue Castle
| native_name = Caisleán na Cnapóige
| location = [[County Clare]], [[Republic of Ireland|Ireland]]
| image = Knappogue Castle.jpg
| image_size = 300px
| caption = Knappogue Castle
| map_type = Ireland
| map_relief =
| map_size =
| map_alt =
| map_caption =
| type =
| coordinates = {{coord|52.793919|-8.831597|type:landmark|display=inline}}
| code =
| built = {{Start date|1467}}
| builder = Seán Mac Conmara (MacNamara)
| materials =
| height =
| used =
| condition =
| ownership = Shannon Heritage
| open_to_public = yes
| occupants =
| battles =
| events =
| image2 =
| caption2 =
}}

'''Knappogue Castle''' ([[Irish language]]: ''Caisleán na Cnapóige''<ref name="Library"/>) is a [[tower house]], built in 1467 and expanded in the mid-19th century, located in the parish of [[Quin, County Clare|Quin]], [[County Clare]], [[Republic of Ireland|Ireland]]. It has been restored and is open to guided tours.

==History==
The original castle was built in 1467 by [[Seán Mac Conmara]] (or [[MacNamara]]), son of Sioda MacNamara, and is a good example of a late medieval [[tower house]]. The castle's name translates as "castle of the place abounding in little hills".<ref name="Library">{{cite web|url= http://www.clarelibrary.ie/eolas/coclare/places/quininterest.htm |title=Quin, Places of Interest|publisher=Clare Library| date= |first= |last=|accessdate=16 August 2013}}</ref>

In 1571, the castle became the seat of the MacNamara (Mac Conmara) sept, the Earls of [[West Clancullen]]. [[Donnchadh Mac Conmara]] was a leader of the [[Irish Rebellion of 1641]] and Knappogue remained in MacNamara hands throughout the [[Irish Confederate Wars]] of the 1640s. After the [[Cromwellian conquest of Ireland]] (1649–53) it was confiscated in accord with the [[Adventurers' Act]] and its new owner was a [[roundhead]], Arthur Smith.<ref name="History">{{cite web|url= http://www.shannonheritage.com/DaytimeAttractions/KnappogueCastleWalledGarden/HistoryofKnappogueCastle/ |title=History of Knappogue Castle|publisher=Shannon Heritage| date= |first= |last=|accessdate=16 August 2013}}</ref>

Arthur Smith occupied the castle from 1659 to 1661. After the monarchy was [[Restoration (Ireland)|restored in 1660]], Knappogue was returned to its MacNamara owners. Eventually, Francis MacNamara, [[High Sheriff of Clare]] in 1789, sold the castle to the Scott family of [[Cahircon]] in 1800; the latter carried out major restoration and extension work.<ref name="Library"/> In 1837, the castle was owned by William Scott{{Who|date=July 2016|reason=Many William Scotts and may not have been any of the famous.}}.<ref name="Estate1">{{cite web|url= http://landedestates.nuigalway.ie:8080/LandedEstates/jsp/property-show.jsp?id=2020|title= Landed estates database: Knappogue Castle|publisher= NUI Galway| date= |first= |last= |accessdate=17 August 2013}}</ref>

In 1855, the castle was acquired by [[Baron Dunboyne|Theobold Fitzwalter Butler, 14th Baron Dunboyne]]. It became the family seat of the Dunboyne family. They continued the restoration work of the Scotts, adding a drawing-room, the long room and a west wing, including the clock tower and the gateway.<ref name="Library"/> The remodelling was done by architects [[James Pain]] and his brother [[George Richard Pain]].<ref name="Architect"/>

During the [[Irish war of independence|War of Independence]] (1919–21), Clare County Council held their meetings at Knappogue Castle where they were guarded by the East Clare Flying Column. Michael Brennan, Commander of the East Clare Brigade also used the castle as his headquarters during that time.<ref name="Library"/><ref>[http://www.clarelibrary.ie/eolas/claremuseum/acquisitions/framed_glazed_photo_east_clare_brigade.htm Framed and Glazed photograph, East Clare Brigade]</ref>

In 1927, Knappogue demesne was purchased by the [[Irish Land Commission]] and the castle became the possession of the Quinn family,<ref name="Library"/> local farmers who allowed it to fall into disrepair.<ref name="History"/> The castle and lands were then purchased in 1966 by [[Mark Edwin Andrews]], former Assistant Secretary of the United States Navy, from Houston, Texas. He and his wife Lavonne (a prominent American architect), in collaboration with what was then Shannon Free Airport Development Company (today Shannon Heritage) and [[Fáilte Ireland|Bord Failte Eireann]] carried out an extensive restoration around 1969.<ref name="Library"/> This was intended to accommodate use of the castle as restaurant and private residence.<ref name="Architect">{{cite web|url= http://www.buildingsofireland.ie/niah/search.jsp?type=record&county=CL&regno=20404217 |title=National Inventory of Architectural Heritage: Knappogue Castle |publisher=Department of Arts, Heritage and the Gaeltacht| date= |first= |last=|accessdate=16 August 2013}}</ref>

Their work returned much of the castle to its former 15th century state while encompassing and retaining later additions that chronicle the continuous occupation of the castle.<ref name="Library"/> The Andrews later leased part of the castle to the Irish Government as a cultural and tourist facility for a nominal rent.<ref name="Library"/>

==Today==
[[Shannon Development]] purchased the castle in 1996. Today, the castle is used as a venue for weddings and medieval banquets and offers guided tours.<ref name="History"/>

Dating from 1817, the {{convert|1.248|acre|ha|adj=on}} garden is now restored to its former state. The walls of the garden have been refurnished with climbing roses, grapevines and many varieties of [[clematis]].

There is also a whiskey named "[[Knappogue Castle Irish Whiskey|Knappogue Castle]]" produced by the Castle Brands company, currently bottling liquor produced by [[Bushmills]].<ref name="Jackson">{{cite book|last=Jackson|first=Michael|title=Whisky - the definite world guide |publisher=Dorling Kindersleigh (Penguin)|year=2005|isbn=978-0-7513-4434-9| pages=}}</ref>{{rp|200}}

==References==
{{reflist}}

==External links==
{{commons category}}
* [http://www.shannonheritage.com/DaytimeAttractions/KnappogueCastleWalledGarden/ Shannon Heritage.com] - official site
* [http://www.buildingsofireland.ie/niah/search.jsp?type=record&county=CL&regno=20404217 National Inventory of Architectural Heritage - Castle]
* [http://www.buildingsofireland.ie/niah/search.jsp?type=record&county=CL&regno=20404219 National Inventory of Architectural Heritage - Gateway]

{{Dalcassians}}
{{Historic Irish houses}}
{{Tourism in County Clare}}

[[Category:Castles in County Clare]]
[[Category:Historic house museums in the Republic of Ireland]]
[[Category:Museums in County Clare]]
[[Category:Gardens in County Clare]]

Carbury Castle

2018-04-05T21:56:47Z

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{{more footnotes|date=January 2018}}
{{Infobox military structure
|name = Carbury Castle
|image = Ruins of Carbury, County Kildare.jpg
|image_size = 300px
|caption = Carbury [[south-west]] [[facade]]
|type = [[Motte and Bailey]]
|built = before 1220
|builder = [[Meiler FitzHenry]]
|materials = Limestone
|used = Until 1600's
|condition = Ruined
|ownership = [[Bermingham]]
|open_to_public = Yes
|battles =
}}
[[File:Windows of Carbury, County Kildare.jpg|thumb|left|Window detail]]
'''Carbury Castle''' is a [[castle]] situated in the townland of [[Carbury]], on the borders of [[Kildare]] and [[Offaly]]. The area is dominated by the ruins of this great [[Tudor mansion]] set atop Carbury Hill, which was also known as [[Fairy Hill]]. The motte on the hill was probably built by [[Meiler FitzHenry]] who was granted the area by Strongbow. It was acquired by the de Berminghams in the 14th century, before being taken by the native Irish in the 15th century. In 1588 it was granted to the Colley family, ancestors of the Dukes of Wellington, who built a large stronghouse in the 17th century.

==History==
The Castle was first built by the [[Norman invasion of Ireland|Normans]] and the motte on the hill was probably built by [[Meiler FitzHenry]]. The central scenic focus of Carbury Hill is the ruins of the Tudor mansion of the Colleys, which was also known as Fairy Hill.<ref>{{cite web|title=County Kildare towns - Carbury |work=Go Ireland.com |url=http://www.countykildare.com/kildare_carbury.htm |accessdate=2007-09-14 |deadurl=yes |archiveurl=https://web.archive.org/web/20071117015703/http://www.countykildare.com/kildare_carbury.htm |archivedate=2007-11-17 |df= }}</ref> who was granted the area by [[Richard de Clare, 2nd Earl of Pembroke|Strongbow]]. It was acquired by the de Berminghams in the 14th century, from whom it passed by inheritance to the Preston family, who held the title [[Baron Gormanston]], before being taken by the native Irish in the 15th century. The 1st Earl of Shrewsbury, John Talbot, later also Earl of Wexford, Earl of Waterford and Baron of Dungarvan, rebuilt Carbury Castle in 1446-1447.<ref>Dublin, 1789, Moore: Archdall and Lodge, The Peerage of Ireland, pp.120-122</ref>

From 23 October 1554 a 21-year lease was granted to Sir [[Henry Colley (died 1584)|Henry Colley]](the [[patrilineal]] ancestor of the [[Dukes of Wellington]]);<ref>Fiant no.53, [[Philip & Mary]], cited in Appendix IV, 9th Report of the Deputy Keeper of Public Records in Ireland.</ref> this was renewed, and the Colley family built a large stronghouse on the hilltop in the 17th century, now a ruin.<ref>{{cite web |title=Carbury Castle, County Kildare |work=Look Around Ireland |url=http://www.lookaroundireland.com/castlesinteractive/ipix/carbury.htm |accessdate=2007-09-14 |deadurl=yes |archiveurl=https://web.archive.org/web/20070830102823/http://www.lookaroundireland.com/castlesinteractive/ipix/carbury.htm |archivedate=2007-08-30 |df= }}</ref>

== References ==

{{reflist}}

Ballinskelligs Castle

2018-04-05T21:46:37Z

Tom.Bot: Task 4: fix :Category:Pages using deprecated image syntax; WP:GenFixes on, using AWB

{{Infobox military structure
|name = Ballinskelligs Castle
|native_name =
|partof =
|location = [[Ballinskelligs]], [[County Kerry|Kerry]]
|image = Castles_of_Munster,_Ballinskelligs,_Kerry_-_geograph.org.uk_-_1392857.jpg
|image_size = 300px
|caption = Ruins of Ballinskelligs Castle.
|map_type = Ireland
|map_size =
|map_alt =
|map_caption = Location within Ireland
|type =
|coordinates = {{coord|51.8200970|-10.2705060|type:landmark|display=inline}}
|code =
|built =
|builder =
|materials =
|height =
|used =
|demolished =
|condition =
|ownership =
|open_to_public =
|controlledby =
|garrison =
|current_commander =
|commanders =
|occupants =
|battles =
|events =
|image2 =
|caption2 =
}}

'''Ballinskelligs Castle''' is a castle located near [[Ballinskelligs]] on the [[Iveragh Peninsula]] in [[County Kerry|Kerry]], [[Republic of Ireland|Ireland]].<ref name=History>{{cite web|title=Ballinskelligs Castle|url=http://ballinskelligscastle.com/history.html|accessdate=20 August 2012|author=Ballinskelligs Castle Committee}}</ref> The castle is situated on the western shore of Ballinskelligs Bay, on a narrow promontory which is subject to heavy erosion.<ref name=Excavation1>{{cite web|last=Sheehan|first=John|title=Ballinskelligs Castle, Ballinskelligs|url=http://www.excavations.ie/Pages/Details.php?Year=&County=Kerry&id=3538|publisher=[[Department of the Environment, Heritage and Local Government]]|accessdate=21 August 2012|year=1988}}</ref> The castle was constructed by the [[MacCarthy dynasty|MacCarthy Mórs]] in the 16th century to protect the bay from pirates, and possibly in order to charge a tariff on incoming trade vessels.<ref name=History />

== References ==
{{reflist}}

{{commons category|Ballinskelligs Castle}}

[[Category:Castles in County Kerry]]

{{Ireland-castle-stub}}
{{Kerry-geo-stub}}

Phytophthora ramorum

2018-03-23T12:30:18Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q3382075}} (3 sig. taxon IDs); WP:GenFixes using AWB

{{Italic title}}
{{Taxobox
| name = ''Phytophthora ramorum
| image = Sudden_Oak_Death.jpg
| image_width = 240px
| image_caption = Canker on an infected oak
| domain = [[Eukaryota]]
| unranked_regnum = [[SAR supergroup|SAR]]
| superphylum = [[Heterokonta]]
| classis = [[Oomycetes]]
| ordo = [[Pythiales]]
| familia = [[Pythiaceae]]
| genus = ''[[Phytophthora]]''
| species = '''''P. ramorum'''''
| binomial = ''Phytophthora ramorum''
| binomial_authority = Werres et al. 2001
}}

'''''Phytophthora ramorum''''' is the [[oomycete]] [[plant pathogen]] known to cause the disease '''sudden oak death''' ('''SOD'''). The disease kills [[oak]] and other species of trees and has had devastating effects on the oak populations in [[California]] and [[Oregon]], as well as being present in [[Europe]]. Symptoms include bleeding [[canker]]s on the tree's [[Trunk (botany)|trunk]] and [[Forest dieback|dieback]] of the [[foliage]], in many cases eventually leading to the death of the tree.

''P. ramorum'' also infects a great number of other plant species, significantly woody ornamentals such as ''[[Rhododendron]]'', ''[[Viburnum]]'', and ''[[Pieris (plant)|Pieris]]'', causing foliar symptoms known as ramorum dieback or ramorum blight. Such plants can act as a source of [[wikt:inoculum|inoculum]] for new infections, with the [[pathogen]]-producing [[spore]]s that can be transmitted by rainsplash and [[rainwater]].

''P. ramorum'' was first reported in 1995, and the origins of the pathogen are still unclear, but most evidence suggests it was repeatedly introduced as an exotic species.<ref>{{cite journal|last=Grünwald|first=N. J.|author2=Garbelotto, M. |author3=Goss, E. M. |author4=Heungens, K. |author5=Prospero, S. |title=Emergence of the sudden oak death pathogen ''Phytophthora ramorum''|journal=Trends in Microbiology|year=2012|volume=20|pages=131–138|url=http://www.cell.com/trends/microbiology/abstract/S0966-842X(11)00227-7|doi=10.1016/j.tim.2011.12.006|pmid=22326131|issue=3}}</ref> Very few control mechanisms exist for the disease, and they rely upon early detection and proper disposal of infected plant material.

==Presence==
The disease is known to exist in California's coastal region between [[Big Sur]] (in [[Monterey County, California|Monterey County]]) and southern [[Humboldt County, California|Humboldt County]]. It is confirmed to exist in all coastal counties in this range, as well as in all immediately inland counties from [[Santa Clara County, California|Santa Clara County]] north to [[Lake County, California|Lake County]]. It has not been found east of the [[California Coast Ranges]], however. It was reported in [[Curry County, Oregon]] (just north of the California border), in 2001. [[Sonoma County, California|Sonoma County]] has been hit hardest, having more than twice the area of new mortality of any other county in California.<ref>[http://www.sonoma-county.org/des/pdf/sodsr_plan.pdf Sonoma County Sudden Oak Death Strategic Response Plan, University of California Cooperative Extension, Sonoma County and the Sonoma County Department of Emergency Services (February, 2008)]</ref>

About the same time, a similar disease in continental [[Europe]] and the [[United Kingdom|UK]] was also identified as ''Phytophthora ramorum''.<ref name="parkelucas"/>

==Hosts and symptoms==

===In North America===
[[File:Sudden oak death IMG 0223.JPG|thumb|right|A hillside in Big Sur, California, devastated by sudden oak death]]
It was first discovered in California in 1995 when large numbers of [[tanoak]]s (''[[Notholithocarpus densiflorus]]'') died mysteriously, and was described as a new species of ''[[Phytophthora]]'' in 2000. It has subsequently been found in many other areas, including [[Great Britain|Britain]], [[Germany]], and some other [[United States|U.S.]] states, either accidentally introduced in nursery stock, or already present undetected.

In tanoaks, the disease may be recognized by [[wilting]] new [[shoot]]s, older leaves becoming pale green, and after a period of two to three weeks, foliage turning brown while clinging to the branches. Dark brown [[sap]] may stain the lower trunk's bark. Bark may split and exude gum, with visible discoloration. After the tree dies back, suckers try to sprout the next year, but their tips soon bend and die. [[Ambrosia beetle]]s (''Monarthrum scutellare'') will most likely infest a dying tree during midsummer, producing piles of fine white dust near tiny holes. Later, [[bark beetle]]s (''Pseudopityophthorus pubipennis'') produce fine, red boring dust. Small black domes, the fruiting bodies of the ''[[Hypoxylon]]'' fungus, may also be present on the [[Bark (botany)|bark]]. Leaf death may occur more than a year after the initial infection and months after the tree has been girdled by beetles. {{Citation needed|date=December 2014}}

In [[coast live oak]]s and [[Quercus kelloggii|Californian black oak]]s, the first symptom is a burgundy-red to tar-black thick sap bleeding from the bark surface. These are often referred to as bleeding [[canker]]s.

In addition to oaks, many other forest species may be hosts for the disease; in fact, it was observed in the USA that nearly all woody plants in some Californian [[forest]]s were susceptible to ''P. ramorum''.<ref>Rizzo, D.M., et al. Phytophthora ramorum and Sudden Oak Death in California 1: Host Reletionships. in Fifth Symposium on Oak Woodlands: Oaks in California's Changing Landscapes. 2001. San Diego, California.</ref> including [[rhododendron]], [[madrone]] (''Arbutus menziesii''), evergreen [[huckleberry]] (''Vaccinium ovatum''), [[California bay laurel]] (''Umbellularia californica''), buckeye (''[[Aesculus californica]]''), [[bigleaf maple]] (''Acer macrophyllum''), [[toyon]] (''Heteromeles arbutifolia''), [[manzanita]] (''Arctostaphylos spp.''), [[coast redwood]] (''Sequoia sempervirens''), [[Douglas fir]] (''Pseudotsuga menziesii''), coffeeberry (''[[Rhamnus californica]]''), [[honeysuckle]] (''Lonicera hispidula''), and Shreve oak (''[[Quercus parvula]]''). ''P. ramorum'' more commonly causes a less severe disease known as ramorum dieback/leaf blight on these [[Host (biology)|host]]s. Characteristic symptoms are dark spots on [[foliage]] and in some hosts the dieback of the stems and twigs.<ref>Werres, S., et al., Phytophthora ramorum sp. nov., a new pathogen on Rhododendron and Viburnun. Mycological Research, 2001. 105(10): p. 1155–1165.</ref> The disease is capable of killing some hosts, such as rhododendron, but most survive. Disease progression on these species is not well documented. Redwoods exhibit needle discoloration and cankers on small branches, with purple lesions on sprouts that may lead to sprout mortality.

===In Europe===
[[File:Phytophtora ramorum.png|thumb|right|Leaf death caused by ''P. ramorum'']]
In Europe, ''Ramorum'' blight was first observed on rhododendron and viburnum in the early 1990s,<ref name="parkelucas">{{Cite journal | doi = 10.1094/PHI-I-2008-0227-01| title = Sudden oak death, ramorum leaf blight, ramorum shoot blight| journal = The Plant Health Instructor| year = 2008| last1 = Parke}}</ref> where it was initially found mainly on container-grown plants in nurseries.<ref name="fera">{{cite web |url=http://www.fera.defra.gov.uk/plants/plantHealth/pestsDiseases/phytophthora/pRamorum/ |title=Phytophthora ramorum |publisher=FERA | accessdate=17 February 2014}}</ref> The principal symptoms were leaf and twig blight.<ref name="eppo">{{cite web |url=http://www.eppo.int/QUARANTINE/Alert_List/fungi/PHYTRA.htm |title=Phytophthora ramorum |publisher=European and Mediterranean Plant Protection Organization | accessdate=17 February 2014}}</ref> By 2007, it had spread throughout nurseries and retail centers in 16 European countries, and had been detected in gardens, parks, and woodlands in at least eight countries.<ref name="parkelucas"/> It has not caused significant harm to European oak species.<ref name="eppo"/>

In 2009, the pathogen was found to be infecting and killing large numbers of Japanese larch trees (''[[Larix kaempferi]]'') in the [[United Kingdom]] at sites in the [[England|English]] counties of [[Somerset]], [[Devon]], and [[Cornwall]].<ref name="bbc1">{{cite news |url=http://www.bbc.co.uk/news/science-environment-22575543 |title= UK Tree health recommendations aim to 'stop the spread|publisher=BBC News |date=20 May 2013 |accessdate=17 February 2014}}</ref> It was the first time in the world that ''Phytophthora ramorum'' had been found infecting this species.<ref name="bbc2">{{cite news |url=http://www.bbc.co.uk/news/science-environment-19167307 |title='Unprecedented threat' for UK trees from pests |publisher=BBC News |date=3 September 2012 |accessdate=17 February 2014}}</ref> Since then, it has also been found extensively in larch plantations in [[Wales]]<ref>{{cite news|publisher=BBC|title=Thousands of Afan Forest trees planted after infected larch|date=21 February 2015|url=http://www.bbc.co.uk/news/uk-wales-south-west-wales-31559342|accessdate=23 February 2015}}</ref> and in southwest [[Scotland]], leading to the deliberate felling of large areas.<ref name="fera"/> The UK [[Forestry Commission]] noted that eradication of the disease would not be possible, and instead adopted a strategy of containing the disease to reduce its spread.<ref name="bbc2"/> Symptoms of the disease on larch trees include dieback of the tree's crown and branches, and a distinctive yellowing or ginger colour beneath the bark.<ref name="bbc2"/> In August 2010, the disease was found in Japanese larch trees in Counties [[Waterford]] and [[County Tipperary|Tipperary]] in [[Ireland]].<ref name="dafm">{{cite news |url=http://www.agriculture.gov.ie/press/pressreleases/2010/august/title,45756,en.html |title=Disease found in Japanese Larch Trees in Ireland |publisher=Department of Agriculture, Food & the Marine |date=17 August 2010 |accessdate=17 February 2014}}</ref> It had spread to Japanese larch plantations across the south of the country by February 2014.<ref>{{cite news |last=Roche |first=Barry |url=http://www.irishtimes.com/news/environment/cork-s-gougane-barra-forest-park-closing-due-to-tree-fungus-1.1643902 |title=Coillte urged to clarify extent of larch disease in Gougane Barra |publisher=Irish Times |date=6 February 2014 |accessdate=22 November 2016}}</ref> [[Coillte]], who owned twenty forests where the disease was present, felled 16,000 trees in one of its forests, having already felled 150 hectares to contain the disease.<ref>{{cite news |last=Roche |first=Barry |title= Cork’s Gougane Barra Forest Park closing due to tree fungus|publisher=Irish Times |date=4 January 2014 |accessdate=22 November 2016}}</ref>

The closely related ''[[Phytophthora kernoviae]]'' causes similar symptoms to ''P. ramorum'', but infects the European beech (''[[Fagus sylvatica]]'').<ref name="rhs">{{cite web |url=https://apps.rhs.org.uk/advicesearch/profile.aspx?pid=329 |title=Phytophthora ramorum and P. kernoviae |publisher=Royal Horticultural Society | accessdate=17 February 2014}}</ref>

==Transmission==
''P. ramorum'' produces both resting [[spore]]s (chlamydospores) and [[zoospore]]s, which have [[flagella]] enabling [[aquatic locomotion|swimming]]. ''P. ramorum'' is spread by air;<ref name="transmission">Davidson, J.M., et al. Phytophthora ramorum and Sudden Oak Death in California: II. Transmission and Survival. in Fifth Symposium on Oak Woodlands: Oaks in California's Changing Landscape. 2001. San Diego, California.</ref> one of the major mechanisms of [[Biological dispersal|dispersal]] is [[rainwater]] splashing spores onto other susceptible plants, and into [[watercourse]]s to be carried for greater distances.<ref name="transmission"/> [[Chlamydospore]]s can withstand harsh conditions and are able to overwinter.<ref name="transmission"/> The pathogen will take advantage of wounding, but it is not necessary for [[infection]] to occur.<ref>Anon, Phytophthora ramorum and Phytophthora kernoviae: Key findings from research, DEFRA, Editor. 2005, DEFRA.</ref>

As mentioned above, ''P. ramorum'' does not kill every plant that can be used as a [[host (biology)|host]], and these plants are most important in the [[epidemiology]] of the disease as they act as sources of [[wikt:inoculum|inoculum]].<ref>Garbelotto, M., et al., Non-oak native plants are the main hosts for sudden oak death pathogen in California. Californian Agriculture, 2003. 57(1): p. 18-23.</ref> In California, [[California bay laurel]] (''Umbellularia californica'') seems to be the main source of inoculum.<ref name="Swiecki, T. J 2007. Pages 181-194">Swiecki, T. J. and E. A. Bernhardt. 2007. Increasing Distance from California Bay Laurel Reduces the Risk and Severity of Phytophthora ramorum Canker in Coast Live Oak. Pages 181-194 in Sudden Oak Death Third Science Symposium. USDA Forest Service, Santa Rosa, CA.</ref> Green waste, such as leaf litter and tree stumps, are also capable of supporting ''P. ramorum'' as a [[saprotroph]] and acting as a source of inoculum. Because ''P. ramorum'' is able to infect many [[ornamental plant]]s, it can be transmitted by ornamental plant movement.

[[Hiker]]s, [[mountain bike]]rs, [[Equestrianism|equestrian]]s, and other [[Seed dispersal#Dispersal by humans|people]] engaged in various outdoor activities may also unwittingly move the pathogen into areas where it was not previously present. Those travelling in an area known to be infested with SOD can help prevent the spread of the disease by cleaning their (and their animals') feet, tyres, tools, camping equipment, etc. before returning home or entering another uninfected area, especially if they have been in muddy soil. Additionally, the movement of firewood<ref>{{cite web|title=Moving Firewood Spreads Forest Pests|url=http://www.dontmovefirewood.org|work=Don't Move Firewood|accessdate=24 October 2011}}</ref> could introduce sudden oak death to otherwise uninfected areas. Both homeowners and travelers are advised to buy and burn local firewood.

==The two mating types==
[[Image:Phytophtora reproduction.png|thumb|Mating structures]]

''P. ramorum'' is [[heterothallic]] and has two [[mating type]]s, A1 and A2, required for sexual reproduction.<ref>{{cite journal|last=Boutet|first=X.|author2=Vercauteren, A. |author3=Heungens, K. |author4=Laurent, F. |author5=Chandelier, A. |title=Oospores progenies from Phytophthora ramorum|journal=Fungal Biology|year=2010|volume=114|pages=369–378|url=http://mw8xt6bj7r.search.serialssolutions.com/?&url_ver=Z39.88-2004&url_ctx_fmt=info:ofi/fmt:kev:mtx:ctx&rft_val_fmt=info:ofi/fmt:kev:mtx:journal&rft.atitle=Oospores%20progenies%20from%20Phytophthora%20ramorum&rft.aufirst=Xavier&rft.aulast=Boutet&rft.date=2010&rft.epage=378&rft.genre=article&rft.issn=1878-6146&rft.issue=4&rft.jtitle=FUNGAL%20BIOLOGY&rft.pages=369-378&rft.spage=369&rft.stitle=FUNGAL%20BIOL-UK&rft.volume=114&rfr_id=info:sid/www.isinet.com:WoK:WOS&rft.au=Vercauteren%2C%20Annelies&rft.au=Heungens%2C%20Kurt&rft.au=Laurent%2C%20Frederic&rft.au=Chandelier%2C%20Anne&rft_id=info:doi/10%2E1016%2Fj%2Efunbio%2E2010%2E02%2E009|doi=10.1016/j.funbio.2010.02.009|pmid=20943147|issue=4}}</ref> Interestingly, the European population is predominantly A1 while both mating types A1 and A2 are found in [[North America]].<ref>{{cite journal|last=Grünwald|first=N. J.|author2=Goss, E. M. |author3=Press, C. M. |title=''Phytophthora ramorum'': a pathogen with a remarkably wide host range causing sudden oak death on oaks and ramorum blight on woody ornamentals|journal=Mol. Plant Pathol.|year=2008|volume=9|pages=729–740|doi=10.1111/j.1364-3703.2008.00500.x|issue=6|pmid=19019002}}</ref> Genetics of the two isolates indicate that they are reproductively isolated.<ref>Ivors, K., et al., AFLP and phylogenetic analyses of North American and European populations of Phytophthora ramorum. Mycological Research, 2004. 108(4): p. 378-392.</ref> On average, the A1 mating type is more virulent than the A2 mating type, but more variation occurs in the pathogenicity of A2 isolates.<ref>Brasier, C., et al. Pathogenicity of Phytophthora ramorum isolates from North America and Europe to bark of European Fagaceae, American Quercus rubra and other forest trees. In Sudden Oak Death Science Symposium. 2002. Marriott Hotel, Monterey.</ref> It is currently not clear whether this pathogen can reproduce sexually in nature and genetic work has suggested that the lineages of the two mating types might be isolated reproductively or geographically given the evolutionary divergence observed.<ref>{{cite journal|last=Grünwald|first=N. J.|author2=Goss, E. M.|title=Evolutionary and Population Genetics of Exotic and Re-emerging Pathogens: Traditional and Novel Tools and Approaches|journal=Annu. Rev. Phytopathol.|year=2011|volume=49|pages=249–267|doi=10.1146/annurev-phyto-072910-095246}}</ref>

==Possible origins==

''P. ramorum'' is a relatively new disease, and several debates have occurred about where it may have originated or how it evolved.

===Introduction as an exotic species===
[[Evidence]] suggests ''P. ramorum'' may be an introduced species, and these introductions occurred separately for the European and North American populations, hence why only one mating type exists on each continent – this is called a [[founder effect]].<ref>Brasier, C., Evolutionary Biology of Phytophthora PART I: Genetic System, Sexuality and the Generation of Variation. Annual Review of Phytopathology, 1992. 30: p. 153-171.</ref> The differences between the two populations are thus caused by adaptation to separate climates. Evidence includes little genetic variability, as ''P. ramorum'' has not had time to diversify since being introduced. Existing variability may be explained by multiple introductions with a few individuals adapting best to their respective environments.<ref>Garbelotto, M., et al. ''Phytophthora ramorum'' and Sudden Oak Death in California: III. Preliminary Studies in Pathogen Genetics. in Fifth Symposium on Oak Woodlands: Oaks in California's Changing Landscape. 2001. San Diego, California.</ref> The behavior of the pathogen in California is also indicative of being introduced; it is assumed that such a high mortality rate of trees would have been noticed sooner if ''P. ramorum'' were [[Indigenous (ecology)|native]].

Where ''P. ramorum'' originated remains unclear, but most researchers feel [[Asia]] is the most likely, since many of the hosts of ''P. ramorum'' originated there.<ref>Martin, F.N. and P.W. Tooley, Phylogenetic relationships of ''Phytophthora ramorum'', ''P. nemorosa'' and ''P. pseudosyringae'', three species recovered from areas in California with sudden oak death. Mycological Research, 2003. 107(12): p. 1379–1391.</ref> Since certain climates are best suited to ''P. ramorum'', the most likely sources are the southern [[Himalayas]], [[Tibet]], or [[Yunnan province]].<ref>Nicholls, H., Stopping the Rot. PLoS Biology, 2004. 2(7): p. 891-895.</ref>

===Hybridization events===
Species of ''[[Phytophthora]]'' have been shown to have evolved by the [[Hybrid (biology)#Interspecific hybrids|interspecific hybridization]] of two different species from the genus.<ref>Brasier, C. and E.M. Hansen, Evolutionary Biology of Phytophthora PART II: Phylogeny, Speciation, and Population Structure. Annual Review of Phytopathology, 1992. 30: p. 173-200.</ref> When a species is introduced into a new [[Natural environment|environment]], it causes [[episodic selection]]. The invading species is exposed to other resident [[taxon|taxa]], and hybridization may occur to produce a new species. If these hybrids are successful, they may outcompete their parent species.<ref>Brasier, C., Episodic Selection as a force in fungal microevolution, with special reference to clonal speciation and hybrid introgression. Canadian Journal of Botany, 1994. 73 (suppl. 1.): p. S1213-S1220.</ref> Thus, ''P, ramorum'' is possibly a hybrid between two species. Its unique [[morphology (biology)|morphology]] does support this. Also, three sequences studied to establish the phylogeny of ''Phytophthora'': ITS, cox II and nad 5, were identical, supporting ''P. ramorum'' having recently evolved.

===A native organism===
''P. ramorum'' may be native to the United States. Infection rates could have previously been at a low level, but changes in the environment caused a change to the [[population]] structure.<ref>Rizzo, D.M., et al., Phytophthora ramorum as the Cause of Extensie Mortality of Quercus spp. and Luthocarpus densiflorus in California. Plant Disease, 2002. 86(3): p. 205-214.</ref> Alternatively, the symptoms of ''P. ramorum'' may have been mistaken for that of other pathogens. When SOD first appeared in the United States, many other pathogens and conditions were blamed before ''P. ramorum'' was found to be the causal agent.<ref>Garbelotto, M., P. Svihra, and D.M. Rizzo, Sudden oak death syndrome fells 3 oak species. Californian Agriculture, 2001. 55(1): p. 9-19.</ref> With many of the most seriously affected plants being in the forest, the likelihood of seeing diseased trees is also low.

==Ecological impacts==
In relation to human ecology, the loss of tanoak as the pathogen spreads to culturally sensitive indigenous lands represents a loss of tanoak acorn as one of the most important traditional and ceremonial foods still used in Northern California such as among [[Yurok people]], [[Hupa]], [[Miwok]], and [[Karuk]] peoples. Similar impact applies to the decline of other native plant species that are traditional food sources in tanoak and oak regimens infected by the pathogen.<ref>Traditional ecological knowledge (TEK)</ref>

In forest ecology, the pathogen contributes to loss of environmental services provided by diversity of plant species and interdependent wildlife.

The mortality caused by this emerging disease is expected to cause many indirect effects. Several predictions of long-term impacts have been discussed in the scientific literature.<ref>{{cite journal | last1 = Rizzo | first1 = D. | last2 = Garbelotto | first2 = M. | last3 = Hansen | first3 = E. | year = 2005 | title = Phytophthora ramorum: Integrative research and management of an emerging pathogen in California and Oregon forests | url = | journal = Annual Review of Phytopathology | volume = 43 | issue = | pages = 309–335 | doi=10.1146/annurev.phyto.42.040803.140418}}</ref> While such predictions are necessarily speculative, indirect impacts occurring on shorter time scales have been documented in a few cases. For instance, one study demonstrated that redwood trees (''[[Sequoia sempervirens]]'') grew faster after neighboring tanoaks were killed by sudden oak death.<ref>Waring, K.M., and O’Hara, K.L. 2008. Redwood/tanoak stand development and response to tanoak mortality caused by Phytophthora ramorum. Forest Ecology and Management. 255: 2650–2658.</ref> Other studies have combined current observations and reconstruction/projection techniques to document short-term impacts while also inferring future conditions. One study used this approach to investigate the effects of SOD on the structural characteristics of redwood forests.<ref>Ramage, B. S., and O’Hara, K. L. 2010. Sudden oak death-induced tanoak mortality in coast redwood forests: Current and predicted impacts to stand structure. Forests. 1: 114-130, [http://www.mdpi.com/1999-4907/1/3/114/].</ref>

Additional long-term impacts of SOD may be inferred from regeneration patterns in areas that have experienced severe mortality. These patterns may indicate which tree species will replace tanoak in diseased areas. Such transitions will be of particular importance in forest types that were relatively poor in tree species diversity before the introduction of SOD, e.g., redwood forest. {{Asof|2011}}, the only study to comprehensively examine regeneration in SOD-impacted redwood forests found no evidence that other broadleaf tree species are beginning to recruit.<ref>{{cite journal | last1 = Ramage | first1 = B.S. | last2 = OʼHara | first2 = K.L. | last3 = Forrestel | first3 = A.B. | year = 2011 | title = Forest transformation resulting from an exotic pathogen: regeneration and tanoak mortality in coast redwood stands affected by sudden oak death | url = http://www.nrcresearchpress.com/doi/abs/10.1139/x11-020 | journal = Canadian Journal of Forest Research | volume = 41 | issue = | pages = 763–772 | doi=10.1139/x11-020}}</ref> Instead, redwood was colonizing most mortality gaps. However, they also found inadequate regeneration in some areas and concluded that regeneration is continuing. Since this study only considered one site in [[Marin County, California]], these results may not apply to other forests. Other impacts to the local ecology include, among others, the residual effects of spraying heavy pesticides (Agrifos) to treat SOD symptoms, and the heavy mortality of the native pollinator community that occurs as a result. Bee hives situated in areas of heavy Agrifos spraying have incurred significant losses of population in direct correlation to the application of these chemicals. Counties such as Napa and Sonoma may be doing significant damage to their native pollinator populations by virtue of adopting broad-based prophylactic pesticide policies. Such damage to the pollinator populations may have tertiary negative effects on the entire local plant community, compounding the loss of biodiversity, and thus environmental value, attributable to SOD.{{Citation needed|date=November 2013}}

==Control==


===Early detection===
Early detection of ''P. ramorum'' is essential for its control. On an individual-tree basis, preventive treatments, which are more effective than therapeutic treatments,<ref name="Garbelotto, M. 2007">Garbelotto, M., D. J. Schmidt, and T. Y. Harnik. 2007. Phosphite Injections and Bark Application of Phosphite + Pentrabark Control Sudden Oak Death in Coast Live Oak. Arboriculture and Urban Forestry 33:8.</ref> depend on knowledge of the pathogen’s movement through the landscape to know when it is nearing prized trees. On the landscape level, ''P. ramorum’s'' fast and often undetectable movement means that any treatment hoping to slow its spread must happen very early in the development of an infestation. Since ''P. ramorum’s'' discovery, researchers have been working on the development of early detection methods on scales ranging from diagnosis in individual infected plants to landscape-level detection efforts involving large numbers of people.

Detecting the presence of ''Phytophthora'' species requires laboratory confirmation. The traditional method of culturing is on a growth medium selective against fungi (and, in some cases, against other oomycetes such as ''[[Pythium]]'' species). Host material is removed from the leading edge of a plant tissue canker caused by the pathogen; resulting growth is examined under a [[microscope]] to confirm the unique [[morphology (biology)|morphology]] of ''P. ramorum''. Successful isolation of the pathogen often depends on the type of host tissue and the time of year that detection is attempted.<ref name="ReferenceA">Kliejunas, J. T. 2007c. Chapter 2: Identification and Distribution. Sudden Oak Death and Phytophthora ramorum: A Summary of the Literature. California Oak Mortality Task Force.</ref>

Because of these difficulties, researchers have developed some other approaches for identifying ''P. ramorum''. The [[ELISA|enzyme-linked immunosorbent assay]] test can be the first step in nonculture methods of identifying ''P. ramorum'', but it can only be a first step, because it detects the presence of [[proteins]] that are produced by all ''Phytophthora'' species. In other words, it can identify to the genus level, but not to the species level. ELISA tests can process large numbers of samples at once, so researchers often use it to screen out likely positive samples from those that are not when the total number of samples is very large.<ref name="ReferenceA"/> Some manufacturers produce small-scale ELISA “field kits” that the homeowner can use to determine if plant tissue is infected by ''Phytophthora''.

[http://grunwaldlab.cgrb.oregonstate.edu/about-us Researchers] have also developed numerous molecular techniques for ''P. ramorum'' identification. These include amplifying [[DNA]] sequences in the [[internal transcribed spacer]] region of the ''P. ramorum'' [[genome]] (ITS [[polymerase chain reaction]], or ITS PCR); real-time PCR, in which DNA abundance is measured in real time during the PCR reaction, using dyes or probes such as SBYR-Green or TaqMan; [[multiplex PCR]], which amplifies more than one region of DNA at the same time; and [[single strand conformation polymorphism]] (SSCP), which uses the ITS DNA sequence amplified by the PCR reaction to differentiate ''Phytophthora'' species according to their differential movement through a gel.<ref name="ReferenceA"/>

Additionally, researchers have begun using features of the DNA sequence of ''P. ramorum'' to pinpoint the minuscule differences of separate ''P. ramorum'' isolates from each other. Two techniques for doing this are [[amplified fragment length polymorphism]], which through comparing differences between various fragments in the sequence has enabled researchers to differentiate correctly between EU and U.S. isolates,<ref name="ReferenceA"/> and the examination of [[Microsatellite (genetics)|microsatellites]], which are areas on the sequence featuring repeating base pairs. When ''P. ramorum'' [[propagules]] arrive in a new geographic location and establish colonies, these microsatellites begin to display [[mutation]] in a relatively short time, and they mutate in a stepwise fashion. Based on this, researchers in California have been able to construct trees, based on microsatellite analyses of isolates collected from around the state, that trace the movement of ''P. ramorum'' from two likely initial points of establishment in [[Marin County, California|Marin]] and [[Santa Cruz County, California|Santa Cruz]] Counties and out to subsequent points.<ref>{{cite journal | last1 = Mascheretti | first1 = S. | last2 = Croucher | first2 = P. J. P. | last3 = Vettraino | first3 = A. | last4 = Prospero | first4 = S. | last5 = Garbelotto | first5 = M. | year = 2008 | title = Reconstruction of the Sudden Oak Death epidemic in California through microsatellite analysis of the pathogen Phytophthora ramorum | url = | journal = Molecular Ecology | volume = 17 | issue = | pages = 2755–2768 | doi=10.1111/j.1365-294x.2008.03773.x}}</ref>

Early detection of ''P. ramorum'' on a landscape scale begins with the observation of symptoms on individual plants (and/or detecting ''P. ramorum'' propagules in watercourses; see below). Systematic ground-based monitoring has been difficult within the range of ''P. ramorum'' because most infected trees stand on a complex mosaic of lands with various ownerships. In some areas, targeted ground-based surveys have been conducted in areas of heavy recreation or visitor use such as parks, trailheads, and boat ramps. In California, when conducting ground-based detection, looking for symptoms on bay laurel is the most effective strategy, since ''P. ramorum'' infection of true oaks and tanoaks is almost always highly associated with bay laurel, the main epidemiological springboard for the pathogen.<ref>Davidson, J. M., D. M. Rizzo, M. Garbelotto, S. Tjosvold, and G. W. Slaughter. 2001. Phytophthora ramorum and Sudden Oak Death in California: II. Transmission and Survival. Pages 741-749 in Fifth Symposium on Oak Woodlands: Oaks in California's Changing Landscape. USDA Forest Service, San Diego, CA.</ref><ref>Davidson, J. M. and C. G. Shaw. 2003. Pathways of Movement for Phytophthora ramorum, the Causal Agent of Sudden Oak Death. Sudden Oak Death Online Symposium. The American Phytopathological Society.</ref><ref>{{cite journal | last1 = Maloney | first1 = P. E. | last2 = Lynch | first2 = S. C. | last3 = Kane | first3 = S. F. | last4 = Jensen | first4 = C. E. | last5 = Rizzo | first5 = D. M. | year = 2005 | title = Establishment of an Emerging Generalist Pathogen in Redwood Forest Communities | doi = 10.1111/j.1365-2745.2005.01031.x | journal = Journal of Ecology | volume = 93 | issue = | page = 6 }}</ref> Moreover, on many sites in California (though not all), ''P. ramorum'' can typically be detected from infected bay laurel tissues via culturing techniques year-round; this is not the case for most other hosts, nor is it the case in Oregon, where tanoak is the most reliable host.<ref>E. Goheen, USDA Forest Service, personal communication</ref>

As part of a nationwide USDA program, a ground-based detection survey was implemented from 2003 to 2006 in 39 U.S. states to determine whether the pathogen was established outside the West Coast areas already known to be infested. Sampling areas were stratified by environmental variables likely to be conducive to pathogen growth and by proximity to possible points of [[wikt:inoculum|inoculum]] introduction such as nurseries. Samples were collected along transects established in potentially susceptible forests or outside the perimeters of nurseries. The only positive samples were collected in California, confirming that ''P. ramorum'' was not yet established in the environment outside the West Coast.<ref>Oak, S. W., A. H. Elledge, E. K. Yockey, W. D. Smith, and B. M. Tkacz. 2008. Phytophthora ramorum Early Detection Suveys for Forests in the United States, 2003–2006. Pages 413-416 in Proceedings of the Sudden Oak Death Science Symposium. USDA Forest Service, Santa Rosa, CA.</ref>

Aerial surveying has proven useful for detection of ''P. ramorum'' infestations across large landscapes, although it is not as “early” a technique as some others because it depends on spotting dead tanoak crowns from fixed-wing aircraft. Sophisticated [[GPS]] and sketch-mapping technology enable spotters to mark the locations of dead trees so that ground crews can return to the area to sample from nearby vegetation.<ref>Mai, J. A., W. Mark, L. Fischer, and A. Jirka. 2006. Aerial and Ground Surveys for Mapping the Distribution of Phytophthora ramorum in California. Pages 345-360 in Proceedings of the Sudden Oak Death Second Science Symposium: The State of Our Knowledge. USDA Forest Service, Monterey, Calif.</ref>

Detection of ''P. ramorum'' in watercourses has emerged as the earliest of early detection methods. This technique employs [[pear]] or [[rhododendron]] baits suspended in the watercourse using ropes, buckets, mesh bags, or other similar devices. If plants in the watershed are infected with ''P. ramorum'', [[zoospores]] of the pathogen (as well as other ''Phytophthora'' spp.) are likely present in adjacent waterways. Under conducive weather conditions, the zoospores are attracted to the baits and infect them, causing lesions that can be isolated to culture the pathogen or analyzed via PCR assay.<ref>Murphy, S. K., C. Lee, J. Bienapfl, Y. Valachovic, and D. M. Rizzo. 2006. Monitoring Phytophthora ramorum Distribution in Streams within Coastal California Watersheds. Page 531 in Proceedings of the Sudden Oak Death Second Science Symposium: The State of Our Knowledge, Monterey, Calif.</ref><ref>Murphy, S. K., C. Lee, Y. Valachovic, J. Bienapfl, W. Mark, A. Jirka, D. R. Owen, T. Smith, and D. M. Rizzo. 2008. Monitoring Phytophthora ramorum Distribution in Streams within Coastal California Watersheds. Pages 409-411 in Proceedings of the Sudden Oak Death Third Science Symposium. USDA Forest Service, Santa Rosa, CA.</ref> This method has detected ''P. ramorum'' at scales ranging from small, intermittent seasonal drainages to the Garcia, Chetco, and South Fork Eel Rivers in California and Oregon (144, 352, and 689 mi2 drainage areas, respectively). It can detect the existence of infected plants in watersheds before any mortality from the infections becomes evident. Of course, it cannot detect the exact locations of those infected plants: at the first sign of ''P. ramorum'' propagules in the stream, crews must scour the watershed using all available means to find symptomatic vegetation.

A less technical means of detecting ''P. ramorum'' at the landscape level involves engaging local landowners across the landscape in the search. Many local county agriculture departments and University of California Cooperative Extension offices in California have been able to keep track of the distribution of the pathogen in their regions through reports and samples brought to them by the public. In 2008, the Garbelotto Laboratory at University of California, Berkeley, along with local collaborators, hosted a series of educational events, called "SOD Blitzes", designed to give local landowners basic information about ''P. ramorum'' and how to identify its symptoms; each participant was provided with a sampling kit, sampled a certain number of trees on his or her property, and returned the samples to the lab for analysis. This kind of [[citizen science]] hopefully can help generate an improved map of ''P. ramorum'' distribution in the areas where the workshops are held.

===Wildland management===
The course that ''P. ramorum'' management should take depends on a number of factors, including the scale of the landscape upon which one hopes to manage it. Management of ''P. ramorum'' has been undertaken at the landscape/ regional level in Oregon in the form of a campaign to completely eradicate the pathogen from the forests in which it has been found (mostly private, but also [[USDA Forest Service]] and USDI [[Bureau of Land Management]] ownership).<ref>{{cite journal | last1 = Goheen | first1 = E. M. | last2 = Hansen | first2 = E. M. | last3 = Kanaskie | first3 = A. | last4 = McWilliams | first4 = M. G. | last5 = Osterbauer | first5 = N. | last6 = Sutton | first6 = W. | year = 2002 | title = Eradication of Sudden Oak Death in Oregon | url = | journal = Phytopathology | volume = 92 | issue = | page = S30 }}</ref><ref>{{cite journal | last1 = Goheen | first1 = E. | last2 = Hansen | first2 = E. | last3 = Kanaskie | first3 = A. | last4 = McWilliams | first4 = M. | last5 = Osterbauer | first5 = N. | last6 = Sutton | first6 = W. | last7 = Rehms | first7 = L. | year = 2004 | title = An Eradication Strategy for Phytophthora ramorum in Oregon Forests | url = | journal = Phytopathology | volume = 94 | issue = | page = S35 }}</ref><ref>A. Kanaskie, N. Osterbauer, M. McWilliams, E. Goheen, E. Hansen, and W. Sutton. 2006. Eradication of Phytophthora ramorum in Oregon Forests — Status after Three Years. Pages 489-490 in Proceedings of the Sudden Oak Death Second Science Symposium: The State of Our Knowledge. USDA Forest Service, Monterey, Calif.</ref> The eradication campaign involves vigorous early detection by airplane and watercourse monitoring, a U.S. Department of Agriculture Animal and Plant Health Inspection Service (USDA APHIS) and Oregon Department of Agriculture-led [[quarantine]] to prevent movement of host materials out of the area where infected trees are found, and immediate removal of ''P. ramorum'' host vegetation, symptomatic or not, within a {{convert|300|ft|m|adj=on}} buffer around each infected tree.

The Oregon eradication effort, which began near the town of [[Brookings, Oregon|Brookings]] in southwest Oregon in 2001, has adapted its management efforts over the years in response to new information about ''P. ramorum''. For example, after inoculation trials of various tree species more clearly delineated which hosts are susceptible, the Oregon cooperators began leaving nonhost species such as [http://oregonstate.edu/trees/conifer_genera/douglas_fir.html Douglas fir] and red alder on site. In another example, after finding that a small percentage of tanoak stumps that were resprouting on the host removal sites were infected with the pathogen—whether these infections were systemic or reached the sprouts from the surrounding environment is unknown—the cooperators began pretreating trees with very small, targeted amounts of [[herbicide]] to kill the root systems of infected tanoaks before cutting them down. The effort has been successful in that while it has not yet completely eradicated the pathogen from Oregon forests, the epidemic in Oregon has not taken the explosive course that it has in California forests.<ref>citation needed</ref>

California, though, faces significant obstacles that preclude it from mounting the same kind of eradication effort. For one thing, the organism was too well established in forests in the [[Santa Cruz, California|Santa Cruz]] and [[San Francisco Bay]] areas by the time the cause of sudden oak death was discovered to enable any eradication effort to succeed. Even in still relatively uninfested areas of the north coast and southern [[Big Sur]], regionally coordinated efforts to manage the pathogen face huge challenges of leadership, coordination, and funding. Nevertheless, land managers are still working to coordinate efforts between states, counties, and agencies to provide ''P. ramorum'' management in a more comprehensive manner.

Several options exist for landowners who want to limit the impacts of SOD death on their properties. None of these options is foolproof, guaranteed to eradicate ''P. ramorum'', or guaranteed to prevent a tree from becoming infected. Some are still in the initial stage of testing. Nevertheless, when used thoughtfully and thoroughly, some of the treatments do improve the likelihood of either slowing the spread of the pathogen or of limiting its impacts on trees or stands of trees. Assuming that the landowner has correctly identified the host tree(s) and symptom(s), has submitted a sample to a local authority to send to an approved laboratory for testing, and has received confirmation that the tree(s) are indeed infected with ''P. ramorum''—or, alternatively, assuming that the landowner knows that ''P. ramorum''-infected trees are nearby and wants to protect the resources on his or her property—he or she can attempt control by cultural (individual-tree), chemical, or silvicultural (stand-level) means.

The best evidence that cultural techniques might help protect trees against ''P. ramorum'' comes from research that has established a correlation between disease risk in coast live oak trees and the trees’ proximity to bay laurel.<ref name="Swiecki, T. J 2007. Pages 181-194"/> In particular, this research found that bay laurel trees growing within 5 m of the trunk of an oak tree were the best predictors of disease risk. This implies that strategic removal of bay laurel trees near coast live oaks might decrease the risk of oak infection. Wholesale removal of bay laurel trees would not be warranted, since the bay laurels close to the oak trees appear to provide the greatest risk factor. Whether the same pattern is true for other oaks or tanoaks has yet to be established. Research on this subject has been started for tanoak, but any eventual cultural recommendations will be more complicated, because tanoak twigs also serve as sources of ''P. ramorum'' inoculum.

A promising treatment for preventing infection of individual oak and tanoak trees—not for curing an already established infection—is a [[phosphonate]] [[fungicide]] marketed under the trade name Agri-fos. Phosphonate is a neutralized form of phosphorous acid that works not by direct antagonism of ''Phytophthora'', but by stimulating various kinds of immune responses on the part of the tree.<ref name="Garbelotto, M. 2007"/> It is mostly environmentally benign if not applied to nontarget plants and can be applied either as an injection into the tree stem or as a spray to the bole. When applying Agri-fos as a spray, it must be combined with an organosilicate [[surfactant]], Pentra-bark, to enable the product to adhere to the bole long enough to be absorbed by the tree. Agri-fos has been very effective in preventing tree infections, but it must be applied when visible symptoms of ''P. ramorum'' on other trees in the immediate neighborhood are still relatively distant; otherwise, the tree to be treated likely is already infected, but visible symptoms have not yet developed (especially true for tanoak).

Trials of silvicultural methods for treating ''P. ramorum'' began in Humboldt County in northwest coastal California in 2006. The trials have taken place on a variety of infested properties both private and public and have generally focused on varying levels and kinds of host removal. The largest ({{convert|50|acre|m2}}) and most replicated trials have involved removal of tanoak and bay laurel by [[chainsaw]] throughout the infested stand, both with and without subsequent underburning designed to eliminate small seedlings and infested leaf litter.<ref>Valachovic, Y., C. Lee, J. Marshall, and H. Scanlon. 2008. Wildland Management of Phytophthora ramorum in Northern California Forests. Pages 305-312 in Sudden Oak Death Third Science Symposium. USDA Forest Service, Santa Rosa.</ref> Other treatments included host removal in a modified “shaded fuelbreak” design in which all bay laurel is removed, but not all tanoaks; bay and tanoak removal using herbicides; and removal of bay laurel alone. The results of these treatments are still being monitored, but repeated sampling has so far detected only very small amounts of ''P. ramorum'' in the soil or on vegetation in the treated sites.

===Nursery management===
Research and development in managing ''P. ramorum'' in [[Plant nursery|nursery]] settings extends from ''P. ramorum'' in the individual plant, to ''P. ramorum'' in the nursery environment, to the pathogen’s movement across state and national borders in infected plants.

An array of studies have tested the curative and protective effects of various chemical compounds against ''P. ramorum'' in plants valued as ornamentals or Christmas trees. Many studies have focused on the four main ornamental hosts of ''P. ramorum'' (''[[Rhododendron]]'', ''[[Camellia]]'', ''[[Viburnum]]'', and ''[[Pieris (plant)|Pieris]]''). Several effective compounds have been found; some of the most effective include [[mefenoxam]], [[metalaxyl]], [[dimethomorph]], and [[fenamidone]]. Many of these studies have converged upon the following conclusions: chemical compounds are, in general, more effective as preventives than in curatives; when used preventively, chemical compounds must be reapplied at various intervals; and chemical compounds can mask the symptoms of ''P. ramorum'' infection in the host plant, potentially interfering with inspections for quarantine efforts. In general, these compounds suppress but do not eradicate the pathogen, and some researchers are concerned that with repeated use the pathogen may become resistant to them. These studies and conclusions are summarized by Kliejunas.<ref>Kliejunas, J. T. 2007b. Chapter 5: Management and Control. Sudden Oak Death and Phytophthora ramorum: A Summary of the Literature. California Oak Mortality Task Force.</ref>

Another area of research and evolving practice deals with eliminating ''P. ramorum'' from nursery environments in which it is established to prevent human-mediated pathogen movement within the ornamental plant trade. One way of approaching this is through a robust quarantine and inspection program, which the various federal and state regulatory agencies have implemented. Under the federal ''P. ramorum'' quarantine program implemented by USDA [[APHIS]], nurseries in California, Oregon, and Washington are regulated and must participate in an annual inspection regimen; nurseries in the 14 infested counties in coastal California, plus the limited infested area in [[Curry County, Oregon|Curry County]], Oregon, must participate in a more stringent inspection schedule when shipping out of this area.<ref>USDA APHIS. 2007. Phytophthora ramorum; Quarantine and Regulations. Pages 8585-8604 7 CFR Part 301. Federal Register, Washington, D.C.</ref>

Much of the research into disinfesting nurseries has focused on the voluntary [[best management practices]] (BMPs) that nurseries can implement to prevent ''P. ramorum’s'' introduction into the nursery and movement from plant to plant. In 2008, a group of nursery industry organizations issued a list of BMPs that includes subsections on pest prevention/management, training, internal/external monitoring/audits, records/traceability, and documentation. The document includes such specific recommendations as “Avoid overhead [[irrigation]] of high-risk plants”; “After every [[crop rotation]], disinfect [[Plant propagation|propagation]] mist beds, sorting area, cutting benches, machines and tools to minimize the spread or introduction of pathogens”; and “Nursery personnel should attend one or more ''P. ramorum ''trainings conducted by qualified personnel or document self-training”.<ref>Suslow, K. 2008. Recommended Industry Best Management Practices for the Prevention of Phytophthora ramorum Introductions in Nursery Operations. Pages 115-128 in Proceedings of the Sudden Oak Death Third Science Symposium. USDA Forest Service, Santa Rosa, CA.</ref><ref>HRI P. ramorum Industry Working Group. 2008. Nursery Industry Best Management Practices for Phytophthora ramorum to Prevent the Introduction or Establishment in California Nursery Operations.</ref>

Research on control of ''P. ramorum'' in nurseries has also focused on disinfesting irrigation water containing ''P. ramorum'' inoculum. Irrigation water can become infested from bay trees in the forest (if the irrigation source is a stream), from bay trees overhanging irrigation ponds, from runoff from infested forests,<ref>Tjosvold, S. A., D. L. K. Chambers, S., and E. Fichtner. 2006. Epidemiology of Phytophthora ramorum Infecting Rhododendrons under Simulated Nursery Conditions. Pages 459-461 in Proceedings of the Sudden Oak Death Science Symposium: The State of Our Knowledge. USDA Forest Service, Monterey, Calif.</ref> or from recirculated irrigation water.<ref>{{cite journal | last1 = Werres | first1 = S. | last2 = Wagner | first2 = S. | last3 = Brand | first3 = T. | last4 = Kaminski | first4 = K. | last5 = Seipp | first5 = D. | year = 2007 | title = Infectivity and Survival of Phytophthora ramorum in Recirculation Water | url = | journal = Plant Disease | volume = 91 | issue = | pages = 1034–1044 | doi=10.1094/pdis-91-8-1034}}</ref> Experiments in Germany with three types of filters—[[slow sand filters]], [[lava filter]]s, and constructed wetlands—showed that the first two removed ''P. ramorum'' from the irrigation water completely, while 37% of the post-treatment water samples from the constructed wetland still contained ''P. ramorum''.<ref>Ufer, T., M. Posner, H.-P. Wessels, S. Wagner, K. Kaminski, T. Brand, and S. Werres. 2008. Four Years Experience with Filtration Systems in Commercial Nurseries for Eliminating Phytophthora Species from Recirculation Water. Page 111 in Proceedings of the Sudden Oak Death Third Science Symposium. USDA Forest Service, Santa Rosa, CA.</ref>

Since ''P. ramorum'' can persist for an undetermined period of time within the soil profile, management programs in nurseries should also deal with delineating the pathogen’s distribution in nursery soil and eliminating it from infested areas. A variety of chemical options has been tested for soil disinfestation, including such chemicals as [[chloropicrin]], [[metham sodium]], [[iodomethane]], and [[dazomet]]. Lab tests indicated that all of these chemicals were effective when applied to infested soil in glass jars. Additionally, tests on volunteer nurseries with infested soil demonstrated that dazomet (trade name Basamid) fumigation followed by a 14-day tarping period successfully removed ''P. ramorum'' from the soil profile.<ref>Yakabe, L. E. and J. D. MacDonald. 2008. Soil Treatments for the Elimination of Phytophthora ramorum from Nursery Beds: Current Knowledge from the Laboratory and the Field. Pages 113-114 in Proceedings of the Sudden Oak Death Third Science Symposium. USDA Forest Service, Santa Rosa, CA.</ref> Other soil disinfestation practices under investigation, or in which interest has been expressed, include steam [[Sterilization (microbiology)|sterilization]], [[Soil solarization|solarization]], and paving of infested areas.

===General sanitation in infested areas===
One of the most important aspects of ''P. ramorum'' control involves interrupting the human-mediated movement of the pathogen by ensuring that infested materials do not move from location to location. While enforceable quarantines perform part of this function, basic cleanliness when working or recreating in infested areas is also important. In most cases, cleanliness practices involve ridding potentially infested surfaces—such as shoes, vehicles, and pets—of foliage and mud before leaving the infested area. The demands of implementing these practices become more complex when large numbers of people are working in infested areas, as in construction, timber harvesting, or [[wildfire suppression]]. The [[California Department of Forestry and Fire Protection]] and USDA Forest Service have implemented guidelines and mitigation requirements for the latter two situations; basic information about cleanliness in ''P. ramorum''-infested areas can be found at the California Oak Mortality Task Force web site ([http://www.suddenoakdeath.org www.suddenoakdeath.org]) under the “Treatment and Management” section (subsection “Sanitation and Reducing Spread”).

==Government agency involvement==
In [[England]] in 2009, the [[Forestry Commission]], [[DEFRA]], the [[Food and Environment Research Agency]], [[Cornwall County Council]], and [[Natural England]] are working together to record the locations and deal with this disease. Natural England is offering grant funding through its [[Environmental Stewardship]], [[Countryside Stewardship]], and [[Environmentally Sensitive Area]] schemes to clear rhododendron.<ref>[http://www.forestry.gov.uk/pramorum Forestry Commission page]</ref> In 2011, the [[Forestry Commission]] started felling {{convert|10,000|acre|km2}} of [[larch]] forest in the south-west of England, as an attempt to halt the spread of the disease.<ref>{{cite news| url=http://www.dailymail.co.uk/news/article-1350843/Britains-forests-10k-acres-trees-cut-stop-pathogen.html | work=Daily Mail | first=Robert | last=Hardman | title=The British Chainsaw Massacre: Desperate bid to cut down 10,000 acres of trees to stop new pathogen | location=London | date=26 January 2011}}</ref> In Northern Ireland at the end of 2011, the Department of Agriculture and Rural Development’s Forest Service began felling 14 hectares of affected Larch woodland at Moneyscalp, on the edge of Tollymore Forest Park in County Down.<ref>http://www.northernireland.gov.uk/index/media-centre/news-departments/news-dard/news-dard-291111-phytophthora-ramorum-disease.htm</ref>

==See also==
*[[Acute oak decline]]
*[[Forest pathology]]
*[[Forest disturbance of invasive insects and diseases in the United States]]

==References==
{{Reflist|2}}

==External links==
{{Commons category|Phytophthora ramorum}}
* [http://www.suddenoakdeath.org/ www.suddenoakdeath.org], California Oak Mortality Task Force
* [http://www.fera.defra.gov.uk/plants/plantHealth/pestsDiseases/phytophthora/pRamorum/ Phytophthora ramorum], UK [[Department for Environment, Food and Rural Affairs]]
* [http://cisr.ucr.edu/sudden_oak_death.html Sudden Oak Death], Center for Invasive Species Research
* [http://www.invasivespeciesinfo.gov/microbes/suddenoak.shtml Species Profile- Sudden Oak Death (''Phytophthora ramorum'')], National Invasive Species Information Center, [[United States National Agricultural Library]]. General information and resources for Sudden Oak Death.
* [http://dontmovefirewood.org/gallery-of-pests/sudden-oak-death-syndrome.html Gallery of Pests — Sudden Oak Death Syndrome], Don't Move Firewood

{{Taxonbar|from=Q3382075}}

{{DEFAULTSORT:Phytophthora ramorum}}
[[Category:Phytophthora|ramorum]]
[[Category:Tree diseases]]
[[Category:Water mould plant pathogens and diseases]]
[[Category:Biota of California]]
[[Category:Natural history of the California chaparral and woodlands]]
[[Category:Quercus]]
[[Category:Eukaryotes described in 1995]]

Acremonium strictum

2018-03-23T11:57:41Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q4675100}} (3 sig. taxon IDs); WP:GenFixes using AWB

{{Taxobox
| regnum = [[fungus|Fungi]]
| divisio = [[Ascomycota]]
| classis = [[Sordariomycetes]]
| ordo = [[Hypocreales]]
| familia = ''[[Incertae sedis]]''
| genus = ''[[Acremonium]]''
| species = '''''A. strictum'''''
| binomial = ''Acremonium strictum''
| binomial_authority = W.Gams (1971)
| synonyms =
*''Cephalosporium acremonium'' [[August Carl Joseph Corda|Corda]] (1839)
*''Haplotrichum acremonium'' (Corda) [[Roscoe Pound|Pound]] & [[Clem.]] (1896)
*''Hyalopus acremonium'' (Corda) M.A.J.Barbosa (1941)
*''Sarocladium strictum'' (Gams) Summerbell (2011)
}}

'''''Acremonium strictum''''' is an environmentally widespread [[saprotroph]] species found in soil, plant debris, and rotting mushrooms.<ref name=Guarro1997/> Isolates have been collected in North and Central America, Asia, Europe and Egypt.<ref name= Schell1996/> ''A. strictum'' is an agent of [[hyalohyphomycosis]] and has been identified as an increasingly frequent human pathogen in immunosuppressed individuals, causing localized, disseminated and invasive infections.<ref name=Schell1996 /><ref name=Sharma2013 /><ref name=Fincher1991 /> Although extremely rare, ''A. strictum'' can infect immunocompetent individuals, as well as neonates.<ref name=Sharma2013 /><ref name=Fincher1991 /> Due to the growing number of infections caused by ''A. strictum'' in the past few years, the need for new medical techniques in the identification of the fungus as well as for the treatment of human infections has risen considerably.<ref name=Perdomo2010 />

''A. strictum'' has been shown to be involved in some myoparasitic relationships, as well as a wide range of plant endophytic and parasitic relationships,<ref name=Rivera2007 /><ref name=Chalfoun2013 /><ref name=Yang2014 /><ref name=Zhou2014 /> and further studies are required to determine ''A. strictum's'' use as a biological control agent and role as a parasite that reduces crop yields. ''A. strictum'' exhibits metabolism of many products that imply future agricultural and pharmaceutical significance.<ref name=Chen2008 /><ref name=Chang2013 />

==General description==

The genus ''[[Acremonium]]'' is a large [[polyphyletic]] genus of approximately 150 species, many of which are derived from a closely related families in the ''Sordariomycetes''.<ref name=Summerbell2011 /> The genus includes many slow growing, simply structured, anamorphic filamentous fungi,<ref name=Sharma2013 /><ref name=Perdomo2010 /><ref name=Summerbell2011 /> typically encountered in wet, cellulose-based building materials suffering form chronic wet conditions.<ref name=Skrobot2014 /> Characteristic morphology in this genus is septate hyphae giving rise to thin, tapered aculeate phialides that are usually unicellular, or weakly branched conidiophores.<ref name=Guarro1997 /><ref name=Perdomo2010 /><ref name=Summerbell2011 /> Human infections, though rare, usually occur in severely immunodeficient individuals.<ref name=Schell1996 /> ''A. strictum'' is mostly known to be involved with myparasitic relationships, as well as being a plant parasite and endophyte.<ref name=Rivera2007 /><ref name=Leslie2008 /><ref name= Goswami2008 /><ref name= Goswami2008 />

===Morphological identification===

====Colonial appearance====

''Acremonium strictum'' grows readily at 30 °C on glucose peptone agar, showing mycelium of approximately 50mm in size in 7 days. Colonies are flat, with smooth, wet, velvety or floccose texture, sometimes resembling thin cottony mounds.<ref name=Campbell2013 /> The colour of mycelia ranges widely from light pink to orange, and sometimes yellow, white or green.<ref name=Perdomo2010 /><ref name=Campbell2013 /> ''A. strictum'' filaments are sometimes bound together into ropes several cells in diameter.<ref name=Guarro1997 />[[Conidium]] grow as wet clusters or dry chains,<ref name=Guarro1997 /> and grains produced are white to pale-yellow, soft and variable in shape.<ref name=Guarro1997 /> Subcultures of the fungus can also be grow within seven days into smooth, moist, pink mycelia that resemble thin cotton.<ref name=Perdomo2010 /><ref name=Novicki2003 />

====Microscopic appearance====

[[File:Conidia and conidiophores of the fungus Acremonium falciforme PHIL 4168 lores.jpg|thumb|Conidia and conidiophores of the fungus Acremonium falciforme PHIL 4168 lores]]

Under the microscope at 30 °C, ''A. strictum'' shows long slender phialides, and conidia are cylindrical or ellipsoidal, formed in slimy bundles at the tips of the phialides. Lower microscopy shows pin-head spore ball formation.<ref name=Campbell2013 />

Species of ''Acremonium'' are morphologically very similar, making identification difficult. Shown in the image is a microscopic image of ''A. falciforme'', an example showing the morphological similarities to ''A. strictum''. Cases involving different species of ''Acremonium'' are often reported as simply as an ''Acremonium'' species, which reduces the amount of accurate information on the clinical presentation of ''A. strictum''.<ref name=Perdomo2010 /> Isolates of phylogenetically remote species of ''Acremonium'' show considerable convergence.<ref name=Summerbell2011 /> As a human pathogen, diagnosis is made in isolation and identification of the fungus from granules in tissue and the presence of hyphae in microscopic examination of cutaneous biopsy and discharge.<ref name=Guarro1997 /><ref name=Sharma2013 />

Genera that are morphologically similar to ''Acremonium'' include ''[[Fusarium]]'', ''[[Phaeoacremonium]]'', ''[[Verticillium]]'', ''[[Phialemonium]]'', and ''[[Lecanicillium]]''.<ref name=Perdomo2010 />

===Genetic identification===

Identification of ''A. strictum'' isolates has shown that this fungus is phenotypically diverse and may vary genetically.<ref name=Novicki2003 /> Due to phylogenetic ambiguities, an unknown proportion of the literature on ''A. strictum'' is based on studies of ''Acremonium sclerotigenum''.<ref name=Summerbell2011 /> The fungus can generally be successfully identified by the nuclear [[Internal transcribed spacer|ITS]] region sequence analysis.<ref name=Perdomo2010 /><ref name=Guarro2009 /> Analysis of the genes for ribosomal large subunit (LSU) and whole small subunit (SSU) also help to elucidate phylogenetic relationships, since these genes are more conserved and less subject to evolutionary changes.<ref name=Summerbell2011 /> The species ''A. strictum'' is separated into three genogroups. Genogroup I is represented by type strain CBS 346.70, genogroup II by UW836 and genogroup III by UWFP940. These genogroups were determined based on [[GenBank]] entries for ''A. strictum''.

==Pathophysiology==

Human infections of ''Acremonium strictum'' are very rare, and usually develop after traumatic inoculation of the fungus.<ref name=Perdomo2010 /> [[Hyalohyphomycosis|Hyalophomycosis]] may occur in immunodeficient individuals, presented in the infected tissue by hyaline or colourless hyphae.<ref name=Guarro1997 /><ref name=Fincher1991 /> [[Peritonitis]] and [[pleuritis]] have resulted from ''A. strictum'' infections,<ref name=Sener2008 /><ref name=Koc2008 /> but [[Skin infection|cutaneous]] and subcutaneous infections of ''A. strictum'' are rarely reported.
* Most human infections have been reported to occur in '''[[Immunodeficiency|immunocompromised patients]]''' and have been presented as localized or disseminated, [[fungemia]], mycetoma or ocular infections,<ref name=Schell1996 /><ref name=Fincher1991 /> and often result in fatal cases.<ref name=Sharma2013 /> ''A. strictum'' may result in invasive infections such as [[pneumonia]], [[arthritis]], [[osteomyelitis]], [[endocarditis]], [[meningitis]] and [[sepsis]] in immunodeficient patients.<ref name=Sharma2013 /><ref name=Sener2008 />
* Infections in '''[[Immunocompetence|immunocompetent individuals]]''' usually follows inoculation during penetration of the extremities and cornea, resulting in localized infections.<ref name=Sharma2013 /> The fungus can also cause onychomycosis, ontomycosis and burn wound infection in immunocompetent individuals.<ref name=Sharma2013 /> Patients with prosthetic valves who are infected with ''A. strictum'' in the region of the valve may suffer from severe inflammation, resulting in sepsis and multi-organ failure.<ref name=Guarro2009 />
* Infections in '''[[Infant|neonates]]''', although rare, can occur and be fatal.<ref name=Fincher1991 />

Many environmental factors such as the density of fungi in soil, rainfall, temperature, humidity and types of vegetation in close contact are relevant in determining the likelihood of acquiring hyalohypjomycosis infection by ''A. strictum''.<ref name= Sharma2013 /> Frequent exposure to contaminated water along with high temperature and humid environments increases the risk of infection.<ref name= Sharma2013 />

==Clinical presentation and treatments==

Clinical presentation of an infection is ill-defined, but most individuals may present a skin rash and flu like symptoms, such as elevated body temperature and fatigue.<ref name=Guarro1997 /><ref name=Sharma2013 /> In more severe infections, such as in immunodeficient individuals, [[peritonitis]] and [[pleuritis]], and may lead to multi-organ failure.<ref name=Sener2008 /><ref name=Koc2008 /> In the case of invasive infections, surgical intervention may be required to remove fungal mass from body tissues.<ref name=Fincher1991 /> Due to limited, ill-defined cases and the variance in clinical presentation and species identification, no optimal treatments are available.<ref name=Schell1996 /> ''A. strictum'' and other ''Acremonium'' species are generally resistant to most [[antifungals]], but antifungal susceptibility testing is recommended to select the most appropriate treatment for the strain of ''A. strictum'' that is the infection agent.<ref name=Sharma2013 /> [[Amphotericin B]] therapy coupled with [[ketoconazole]] is usually recommended as the best available treatment.<ref name=Schell1996 /><ref name=Sharma2013 />

===Biological control===

It has been shown that seedlings infected with ''A. strictum'' have high mortality rates. It would be agriculturally significant to identify biological control agents for this fungus. Aerial parts of ''[[Cymbopogon schoenanthus]]'', ''[[Hyptis|Hyptis spicigera]]'', ''[[Lantana camara]]'' and ''[[Ocimum americanum]]'' were collected and air-dried for four days. After drying the plants, essential oils were extracted from the materials. A variety of seeds inoculated with fungi, some cohorts with ''A. strictum''. The oils were applied to the infected seeds. After allowing seedlings to develop, it was found that the oils in combination inhibited ''A. strictum'' mycelial growth significantly.<ref name=Zida2008 />

==Fungal interactions==

===''Helminthosporium solani''===

''Acremonium strictum'' is generally known as a mycoparasite, as shown in its antagonistic relationship with ''[[Helminthosporium solani]]''. ''H. solani'' is a potato ([[Potato|''Solanum tuberosum'']]) associated fungus, that has caused extreme and widespread losses in all market classes of potatoes since emerging in the United States. Commonly referred to as silver scurf, ''H. solani'' causes blemishes that decreases the quality of the crop, making it unfit for marketing. In more severe cases, ''H. solani'' causes weight loss in potatoes and creates lesions in the periderm, creating entry points for other tuber pathogens. In pure cultures of ''H. solani'', isolates show white sectoring and rings, differential coloration and reduced sporulation in culture. Upon infection of ''A. strictum'', cultures of ''H. solani'' were uniformly black, without white sectors or rings. ''A. strictum'' was able to significantly reduce sporulation of ''H. solani'' by 30%, spore germination by 20%, and mycelial growth 8% in culture. This evidence suggests that ''A. strictum'' may be used as a biological control agent against ''H. solani'', which would greatly increase potato crop yields.<ref name=Rivera2007 />

==Plant interactions==

===Stalk rot===
''Acremonium strictum'' is pathogenic to many monocotyledonous and dicotyledonous crops, causing leaf desiccation on one side of the [[Leaf|midrib]] of these plants, plant wilt and abnormal, discoloured [[Vascular tissue|vasculature]] of the stalk near the soil line. Vasculature of the plant forms orange, red and brown bundles, usually resulting in death. Infection of ''A. strictum'' is systemic, and the fungus can be isolated from all tissues of the plant. Isolates have been found in plant seeds, which is probably the route of dissemination of the fungus. Crops affected by ''A. strictum'' include ''[[Acacia]]'', [[Alder|''Alnus'']], ''[[Ficus]]'', [[Glycine (plant)|''Glycine'']], ''[[Gossypium]]'', [[Wheat|''Triticum'']] and ''[[Zea (plant)|Zea]]''. Because of its ubiquitous presence in soils, ''A. strictum'' negatively impacts many agricultural plants, although more research is needed to investigate the parasitic interactions and develop strategies for its biological control.<ref name=Leslie2008 />

===Root knot nematode===
''[[Meloidogyne incognita]]'' is a polyphagous nematode that severely damages tomato crops by causing lesions in the roots by using a [[Stylet (anatomy)|stylet]], which allows other soil-dwelling fungal parasites to infect the host plant and cause complex disease interactions. ''A. strictum'' is reported to be a nematode egg parasite, as the eggs of ''M. incognita'' infested plants were found to be empty under ''A. strictum'' treatment. This treatment of ''A. strictum'' coupled with ''[[Trichoderma harzianum]]'' was found to be a very promising combination in the control of ''M. incognita'' in tomato plants.<ref name= Goswami2008 />

===''Fragaria ananassa''===
It demonstrates a complicated relationship with strawberry host [[Strawberry|''Fragaria ananassa'']],<ref name=Chalfoun2013 /> in which the fungus may cause lesions and small necrotic, light-brown spots in leaves and petioles which increase as the disease progresses, adversely affecting strawberry crop.<ref name=Racedo2013 /> Eventually the necrotic regions expand and cause the plant to wilt, but crown rot is not observed at any stage of the infection.<ref name=Racedo2013 /> Although it appears to have a parasitic relationship with ''Fragaria ananassa'', it also produces an elicitor protein, AsES, which provides systemic protection against [[anthracnose]] disease in strawberry host ''Fragaria ananassa'', which shows a symbiotic relationship between the strawberry plant and ''Acremonium strictum''.<ref name=Chalfoun2013 />

===''Atractylodes lancea''===
''[[Atractylodes lancea]]'' is a medicinal herb that grows in central China. ''A. strictum'' acts as a fungal endophyte and interacts with ''A. lancea'' in drought conditions and confers tolerance in moderate drought. Under mild drought conditions, ''A. strictum'' enhanced leaf soluble sugars, proteins, proline and antioxidant enzyme activity, which decreased the degree of plasmalemma oxidation. This increased ''A. lancea'' abscisic acid level and root:shoot ratio. While ''A. strictum'' may alleviate the effects of a mild to moderate drought, benefits of this endophytic relationship are constrained by drought degree, as there were no significant effects of ''A. strictum'' on ''A. lancea'' during periods of regular watering or severe drought.<ref name=Yang2014 />

===''Maclura cochinchinensis''===
In ''[[Maclura cochinchinensis]]'', ''Acremonium strictum'' acts as an [[Endophyte|endophytic]] fungi that infects primarily the leaves of the plant. In this relationship, ''A. strictum'' was found to provide and mediate a protective response against herbivorous insects.<ref name=Zhou2014 />

==Natural metabolites==

===Acremostrictin===

Acremostrictin can be isolated from certain strains of ''A. strictum'' and is characterized as a highly oxygenated, tricyclic lactone metabolite.<ref name=Julianti2011 /> This compound exhibits week antibacterial properties against the bacterium ''[[Micrococcus luteus]]'', ''[[Salmonella typhimurium]]'' and ''[[Proteus vulgaris]]''. However, it had no effect on ''[[Bacillus subtilis]]'', ''[[Staphylococcus aureus]]'' and ''[[Escherichia coli]]''.<ref name=Julianti2011 /> Acremostrictin has been shown to have concentration-dependent antioxidant activity, which conferred protection against [[oxidative stress]] induced cell death. Acremostrictin was shown to inhibit H2O2-induced death of human keratinocyte HaCaT cells. When extracted and isolated by filtration, acremostrictin presents as a colorless crystal solid.<ref name=Julianti2011 />

===AsES===

AsES protein is an extracellular elicitor protein produced by ''A. strictum'' that provides complete systemic protection against [[anthracnose]], cause by the fungal species ''[[Colletotrichum]]'', in the natural host ''Fragaria ananassa'' as well as the non-natural host ''[[Arabidopsis thaliana]]''. Anthracnose can affect all plant tissues, and appears as irregular and black leaf spot, flower blight, and fruit and crown rot, which results in serious losses in plant and fruit production. AsES has [[Proteolysis|proteolytic]] activity that appears to elicit an immune response in these species that results in the accumulation of reactive oxygen species and the expression of defence related genes like [[Pathogenesis-related protein|PR1]] and [[Chitinase|Chi2-1]]. Because it has been shown to provide the same systemic protection in non-natural hosts, this natural metabolite of ''A. strictum'' may be considered as a possible strategy for controlling anthracnose disease in plants.<ref name=Chalfoun2013 />

===Cephalosporins===
''A. strictum'' produces some types of [[cephalosporin]]s a group of antibiotics.

==Industrial Uses==

===BMOs===

Biogenic Mn oxides (BMOs) are naturally occurring Mn oxides that have the ability to oxidize various redox-sensitive elements. ''A. strictum'' is a Mn(II)-oxidizing fungus that forms BMOs through the action of Mn(II) oxidase. In the presence of BMOs in buffer solutions with no additional nutrients, ''A. strictum'' is capable of sequestering high Mn(II) concentrations for at least 8 days, in which the amount of dissolved Mn(II) decreases rapidly in several hours and is converted to oxidized Mn(II). Deaeration of the buffer solution with N2 gas purging suppressed Mn(II) conversion, but this suppression is easily rescued by aeration, implying that dissolved oxygen is required for the Mn(II) sequestration and oxidation process. Adding [[Sodium azide|NaN3]], a toxic substance, also significantly reduces the sequestration rates of the fungal BMOs. Heat treatments revealed that temperatures below 85 °C do not alter the conformation of the Mn(II) oxidase in the BMOs. Freezing the fungal BMOs at -80 °C for 4 weeks did not affect the Mn(II) ability, and the reducible Mn was still dominated in solution. This makes fungal BMOs an effective Mn(II) sequestering material if needed. For example, it can be used for the continuous removal of Mn(II) from Mn(II) contaminated water without the need for any additives other than dissolved oxygen. The product is an oxide phase Mn(II) that would provide additional affinity for other toxic elements and thus prove as an effective method of water cleansing. Enzymatically active fungal BMOs can be harvested under specific cultivation conditions and remain active even under circumstances that would be unfavourable for fungal growth.<ref name=Chang2013 />

===Ginsenoside analogs===

Fermentation of ginsenoside Rb(1) with ''A. strictum'' yields three new compounds — 12β-hydroxydammar-3-one-20 (S)-O-β-D-glucopyranoside, 12β, 25-dihydroxydammar-(E)-20(22)-ene-3-O-β-D -glucopyranosyl-(1→2)-β-D -glucopyranoside, and 12β, 20 (R), 25-trihydroxydammar-3-O-β-D -glucopyranosyl-(1→2)-β-D -glucopyranoside — as well as five known compounds — ginsenoside Rd, gypenoside XVII, ginsenoside Rg, ginsenoside F, and compound K. Many of these compounds are metabolites of ginsenoside Rb(1) in mammals, suggesting that fermentation of ginsenoside Rb(1) in ''A. strictum'' may be similar to mammalian metabolism and may be a useful agent for generating specific metabolites or related ginsenoside analogs, which can be later isolated for structural elucidation and use in pharmaceutical research.<ref name=Chen2008 />

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}}

==External links==
*{{IndexFungorum|308201}}

{{Taxonbar|from=Q4675100}}

[[Category:Hypocreales incertae sedis]]
[[Category:Plant pathogens and diseases]]
[[Category:Fungi described in 1839]]

Sparrige Flockenblume

2018-03-23T11:32:15Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q4115083}} (9 sig. taxon IDs); WP:GenFixes using AWB

{{Italic title}}
{{taxobox
|name = Diffuse knapweed
|image = Centaurea diffusa1.jpg
|image_caption = ''Centaurea diffusa'' in flower
|regnum = [[Plantae]]
|unranked_divisio = [[Angiosperms]]
|unranked_classis = [[Eudicots]]
|unranked_ordo = [[Asterids]]
|ordo = [[Asterales]]
|familia = [[Asteraceae]]
|tribus = [[Cynareae]]
|genus = ''[[Centaurea]]''
|species = '''''C. diffusa'''''
|binomial = ''Centaurea diffusa''
|binomial_authority = Lam., Encycl. Méth. Bot., 1: 675-676, 1785 [http://www.botanicus.org/page/717267]
}}

'''''Centaurea diffusa''''', also known as '''diffuse knapweed''', '''white knapweed''' or '''tumble knapweed''', is a member of the genus ''[[Centaurea]]'' in the family [[Asteraceae]]. This species along with ''[[Centaurea stoebe]]'' are the stereotypical "tumbleweed" of the West- breaking off at the top of the roots which facilitates its seeds spreading far and wide, but arean't actually native to the North American continent, but to a similar dry climate in the Mediterranean.

== Description ==
Diffuse knapweed is an [[annual plant|annual]] or [[biennial plant]], generally growing to between 10 and 60 cm in height. It has a highly branched stem and a large [[taproot]], as well as a basal rosette of [[leaf|leaves]] with smaller leaves alternating on the upright stems. [[Flower]]s are usually white or pink and grow out of urn-shaped [[Head (botany)|heads]] carried at the tips of the many branches. Diffuse knapweed often assumes a short rosette form for one year, reaching maximum size, then rapidly growing and flowering during the second year. A single plant can produce approximately 18,000 [[seeds]].<ref>Harris, P., and R. Cranston. 1979. An economic evaluation of control methods for diffuse and spotted knapweed in western Canada. Canadian Journal of Plant Science 59:375-382.</ref>
[[File:Centaurea diffusa 3.jpg|thumb|left|Centaurea diffusa basal rosette, first year plant]]

== Synonyms ==
* ''Centaurea microcalathina'' [[Tarassov]]
* ''Centaurea cycladum'' [[Heldr.]]
* ''Centaurea parviflora'' [[Sibth.]] & [[Sm.]], ''non [[Desf.]]
* ''Centaurea comperiana'' [[Steven]]<ref>[http://rbg-web2.rbge.org.uk/cgi-bin/nph-readbtree.pl/feout?FAMILY_XREF=&GENUS_XREF=Centaurea&SPECIES_XREF=diffusa&TAXON_NAME_XREF=&RANK= In Tutin & ''alt.'', Flora Europea]</ref>

== Distribution ==
It is native to [[Asia Minor]] ([[Turkey]], [[Syria]]), the [[Balkans]], ([[Bulgaria]], [[Greece]], [[Romania]]), [[Ukraine]], and southern [[Russia]].

[[File:Centaurea diffusa 5.jpg|thumb|Centaurea diffusa next to the Columbia River, Douglas County Washington]]

==An invasive species==
Diffuse knapweed is considered an [[invasive species]] in some parts of [[North America]], having established itself in many areas of the continent. ''C. diffusa'' was first identified from North America in 1907 when it was found in an [[alfalfa]] field in [[Washington (U.S. state)|Washington]] state. The seeds had presumably{{citation needed|date=April 2014}} been transported in an impure alfalfa seed shipment coming from somewhere in the species native range. Now present in at least 19 states in the [[United States]], it has naturalized in all contiguous states west of the [[Rocky mountains|Rockies]] and additionally in [[Connecticut]], [[Massachusetts]], and [[New Jersey]]. Portions of western [[Canada]] have also been colonized by this plant.

Areas in which diffuse knapweed has been established generally are plains rangelands or forest benchlands. Land that has been recently disturbed—by human or natural processes—is favored for the establishment of diffuse knapweed. It grows in semi-arid and arid environments and seems to favor light, dry, porous [[soil]]s. Areas with large amounts of shade or high levels of water discourage diffuse knapweed growth.

Dispersion occurs in the following ways:
* Agriculture – [[alfalfa]] contaminated with diffuse knapweed seed can promote the spread of diffuse knapweed;
* Wildlife – wild animals eating the seeds or transporting the seeds on fur;
* Wind – seeds blown out of their capsules held on the plant are distributed over a short range, but when dried out the plant may become a [[tumbleweed]], rolling for great distances and releasing seeds along the way;
* Water – waterways carry seeds in their flow for long distances before depositing them onto a shore where they germinate.

Wind is the primary means by which diffuse knapweed seeds are spread.

===Effects===
By 1998 diffuse knapweed had naturalized over {{convert|26640|km2|sqmi}} in the western US, and was increasing its range at a rate of 18% annually. {{Citation needed|date=July 2007}} Diffuse knapweed can establish itself in grassland, scrubland and riparian environments. {{Citation needed|date=July 2007}} It has little value as feed for [[livestock]], as its thistles can damage the mouth and digestive tract of animals that attempt to feed on it. A study in 1973 concluded that ranches lost approximately [[United States dollar|US$]]20/km² (8 cents per acre) of diffuse knapweed due to decreased grazing area. {{Citation needed|date=July 2007}} In an agricultural setting, it can greatly reduce crop yield and purity. {{Citation needed|date=July 2007}}

===Control===
Effective control of diffuse knapweed requires a fusion of well-executed land management, biological control, physical control, chemical control and reestablishment of the native species. Any method of control must ensure that the root is removed or the plant will grow back. Additionally, native plant growth in areas where diffuse knapweed has been removed should be encouraged to prevent reestablishment.

====Biological control====
[[Biological control]] involves the introduction of organisms, usually natural competitors of the invasive species, into the invaded environment in order to control the invasive species. Since 1970, 12 insects have been released to control diffuse knapweed. Of these 12, 10 have become established, and 4 are widely established (''[[Urophora affinis]]'' and ''[[Urophora quadrifasciata]]'', the root boring beetle, ''[[Sphenoptera jugoslavica]]'', and the weevil ''[[Larinus minutus]]'').<ref name="Myers">{{cite web |url=http://culter.colorado.edu:1030/~tims/Myers09.pdf |title = Successful biological control of diffuse knapweed, ''Centaurea diffusa'', in British Columbia, Canada |author = Judith H. Myers |publisher = Biological Control |year = 2009}}</ref> Research based on simulation models have shown that for biocontrol agents to be effective, they must kill their host, otherwise plants can compensate by having increased seedling survival.<ref>Myers, J.H., Risley, C., 2000. Why reduced seed production is not necessarily translated into successful biological weed control. In: Spencer, N. (Ed.), Proceedings X. International Symposium Biological Control of Weeds. Montana State University, Bozeman, MO, pp. 569–581.</ref>

Some of the more commonly utilized biocontrol agents are:
*[[Lesser knapweed flower weevil]] (''[[Larinus minutus]]''). Individuals of this species lay their eggs on the seed heads of both diffuse and [[spotted knapweed]]. When the larvae emerge from the [[Egg (biology)|egg]]s, they feed upon the seeds of their host plant. As the females of this species can create from 28 to 130 eggs and each larva can consume an entire seed head, an adequate population of ''Larinus minutus'' can devastate entire stands of knapweed. The adult weevils feed upon the stems, branches, leaves and undeveloped flower buds. It is native to Greece and is now found in Montana, Washington, Idaho and Oregon.<ref>Groppe, K. 1990. ''Larinus minutus'' Gyll. (Coleoptera: Curculionidae), a suitable candidate for the biological control of diffuse and spotted knapweed in North America. In: Final Report C.A.B International Institute of Biological Control. Delemont, Switzerland, p. 30.</ref>
*[[Knapweed root weevil]] (''[[Cyphocleonus achates]]''). Knapweed root weevils lay approximately 50 to 70 eggs on either diffuse or spotted knapweed. As the name suggests, the larvae produced burrow into the root where they metamorphose into adult form. At this point, they will tunnel through the root to the surface where they will feed on the leaves of knapweed plants. It is native to Austria, Greece, Hungary and Romania and has been introduced to Idaho, Montana, Washington and [[Oregon]].
*[[Agapeta zoegana|Yellow-winged knapweed root moth]] (''[[Agapeta zoegana]]''). This [[moth]] with root-mining [[larva]]e attacks diffuse and spotted knapweed in parts of the United States where it was introduced to control these plants.
*Two species of Tephritid flies ''[[Urophora affinis]]'' and ''[[Urophora quadrifasciata]]''.<ref name="Myers"/> Females lay their eggs on flower buds and larvae form galls on the flower head.<ref name="Myers"/>

====Physical control====
Physical control of diffuse knapweed primarily comprises cutting, digging or burning to remove the plants.
* '''Cutting''' - While cutting the aboveground portion of diffuse knapweed will greatly decrease the spread of seeds, it does not remove the root. With only its root still intact, diffuse knapweed can survive and continue to grow. For a program of cutting to be effective, it must be long-term so that the effect of reduced seed spreading can be realized.
* '''Digging''' - this removes both the portion above ground and the root of diffuse knapweed and has shown to be very effective; if the plant is properly disposed of, it can neither regrow nor spread its seeds. The largest problem with digging knapweed is that it is extremely labor-intensive. Additionally, the recently vacated soil should be planted with a native species to avoid knapweed reintroducing itself in the disturbed soil.
* '''Burning''' - setting fire to a crowd of knapweed, if the fire is sufficiently severe, can successfully destroy the above ground and belowground sections of diffuse knapweed. However, precautions must be taken to first ensure that the fire is properly contained and that a new plant community is established to prevent the reintroduction of diffuse knapweed.

====Chemical control====
Chemical control involves the use of [[herbicide]]s to control diffuse knapweed. The herbicide Tordon (picloram) is recognized{{By whom|date=September 2016}} as the most effective, but it is common to use multiple herbicides in order to reduce strain on local grasses. The herbicides 2,4-D, dicamba, and [[glyphosate]] are also effective for control. In order to be most effective, it must be applied before the knapweed plants have released their seeds, regardless of which herbicide is used. Ongoing research at the University of Colorado suggests that Tordon treatment does not contribute to long-term reductions of exotic species cover and may contribute to recruitment of other invasive species, such as redstem filaree and Japanese brome, which quickly take the place of herbicide-treated diffuse knapweed.{{Citation needed|date=September 2016}}

===Human influence on invasion===
One of the first influences humans had on diffuse knapweed was to inadvertently introduce it to North America. Diffuse knapweed has reached a level of coverage in that area that dwarfs its native range.{{Citation needed|date=July 2007}}

Diffuse knapweed is known to establish more easily and effectively in recently disturbed environments.{{Citation needed|date=July 2007}} Disturbed environments generally present low environmental stress because more resources are available than are being used. These available resources often allow the establishment of an invasion in an ecological community.{{Citation needed|date=July 2007}} The concentration of diffuse knapweed in such an area is often linked to the level of soil disturbance.{{Citation needed|date=July 2007}} Human disturbances often lead to less [[species diversity]] in a community.{{Citation needed|date=July 2007}} In turn, less species diversity can lead to unused resources, which allow invasive species to more readily establish. {{Citation needed|date=July 2007}} Areas such as fallow land, ditches, rangelands, residential and industrial districts and roadsides are all disturbed habitats where diffuse knapweed frequently establishes. {{Citation needed|date=July 2007}} Additionally, the removal of foliage and other ground cover increases the likelihood that seeds will come in contact with the soil and [[germinate]].

The largest impact of humans on diffuse knapweed is certainly due to our efforts in controlling and eradicating its invasive populations. The several methods outlined in the control section represent a small sample of literally hundreds of approaches being tried with varying levels of effectiveness. Besides reducing the spread of diffuse knapweed, we are also providing selective pressure against the individuals that cannot withstand a certain method of control. Selective pressure, given sufficient time, can cause the [[adaptation]] or [[evolution]] of invasive species such as diffuse knapweed. If an individual diffuse knapweed plant survives control efforts because of a trait it possesses, its progeny will make up a greater portion of the population than the plants that succumbed to the control.

===Toward an integrated control strategy===
To successfully control diffuse knapweed, an understanding of the mechanism that allows it to be invasive must be developed. Isolating the reason for its invasiveness would allow control methods designed to specifically target the effectiveness of that mechanism to be developed. Additionally, precautions designed to minimize the invisibility of at-risk environments could be carried out.

===Summary===
The success of diffuse knapweed must be attributed to a combination of several mechanisms. Its invasiveness is due to a mix of [[allelopathy]], ERH and superior resource competition. {{Citation needed|date=July 2007}} However, the most importance must be attributed to the ERH because diffuse knapweed, while a very effective invasive species in its novel environment, is non-invasive and doesn’t establish monocultures in its native range. It is the differences, [[Biotic component|biotic]] and [[abiotic]], between its novel and native surroundings that cause it to be invasive. {{Citation needed|date=July 2007}}

To demonstrate that the ERH applies to diffuse knapweed, it is essential to show that the absence of natural enemies has a significant positive effect on its success. {{Citation needed|date=July 2007}} One way to show this is to observe the effect of introducing some of diffuse knapweed’s natural enemies into its novel environment. If diffuse knapweed, which generally thrives in its invaded environment, is significantly inhibited through the introduction of natural enemies, it can be concluded that diffuse knapweed is more competitive in the absence of its natural enemies. A recent effort at biocontrol of diffuse knapweed in Idaho’s [[Camas County, Idaho|Camas County]] effectively reduced {{convert|80|km2|acre}} of knapweed to minimal levels through the release of the lesser knapweed flower weevil and the knapweed root weevil. Since both of the insects released are natural competitors of diffuse knapweed, and since this and other similar efforts at [[biocontrol]] have been successful, there is significant evidence that diffuse knapweed benefits from the absence of its natural enemies.

Another aspect of diffuse knapweed’s success relies on the effect of its allelopathic chemicals in its novel environment. {{Citation needed|date=July 2007}} Although there is still debate concerning the effectiveness of allelopathic chemicals in the field, the evidence of allelopathic effects demonstrated in a laboratory setting and its propensity to establish monocultures support the importance of allelopathy to diffuse knapweed’s success.

Curiously, diffuse knapweed’s allelopathic chemicals were shown to have a deleterious effect on the North American competitors but were beneficial to its native competitors. {{Citation needed|date=July 2007}} While diffuse knapweed’s native competitors are able to compete more effectively in the presence of allelopathic chemicals, the novel competitor’s fitness is decreased. This situation provides an example of the effectiveness of the allelopathy mechanism benefiting from the ERH. The increased effectiveness of allelopathic chemicals cause diffuse knapweed to experience less competitive pressure. As a result, diffuse knapweed is able to establish more predominantly in this new area.

Another connection between allelopathy and the ERH is the fact that concentrations of allelopathic chemicals were found to increase when diffuse knapweed was planted in North American soil as opposed to [[Eurasia]]n soil. {{Citation needed|date=July 2007}} This effect is probably due to the absence of unfavorable soil conditions or soil microorganisms that exist in its native environment. As a result, the allelopathic chemicals will be able to reach higher concentrations, spread farther and therefore be more effective. By effecting more neighboring plants, the favorable changes in soil condition contribute to the success of diffuse knapweed.

Besides the advantages that diffuse knapweed gains from the ERH and allelopathy, it also possesses several characteristically invasive traits. {{Citation needed|date=July 2007}} One factor leading to the superior resource competition of diffuse knapweed is its ability to exist in drought conditions. This advantage allows diffuse knapweed to devote its resources to competition while its neighbors are conserving resources to survive. {{Citation needed|date=July 2007}} The high number of seeds produced by diffuse knapweed is also a common trait of invasive plants. {{Citation needed|date=July 2007}} A higher density of knapweed will not only increase the concentration of allelopathic chemicals in the soil but will also restrict the nutrients available to native plants. {{Citation needed|date=July 2007}} Unfortunately, very little research has been conducted to determine the relative competitive ability between diffuse knapweed and its novel competitors. However, tests conducted on the effect of diffuse knapweed on North American grasses in the absence on allelopathic chemicals demonstrated that the fitness of these grasses declined in the presence of diffuse knapweed. {{Citation needed|date=July 2007}} Regrettably, we cannot decide if diffuse knapweed is, for general purposes, a better competitor from this data alone. Comparisons of the deleterious effects between these and other pairs of competitors to arrive at a conclusion.

Diffuse knapweed is successful in its novel range primarily because the organisms and conditions that prevent it from becoming invasive in its native environment are absent. {{Citation needed|date=July 2007}} It follows that the introduction of species from its native habitat would be an effective method of control. However, the introduction of a non-native organism has the potential to result in another invasive species outbreak. Therefore, any method of biological control must be preceded by analysis of possible effects.

== References == 
{{Reflist}}

==Sources==
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# [http://www.agf.gov.bc.ca/cropprot/weedguid/diffknap.htm Diffuse Knapweed (Centaurea diffusa)]
# H. G. Baker (1974). ''Annual Review of Ecology and Systematics'' 5: 1.
# K. Bossick, ''Wood River Journal''. A16 (2004).
# R. M. Callaway, W. M. Ridenour (2004). ''Front Ecol Environment'' 2 (8): 436.
# R. M. Callaway, E. T. Aschehoug (2000). ''Science'' 290: 521.
# A. T. Carpenter & T. A. Murray. ELEMENT STEWARDSHIP ABSTRACT for ''Centaurea diffusa'' Lamarck diffuse knapweed. [https://web.archive.org/web/20041126162621/http://tncweeds.ucdavis.edu/esadocs/documnts/centdif.html Online].
# C. Chou (1999). ''Crit. Rev. Plant Sci.'' 18: 609.
# D. R. Clements ''et al.''. ''[[Agric Ecosyst Environ]]'' (in press, corrected proof).
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# [http://www.issg.org/database/species/ecology.asp?si=313&fr=1&sts=sss L. Larson. ''Centaurea diffusa''].
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==External links==
*This article is extracted ''p.p.max.'' from homonym one in Bugwood Wiki [http://wiki.bugwood.org/Centaurea_diffusa].
* [http://www.invasivespeciesinfo.gov/plants/diffknapweed.shtml Species Profile- Diffuse Knapweed (''Centaurea diffusa'')], National Invasive Species Information Center, [[United States National Agricultural Library]]. Lists general information and resources for Diffuse Knapweed.

{{Commons category|Centaurea diffusa}}
{{Wikispecies|Centaurea diffusa}}

{{Taxonbar|from=Q4115083}}

[[Category:Centaurea|diffusa]]
[[Category:Invasive plant species]]
[[Category:Tumbleweeds|Centaurea diffusa]]

Brodiaea

2018-03-23T04:42:57Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q1894962}} (9 sig. taxon IDs); WP:GenFixes using AWB

{{Italic title}}
{{Automatic taxobox
|taxon = Brodiaea
|image = Brodiaea californica ssp leptandra 1.jpg
|image_caption = ''[[Brodiaea californica]]'' ssp. ''leptandra''
|name=cluster-lily
|authority = [[James Edward Smith|Sm.]]
|type_species = ''[[Brodiaea coronaria]]''
|type_species_authority =(Salisb.) [[Willis Linn Jepson|Jeps.]]
|synonyms = ''Hookera'' Salisb., rejected name
|synonyms_ref = <ref name="melany"/>
|}}

'''''Brodiaea''''' {{IPAc-en|ˌ|b|r|oʊ|ˈ|d|iː|ə|,_|ˌ|b|r|oʊ|d|i|ˈ||iː|ə}},{{citation needed|date=July 2016}} also known by the [[common name]] '''cluster-lilies''', is a [[monocot]] [[genus]] of [[flowering plant]]s in the {{C|Themidaceae}} family, in the [[Asparagales]] order.<ref>[http://ucjeps.berkeley.edu/cgi-bin/get_IJM.pl?key=93795 Jepson Herbarium, Jepson eFlora: Key to Themidaceae] . accessed 29 April 2016.</ref><ref name=jepsoneflora>[http://ucjeps.berkeley.edu/eflora/eflora_display.php?tid=9493 Jepson Herbarium, Jepson eFlora: ''Brodiaea'', family Themidaceae] . accessed 29 April 2016.</ref><ref>[http://www.calflora.org/cgi-bin/specieslist.cgi?where-family=THEMIDACEAE Calflora Database: Themidaceae genera] . accessed 1 May 2016.</ref><ref name=calflora>[http://www.calflora.org/cgi-bin/specieslist.cgi?where-genus=Brodiaea Calflora: ''Brodiaea'', family Themidaceae] . accessed 1 May 2016.</ref><ref>{{cite journal|last1=Pires|first1=J. C.|last2=Sytsma|first2=K. J.|title=A phylogenetic evaluation of a biosystematic framework: ''Brodiaea'' and related petaloid monocots ('''Themidaceae''')|journal=[[American Journal of Botany]]|date=1 August 2002|volume=89|issue=8|pages=1342–1359|doi=10.3732/ajb.89.8.1342|pmid=21665737}}</ref>

It was formerly classified within the [[Brodiaeoideae]] subfamily of the [[Asparagaceae]] family, in the Asparagales order.<ref>{{Cite web |last=Stevens |first=P.F. |title=Angiosperm Phylogeny Website: Asparagales: Brodiaeoideae |url=http://www.mobot.org/mobot/research/apweb/orders/asparagalesweb.htm#Themidaceae }}</ref><ref name="FNA">{{cite book
| last = Pires
| first = J. Chris
| chapter = 63. ''Brodiaea'' Smith
| title = Flora of North America
| volume =26
| pages =Page 20, 53, 55, '''321''', 326, 328, 331, 332, 336, 3
| publisher = [[Oxford University Press]]
| location = New York & Oxford
| url =http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=104654
| accessdate = 2008-06-27 }}</ref> The [[USDA]] Plants Database currently classifies the genus 'Brodiaea'' in the family [[Liliaceae]].<ref>[http://plants.usda.gov/java/ClassificationServlet?source=profile&symbol=BRODI&display=31 USDA Plants Database: Classification of ''Brodiaea''] . accessed 1 May 2016.</ref>

''Brodiaea'' species occur along the [[Pacific Coast]] region of [[North America]], from [[British Columbia]] throughout [[California]] into the [[Baja California Peninsula]].<ref>[http://plants.usda.gov/core/profile?symbol=BRODI USDA Plants: Distribution map of ''Brodiaea'']</ref> They are especially common in [[northern California]].<ref name="melany">[http://apps.kew.org/wcsp/namedetail.do?name_id=301035 Kew World Checklist of Selected Plant Families]</ref>

==Description==

''Brodiaea'' species are [[Herbaceous plant|herbaceous]] [[Perennial plant|perennials]], growing from [[corm]]s. Between one and six narrow leaves are produced from the corm. The bare flowering stem ([[Scape (botany)|scape]]) carries an [[umbel]] of flowers. Individual flowers have six blue to purple [[tepal]]s, joined at the base to form a tube with free lobes at the mouth. The outer three tepal lobes are narrower than the inner three.<ref name=FNA/>

In almost all species, inside the tepals and joined to their bases are three sterile stamens ([[staminode]]s), resembling small petals, each opposite one of the outer tepals. Three normal [[stamen]]s are also joined to the bases of the tepals and are placed opposite the inner ones. The base of the filaments of the stamens may be expanded into various shapes, such as flaps or wings. The size and shape of the staminodes and of the structures at the base of the filaments are important diagnostic characters. The [[Gynoecium#Pistils|compound pistil]] is formed of three [[carpel]]s forming a superior [[Ovary (botany)|ovary]] with three locules. The style which emerges between the three stamens has a three-lobed [[Stigma (botany)|stigma]]. The seeds are black.<ref name=FNA/>

==Taxonomy==

===Nomenclature===
The origin of the scientific name of the genus is somewhat tangled. Specimens of what is now called ''Brodiaea'' were first collected by [[Archibald Menzies]], botanist to the [[Vancouver Expedition]], in 1792. Menzies collected the plant from the vicinity of the [[Strait of Georgia]], named "New Georgia" by [[George Vancouver]]. The first published reference to the plant did not give it a name. This was in [[James Edward Smith]]'s 1807 ''An introduction to physiological and systematical botany'', where Smith used it to argue that the [[tepal]]s of [[lilioid monocot|liliaceous]] plants are [[sepal]]s rather than [[petal]]s: {{quote|"I cannot conceal a recent discovery which strongly confirms the opinion of my acute and candid friend. Two species of a new genus, found by Mr. Menzies on the West coast of North America, have beautiful liliaceous flowers like an Agapanthus, with six internal petals besides!"<ref name="Smith 1807">{{cite book | first = James Edward | last = Smith | authorlink = James Edward Smith | year = 1807 | title = [[Wikisource:An introduction to physiological and systematical botany|An introduction to physiological and systematical botany]] | page = [[Wikisource:An introduction to physiological and systematical botany#261|261]]}}</ref>}}

The following year, early in 1808, [[Richard Salisbury]] published a description of the first ''Brodiaea'' species in ''[[The Paradisus Londinensis]]'', naming it ''Hookera coronaria'', the genus name being in honour of the illustrator [[William Hooker (botanical illustrator)|William Hooker]].<ref>{{Cite book |last=Salisbury |first=R.S. |chapter=''Hookera coronaria'' |title=The Paradisus Londinensis |others=t. 98 |year=1808 |url=https://www.biodiversitylibrary.org/page/36898369 |accessdate=2013-09-11 }}</ref> Shortly afterwards, Smith named a moss genus ''[[Hookeria]]'', and in April 1808, he read to the [[Linnean Society of London]] a formal description of a new genus, based on the same species as Salisbury's ''Hookera coronaria'', naming the genus ''Brodiaea'' in honour of Scottish botanist [[James Brodie (botanist)|James Brodie]].<ref name="Smith 1811">{{cite journal | first = James Edward | last = Smith | year = 1810 | title = [[Wikisource:Transactions of the Linnean Society of London/Volume 10/Characters of a new Liliaceous Genus called Brodiaea|Characters of a new Liliaceous Genus called Brodiæa]] | journal = [[Wikisource:Transactions of the Linnean Society of London|Transactions of the Linnean Society of London]] | volume = [[Wikisource:Transactions of the Linnean Society of London/Volume 10|X]] | pages = 1–5}}</ref> Formal publication did not occur, however, until Smith's presentation went to print in 1810. [[George Simonds Boulger|George Boulger]], writing in the ''Dictionary of National Biography'', says that Smith's actions were deliberately intended to deprive Salisbury of credit for the genus.<ref name=DNB00>{{cite DNB|wstitle=Salisbury, Richard Anthony |first=George Simonds |last=Boulger |authorlink=George Simonds Boulger |volume=50 }}</ref>

If this was Smith's intention it was successful, since although Salisbury's genus name ''Hookera'' has [[Priority (nomenclature)|priority]] over Smith's name ''Brodiaea'', names as similar as ''Hookera'' and ''Hookeria'' are considered to be confusing and a formal proposal to [[Conserved name|conserve]] the names ''Brodiaea'' and ''Hookeria'' over the name ''Hookera'' was accepted.<ref name=RickStaf59>{{Cite journal |last=Rickett |first=H.W. |last2=Stafleu |first2=F.A. |year=1959 |title=Nomina generica conservanda et rejicienda spermatophytorum |journal=Taxon |volume=8 |issue=7 |pages=213–243 |doi=10.2307/1217883 |lastauthoramp=yes }}</ref> ''Brodiaea'' is thus a "conserved name" or "''nomen conservandum''", shown by the abbreviation "''nom. cons.''" after the name in botanical sources. The [[type (biology)|type]] species is now ''[[Brodiaea coronaria]]'', and the original type, ''Brodiaea grandiflora'' Sm., is an illegitimate name.<ref name=ing>{{cite web |url = http://botany.si.edu/ing/INGsearch.cfm?searchword=Brodiaea |title = ''Brodiaea'' |accessdate = 2008-06-27 |year = 1996 |work = Index Nominum Genericorum (Plantarum)|publisher = [[International Association for Plant Taxonomy]]|editor=Farr, E. R. and G. Zijlstra}}</ref>

===Phylogeny and classification===
''Brodiaea'' belongs to a group of 12 genera whose [[Phylogenetic tree|affinities]] were the subject of much controversy until the end of the 20th century. Salisbury treated them as a family which he named Themidaceae. Others placed this group at lower [[taxonomic rank]] and usually included them in [[Liliaceae]], [[Alliaceae]], or [[Amaryllidaceae]]. [[Molecular phylogenetic]] studies confirmed the suspicions of many that this group was misplaced, and consequently, the [[Family (biology)|family]] Themidaceae was resurrected in 1996.<ref name="fay1996">{{cite journal | last1 = Fay | first1 = Michael F. | last2 = Chase | first2 = Mark W. | year = 1996 | title = Resurrection of Themidaceae for the ''Brodiaea'' alliance, and recircumscription of Alliaceae, Amaryllidaceae, and Agapanthoideae | url = | journal = Taxon | volume = 45 | issue = 3| pages = 441–451 | doi=10.2307/1224136}}</ref> When the [[Angiosperm Phylogeny Group]] published the [[APG II system]] in 2003, Themidaceae was accepted as an optional family for those who wanted to [[Circumscription (taxonomy)|circumscribe]] families narrowly in the [[Order (biology)|order]] [[Asparagales]]. When the [[APG III system]] was published in 2009, the former Themidaceae was treated as a [[subfamily]], Brodiaeoideae, of the family [[Asparagaceae]] [[sensu lato]].<ref name="crf2009">{{cite journal | last1 = Chase | first1 = Mark W. | last2 = Reveal | first2 = James L. | last3 = Fay | first3 = Michael F. | year = | title = A subfamilial classification for the expanded asparagalean families Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae | url = | journal = Botanical Journal of the Linnean Society | volume = 161 | issue = 2| pages = 132–136 | doi=10.1111/j.1095-8339.2009.00999.x}}</ref>

Some sources, such as [[Integrated Taxonomic Information System|ITIS]], continue to use the [[polyphyletic]] [[Taxon|groups]] of obsolete [[Plant taxonomy|taxonomic]] systems.<ref>{{ITIS |id=42803 |taxon=''Brodiaea coronaria'' (Salisb.) Engl. |accessdate=27 June 2008}}</ref> Other sources, such as the [[Angiosperm Phylogeny Website]] mostly follow the Angiosperm Phylogeny Group.<ref name=APW_Asp>{{cite web
| url = http://www.mobot.org/MOBOT/Research/APweb/orders/asparagalesweb.htm#Asparagales
| title = Asparagales Bromhead
| accessdate = 2008-06-27
| last = Stevens |first=P.F.
| work = Angiosperm Phylogeny Website
| publisher = [[Missouri Botanical Garden]]
}}</ref>

Brodiaea (or brodeia<ref>[http://www.jpbeardsley.com//newplantslist.doc Encyclopedia of Herbs]</ref>) is also used as a common name to refer to three genera, ''Brodiaea'', ''[[Dichelostemma]]'', and ''[[Triteleia]]''. The latter two genera were once included as part of the genus ''Brodiaea''.<ref name="GRIN">{{cite web
| url = http://www.ars-grin.gov/cgi-bin/npgs/html/splist.pl?1721
| title = Species Records of ''Brodiaea''
| accessdate = 2008-06-27
| author = Germplasm Resources Information Network (GRIN)
| authorlink = Germplasm Resources Information Network
| date = 2007-10-05
| work = Taxonomy for Plants
| publisher = [[United States Department of Agriculture|USDA]], [[Agricultural Research Service|ARS]], National Genetic Resources Program, National Germplasm Resources Laboratory, Beltsville, Maryland
}}</ref>
The [[monophyly]] of ''Brodiaea'' as presently defined is not entirely certain. It might be [[Paraphyly|intermixed]] with ''Dichelostemma''.<ref name="pires2001">{{cite journal | last1 = Pires | first1 = J. Chris | last2 = Fay | first2 = Michael F. | last3 = Davis | first3 = Warren S. | last4 = Hufford | first4 = Larry | last5 = Rova | first5 = Johan | last6 = Chase | first6 = Mark W. | last7 = Sytsma | first7 = Kenneth J. | year = 2001 | title = Molecular and morphological phylogenetic analyses of Themidaceae (Asparagales) | url = | journal = Kew Bulletin | volume = 56 | issue = 3| pages = 601–626 | doi=10.2307/4117686}}</ref>

===Species===
{{As of|2013|September}}, the [[World Checklist of Selected Plant Families]] lists the following 17 species.<ref name="WCSP_Brodiaea">Search for "Brodiaea", {{Cite web |title=World Checklist of Selected Plant Families |publisher=[[Royal Botanic Gardens, Kew]] |url=http://apps.kew.org/wcsp/ |accessdate=2013-09-12 }}</ref><ref name=calflora/><ref name=jepsoneflora/><ref name=usdataxa>[http://plants.usda.gov/core/profile?symbol=BRODI USDA Plants Profile for ''Brodiaea'' − "Subordinate Taxa" tab] . accessed 29 April 2016.</ref><ref>[https://www.gbif.org/species/101317597 US National Plant Germplasm System−GRIN Taxonomy of ''Brodiaea''— with Subordinate Taxa list/links] . accessed 1 May 2016.</ref> English common names are from the ''Flora of North America''.<ref name="FNA"/>

# ''[[Brodiaea appendiculata]]'' Hoover - appendage cluster-lily - central California
# ''[[Brodiaea californica]]'' Lindl. ex Lem. - California cluster-lily - northern California, southwestern Oregon
# ''[[Brodiaea coronaria]]'' (Salisb.) Jeps. - harvest cluster-lily; Californian hyacinth - British Columbia, Washington, Oregon, much of California
# ''[[Brodiaea elegans]]'' Hoover - elegant cluster-lily - western Oregon, most of California
# ''[[Brodiaea filifolia]]'' S.Watson - threadleaf cluster-lily - southern California
# ''[[Brodiaea insignis]]'' (Jeps.) Niehaus - Kaweah cluster-lily - [[Tulare County, California|Tulare County]]
# ''[[Brodiaea jolonensis]]'' Eastw. - chaparral cluster-lily - southern California, northern [[Baja California]]
# ''[[Brodiaea kinkiensis]]'' Niehaus - San Clemente Island cluster-lily - [[San Clemente Island]]
# ''[[Brodiaea matsonii]]'' R.E.Preston - [[Shasta County, California|Shasta County]]
# ''[[Brodiaea minor]]'' (Benth.) S.Watson (syn. ''Brodiaea purdyi'' Eastw.) - vernalpool cluster-lily - northern California
# ''[[Brodiaea nana]]'' Hoover - northern California
# ''[[Brodiaea orcuttii]]'' (Greene) Baker - Orcutt's cluster-lily - southern California, northern [[Baja California]]
# ''[[Brodiaea pallida]]'' Hoover - Chinese Camp cluster-lily - [[Calaveras County, California|Calaveras]] and [[Tuolumne County, California|Tuolumne]] Counties
# ''[[Brodiaea santarosae]]'' T.J.Chester - Santa Rosa basalt brodiaea - [[Riverside County, California|Riverside]] and [[San Diego County, California|San Diego]] Counties
# ''[[Brodiaea sierrae]]'' R.E.Preston - [[Butte County, California|Butte]], [[Yuba County, California|Yuba]] and [[Nevada County, California|Nevada]] Counties
# ''[[Brodiaea stellaris]]'' S.Watson - starflower cluster-lily - [[Sonoma County, California|Sonoma]], [[Mendocino County, California|Mendocino]] and [[Humboldt County, California|Humboldt]] Counties
# ''[[Brodiaea terrestris]]'' Kellogg - dwarf cluster-lily - southwestern Oregon and much of coastal and southern California

;formerly included<ref name="WCSP_Brodiaea"/>
Numerous other names have been coined using the name ''Brodiaea,'' referring to species now regarded as better suited to other genera ''([[Androstephium]] [[Beauverdia]] [[Dandya (plant)|Dandya]] [[Dichelostemma]] [[Leucocoryne]] [[Nothoscordum]] [[Tristagma]] [[Triteleia]] [[Triteleiopsis]]).''

==Distribution and habitat==

''Brodiaea'' species are confined to western North America, from [[British Columbia]] in the north, through the [[West Coast of the United States]] region, to northwestern [[Mexico]] in the south.<ref name=usdataxa/><ref name=WCSP_301035>{{Cite web |title=''Brodiaea''|work=World Checklist of Selected Plant Families |publisher=[[Royal Botanic Gardens, Kew]] |url=http://apps.kew.org/wcsp/namedetail.do?name_id=301035 |accessdate=2013-09-12}}</ref> The majority of species are [[endemic]] to [[California]].<ref name=calflora/><ref name=jepsoneflora/>

Many are adapted to [[serpentine soil]]s or other soils with particular chemical compositions, resulting in limited distributions and several rare and endangered species.<ref name=FNA/> An example is ''[[Brodiaea pallida]]'', known only from two populations along the border between [[Tuolumne County, California|Tuolumne County]] and [[Calaveras County, California]].<ref name=five>{{cite web |author=USFWS |year=2007 |url=http://ecos.fws.gov/docs/five_year_review/doc1866.pdf |title=Five Year Review: ''B. pallida'' |accessdate=2013-09-12 }}</ref>

==Cultivation==
A number of species of ''Brodiaea'' are in cultivation. Species such as ''[[Brodiaea californica|B. californica]]'' and ''[[Brodiaea coronaria|B. coronaria]]'' are recommended for sunny positions in the garden, where they extend the flowering season of most [[ornamental bulb]]s, flowering in early summer rather than in spring. The flower heads ([[umbel]]s) of larger species can be dried for use as winter decorations. Smaller species, such as ''[[Brodiaea terrestris|B. terrestris]]'', may be grown in a [[bulb frame]] or [[alpine house]].<ref>{{Cite book |last=Mathew |first=Brian |year=1987 |title=The Smaller Bulbs |publication-place=London |publisher=B.T. Batsford |isbn=978-0-7134-4922-8 }} pp. 19–21.</ref>

==References==
[[Image:Brodiaea.jpg|thumb|right|240px|Brodiaea sp.]]
{{Reflist|26em}}

== Bibliography ==
{{refbegin}}
*
{{refend}}

==External links==
{{Wikisource-inline|Characters of a new Liliaceous Genus called Brodiaea}}
{{Wikispecies}}
*[http://www.calflora.org/cgi-bin/specieslist.cgi?where-genus=Brodiaea Calflora Dayabase: ''Brodiaea'' species]
*[http://ucjeps.berkeley.edu/eflora/eflora_display.php?tid=9493 Jepson Manual eFlora (TJM2) treatment of ''Brodiaea'']
*[http://plants.usda.gov/java/ClassificationServlet?source=profile&symbol=BRODI&display=31 USDA Plants Profile for ''Brodiaea'' species (brodiaea)]
{{Commons category|position=left|Brodiaea| Brodiaea}}

{{Taxonbar|from=Q1894962}}

[[Category:Brodiaea| 01]]
[[Category:Asparagaceae genera]]
[[Category:Flora of California]]

Ranavirus

2018-03-23T04:28:34Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q3418768}} (1 sig. taxon ID); WP:GenFixes, using AWB

{{Use dmy dates|date=April 2017}}
{{Taxobox
| name = Ranavirus
| image =CSIRO ScienceImage 2010 Ranavirus Pathogen.jpg
| image_caption =[[Transmission electron micrograph]] of ranaviruses (dark hexagons) gathering at the cell border and leaving the cell via a process called "[[budding]]".
| virus_group = i
| ordo = ''[[Incertae sedis]]''
| familia = [[Iridoviridae]]
| genus = '''''Ranavirus'''''
| subdivision_ranks = Type 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.]]

'''''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|language=en}}</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]].

==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. 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>

== 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>

==Evolution==
[[File:Reptiles, Amphibians in US Succumbing to Deadly Ranavirus.ogv|thumb|thumbtime=1:05|[[VOA]] report about ''Ranavirus'']]
The ranaviruses appear to have evolved from a fish virus which subsequently infected amphibians and reptiles.<ref name=Jancovich2010>{{cite journal | last1 = Jancovich | first1 = JK | last2 = Bremont | first2 = M | last3 = Touchman | first3 = JW | last4 = Jacobs | first4 = BL | year = 2010 | title = Evidence for multiple recent host species shifts among the Ranaviruses (family ''Iridoviridae'') | url = | journal = J Virol | volume = 84 | issue = 6| pages = 2636–2647 | doi = 10.1128/JVI.01991-09 | pmid = 20042506 | pmc = 2826071 }}</ref>

==Hosts==
Several reptile species are known to be affected:

*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>
*Burmese star tortoises (''[[Geochelone platynota]]'')
*Leopard tortoise (''[[Geochelone pardalis]]'')<ref name=Benetka2007>{{cite journal|author1=Benetka V.|year=2007|title=First report of an iridovirus (genus ''Ranavirus'') infection in a leopard tortoise (''Geochelone pardalis pardalis'')|journal= Vet Med Austria |volume=94|pages=243–248|url=http://www.schildkroeten-sfb.ch/fileadmin/docs/news/729-pantherschildkroete.pdf}}</ref>
*Gopher tortoises (''[[Gopherus polyphemus]]'')
*Mountain lizard (''[[Lacerta monticola]]'')<ref name=Alves2011>{{cite journal|title=New viruses from ''Lacerta monticola'' (Serra da Estrela, Portugal): Further evidence for a new group of nucleo-cytoplasmic large deoxyriboviruses (NCLDVs)|journal=Microscopy and microanalysis : the official journal of Microscopy Society of America, Microbeam Analysis Society, Microscopical Society of Canada |volume=17 |issue=1 |pages=101–8 |doi=10.1017/S143192761009433X |pmid=21138619|year=2011 |author1=De Matos |first1=A. P. |last2=Caeiro |first2=M. F. |last3=Papp |first3=T |last4=Matos |first4=B. A. |last5=Correia |first5=A. C. |last6=Marschang |first6=R. E. |bibcode=2011MiMic..17..101A }}</ref>
*Eastern box turtles (''[[Terrapene carolina carolina]]'')<ref name=Mao1997>{{cite journal | last1 = Mao | first1 = J | last2 = Hedrick | first2 = RP | last3 = Chinchar | first3 = VG | year = 1997 | title = Molecular characterization, sequence analysis, and taxonomic position of newly isolated fish iridoviruses | url = | journal = Virology | volume = 229 | issue = 1| pages = 212–220 | doi = 10.1006/viro.1996.8435 | pmid = 9123863 }}</ref>
*Florida box turtles (''[[Terrapene carolina bauri]]'')
*Western ornate box turtles (''[[Terrapene ornata]]'')<ref name=Johnson2007>{{cite journal|title= Experimental transmission and induction of ranaviral disease in Western Ornate box turtles (''Terrapene ornata ornata'') and red-eared sliders (''Trachemys scripta elegans'')|journal=Veterinary Pathology|volume=44|issue=3|pages=285–97|pmid=17491069|year=2007|author1=Johnson|first1=A. J.|last2=Pessier|first2=A. P.|last3=Jacobson|first3=E. R.|doi=10.1354/vp.44-3-285}}</ref>
*Spur-thighed tortoises (''[[Testudo graeca]]'')<ref name=Blahak2010>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</ref>
*Hermann’s tortoises (''[[Testudo hermanni]]'')
*Egyptian tortoises (''[[Testudo kleinmanni]]'')
*Russian tortoises (''[[Testudo horsfieldii]]'')
*Marginated tortoises (''[[Testudo marginata]]'')
*Red-eared sliders (''[[Trachemys scripta elegans]]'')<ref name="Johnson2007" />
*Chinese softshell turtles (''[[Trionyx sinensis]]'')<ref name=Chen1999>{{cite journal|title= A new iridovirus isolated from soft-shelled turtle|journal=Virus research|volume=63|issue=1–2|pages=147–51|pmid=10509727|doi=10.1016/S0168-1702(99)00069-6|year=1999|author1=Chen|first1=Z. X.|last2=Zheng|first2=J. C.|last3=Jiang|first3=Y. L.}}</ref>
*Gecko (''[[Uroplatus fimbriatus]]'')<ref name=Marschang2005>{{cite journal|title=Isolation of a ranavirus from a gecko (''Uroplatus fimbriatus'')|journal=Journal of zoo and wildlife medicine : official publication of the American Association of Zoo Veterinarians|volume=36|issue=2|pages=295–300|jstor=20096453|pmid=17323572|year=2005|author1=Marschang|first1=R. E.|last2=Braun|first2=S|last3=Becher|first3=P|doi=10.1638/04-008.1}}</ref>

==Taxonomy==
<big>'''Group: dsDNA'''</big>
{{Collapsible list|title= <big>Order: Unassigned</big>
|1={{Collapsible list| framestyle=border:none; padding:1.0em;|title=Family: [[Iridoviridae]]
|1={{hidden begin|title=Genus: Ranavirus}}
*[[Ambystoma tigrinum virus]]
*[[Bohle iridovirus]]
*[[Common midwife toad virus]]
*[[Epizootic haematopoietic necrosis virus]]
*[[European catfish virus]]
*'''''[[Frog virus 3]]'''''
*[[Santee-Cooper ranavirus]]
*[[Singapore grouper iridovirus]]
{{hidden end}}
}}
}}<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/>

==Structure==
Ranaviruses are large [[icosahedral]] DNA viruses measuring approximately 150 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).
{| class="wikitable sortable" style="text-align:center"
|-
! 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).<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|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 [[circularly permuted]] and exhibits [[terminally redundant DNA]].<ref name=Goorha/>
{| class="wikitable sortable" style="text-align:center"
|-
! 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.<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}}</ref>

==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}}</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}}</ref>

==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>

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

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

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

==External links==
{{Commons category|Ranavirus}}
{{Wikispecies-inline|List of viruses}}
* [https://talk.ictvonline.org/ictv-reports/ictv_online_report/dsdna-viruses/w/iridoviridae ICTV Online (10th) Report: Iridoviridae]
* [http://viralzone.expasy.org/all_by_species/585.html '''Viralzone''': Ranavirus]
* [http://www.ranavirus.net Ranavirus Research Project]
* [http://www.jcu.edu.au/school/phtm/PHTM/frogs/otherdiseases-viruses.htm Viral Diseases of Amphibians]

{{Baltimore classification}}

{{Taxonbar|from=Q3418768}}

[[Category:Iridoviridae]]
[[Category:Fish viral diseases]]
[[Category:Articles containing video clips]]

Tomato-bushy-stunt-Virus

2018-03-23T04:17:19Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q16861030}} (1 sig. taxon ID); WP:GenFixe using AWB

{{Use dmy dates|date=April 2017}}
{{Taxobox
| image = File:2tbv capsid ray.png
| image_caption = The capsid of the tomato bushy stunt virus, with the three symmetrically distinct coat protein (p41) monomers colored in orange, green, and blue.<ref name="hopper_1984">{{cite journal|last1=Hopper|first1=P|last2=Harrison|first2=SC|last3=Sauer|first3=RT|title=Structure of tomato bushy stunt virus. V. Coat protein sequence determination and its structural implications.|journal=Journal of Molecular Biology|date=25 August 1984|volume=177|issue=4|pages=701–13|pmid=6481803|doi=10.1016/0022-2836(84)90045-7}}</ref>
| virus_group = iv
| ordo = ''Unassigned''
| familia = ''[[Tombusviridae]]''
| genus = ''Tombusvirus''
| species = '''Tomato bushy stunt virus'''
}}

'''Tomato bushy stunt virus''' (TBSV) is a [[virus]] that is the [[type species]] of the [[tombusvirus]] family.<ref name="MahyRegenmortel2009">{{cite book|last1=Mahy|first1=Brian W. J.|last2=Regenmortel|first2=Marc H. V. Van|title=Desk Encyclopedia of Plant and Fungal Virology|url=https://books.google.com/books?id=8Mnnr-_mtIwC&pg=PA445|accessdate=4 December 2012|date=2009-10-15|publisher=Academic Press|isbn=978-0-12-375148-5|pages=445–}}</ref> It was first reported in [[tomato]]es in 1935 and primarily affects vegetable crops, though it is not generally considered an economically significant plant pathogen. Depending upon the host, TBSV causes stunting of growth, leaf mottling, and deformed or absent fruit. The virus is likely to be soil-borne in the natural setting, but can also transmitted mechanically, for example through contaminated cutting tools. TBSV has been used as a [[model organism|model system]] in [[virology]] research on the life cycle of [[plant virus]]es, particularly in experimental infections of the model host plant ''[[Nicotiana benthamiana]]''.<ref name="scholthof_2006">{{cite journal|last1=Scholthof|first1=Herman B.|title=The Tombusvirus-encoded P19: from irrelevance to elegance|journal=Nature Reviews Microbiology|date=6 March 2006|volume=4|issue=5|pages=405–411|doi=10.1038/nrmicro1395}}</ref><ref name="yamamura_2005">{{cite journal|last1=Yamamura|first1=Y|last2=Scholthof|first2=HB|title=Tomato bushy stunt virus: a resilient model system to study virus-plant interactions.|journal=Molecular plant pathology|date=1 September 2005|volume=6|issue=5|pages=491–502|pmid=20565674|doi=10.1111/j.1364-3703.2005.00301.x}}</ref>

==Host range==
[[File:Nicoatiana benthamiana plant.jpg|thumb|right|''N. benthamiana'', a common experimental host for TBSV.]]
TBSV has a broad [[host range]] under experimental conditions and has been reported to infect over 120 plant species spanning 20 families. However, under natural conditions its range is much narrower and generally comprises crop [[vegetable]]s and ornamental plants. It was first identified in [[tomato]] plants and also has been documented to affect [[apple]], [[artichoke]], [[cherry]], [[grapevine]], [[hops]], and [[Black pepper|pepper]]. Although it causes significant loss of yield in tomato plants, it is not considered an economically significant pathogen.<ref name=yamamura_2005 /><ref name=martelli_2001>{{Cite web|url=http://www.dpvweb.net/dpv/showdpv.php%3Edpvno=382|title=Tomato bushy stunt virus|last=Martelli|first=G.P.|last2=Russo|first2=M.|date=December 2001|website=Descriptions of Plant Viruses|publisher=Association of Applied Biologists|access-date=15 December 2016|last3=Rubino|first3=L.}}</ref> It is, however, a very well-established [[model organism|model system]] for the study of plant viruses, usually through experimental infection of ''[[Nicotiana benthamiana]]'' or ''[[Nicotiana clevelandii]]'', relatives of [[tobacco]] plants in which TBSV can cause systemic infection. Notably, the common model plant ''[[Arabidopsis thaliana]]'' is not a host.<ref name=scholthof_2006 /><ref name=yamamura_2005 /> TBSV can also replicate in [[yeast]] in laboratory conditions.<ref name=panavas_2003>{{Cite journal|last=Panavas|first=Tadas|last2=Nagy|first2=Peter D|date=2003-09-15|title=Yeast as a model host to study replication and recombination of defective interfering RNA of Tomato bushy stunt virus|url=http://www.sciencedirect.com/science/article/pii/S0042682203004367|journal=Virology|volume=314|issue=1|pages=315–325|doi=10.1016/S0042-6822(03)00436-7}}</ref>

==Signs==
The signs of TBSV are host-dependent. Local infections can cause [[necrosis|necrotic]] or [[chlorosis|chlorotic]] lesions. Systemic infections can cause stunted growth, deformed or absent fruit, and damaged leaves; in agricultural settings yield can be significantly reduced. The stunted, "bushy" appearance of the tomato plants in which the virus was first discovered gave the pathogen its name. In some hosts, most notably ''[[N. benthamiana]]'', TBSV can cause lethal systemic necrosis.<ref name=yamamura_2005 /><ref name=martelli_2001 />

==Transmission==
TBSV is thought to be passively transmitted in the wild, primarily through soil or water. There are no known [[vector (epidemiology)|vector]] organisms; transmission by [[aphid]]s, [[mite]]s, and the [[fungus]] ''[[Olpidium brassicae]]'' has specifically been ruled out.<ref name=martelli_2001 /> However, the closely related tombusvirus [[Cucumber necrosis virus]] (CNV) has been observed to be transmitted by ''[[Olpidium bornovanus]]'' [[zoospore]]s, so transmission of TBSV by as-yet unknown vector remains a possibility.<ref name=yamamura_2005 /> TBSV can also be transmitted through [[seed]] or by mechanical inoculation.<ref name=yamamura_2005 /><ref name=martelli_2001 /> In experimental tests, the virus can survive passage through the human [[digestive system]] if consumed in food and will remain infectious; it has been hypothesized that spread through [[sewage]] could occur.<ref name=balique_2015>{{Cite journal|last=Balique|first=Fanny|last2=Lecoq|first2=Hervé|last3=Raoult|first3=Didier|last4=Colson|first4=Philippe|date=2015-04-20|title=Can Plant Viruses Cross the Kingdom Border and Be Pathogenic to Humans?|url=http://www.mdpi.com/1999-4915/7/4/2074|journal=Viruses|language=en|volume=7|issue=4|pages=2074–2098|doi=10.3390/v7042074|pmc=4411691|pmid=25903834}}</ref>

==Distribution and management==
TBSV is distributed fairly widely across central and western Europe, north Africa, and North and South America.<ref name=yamamura_2005 /><ref name=martelli_2001 /> No specific control measures are recommended for the virus, though [[pest management]] guidelines distributed by the University of California recommend avoiding fields with a history of TBSV or using long [[crop rotation]]s.<ref name=ipm>{{Cite web|url=http://ipm.ucanr.edu/PMG/r783102411.html|title=Integrated Pest Management Program|last=|first=|date=December 2013|website=UC Pest Management Guidelines|publisher=University of California Division of Agriculture and Natural Resources|access-date=16 December 2016}}</ref>

==Taxonomy==
TBSV is the [[type species]] of the [[tombusvirus]] [[genus (biology)|genus]] in the family ''[[Tombusviridae]]''.<ref name=ictv>{{Cite web|url=http://ictvonline.org/taxonomyHistory.asp?taxnode_id=20154425&taxa_name=Tomato%20bushy%20stunt%20virus|title=ICTV Taxonomy History for Tomato bushy stunt virus|last=|first=|date=July 2015|website=International Committee on Taxonomy of Viruses|publisher=|access-date=16 December 2016}}</ref> Both the genus and the family derive their names from an abbreviation of "tomato bushy stunt virus".<ref name=harrison_1971>{{Cite journal|last=Harrison|first=B. D.|last2=Finch|first2=J. T.|last3=Gibbs|first3=A. J.|last4=Hollings|first4=M.|last5=Shepherd|first5=R. J.|last6=Valenta|first6=V.|last7=Wetter|first7=C.|date=1971-08-01|title=Sixteen groups of plant viruses|url=http://www.sciencedirect.com/science/article/pii/0042682271903369|journal=Virology|volume=45|issue=2|pages=356–363|doi=10.1016/0042-6822(71)90336-9}}</ref>

==Structure==
TBSV is an [[viral envelope|unenveloped]] [[icosahedral]] virus with a [[triangulation number|T=3]] [[viral capsid]] composed of 180 subunits of a single [[capsid protein]]. Its structure was studied extensively by [[X-ray crystallography]] from the late 1950s; its icosahedral symmetry was first identified by structural biologist [[Donald Caspar]], who also pioneered the study of the [[tobacco mosaic virus]].<ref name=rossmann_2013>{{Cite journal|last=Rossmann|first=Michael G.|date=2013-05-01|title=Structure of viruses: a short history|url=https://www.cambridge.org/core/journals/quarterly-reviews-of-biophysics/article/div-classtitlestructure-of-viruses-a-short-historydiv/F2A3C696CA5E684A6B3E4F670F80104A|journal=Quarterly Reviews of Biophysics|volume=46|issue=2|pages=133–180|doi=10.1017/S0033583513000012|issn=0033-5835}}</ref> A near-atomic-resolution map was obtained in 1978 by a research team including [[Stephen C. Harrison]].<ref name="harrison_1978">{{cite journal|last1=Harrison|first1=S. C.|last2=Olson|first2=A. J.|last3=Schutt|first3=C. E.|last4=Winkler|first4=F. K.|last5=Bricogne|first5=G.|title=Tomato bushy stunt virus at 2.9 Å resolution|journal=Nature|date=23 November 1978|volume=276|issue=5686|pages=368–373|doi=10.1038/276368a0}}</ref><ref name="HellemansBunch1988">{{cite book|last1=Hellemans|first1=Alexander|last2=Bunch|first2=Bryan H.|title=The timetables of science: a chronology of the most important people and events in the history of science|url=https://books.google.com/books?id=CpuWdyVKrAgC|accessdate=4 December 2012|year=1988|publisher=Simon and Schuster|isbn=978-0-671-62130-8}}{{Page needed|date=January 2016}}</ref>

==Genome and protein complement==
TBSV is a [[positive-sense single-stranded RNA virus]] with a linear [[genome]] of ~4800 [[nucleotide]]s.<ref name=hearne_1990>{{Cite journal|last=Hearne|first=Patrick Q.|last2=Knorr|first2=David A.|last3=Hillman|first3=Bradley I.|last4=Morris|first4=Thomas J.|date=1990-07-01|title=The complete genome structure and synthesis of infectious RNA from clones of tomato bushy stunt virus|url=http://www.sciencedirect.com/science/article/pii/0042682290904687|journal=Virology|volume=177|issue=1|pages=141–151|doi=10.1016/0042-6822(90)90468-7}}</ref><ref name="WagnerHewlett2007">{{cite book|last1=Wagner|first1=Edward K.|last2=Hewlett|first2=Martinez J.|last3=Bloom|first3=David C.|author4=David Camerini|title=Basic Virology|url=https://books.google.com/books?id=vYJBuebvgIcC&pg=PA268|accessdate=4 December 2012|date=2007-11-06|publisher=John Wiley & Sons|isbn=9781405147156|pages=268–}}</ref> The genome contains five [[gene]]s that encode a [[replicase]] composed of two proteins (p33 and p92), a capsid protein (called CP or p41), and two additional proteins, the [[RNA silencing suppressor p19]] and [[movement protein p22]].<ref name=yamamura_2005 /> These two proteins are expressed from [[overlapping gene]]s arranged so that the [[open reading frame]] of p19 is completely within the ORF of p22.<ref name=wu_2013>{{Cite journal|last=Wu|first=Baodong|last2=Grigull|first2=Jörg|last3=Ore|first3=Moriam O.|last4=Morin|first4=Sylvie|last5=White|first5=K. Andrew|date=2013-05-23|title=Global Organization of a Positive-strand RNA Virus Genome|url=http://journals.plos.org/plospathogens/article?id=10.1371/journal.ppat.1003363|journal=PLOS Pathogens|volume=9|issue=5|pages=e1003363|doi=10.1371/journal.ppat.1003363|issn=1553-7374|pmc=3662671|pmid=23717202}}</ref> The genome contains one additional possible gene, called pX, of unknown function.<ref name=yamamura_2005 />

===p33 and p92===
Together p33 and p92 comprise the viral [[replicase]] complex. P33 is smaller and p92 is produced through [[ribosome|ribosomal]] read-through of the p33 [[stop codon]], resulting in a shared [[N-terminal]] amino acid sequence and a large excess of p33 relative to p92. P33 proteins cooperatively bind single-stranded nucleic acids, while the p92 protein is a [[RNA-dependent RNA polymerase]] (RdRp). Both are [[essential gene|essential]] to viral proliferation. Both proteins are associated with [[cell membrane]]s.<ref name=yamamura_2005 />

===p41 (capsid protein)===
The viral [[capsid protein]] CP, or p41, is a double [[jelly roll fold|jelly roll]] protein that assembles into an [[icosahedral]] capsid containing 180 copies of the protein. Formation of [[virion]]s is not always necessary for localized spread of the virus into neighboring plant cells, because [[ribonucleoprotein]] particles containing viral genetic material can spread to immediate neighbors through [[plasmodesmata]]. However, the capsid protein is required for systemic infection.<ref name=yamamura_2005 />

===p19===
[[File:1R9F tombusvirus p19.png|thumb|right|The p19 protein in complex with double-stranded RNA.]]
The [[RNA silencing suppressor p19|p19 protein]] is a [[pathogenicity factor]] and functions by suppressing the [[RNA silencing]] pathway, a common form of antiviral defense. The p19 protein binds [[short interfering RNA]]s and prevents their incorporation into the [[RNA-induced silencing complex]] (RISC), thereby allowing viral propagation in the host plant.<ref name=scholthof_2006 /><ref name=scholthof_1995>{{cite journal |last1=Scholthof |first1=Herman B. |last2=Scholthof |first2=Karen-Beth G. |last3=Kikkert |first3=Marjolein |last4=Jackson |first4=A.O. |title=Tomato Bushy Stunt Virus Spread Is Regulated by Two Nested Genes That Function in Cell-to-Cell Movement and Host-Dependent Systemic Invasion |journal=Virology |volume=213 |issue=2 |pages=425–38 |year=1995 |pmid=7491767 |doi=10.1006/viro.1995.0015 }}</ref><ref>{{cite journal |last1=Jones |first1=Richard W. |last2=Jackson |first2=A.O. |last3=Morris |first3=Thomas J. |title=Defective-interfering RNAs and elevated temperatures inhibit replication of tomato bushy stunt virus in inoculated protoplasts |journal=Virology |volume=176 |issue=2 |pages=539–45 |year=1990 |pmid=2345965 |doi=10.1016/0042-6822(90)90024-L }}</ref> The presence of p19 is necessary for systemic infection or for lethal infection in some hosts; in the experimental host ''[[N. benthamiana]]'', p19 largely mediates the lethal systemic [[necrosis]] that is the outcome of TBSV infection.<ref name=yamamura_2005 /><ref name=scholthof_1995 />

===p22===
The p22 protein is a [[movement protein]] that is required for the virus to spread from cell to cell. P22 is an [[RNA-binding protein]] that is associated with the [[cell wall]] and facilitates movement of viral genetic material from one cell to its neighbor through interconnecting [[plasmodesmata]].<ref name=yamamura_2005 />

==Replication==
A TBSV [[virion]] contains one copy of its [[positive-sense single-stranded RNA virus|positive-sense single-stranded RNA]] [[genome]], which is linear and lacks a [[3']] [[polyadenine tail]] or [[5' cap]]. Nevertheless, the p33 and p92 proteins are [[translation (biology)|translated]] directly from genomic RNA. When the genome is replicated, two [[subgenomic RNA]] molecules are produced that act as [[messenger RNA]]; one from which the p41 (CP) gene is expressed, and one from the p19 and p22 genes are expressed. The [[overlapping gene|overlapping]] p19 and p22 genes are both translated through the effects of [[leaky scanning]].<ref name=yamamura_2005 /> Several long-distance interactions between linearly well-separated areas of the genome have been identified with functional importance in ensuring efficient replication.<ref name=wu_2013 />

==Defective interfering RNA==
[[Defective interfering particle|Defective interfering RNA]] (DI) molecules are RNAs that are produced from the viral genome but are not competent to infect cells on their own; instead they require [[coinfection]] with an intact "helper" virus. TBSV infections often produce significant numbers of DIs from consistent parts of the genome under experimental conditions, but this behavior has not been observed in the wild. Their production is likely to be host specific. Infections that give rise to DIs usually have milder signs.<ref name=yamamura_2005 /><ref name=scholthof_1995_2>{{Cite journal|last=Scholthof|first=Karen-Beth G.|last2=Scholthof|first2=Herman B.|last3=Jackson|first3=Andrew O.|date=1995-08-01|title=The Effect of Defective Interfering RNAs on the Accumulation of Tomato Bushy Stunt Virus Proteins and Implications for Disease Attenuation|url=http://www.sciencedirect.com/science/article/pii/S0042682285714109|journal=Virology|volume=211|issue=1|pages=324–328|doi=10.1006/viro.1995.1410}}</ref>

==References==
{{Reflist|30em}}

{{Taxonbar|from=Q16861030}}

[[Category:Tomato pathogens and pests]]
[[Category:Viral plant pathogens and diseases]]
[[Category:Tombusviridae]]

Wurzeltrüffeln

2018-03-23T04:06:58Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q874176}} (3 sig. taxon IDs); WP:GenFixes, using AWB

{{Taxobox
| image = Rhizopogon rubescens.jpg
| image_caption = ''Rhizopogon rubescens''
| regnum = [[Fungi]]
| divisio = [[Basidiomycota]]
| classis = [[Agaricomycetes]]
| ordo = [[Boletales]]
| subordo = [[Suillineae]]
| familia = [[Rhizopogonaceae]]
| genus = '''''Rhizopogon'''''
| genus_authority = [[Elias Magnus Fries|Fr.]] (1817)
| type_species = ''Rhizopogon luteolus''
| type_species_authority = Fr. & Nordholm (1817)
| subdivision_ranks = Species
| subdivision =
}}

'''''Rhizopogon''''' is a [[genus]] of [[ectomycorrhizae|ectomycorrhizal]] [[Basidiomycetes]] in the family [[Rhizopogonaceae]]. Species form [[sporocarp (fungi)|hypogeous sporocarps]] commonly referred to as "[[false truffle]]s". The general [[morphology (biology)|morphological]] characters of ''Rhizopogon'' sporocarps are a [[wikt:simplex|simplex]] or [[wikt:duplex|duplex]] [[peridium]] surrounding a [[locule|loculate]] [[gleba]] that lacks a [[sporangium#Internal structures|columnella]]. [[Basidiospores]] are produced upon [[basidia]] that are borne within the fungal [[hymenium]] that coats the interior surface of gleba locules. The peridium is often adorned with thick mycelial cords, also known as [[rhizomorph]]s, that attach the sporocarp to the surrounding [[substrate (biology)|substrate]]. The scientific name ''Rhizopogon'' is Greek for 'root' (Rhiz-) 'beard' (-pogon) and this name was given in reference to the rhizomorphs found on sporocarps of many species.

''Rhizopogon'' species are primarily found in ectomycorrhizal association with trees in the family [[Pinaceae]] and are especially common [[wikt:symbiont|symbionts]] of [[pine]], [[fir]], and [[Douglas fir]] trees. Through their ectomycorrhizal relationships ''Rhizopogon'' are thought to play an important role in the ecology of [[coniferous forests]]. Recent [[wikt:micromorphological|micromorphological]] and [[molecular phylogenetics|molecular phylogenetic]] study has established that ''Rhizopogon'' is a member of the [[Boletales]], closely related to ''[[Suillus]]''.<ref>{{cite journal |author1=Manfred Binder |author2=David S. Hibbett |lastauthoramp=yes |year=2006 |title=Molecular systematics and biological diversification of Boletales |journal=[[Mycologia]] |volume=98 |issue=6 |pages=971–981 |doi=10.3852/mycologia.98.6.971 |pmid=17486973}}</ref>

==Taxonomy and diversity==
[[File:2009-09-17 Rhizopogon obtextus 76099.jpg|thumb|left|alt=An image of ''Rhizopogon luteolus'' (''=obtextus'') showing rhizomorphs with attached substrate.|A sporocarp of ''Rhizopogon luteolus'' (''=obtextus'') showing rhizomorphs with adhering substrate]]

===Historical classification===
The genus ''Rhizopogon'' occurs throughout the natural and introduced ranges of family Pinaceae trees. Though this range covers much of the [[temperate climate|northern temperate zones]], the diversity of ''Rhizopogon'' species is well characterized only in [[North America]] and [[Europe]]. There are currently over 150 recognized species of ''Rhizopogon''. The morphology of ''Rhizopogon'' species is highly cryptic and characters vary greatly throughout sporocarp maturity. This has led to the description of multiple species from various developmental stages of a single fungus.

The genus ''Rhizopogon'' was first described from Europe by [[Elias Magnus Fries]] in 1817.<ref>{{cite book |last=Fries |first=Elias Magnus |year=1817 |title=Symbolae Gasteromycorum |location=Lundae |publisher= Ex officina Berlingiana}}</ref> The North American [[monograph#Usage#Taxonomy (systematic biology)|monograph]] was produced by [[Alexander H. Smith]] in 1966<ref>{{cite journal |vauthors=Smith AH, Zeller SM | year=1966 |title=A Preliminary Account of the North American Species of ''Rhizopogon'' |journal=Memoirs of the New York Botanical Garden |volume=14 |issue=2 |pages=1–178 | url= http://name.umdl.umich.edu/AJN6347.0001.001}}</ref> with second author credits given [[wikt:posthumous|posthumously]] to [[Sanford Myron Zeller]] due to his contributions to the study of the genus. A European monograph of ''Rhizopogon'' has also been published.<ref>{{cite book |last=Martín |first=MP |year=1996 |title=The Genus Rhizopogon in Europe |location=Barcelona, Spain |publisher= BCG | pages= 173 p. | isbn= 8992161700}}</ref> In the recent past, molecular phylogenetic methods have allowed the revision of the taxonomic concepts of the genus ''Rhizopogon''<ref name="grubisha">{{cite journal |vauthors=Grubisha LC, Trappe JM, Molina R, Spatafora JW |year=2002 |title=Biology of the ectomycorrhizal genus ''Rhizopogon''. VI. Re-examination of infrageneric relationships inferred from phylogenetic analyses of ITS sequences |journal=[[Mycologia]] |volume=94 |issue=4 |pages=607–619| pmid=21156534 |doi=10.2307/3761712}}</ref>

===Modern classification===

Modern taxonomic concepts of the genus ''Rhizopogon'' recognize five subgenera of ''Rhizopogon''.<ref name="grubisha" /> These are subgenus ''[[Rhizopogon subgenus Rhizopogon|Rhizopogon]]'', subgenus ''[[Rhizopogon subgenus Versicolores|Versicolores]]'', subgenus ''[[Rhizopogon subgenus Villosuli|Villosuli]]'', subgenus ''[[Rhizopogon subgenus Amylopogon|Amylopogon]]'', and subgenus ''[[Rhizopogon subgenus Roseoli|Roseoli]]''.

==Ecology==
[[File:2010-05-14 Rhizopogon roseolus 2.jpg|thumb|right|alt=An image of ''Rhizopogon roseolus'' showing a close up of gleba locules.|A sporocarp of ''Rhizopogon roseolus'' in cross section showing a close up of the gleba locules]]

===Mammalian diet and spore dispersal===
''Rhizopogon'' species have been established as a common component in the diet of many small mammals <ref>{{cite journal |vauthors=Maser C, Maser Z |year=1988 |title=Interactions among squirrels, mycorrhizal fungi, and coniferous forests in Oregon |journal=Western North American Naturalist |volume=48 |issue=3 |pages=358–369}}</ref><ref>{{cite journal |vauthors=Izzo AD, Meyer M, Trappe JM, North M, Bruns TD |year=2005 |title=Hypogeous ectomycorrhizal fungal species on roots and in small mammal diet in a mixed conifer forest |journal=Forest Science |volume=51 |issue=3 |pages=243–254}}</ref> as well as deer<ref>{{cite journal |vauthors=Ashkannehhad S, Horton TR|lastauthoramp=yes |year=2006 |title=Ectomycorrhizal ecology under primary succession on coastal sand dunes: interactions involving ''Pinus contorta'', suilloid fungi and deer |journal=[[New Phytologist]] |volume=169 |issue=2 |pages=345–354| doi=10.1111/j.1469-8137.2005.01593.x | pmid=16411937}}</ref> in Western North America. The viability of ''Rhizopogon'' spores is maintained <ref name="colgan">{{cite journal |vauthors=Colgan III W, Claridge AW |year=2002 |title=Mycorrhizal effectiveness of ''Rhizopogon'' spores recovered from faecal pellets of small forest-dwelling mammals |journal=[[Mycological Research]] |volume=106 |issue=3 |pages=314–320| doi=10.1017/S0953756202005634}}</ref><ref>{{cite journal |vauthors=Kotter M, Farentinos RC |year=1984 |title=Formations of Ponderosa pine ectomycorrhizae after inoculation with feces of tassel-earred squirrels |journal=[[Mycologia]] |volume=76 |issue=2 |pages=758–760 |doi=10.2307/3793237 }}</ref> and may even be increased after mammalian gut passage,<ref name="colgan" /> making mammals an important [[dispersal vector]] for ''Rhizopogon''.

===Disturbance ecology===
''Rhizopogon'' species are common members of the fungal communities that colonize the roots of trees during seedling establishment and persist into old growth stands.<ref>{{cite journal |vauthors=Twieg BD, Durall DM, Simard SW |year=2007 |title=Ectomycorrhizal fungal succession in mixed temperate forests|journal=[[New Phytologist]] |volume=176 |issue=2 |pages=437–447| doi=10.1111/j.1469-8137.2007.02173.x | pmid=17888121 }}</ref><ref>{{cite book |last1=Molina |first1=R |last2=Trappe |first2=JM |last3=Grubisha |first3=LC |last4=Spatafora |first4=JW |editor1-last= Cairney |editor1-first=JWG |editor2-last= Chambers |editor2-first=SM |year=1999 |title=Ectomycorrhizal Fungi Key Genera in Profile | chapter=''Rhizopogon'' |location= Heidelberg |publisher= Springer Berlin | pages= 129–161 |doi=10.1007/978-3-662-06827-4_5 |isbn= 978-3-642-08490-4 }}</ref> ''Rhizopogon'' spores are long lived in soil and the spores of some species can persist for at least four years with an increase in viability over time.<ref>{{cite journal |author= Bruns, TD |author2= Peay KG |author3= Boynton PJ |author4= Grubisha LC |author5= Hynson NA |author6= Nguyen NH |author7= Rosenstock NP |year=2009 |title=Inoculum potential of ''Rhizopogon'' spores increases with time over the first 4 yr of a 99-yr spore burial experiment |journal=[[New Phytologist]] |volume=181 |issue=2 |pages=463–470 |doi=10.1111/j.1469-8137.2008.02652.x | pmid= 9121040}}</ref> ''Rhizopogon'' seems to be especially common upon the roots of establishing tree seedlings following disturbance such as fire<ref name="baar">{{cite journal |author1=Baar J. |author2=Horton T.R. |author3=Kretzer A.M. |author4=Bruns T.D. |year=1999 |title=Mycorrhizal colonization of ''Pinus muricata'' from resistant propagules after a stand-replacing wildfire |journal=[[New Phytologist]] |volume=143 |pages=409–418 |issue=2 |doi=10.1046/j.1469-8137.1999.00452.x}}</ref> or logging.<ref>{{cite journal |vauthors=Luoma DL, Stockdale CA, Molina R, Eberhart JL |year=2006 |title=The spatial influence of ''Pseudotsuga menziesii'' retention trees on ectomycorrhiza diversity |journal=[[Canadian Journal of Forest Research]] |volume=36 |issue=10 |pages=2561–2573| doi=10.1139/x06-143 }}</ref> ''Rhizopogon'' are also abundant colonizers of pot cultivated<ref name="baar" /><ref>{{cite journal |vauthors=Taylor DL, Bruns TD |year=1999 |title=Community structure of ectomycorrhizal fungi in a ''Pinus muricata'' forest: minimal overlap between the mature forest and resistant propagule communities |journal=[[Molecular Ecology]] |volume=8 |issue=11 |pages=1837–1850| doi=10.1046/j.1365-294x.1999.00773.x}}</ref><ref>{{cite journal |vauthors=Kjøller R, Bruns TD |year=2003 |title=''Rhizopogon'' spore bank communities within and among California pine forests |journal=[[Mycologia]] |volume=95 |issue=4 |pages=603–613| pmid=21148969 |doi=10.2307/3761936}}</ref><ref>{{cite journal |vauthors=Murata M, Kinoshita A, Nara K |year=2013 |title=Revisiting the host effect on ectomycorrhizal fungal communities: implications from host–fungal associations in relict ''Pseudotsuga japonica'' forests |journal=Mycorrhiza |volume=23 |issue=8 |pages=641–653| doi=10.1007/s00572-013-0504-0 | pmid=23702643}}</ref> and field cultivated <ref name="baar" /> conifer seedlings growing in soil from conifer stands that lacked observations of ''Rhizopogon'' upon the roots of mature trees. These finding suggest that ''Rhizopogon'' species are an important factor in the recovery of conifer forests following disturbance.

==Species==
{{div col|cols=2}}
*''[[Rhizopogon albidus]]''
*''[[Rhizopogon ater]]''
*''[[Rhizopogon amylopogon]]''
*''[[Rhizopogon atroviolaceus]]''
*''[[Rhizopogon brunneniger]]''
*''[[Rhizopogon ellenae]]''
*''[[Rhizopogon evadens]]''
*''[[Rhizopogon fulvigleba]]''
*''[[Rhizopogon fuscorubens]]''
*''[[Rhizopogon hawkerae]]''
*''[[Rhizopogon luteolus]]''
*''[[Rhizopogon nigrescens]]''
*''[[Rhizopogon occidentalis]]''
*''[[Rhizopogon ochraceorubens]]''
*''[[Rhizopogon parksii]]''
*''[[Rhizopogon parvisporus ]]''
*''[[Rhizopogon pedicellus]]''
*''[[Rhizopogon roseolus]]''
*''[[Rhizopogon salebrosus]]''
*''[[Rhizopogon subareolatus]]''
*''[[Rhizopogon subaustralis]]''
*''[[Rhizopogon subcaerulescens]]''
*''[[Rhizopogon subpurpurascens]]''
*''[[Rhizopogon subsalmonius]]''
*''[[Rhizopogon succosus]]''
*''[[Rhizopogon togasawariana]]''
*''[[Rhizopogon truncatus]]''
*''[[Rhizopogon vesiculosus]]''
*''[[Rhizopogon villosulus]]''
*''[[Rhizopogon vinicolor]]''
*''[[Rhizopogon vulgaris]]''
{{div col end}}

==[[Ethnomycology]]==

===Forestry===
The first intentional use of ''Rhizopogon'' species in forestry occurred in the early part of the 20th century when ''[[Rhizopogon luteolus]]'' was deliberately introduced into ''[[Pinus radiata]]'' plantations in [[Western Australia]] after it was observed to improve tree growth.<ref>{{cite journal |author= Kessel SL |year=1927 |title=Soil organisms. The dependence of certain pine species on a biological soil factor. |journal=Empire Forestry Journal |volume=6 |pages=70–74}}</ref> Since that time, ''Rhizopogon'' species have been widely studied as a component of managed forests. ''Rhizopogon'' species have been noted as common members of the ectomycorrhizal community colonizing tree roots of pine and Douglas-fir timber plantations.<ref>{{cite journal |vauthors=Molina R, Trappe JM |year=1994 |title=Biology of the ectomycorrhizal genus, ''Rhizopogon'' I. Host associations, host-specificity and pure culture syntheses |journal=[[New Phytologist]] |volume=126 |issue=4 |pages=653–675| doi=10.1111/j.1469-8137.1994.tb02961.x}}</ref> Naturally occurring ''Rhizopogon roseolus'' (''=rubescens'') spores have been shown to out-compete the spores of other ectomycorrhizal fungi in pine plantations even when competing spores were directly inoculated onto seedlings.<ref>{{cite journal |vauthors=Karkouri KE, Martin F, Mousain D |year=2002 |title=Dominance of the mycorrhizal fungus ''Rhizopogon rubescens'' in a plantation of ''Pinus pinea'' seedlings inoculated with ''Suillus collinitus'' |journal=Annals of Forest Science |volume=59 |issue=2 |pages=197–204| doi=10.1051/forest:2002006}}</ref> The survival rate and performance of pine<ref>{{cite journal |vauthors=Steinfield D, Amaranthus M, Cazares E |year=2003 |title=Survival of Ponderosa pine (''Pinus ponderosa'' Dougl. ex Laws) seedlings outplanted with ''Rhizopogon'' mycorrhizae inoculated with spores at the nursery |journal=Journal of Arborculture |volume=29 |issue=4 |pages=4197–208}}</ref> and Douglas-fir<ref>{{cite journal |vauthors=Castellano MA, Trappe JM |year=1985 |title=Ectomycorrhizal formation and plantation performance of Douglas-fir nursery stock inoculated with ''Rhizopogon'' spores |journal=[[Canadian Journal of Forest Research]] |volume=15 |issue=4 |pages=613–617 |doi=10.1139/x85-100}}</ref> plantation seedlings are increased after inoculation with ''Rhizopogon'' species.

===Gastronomy===

Though many species of ''Rhizopogon'' are considered edible, most are not held in high culinary esteem.<ref>{{cite book |last1=Trappe |first1=M |last2=Evans |first12=F |last3=Trappe |first3=J |year=2007 |title=Field guide to North American Truffles |location=Berkeley, CA |publisher= Ten Speed Press | pages= 136 p. |isbn= 1580088627 }}</ref> A notable exception is ''[[Rhizopogon roseolus]]'' (''=rubescens'') which is considered a delicacy in east Asia and especially in [[Japan]] where it is traditionally known as Shoro.<ref name="yun">{{cite journal |vauthors=Yun W, Hall IR |year=2004 |title=Edible ectomycorrhizal mushrooms: challenges and achievements |journal=[[Canadian Journal of Botany]] |volume=82 |issue=8 |pages=1063–1073 |doi=10.1139/b04-051}}</ref> Techniques for the commercial cultivation of this fungus in pine plantations have been developed and applied with successful results in Japan and [[New Zealand]].<ref name="yun" />

==References==
{{Reflist|colwidth=30em}}

==External links==
{{Commons category|Rhizopogon}}
*[http://www.indexfungorum.org/Names/genusrecord.asp?RecordID=19297 Index Fungorum]
*[http://quod.lib.umich.edu/cgi/t/text/text-idx?c=fung1tc;cc=fung1tc;view=toc;idno=AJN6347.0001.001 A preliminary account of the North American species of ''Rhizopogon''] by Alexander H. Smith and S. M. Zeller, 1966. (Full text of monograph.)

{{Taxonbar|from=Q874176}}

[[Category:Rhizopogonaceae]]
[[Category:Boletales genera]]

Tombusviridae

2018-03-23T03:51:50Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q3117439}} (2 sig. taxon IDs); WP:GenFixes using AWB

{{Use dmy dates|date=April 2017}}
{{taxobox
| virus_group = iv
| familia = '''''Tombusviridae'''''
| subdivision_ranks = Genera
| subdivision =
*''[[Betanecrovirus]]''
*''[[Aureusvirus]]''
*''[[Machlomovirus]]''
*''[[Panicovirus]]''
*''[[Alphanecrovirus]]''
*''[[Umbravirus]]''
*''[[Avenavirus]]''
*''[[Dianthovirus]]''
*''[[Carmovirus]]''
*''[[Macanavirus]]''
*''[[Zeavirus]]''
*''[[Tombusvirus]]''
*''[[Gallantivirus]]''
}}

'''''Tombusviridae''''' is a family of single-stranded [[sense (molecular biology)|positive sense]] [[RNA]] plant [[virus]]es. There are currently 71 species in this family, divided among 13 genera.<ref name=ViralZone>{{cite web|title=Viral Zone|url=http://viralzone.expasy.org/all_by_species/53.html|publisher=ExPASy|accessdate=15 June 2015}}</ref><ref name=ICTV>{{cite web|last1=ICTV|title=Virus Taxonomy: 2014 Release|url=http://ictvonline.org/virusTaxonomy.asp|accessdate=15 June 2015}}</ref> The name is derived from the type species of the ''[[Tombusvirus]]'' genus, [[Tomato bushy stunt virus]] (TBSV).<ref>Habili, N. and Symons, R. H. (1989). [http://nar.oxfordjournals.org/cgi/content/abstract/17/23/9543 Evolutionary relationship between luteoviruses and other RNA plant viruses based on sequence motifs in their putative RNA polymerases and nucleic acid helicases.] ''Nucleic Acids Research'' '''17''':23, 9543–55</ref>

==Genome==
All Tombusviridae have a non-segmented linear [[genome]], with the exception of Dianthoviruses, whose genome is bipartite.<ref name="wiley">[http://mrw.interscience.wiley.com/emrw/9780470015902/els/article/a0000756/current/html Wiley InterScience Encyclopedia of Life Sciences: Tombusviridae]</ref> The genome is approximately 4.6–4.8kb in length, lacks a [[5' cap]] and a poly(A) tail, and it encodes 4–6 [[open reading frame|ORF]]s. The polymerase encodes an [[stop codon|amber stop codon]] which is the site of a readthrough event within ORF1, producing two products necessary for replication. There is no [[helicase]] encoded by the virus.

==Structure==
The RNA is encapsulated in an [[icosahedron|icosahedral]] (T=3) [[capsid]], composed of 180 units of a single coat protein 27–42K in size; the [[virion]] measures 28–35 nm in diameter, and it is not enveloped.<ref name=ViralZone /><ref name="ictvdb">[https://www.ncbi.nlm.nih.gov/ICTVdb/ICTVdB/index.htm ICTVdB—The Universal Virus Database, version 3] 00.074. ''Tombusviridae''</ref>

{| class="wikitable sortable" style="text-align:center"
|-
! Genus !! Structure || Symmetry !! Capsid !! Genomic arrangement !! Genomic segmentation
|-
|Tombusvirus||Icosahedral||T=3||Non-enveloped||Linear||Monopartite
|-
|Gallantivirus||Icosahedral||T=3||Non-enveloped||Linear||Monopartite
|-
|Macanavirus||Icosahedral||T=3||Non-enveloped||Linear||Monopartite
|-
|Dianthovirus||Icosahedral||T=3||Non-enveloped||Linear||Bipartite
|-
|Carmovirus||Icosahedral||T=3||Non-enveloped||Linear||Monopartite
|-
|Alphanecrovirus||Icosahedral||T=3||Non-enveloped||Linear||Monopartite
|-
|Avenavirus||Icosahedral||T=3||Non-enveloped||Linear||Monopartite
|-
|Panicovirus||Icosahedral||T=3||Non-enveloped||Linear||Monopartite
|-
|Betanecrovirus||Icosahedral||T=3||Non-enveloped||Linear||Monopartite
|-
|Aureusvirus||Icosahedral||T=3||Non-enveloped||Linear||Monopartite
|-
|Umbravirus||Icosahedral||T=3||Non-enveloped||Linear||Monopartite
|-
|Machlomovirus||Icosahedral||T=3||Non-enveloped||Linear||Monopartite
|-
|Zeavirus||Icosahedral||T=3||Non-enveloped||Linear||Monopartite
|}

==Life cycle==
Viral replication is cytoplasmic, and is lysogenic. Entry into the host cell is achieved by penetration into the host cell. Replication follows the positive stranded RNA virus replication model. Positive stranded RNA virus transcription, using the premature termination model of subgenomic RNA transcription is the method of transcription. Translation takes place by leaky scanning, −1 ribosomal frameshifting, viral initiation, and suppression of termination. The virus exits the host cell by tubule-guided viral movement. Plants serve as the natural host. Transmission routes are mechanical, seed borne, and contact.<ref name=ViralZone />

Viruses in this family are primarily soil-borne, some transmitted by fungal species of the order [[Chytridiales]], others by no known vector. Virions may spread by water, root growth into infected soil, contact between plants, pollen, or seed, depending on the virus species. These viruses may be successfully transmitted by [[grafting]] or mechanical inoculation, and both the virion and the genetic material alone are ineffective.<ref name="ictvdb" />

{| class="wikitable sortable" style="text-align:center"
|-
! Genus !! Host details !! Tissue tropism !! Entry details !! Release details !! Replication site !! Assembly site !! Transmission
|-
|Tombusvirus||Plants||None||Viral movement; mechanical inoculation||Viral movement||Cytoplasm||Cytoplasm||Mechanical: contact; seed
|-
|Gallantivirus||Plants||None||Viral movement; mechanical inoculation||Viral movement||Cytoplasm||Cytoplasm||Mechanical: contact; seed
|-
|Macanavirus||Plants||None||Viral movement; mechanical inoculation||Viral movement||Cytoplasm||Cytoplasm||Mechanical: contact; seed
|-
|Dianthovirus||Plants||None||Viral movement; mechanical inoculation||Viral movement||Cytoplasm||Cytoplasm||Mechanical: contact; seed
|-
|Carmovirus||Plants||None||Viral movement; mechanical inoculation||Viral movement||Cytoplasm||Cytoplasm||Mechanical: contact; seed
|-
|Alphanecrovirus||Plants||None||Viral movement; mechanical inoculation||Viral movement||Cytoplasm||Cytoplasm||Mechanical: contact; seed
|-
|Avenavirus||Plants||None||Viral movement; mechanical inoculation||Viral movement||Cytoplasm||Cytoplasm||Mechanical: contact; seed
|-
|Panicovirus||Plants: panicae||None||Viral movement; mechanical inoculation||Viral movement||Cytoplasm||Cytoplasm||Mechanical: contact; seed
|-
|Betanecrovirus||Plants||None||Viral movement; mechanical inoculation||Viral movement||Cytoplasm||Cytoplasm||Mechanical: contact; seed
|-
|Aureusvirus||Plants||None||Viral movement; mechanical inoculation||Viral movement||Cytoplasm||Cytoplasm||Mechanical: contact; seed
|-
|Umbravirus||Plants||None||Viral movement; mechanical inoculation||Viral movement||Cytoplasm||Cytoplasm||Mechanical: contact; seed
|-
|Machlomovirus||Plants||None||Viral movement; mechanical inoculation||Viral movement||Cytoplasm||Cytoplasm||Mechanical: contact; seed
|-
|Zeavirus||Plants||None||Viral movement; mechanical inoculation||Viral movement||Cytoplasm||Cytoplasm||Mechanical: contact; seed
|}

==Replication==
Members of Tombusviridae [[Replicate (biology)|replicate]] in the cytoplasm, by use of negative strand templates. The replication process leaves a surplus of positive [[Sense (molecular biology)|sense]] (+)RNA strands, and it is thought that not only does the viral RNA act as a template for replication, but is also able to manipulate and regulate [[RNA synthesis]].{{Citation needed|date=March 2009}}

The level of RNA synthesis has been shown to be affected by the [[cis-acting]] properties of certain elements on the RNA (such as [[Red clover necrotic mosaic virus translation enhancer elements|RNA1 and 2]]<ref>{{cite journal |vauthors=Lommel SA, Weston-Fina M, Xiong Z, Lomonossoff GP |title=The nucleotide sequence and gene organization of red clover necrotic mosaic virus RNA-2 |journal=Nucleic Acids Res. |volume=16 |issue=17 |pages=8587–602 |date=September 1988 |pmid=3047682 |pmc=338578 |doi= 10.1093/nar/16.17.8587|url=}}</ref><ref>{{cite journal |vauthors=Mizumoto H, Tatsuta M, Kaido M, Mise K, Okuno T |title=Cap-independent translational enhancement by the 3' untranslated region of red clover necrotic mosaic virus RNA1 |journal=J. Virol. |volume=77 |issue=22 |pages=12113–21 |date=November 2003 |pmid=14581548 |pmc=254280 |doi= 10.1128/JVI.77.22.12113-12121.2003|url=}}</ref>), which include core promoter sequences which regulate the site of initiation for the complementary RNA strand synthesis. This mechanism is thought to be recognised by RNA-dependent [[RNA polymerase]], found encoded within the genome.{{Citation needed|date=March 2009}}

Tombusviridae have been found to co-opt [[GAPDH]], a host metabolic enzyme, for use in the replication center. GAPDH may bind to the (−)RNA strand and keep it in the replicase complex, allowing (+)RNA strands synthesized from it to be exported and accumulate in the host cell. Downregulation of GAPDH reduced viral RNA accumulation, and eliminated the surplus of (+)RNA copies.<ref>Wang, R. and Nagy, P. (2008) Tomato bushy stunt virus Co-Opts the RNA-Binding Function of a Host Metabolic Enzyme for Viral Genomic RNA Synthesis. ''Cell Host & Microbe'' '''3''':3 178–187</ref>

==Notes==
Research has shown that infection of plants from tombusviruses contain defective interfering RNAs that are born directly from the viruses RNA genome, and no host genome. Viral DI RNAs with their small size and cis-acting elements are good templates both ''in vivo'' and ''in vitro'' on which to study RNA replication.{{Citation needed|date=March 2009}}

Sub-genomic RNA is used in the synthesis of some proteins; they are generated by premature termination of (−)strand synthesis. sgRNAs and sgRNA negative-sense templates are found in infected cells.<ref name="ictvdb" />

==Taxonomy==
<big>'''Group: ssRNA(+)'''</big>
{{Collapsible list|title= <big>Order: Unassigned</big>
|1={{Collapsible list| framestyle=border:none; padding:1.0em;|title=Family: Tombusviridae
|1={{hidden begin|title=Genus: [[Alphanecrovirus]]}}
*[[Olive latent virus 1]]
*[[Olive mild mosaic virus]]
*'''''[[Tobacco necrosis virus A]]'''''
{{hidden end}}
|2={{hidden begin|title=Genus: [[Aureusvirus]]}}
*[[Cucumber leaf spot virus]]
*[[Johnsongrass chlorotic stripe mosaic virus]]
*[[Maize white line mosaic virus]]
*'''''[[Pothos latent virus]]'''''
{{hidden end}}
|3={{hidden begin|title=Genus: [[Avenavirus]]}}
*'''''[[Oat chlorotic stunt virus]]'''''
{{hidden end}}
|4={{hidden begin|title=Genus: [[Betanecrovirus]]}}
*[[Beet black scorch virus]]
*[[Leek white stripe virus]]
*'''''[[Tobacco necrosis virus D]]'''''
{{hidden end}}
|5={{hidden begin|title=Genus: [[Carmovirus]]}}
*[[Ahlum waterborne virus]]
*[[Angelonia flower break virus]]
*[[Bean mild mosaic virus]]
*[[Calibrachoa mottle virus]]
*[[Cardamine chlorotic fleck virus]]
*'''''[[Carnation mottle virus]]'''''
*[[Cowpea mottle virus]]
*[[Cucumber soil-borne virus]]
*[[Hibiscus chlorotic ringspot virus]]
*[[Honeysuckle ringspot virus]]
*[[Japanese iris necrotic ring virus]]
*[[Melon necrotic spot virus]]
*[[Nootka lupine vein clearing virus]]
*[[Pea stem necrosis virus]]
*[[Pelargonium flower break virus]]
*[[Saguaro cactus virus]]
*[[Soybean yellow mottle mosaic virus]]
*[[Turnip crinkle virus]]
*[[Weddel waterborne virus]]
{{hidden end}}
|6={{hidden begin|title=Genus: [[Dianthovirus]]}}
*'''''[[Carnation ringspot virus]]'''''
*[[Red clover necrotic mosaic virus]]
*[[Sweet clover necrotic mosaic virus]]
{{hidden end}}
|7={{hidden begin|title=Genus: [[Gallantivirus]]}}
*'''''[[Galinsoga mosaic virus]]'''''
{{hidden end}}
|8={{hidden begin|title=Genus: [[Macanavirus]]}}
*'''''[[Furcraea necrotic streak virus]]'''''
{{hidden end}}
|9={{hidden begin|title=Genus: [[Machlomovirus]]}}
*'''''[[Maize chlorotic mottle virus]]'''''
{{hidden end}}
|10={{hidden begin|title=Genus: [[Panicovirus]]}}
*[[Cocksfoot mild mosaic virus]]
*'''''[[Panicum mosaic virus]]'''''
*[[Thin paspalum asymptomatic virus]]
{{hidden end}}
|11={{hidden begin|title=Genus: [[Tombusvirus]]}}
*[[Artichoke mottled crinkle virus]]
*[[Carnation Italian ringspot virus]]
*[[Cucumber Bulgarian virus]]
*[[Cucumber necrosis virus]]
*[[Cymbidium ringspot virus]]
*[[Eggplant mottled crinkle virus]]
*[[Grapevine Algerian latent virus]]
*[[Havel River virus]]
*[[Lato River virus]]
*[[Limonium flower distortion virus]]
*[[Moroccan pepper virus]]
*[[Neckar River virus]]
*[[Pelargonium leaf curl virus]]
*[[Pelargonium necrotic spot virus]]
*[[Petunia asteroid mosaic virus]]
*[[Sitke waterborne virus]]
*'''''[[Tomato bushy stunt virus]]'''''
{{hidden end}}
|12={{hidden begin|title=Genus: [[Umbravirus]]}}
*[[Carrot mottle mimic virus]]
*'''''[[Carrot mottle virus]]'''''
*[[Groundnut rosette virus]]
*[[Lettuce speckles mottle virus]]
*[[Pea enation mosaic virus 2]]
*[[Tobacco bushy top virus]]
*[[Tobacco mottle virus]]
{{hidden end}}
|13={{hidden begin|title=Genus: Unassigned}}
*[[Chenopodium necrosis virus]]
*[[Elderberry latent virus]]
*[[Pelargonium chlorotic ring pattern virus]]
*[[Pelargonium line pattern virus]]
*[[Pelargonium ringspot virus]]
*[[Rosa rugosa leaf distortion virus]]
*[[Trailing lespedeza virus 1]]
{{hidden end}}
|14={{hidden begin|title=Genus: [[Zeavirus]]}}
*'''''[[Maize necrotic streak virus]]'''''
{{hidden end}}
}}
}}<ref name=ICTV />

[[Pelarspovirus]] is an additional genus that has been proposed.<ref name="Castaño2009">Castaño A, Ruiz L, Hernández C (2009) Insights into the translational regulation of biologically active open reading frames of Pelargonium line pattern virus. Virology 386(2):417–426</ref>

==References==
{{Reflist|2}}

==External links==
* [http://viralzone.expasy.org/all_by_species/53.html '''Viralzone''': Tombusviridae]
* [http://ictvonline.org/virusTaxonomy.asp '''ICTV''']
{{Baltimore classification}}

{{Taxonbar|from=Q3117439}}

[[Category:Tombusviridae]]
[[Category:Viral plant pathogens and diseases]]
[[Category:Virus families]]

Lymphocystivirus

2018-03-23T03:48:01Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q6708248}} (1 sig. taxon ID); WP:GenFixes, using AWB

{{Use dmy dates|date=April 2017}}
{{taxobox
| virus_group = i
| familia = ''[[Iridoviridae]]''
| genus = '''''Lymphocystivirus'''''
| subdivision_ranks = Type Species
| subdivision =
*''Lymphocystis disease virus 1''
}}

'''''Lymphocystivirus''''' is a genus of [[virus]]es, 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|language=en}}</ref> Fish serve as natural hosts. There is currently only one species in this genus: the type species ''Lymphocystis disease virus 1''.<ref name=ICTVReport/> Diseases associated with this genus include: tumor-like growths on the skin.<ref name=ICTVReport/><ref name=ViralZone>{{cite web|title=Viral Zone|url=http://viralzone.expasy.org/all_by_species/583.html|publisher=ExPASy|accessdate=15 June 2015}}</ref>

==Hosts==
'''Lymphocystivirus''' is one of five [[genus|genera]] of [[virus]]es within the viral family ''[[Iridoviridae]]'', and one of three genera within this family which infect [[teleostei|teleost]] [[fish]]es, along with ''[[Megalocytivirus]]'' and ''[[Ranavirus]]''.<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> Lymphocystiviruses infect more than 140 [[freshwater]] and [[seawater|marine]] species,<ref name=Essbauer>{{cite journal | last1 = Essbauer | first1 = S | last2 = Ahne | first2 = W | title = Viruses of Lower Vertebrates | journal = Journal of Veterinary Medicine B, Infectious Diseases and Veterinary Public Health| volume = 48 | issue = 6 | pages = 403–475 | year = 2001 | pmid = 11550762 | doi = 10.1046/j.1439-0450.2001.00473.x}}</ref> spanning at least 42 host families worldwide,<ref name=Sheng>{{cite journal | last1 = Sheng | first1 = X | last2 = Xing | first2 = J | last3 = Zhan | first3 = W | last4 = Wang | first4 = Y | title = Comparative studies on histopathological features of lymphocystis disease in four species of marine fish | journal = Journal of Fishery Sciences of China | issue = 5 | pages = | year = 2007 |url=http://en.cnki.com.cn/Article_en/CJFDTOTAL-ZSCK200705021.htm}}</ref> causing the chronic, self-limiting clinical disease, [[lymphocystis]].
While lymphocystis does not cause mass mortality events like megalocytiviruses and ranaviruses, fish with lymphocystis exhibit grossly visible papilloma-like skin lesions which substantially reduce their commercial value.<ref name=Cano>{{cite journal | last1 = Cano | first1 = I | last2 = Ferro | first2 = P | last3 = Alonso | first3 = MC | last4 = Bergmann | first4 = SM |last5 = Romer-Oberdorfer | first5 = A |last6 = Garcia-Rosado | first6 = E |last7 = Castro | first7 = D |last8 = Borrego | first8 = JJ |title = Development of molecular techniques for detection of lymphocystis disease virus in different marine fish species | journal = Journal of Applied Microbiology | volume = 102 | pages = 32–40 | year = 2006 | doi=10.1111/j.1365-2672.2006.03066.x}}</ref><ref name=Hossain>{{cite journal | last1 = Hossain | first1 = M | last2 = Kim | first2 = SR | last3 = Oh | first3 = MJ | title = The lymphocystis diseases in the Olive flounder, ''Paralichthys olivaceous'' | journal = University Journal of Zoology, Rajshahi University | volume = 26 | pages = 59–62 | year = 2007 | doi=10.3329/ujzru.v26i0.700}}</ref> No vaccines are currently available for lymphocystis viruses.<ref name=Cano/>

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

The genus ''Lymphocystivirus'' has only one species: ''Lymphocystis disease virus 1'' (LCDV-1), but [[Lymphocystis disease virus 2]] (LCD-2) is another virus that has been proposed as a second member.<ref name=Essbauer/> LCDV-1 infects [[European flounder]] ''[[Platichthys flesus]]'' and European plaice (''[[Pleuronectes platessa]]''). LCDV-2 infects the common dab (''[[Limanda limanda]]'').<ref name=Essbauer/> A third species [[Lymphocystis disease virus C]] has also been proposed based on the relative lack of sequence similarity of an isolate from the olive flounder (''[[Paralichthys olivaceous]]'') to LCDV-1 and LCDV-2.<ref name=Cano/>

==LCDV genome==
Lymphocystiviruses are Group I viruses with a dsDNA [[genome]]. The LCDV-1 genome is approximately 102.7 kilobase pairs (kbp) in length, with 195 potential [[open reading frames]] (ORF), and codes for two DNA-dependent [[RNA polymerase]] subunits, a [[DNA methyltransferase]], a [[DNA polymerase]], a guanosine triphosphate phosphohydrolase ([[GTPase]]), a [[helicase]], [[protein kinase]]s, a ribonucleoside diphosphate reductase, and [[zinc-finger protein]]s, among others.<ref name=Tidona>{{cite journal | last1 = Tidona | first1 = CA | last2 = Darai | first2 = G | title = The complete DNA sequence of LCDV | journal = Virology | volume = 230 | issue = 2 | pages = 207–216 | year = 1997 | pmid = 9143276 | doi = 10.1006/viro.1997.8456}}</ref> The LCDV-2 genome is similar to that of LCDV-1 but is slightly smaller, approximately 98 kilobase pairs (kbp) in length.<ref name=vanRegenmortel>{{cite book | last1 = vanRegenmortel | first1 = MHV | last2 = Fauquet | first2 = CM | last3 = Bishop | first3 = DHL | last4 = Carstens | first4 = EB | last5 = Estes | first5 = MK |last6 = Lemon | first6 = SM | last7 = Maniloff | first7 = J| last8 = Mayo | first8 = MA | last9 = McGeoch | first9 = DJ | last10 = Pringle | first10 = CR | last11 = Wickner| first11 = RB | title = ''Virus Taxonomy, Seventh Report of the International Committee on the Taxonomy of Viruses''| journal = Journal of fish diseases | publisher = Academic Press| location = New York, NY, USA|year = 2000 }}</ref>

==Structure==
Viruses in the genus ''Lymphocystivirus'' are enveloped, with icosahedral and polyhedral geometries, and T=189-217 symmetry. The diameter is around 120-350 nm. Genomes are linear, around 100kb in length.<ref name=ICTVReport/><ref name=ViralZone />

{| class="wikitable sortable" style="text-align:center"
|-
! Genus !! Structure || Symmetry !! Capsid !! Genomic arrangement !! Genomic segmentation
|-
|Lymphocystivirus||Polyhedral||T=189-217||||Linear||Monopartite
|}

==Life cycle==
Lymphocystiviruses attach to the host cell and enter by [[receptor-mediated endocytosis]] similar to other iridoviruses.<ref>Eaton HE, Ring BA, Brunetti CR (2010). "The genomic diversity and phylogenetic relationship in the family ''Iridoviridae''." ''Viruses.'' 2:1458-1475. http://www.mdpi.com/1999-4915/2/7/1458/pdf</ref> Viral particles are uncoated and move to the [[cell nucleus|nucleus]] of the cell, where [[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 moves to the cytoplasm for the second stage of DNA replication, which results in the formation of DNA [[concatemers]].<ref name=Goorha/> The concatameric viral DNA is subsequently packaged via a [[headful mechanism]] into virions.<ref name=Chinchar>Chinchar VG, Essbauer S, He JG, Hyatt A, Miyazaki T, Seligy V, Williams T (2005). "Family ''Iridoviridae'' 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 lymphocystis viral genome is [[circularly permuted]] with [[terminally redundant DNA]].<ref name=Goorha/> DNA-templated transcription is the method of transcription. Fish serve as the natural host.<ref name=ICTVReport/><ref name=ViralZone />

{| class="wikitable sortable" style="text-align:center"
|-
! Genus !! Host details !! Tissue tropism !! Entry details !! Release details !! Replication site !! Assembly site !! Transmission
|-
|Lymphocystivirus||Fish||None||Cell receptor endocytosis||Lysis; budding||Nucleus||Cytoplasm||Unknown
|}

==Pathogenesis==
Lymphocystis disease is a chronic disease that rarely causes mortality.<ref name=Essbauer/> Infection causes transformation and hypertrophy (approximately 1000x) of cells in the dermis, forming grossly visible lymphocystis nodules, as well as transformation and hypertrophy in cells of the connective tissues of various internal organs.<ref name=Essbauer/> [[Fibroblasts]] and [[osteoblasts]] are specifically targeted by the virus.<ref name=Essbauer2/> Lymphocystis viruses are not easily grown in [[cell culture]],<ref name=Essbauer/> placing limitations on ''[[in vitro]]'' molecular pathogenesis experiments.

==Diagnostic pathology==
As lymphocystis viruses are not easily grown in cell culture,<ref name=Essbauer/> [[diagnosis]] is based on [[clinical signs]], [[gross pathology]], [[histopathology]], [[serology]], and/or [[polymerase chain reaction]] (PCR)-based [[molecular biology|molecular]] [[assay]]s.

===Gross pathology===
The pathology of lymphocystis consists of papilloma-like skin lesions composed of greatly hypertrophied infected host cells embedded in extracellular matrix, sometimes called lymphocystis tumor cells, which are grossly evident as white spots on the skin and fins of infected fish.<ref name=Harikrishnan>{{cite journal | last1 = Harikrishnan | first1 = R | last2 = Kim | first2 = MC | last3 = Kim | first3 = JS | last4 = Balasundaram | first4 = C | last5 = Heo | first5 = MS |title = Immune enhancement of chemotherapeutants on lymphocystis disease virus (LDV) infected Paralichthys olivaceous | journal = Fish and Shellfish Immunology | volume = 29 | pages = 862–867 | year = 2010 | pmid = 20688171 | doi = 10.1016/j.fsi.2010.07.032}}</ref> These lesions proliferate as epithelial tumors in some cases.<ref name=Samalecos>{{cite journal | last1 = Samalecos | first1 = CP | title = Analysis of the structure of fish lymphocystis disease virions from skin tumours of pleuronectes | journal = Archives of Virology | volume = 91 | pages = 1–10 | year = 1986 | pmid = 3753198 | issue=1-2 | doi=10.1007/bf01316723}}</ref>

===Histopathology===
In a recent comparison of lymphocystis histopathology of four unrelated marine species, [[lesion]]s consistently associated with lymphocystis included [[hypertrophy|hypertrophied]] cells displaying irregular nuclei, [[basophilic]] [[cytoplasm]]ic [[inclusion bodies]] that stained positively via [[Feulgen and Mann's reaction]] and [[Periodic acid-Shiff]] (PAS)-positive [[hyaline capsules]].<ref name = Sheng/> Hyaline capsules arise from the extracellular matrix that is produced by the infected cells,<ref name=Essbauer2/> and are composed of sulphated and carboxylated glycoproteins (acid mucopolysaccharides).<ref name=Essbauer2/> In contrast, the inclusion body shape, distribution of viral particles within the cytoplasm and overall appearance of lymphocystis nodules varied by species.<ref name = Sheng/> The species examined in this study included the white-spotted puffer (''[[Arothron hispidus]]''), the Japanese sea bass (''[[Lateolabrax japonicus]]''), olive flounder (''[[Paralichthys olivaceus]]'') and the "sting fish" or Schlegel's black rockfish (''[[Sebastes schegeli]]'') <ref name = Sheng/>

===Serology===
Several serologic assays have been developed to identify LCDV infections, including [[flow cytometry]],<ref name=Garcia-Rosado>{{cite journal | last1 = Garcia-Rosado | first1 = E | last2 = Castro | first2 = D | last3 = Cano | first3 = I | last4 = Perez-Prieto | first4 = SI |last5 = Borrego | first5 = JJ | last6 = Borrego | first6 = JJ |title = Serological techniques for detection of lymphocystis virus in fish | journal = Aquatic Living Resources | volume = 15 | pages = 179–185 | year = 2002 | doi=10.1016/s0990-7440(02)01174-9}}</ref> [[immunoblot]],<ref name=Garcia-Rosado/><ref name=Cano2>{{cite journal | last1 = Cano | first1 = I | last2 = Alonso | first2 = MC | last3 = Garcia-Rosado | first3 = E | last4 = RodriguezSaint-Jean | first4 = S |last5 = Castro | first5 = D | last6 = Borrego | first6 = JJ |title = Detection of lymphocystis disease virus (LCDV) in asymptomatic cultured gilt-head seabream (''Spartus aurata'', L) using an immunoblot technique | journal = Veterinary Microbiology| volume = 113 | pages = 137–141 | year = 2006 | pmid = 16298500 | doi = 10.1016/j.vetmic.2005.10.024}}</ref> and [[immunofluorescence]].<ref name=Garcia-Rosado/> However, PCR-based molecular assays are more practical for most applications.<ref name=Cano/>

===Electron microscopy===
[[Transmission electron microscopy]] (TEM) of infected cells reveals cytoplasmic virus particles typically measuring from 198-227 nm in diameter<ref name=Essbauer/> (in some cases as large as 380 nm)<ref name=Essbauer/><ref name=Essbauer2>{{cite journal | last1 = Essbauer | first1 = S | last2 = Fischer | first2 = U | last3 = Bergmann | first3 = S | last4 = Ahne | first4 = W | title = Investigations on the ORF 167L of Lymphocystis Disease Virus (''Iridoviridae'')| journal = Virus Genes | volume = 28 | issue = 1 | pages = 19–39| year = 2004 | pmid = 14739649 | doi = 10.1023/B:VIRU.0000012261.96217.fe}}</ref> and electron-dense substances in the [[perinuclear space]].<ref name = Sheng/>

===Molecular pathology===
Published PCR primers and protocol are available to amplify a portion of the LCDV-1 MCP.<ref name=Cano/> When the PCR diagnostic assay is combined with [[slot blot]], diagnostic [[sensitivity and specificity|sensitivity]] is increased, facilitating the diagnosis of asymptomatic LCDV-1 infections.<ref name=Cano/>

== References ==
{{Reflist}}

== External links ==
* [https://talk.ictvonline.org/ictv-reports/ictv_online_report/dsdna-viruses/w/iridoviridae '''ICTV Online (10th) Report''': Iridoviridae]

{{Taxonbar|from=Q6708248}}

[[Category:Articles created via the Article Wizard]]
[[Category:Iridoviridae]]
[[Category:Fish viral diseases]]

Kalifornische Steineiche

2018-03-23T03:40:00Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q514052}} (9 sig. taxon IDs); WP:GenFixes, using AWB

{{Italic title}}
{{taxobox
| name = Coast live oak
| image = Quercus agrifolia foliage.jpg
| image_caption = Coast live oak foliage with new spring growth
| status = LC
| status_system = IUCN3.1
| status_ref = <ref>{{cite web
| url = http://www.iucnredlist.org/details/194049/0
| title = Quercus agrifolia
| date = 2016
| website = iucnredlist.org
| publisher = iucnredlist
| access-date = 21 November 2017
| quote = data}}</ref>
| regnum = [[Plant]]ae
| unranked_divisio = [[Angiosperms]]
| unranked_classis = [[Eudicots]]
| unranked_ordo = [[Rosids]]
| ordo = [[Fagales]]
| familia = [[Fagaceae]]
| genus = ''[[oak|Quercus]]''
| sectio = ''[[Erythrobalanus|Lobatae]]''
| species = '''''Q. agrifolia'''''
| binomial = ''Quercus agrifolia''
| binomial_authority = [[Luis Née|Née]] 1801
| range_map = Quercus agrifolia range map 1.png
| range_map_caption = Natural range
| synonyms_ref = <ref>[http://www.theplantlist.org/tpl1.1/record/kew-171483 The Plant List, ''Quercus agrifolia'' Née ]</ref>
| synonyms = {{collapsible list|bullets = true
| ''Quercus acroglandis'' Kellogg
| ''Quercus acutiglandis'' Sarg.
| ''Quercus agrifolia'' var. ''frutescens'' Engelm.
| ''Quercus agrifolia'' var. ''oxyadenia'' (Torr.) J.T.Howell
| ''Quercus oxyadenia'' Torr.
| ''Quercus pricei'' Sudw.
}}
}}

'''''Quercus agrifolia''''', the '''California live oak'''<ref>{{PLANTS|id=QUAG|taxon=Quercus agrifolia|accessdate=30 July 2015}}</ref> or '''coast live oak''', is a highly variable, often [[shrubby]] [[evergreen]] [[oak]] tree, a type of [[live oak]], native to the [[California Floristic Province]]. It grows [[Western California|west]] of the [[Sierra Nevada (U.S.)|Sierra Nevada]] mountain range from [[Mendocino County, California|Mendocino County]], California, south to northern [[Baja California]] in Mexico.<ref>[http://www.calflora.org/cgi-bin/species_query.cgi?where-calrecnum=6983 Calflora taxon report, University of California, ''Quercus agrifolia'' Nee, California live oak, coast live oak ]</ref> It is classified in the [[Erythrobalanus|red oak]] [[Section (botany)|section]] of live oaks (''Quercus'' sect. ''Lobatae'').<ref name=tammy/>

This species is commonly [[sympatry|sympatric]] with [[Quercus chrysolepis|canyon live oak]] (''Q. chrysolepis''), and the two may be hard to distinguish because their [[spinose leaves]] are superficially similar.

==Description==
[[File:cloe06.jpg|thumb|left|Coast live oak, Sonoma County]]
Coast live oak typically has a much-branched trunk and reaches a mature height of {{convert|10|–|25|m|ft|sp=us}}. Some specimens may attain an age exceeding 250 years, with trunk diameters up to {{convert|3|or|4|m|ft|sp=us|0|spell=in}}, such as those on the [[Filoli]] estate in [[San Mateo County, California|San Mateo County]].<ref name=tammy/><ref>[https://www.biodiversitylibrary.org/page/8009573#page/272/mode/1up Née, Luis 1801 Anales de Ciencias Naturales 3(9): 271–272] [[Diagnosis (taxonomy)|diagnosis]] in Latin, commentary in Spanish</ref>

The trunk, particularly for older individuals, may be highly contorted, massive and gnarled. The crown is broadly rounded and dense, especially when aged 20 to 70 years; in later life the trunk and branches are more well defined and the leaf density lower.<ref name=tammy/>

===Leaves===
The [[leaf|leaves]] are dark green, oval, often [[wikt:convex|convex]] in shape, {{convert|2|–|7|cm|in|abbr=on}} long and {{convert|1|–|4|cm|in|abbr=on}} broad; the leaf margin is spiny-toothed (spinose), with sharp thistly fibers that extend from the lateral leaf veins. The outer layers of leaves are designed for maximum [[sun|solar]] absorption, containing two to three layers of [[photosynthetic]] cells.<ref name=tammy/>

These outer leaves are deemed to be small in size to more efficiently re-radiate the heat gained from solar capture. Shaded leaves are generally broader and thinner, having only a single layer of photosynthetic cells. The convex leaf shape may be useful for interior leaves which depend on capturing reflected light scattered in random directions from the outer canopy.<ref name=tammy>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233501005 Flora of North America: ''Quercus agrifolia'']</ref>

===Inflorescence and acorns===
[[File:cloacorne.jpg|thumb|left|Acorns and leaves]]
The [[flower]]s are produced in early-to-mid spring; the male flowers are pendulous [[catkin]]s {{convert|5|–|10|cm|in|abbr=on}} long, the female flowers inconspicuous, less than {{convert|0.5|cm|in|abbr=on}} long, with 1-3 clustered together. The [[fruit]] is a slender reddish brown [[acorn]] {{convert|2|-|3.5|cm|in|abbr=on}} long and {{convert|1|-|1.5|cm|in|abbr=on}} broad, with the basal quarter enclosed in a [[Calybium and cupule|cupule]]; unusually for a red oak, the acorns mature about 7–8 months after pollination (most red oak acorns take 18 months to mature).<ref name=tammy/>

===Recognized varieties===
There are two [[variety (biology)|varieties]] of ''Quercus agrifolia'':
*''Quercus agrifolia'' var. ''agrifolia''. Throughout the range of the species. Leaves that are glabrous to slightly hairy on the abaxial side, especially near the leaf vein axils. Hybrids with ''[[Quercus kelloggii|Q. kelloggii]]'', ''Q. parvula'' var. ''shevei'', and ''[[Quercus wislizeni|Q. wislizenii]]'' are known.
*''Quercus agrifolia'' var. ''oxyadenia''. Southwesternmost California (San Diego area), Baja California. Leaves that are [[tomentose]] abaxially, with densely interwoven hairs. It prefers granitic soils; hybrids with ''Q. kelloggii'' known.

===Hybridity===
Several hybrids between coast live oak and other red oak species have been documented. Hybrids with [[interior live oak]] (''Q. wislizenii'') are known in many areas in northern California. Coast live oak also hybridizes with [[Quercus wislizeni|Shreve oak]] (''Q. parvula'' var. ''shrevei''). All these oak species show evidence of [[introgression]] with one another.

==Etymology==
The name ''Quercus agrifolia'' literally means "field-leaved oak," from the Latin "quercus," meaning "oak," "agri" meaning "field," and folia, meaning "leaved." This species is sometimes known by the name "California live oak".

==Habitat and ecology==
[[File:Quercus Agrifolia.jpg|right|thumb|250px|Coast live oak off [[U.S. Route 101 in California|California 101]], central coast.]]
Coast live oak is the only California native oak that actually thrives in the coastal environment, although it is rare on the immediate shore; it enjoys the mild winter and summer climate afforded by ocean proximity, and it is somewhat tolerant of aerosol-borne sea salt. The coastal fog supplies relief from the rainless California summer heat.

It is the dominant overstory plant of the coast live oak [[woodland]] habitat, often joined by [[Umbellularia|California bay laurel]] and [[Aesculus californica|California buckeye]] north of [[Big Sur]]. Associated understory plants include [[toyon]], various [[manzanita]]s and [[Toxicodendron diversilobum|western poison-oak]].

Normally the tree is found on well drained soils of coastal hills and plains, often near year round or [[perennial stream]]s. It may be found in several natural communities including coast live oak woodland, [[Engelmann oak]] woodland, [[valley oak]] woodland and both northern and southern mixed evergreen forests. While normally found within {{convert|100|km|mi|abbr=off|sp=us}} of the [[Pacific Ocean]] at elevations less than {{convert|700|m|ft|abbr=off|sp=us}}, in southern California it occasionally occurs at up to {{convert|1,500|m|ft|abbr=off|sp=us}} in altitude.

The California oak moth (''[[Phryganidia californica]]'') caterpillar subsists entirely on living and fallen leaves of the Coast Live Oak. In 8-10 year cycles, the caterpillar will appear in sufficient abundance to denude healthy trees. The trees recover, and botanists speculate that the species provide mutual benefit, possibly in the form of fertilizer for the oak.<ref>http://trees.stanford.edu/ENCYC/QUERCUS.htm</ref> The coast live oak is also the only known foodplant of ''[[Chionodes vanduzeei]]'' [[caterpillar]]s.

==Economic usage==

===Historical usage===
[[File:Los Encinos live oak.jpg|right|thumb|250px|Coast live oak at [[Rancho Los Encinos]] in the San Fernando Valley]]
At least twelve distinct cultures of [[Native Americans in the United States|Native Americans]] are known to have consumed the acorns as a dietary staple.{{citation needed|date=May 2012}} In the 18th century [[Spanish colonization of the Americas|Spaniards]] in the [[San Fernando Valley]] used the wood for charcoal to fire [[kiln]]s in making [[adobe]]. Later this form of charcoal would be utilized in the baking, [[gunpowder]] and [[electric power]] industries.

In the 18th and 19th centuries [[shipbuilder]]s sought out the odd angular branches to make special joints. [[Settler|Pioneer]]s moving west would harvest small amounts for making farm implements and wagon wheels, but the greatest impact was the wholesale clearing of oak woodlands to erect sprawling cities such as [[San Diego]] and [[San Francisco]]. The irregular shape often let the tree escape widespread harvest for building timbers, and also led the early settlers to endow the coast live oak with mystical qualities. Its stateliness has made it a subject of historical [[Landscape art|landscape painter]]s throughout California modern history since the mid-19th century.

===Modern usage===
Coast live oak has also become a common addition to western USA [[landscaping]]. It is however sensitive to changes in grading and drainage; in particular, it is important to respect the root crown level and avoid adding soil near the trunk when construction or [[landscaping]] occurs.

Also, if incorporating it into a landscaping scheme with artificial irrigation, it is important to avoid regular watering within the oak's [[Tree canopy|drip line]] (canopy), since wet soil in the summer increases infection rates by soil-borne ''[[Phytophthora]]'' diseases like [[sudden oak death]].<ref name="Sudden Oak Death: Management Strategies">{{cite web|url=http://www.plantmanagementnetwork.org/php/shared/sod/|title=Sudden Oak Death and Associated Diseases Caused by Phytophthora ramorum|last=J. M. Davidson|date=7 July 2003|publisher=Plant Management Network|accessdate=12 January 2010}}</ref>

==Geographical monikers==
The coast live oak, especially in its [[Spanish Language|Spanish]] forms ''encino'' or ''encina'', ''encinitas'' "little oaks", and ''encinal'' "oak grove", gave its name to seven land grants across California and to many communities and geographic features.

These include [[Rancho Los Encinos]], the community of [[Encino, Los Angeles, California|Encino]] near Los Angeles, [[Encinitas, California|Encinitas]] near [[San Diego, California|San Diego]], and ''Encinal del Temescal'', now the city of [[Oakland, California|Oakland]].<ref>Gudde, Erwin, and William Bright, ''California Place Names'', University of California Press, 4th edition, 1998, {{ISBN|0-520-21316-5}}, p. 123-124</ref>

''Paso Robles'' ('Pass of the Oaks') also refers to oaks as a geographical place name.

==References==
{{Reflist}}

==Further reading==
{{Portal|Trees}}
{{Commons category|Quercus agrifolia}}

*Jepson Flora Project: [http://ucjeps.berkeley.edu/cgi-bin/get_JM_treatment.pl?Quercus+agrifolia ''Quercus agrifolia''], [http://ucjeps.berkeley.edu/cgi-bin/get_JM_treatment.pl?4316,4326,4328,4329 var. ''agrifolia''], [http://ucjeps.berkeley.edu/cgi-bin/get_JM_treatment.pl?4316,4326,4328,4330 var. ''oxyadenia'']
*Balls, E. K. (1972). ''Early Uses of California Plants''. University of California Press, Berkeley.
*Pavlik, B. M., Muick, P., Johnson, S., & Popper, M. (1991). ''Oaks of California''. Cachuma Press {{ISBN|0-9628505-1-9}}.
*Sawyer, John O., & Keeler-Wolf, Todd. (1995) ''A manual of California Vegetation.'' California Native Plant Society, page 241.
*[http://wildflower.utexas.edu/plants/result.php?id_plant=QUAG Native Plant Information Network]{{dead link|date=July 2016 |bot=InternetArchiveBot |fix-attempted=yes }} More information on ''Quercus agrifolia''
*[http://www.laspilitas.com/groups/oaks/california_oak1.html Las Pilitas: California Oaks]
*[https://web.archive.org/web/20070706233833/http://www.cnr.vt.edu/dendro/dendrology/Syllabus2/factsheet.cfm?ID=548 Virginia Tech Forestry Department: Coast Live Oak]
*[http://plants.usda.gov/java/profile?symbol=QUAG USDA Plants Database: ''Quercus agrifolia'' Née]
*[http://www.plantmaps.com/nrm/quercus-agrifolia-california-live-oak-native-range-map.php Interactive Distribution Map for ''Quercus agrifolia'']

{{Taxonbar|from=Q514052}}

{{DEFAULTSORT:Quercus Agrifolia}}
[[Category:Quercus|agrifolia]]
[[Category:Flora of California]]
[[Category:Trees of Baja California]]
[[Category:Natural history of the California chaparral and woodlands]]
[[Category:Natural history of the California Coast Ranges]]
[[Category:Natural history of the Central Valley (California)]]
[[Category:Plants described in 1801]]
[[Category:Natural history of the Channel Islands of California]]
[[Category:Natural history of the San Francisco Bay Area]]
[[Category:Natural history of the Santa Monica Mountains]]
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Coccolithovirus

2018-03-23T03:10:42Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q962359}} (1 sig. taxon ID); WP:GenFixes, using AWB

{{Use dmy dates|date=April 2017}}
{{taxobox
| familia = ''[[Phycodnaviridae]]''
| genus = '''''Coccolithovirus'''''
| virus_group = i
| subdivision_ranks = Type Species/ Strains
| subdivision =
*''Emiliania huxleyi virus 86''
*EhV-84 (1999 EC)
*EhV-88 (1999 EC)
*EhV-201 (2001 EC)
*EhV-202 (2001 EC)
*EhV-203 (2001 EC)
*EhV-207 (2001 EC)
*EhV-208 (2001 EC)
*EhV-18 (2008 EC)
*EhV-156 (2009 EC)
*EhV-164 (2008 Scotland)
*EhV-145 (2008 Scotland)
*EhV-99B1 (1999 Norway)
*EhV-163 (2000 Norway)
}}

'''''Coccolithovirus''''' is a genus of giant double-stranded [[DNA virus]], in the family [[Phycodnaviridae]]. Algae, specifically ''[[Emiliania huxleyi]]'', a species of [[coccolithophore]]{{Ref|virology}}, serve as natural hosts. There is currently only one species in this genus: ''Emiliania huxleyi virus 86''.<ref name=ViralZone>{{cite web|title=Viral Zone|url=http://viralzone.expasy.org/all_by_species/589.html|publisher=ExPASy|accessdate=15 June 2015}}</ref><ref name=ICTV>{{cite web|last1=ICTV|title=Virus Taxonomy: 2014 Release|url=http://ictvonline.org/virusTaxonomy.asp|accessdate=15 June 2015}}</ref>

==Taxonomy==
Group: dsDNA
*Order: Unassigned
**Genus: Coccolithovirus
***''Emiliania huxleyi virus 86''

==Structure==
Coccolithoviruses are enveloped, icosahedral and have a diameter ranging from 100–220 nm. Their genomes are linear, between 410–415kb in length and predict to encode for approximately 472 proteins.<ref name=ViralZone />

{| class="wikitable sortable" style="text-align:center"
|-
! Genus !! Structure || Symmetry !! Capsid !! Genomic arrangement !! Genomic segmentation
|-
|Coccolithovirus||Icosahedral||T=169||Enveloped||Linear||Monopartite
|}

==Life cycle==
Coccolithoviruses are part of the family of ''Phycodnaviridae'', one of the five families that belong to a large and phylogenetically diverse group of viruses known as nucleocytoplasmic large dsDNA viruses ([[Nucleocytoplasmic large DNA viruses|NCLDVs]]). These viruses either replicate exclusively in the cytoplasm of the host cell or start their life cycle in the host nucleus but complete it in the cytoplasm. In the case of EhV-86 the infection strategy is not fully understood but Mackinder et al. (2009)<ref>Mackinder LCM, Worthy CA, Biggi G, Hall M, Ryan KP, Varsani A, Harper GM, Wilson WH, Brownlee C, Schroeder DC. (2009) A unicellular algal virus, Emilienia huxleyi virus 86, exploits an animal-like infection strategy. Journal of general Virology 90:2306–2316.</ref> have proposed the following model: The virus enters the host cell via endocytosis, followed by fusion of its lipid membrane with the host vacuole membrane and the release of its nucleoprotein core into the cytoplasm. Alternatively the virus membrane could fuse directly with the host plasma membrane. The virus genome is then released from the capsid into the nucleus, where it is replicated by the viral DNA polymerase. The replicated genome is packed into assembled capsids in the cytoplasm and the newly formed (up to 400–1000) virions are thought to be transported to the plasma membrane and released by a controlled budding mechanism, which leads to the cellular breakdown of the host cell.

During G2 and M stage of the life cycle, the coccosphere is incomplete and the exposure of the plasma membrane to the virus is increased. Even with an intact coccosphere infection can occur due to naturally occurring gaps between the coccoliths.<ref>Paasche E. (2001) A review of the coccolithophorid Emiliania huxleyi (Prymnesiophyceae), with particular reference to growth, coccolith formation, and calcification-photosynthesis interactions. Phycologia 40(6):503–52.</ref>

''E. huxleyi'' is known for forming seasonal algal blooms, that can reach 250,000 km2, during which cell density in the upper 200 m increases from 103 to 105 cells per mL seawater.<ref>Schroeder DC, Oke J, Hall M, Malin G, Wilson WH. (2003) Viral succession observed during an Emilinaia huxleyi bloom. Applied and Environmental Microbiology 69:2484–2490.</ref> These algae blooms collapse usually after 5–8 days and several studies have shown that bloom termination is intrinsically linked to infection by coccolithoviruses.<ref>Wilson WH, Tarran GA, Schroeder D, Cox M, Oke J, Malin G. (2002) Isolation of viruses responsible for the demise of an Emiliania huxleyi bloom in the English Channel. Journal of the Marine Biological Association of the United Kingdom 82:369–377.</ref> Transmission of viruses between algal hosts occurs via passive diffusion. Furthermore, EhV DNA was also detected in copepods, leading to the proposal that viruses are further dispersed by virus-carrying zooplankton.<ref>Frada MJ, Schatz D, Farstey V, Ossolinski JE, Sabanay H, Ben-Dor S, Koren I, Vardi A. (2014) Zooplankton May Serve as Transmission Vectors for Viruses Infecting Algal Blooms in the Ocean. Current Biology 24:2592–2597.</ref>

==Genome==
To date 14 EhV strains have been isolated between 1999 and 2008 primarily from the English Channel (EC) but also from the Norwegian and Scotland's Coast.<ref>Nissimov JI, Napierb JA, Kimmance SA, Allen MJ. (2014) Permanent draft genomes of four new coccolithoviruses: EhV-18, EhV-145, EhV-156 and EhV-164. Marine Genomics 15:7–8.</ref><ref>Nissimov JI, Worthy CA, Rooks P, Napier JA, Kimmance SA, Henn MR, Ogata H, Allen MJ. (2011) Draft genome sequence of the Coccolithovirus EhV-84. Standards in Genomic Science 5:1–11.</ref><ref>Nissimov JI, Worthy CA, Rooks P, Napier JA, Kimmance SA, Henn MR, Ogata H, Allen MJ. (2012) Draft Genome Sequence of Four Coccolithopviruses: Emiliania huxleyi Virus EhV-88, EhV-201, EhV-207 and EhV-208. Journal of Virology 86(5):2896–2897.</ref><ref>Pagarete et al. 2012, Allen et al. 2006d, Nissimov JI, Worthy CA, Rooks P, Napier JA, Kimmance SA, Henn MR, Ogata H, Allen MJ. (2012) Draft Genome Sequence of the Coccolithopvirus Emiliania huxleyi Virus 202. Journal of Virology 86(4):380–2381.</ref><ref>Nissimov JI, Worthy CA, Rooks P, Napier JA, Kimmance SA, Henn MR, Ogata H, Allen MJ. (2011) Draft Genomic Sequence of the Coccolithovirus Emiliania huxleyi Virus 203. Journal of Virology 85(24):13468–13469.</ref> Although partial sequences of all these 14 strains are available due to the highly repetitive nature of the genome EhV-86 is the only strain that has been fully sequenced.<ref>Wilson WH, Schroeder DC, Allen MJ, Holden MTG, Parkhill J, Barrell BG, Churcher C, Hamlin N, Mungall K, Norbertczak H, Quail MA, Price C, Rabbinowitsch E, Walker D, Craigon M, Roy D, Ghazal P. (2005) Complete Genome Sequence and Lytic Phase Transcription Profile of a Coccolithovirus. Science 309:1090–1092</ref> The sequencing of EhV-86 revealed a circular [[genome]] of 407,339 bp length with 472 predicted coding sequences (CDs). Remarkably 80% of these putative genes have no database homologs to date. Those that could be assigned a function due to sequence similarity or protein domain matches include DNA and RNA polymerase subunits, eight proteases as well as at least four genes that encode proteins involved in sphingolipid biosynthesis. These were shown to have been acquired from the host via horizontal gene transfer.<ref>Monier A, Pagarete A, De Vargas C, Allen MJ, Read B, Claverie J, Ogata H, De Vargas C. (2009) Horizontal gene transfer of an entire metabolic pathway between a eukaryotic alga and its DNA virus. Genome Research 19:1441–1449.</ref>

Furthermore, the EhV-86 genome revealed three distinct families (A, B, C) of repetitive regions within the genome.<ref>Allen MJ, Schroeder DC, Wilson WH. (2006) Preliminary characterisation of repeat families in the genome of EhV-86, a giant algal virus that infects the marine microalga Emiliania huxleyi. Archives of Virology 151:525–535.</ref> Family C consists of AT-rich repeats that are non-coding and that are probably part of the origin of replication (ORF). Family B are GC-rich repeats that are found in protein products of eight predicted CDSs. Family A homologous regions vary in size between 30–300 bp and are found in a 104 kbp (200–304 kbp) section of the genome, that contains no gene homologs of known function in the current data bases. Family A repeat units are non-coding and characterised by a nanomer (GTTCCC(T/C)AA) that in total, appears at 106 locations within this region. This sequence is found directly upstream of 86 CDSs and is likely to play a role in controlling the expression of associated CDSs.

During infection a distinct expression pattern of viral genes was described that could be divided into three phases according to CDS expression.<ref>Allen MJ, Forster T, Schroeder DC, Hall M, Roy D, Ghazal P, Wilson WH. (2006) Locus specific gene expression pattern suggests a unique propagation strategy for a giant algal virus. Journal of virology 80: 7699–7705.</ref> One hour post infection, 39 viral genes were transcribed, followed by a further 194 genes after 2 h and 71 genes after 4 h. All 39 genes that are expressed 1 h post infection are located in the 104 kbp region and have the nanomer directly upstream of the start codon. Since the expression of viral RNA polymerase was not detected 1 h after infection, it has yet to be established whether the promoter is recognised by a packaged viral RNA polymerase or by the host RNA polymerase. Proteomic analysis of the EhV-86 virion did, however, fail to detect any major RNA polymerase subunits.<ref>Allen MJ, Howard JA, Lilley KS, Wilson WH. (2008) Proteomic analysis of the EhV-86 virion. Proteome Science 6(11).</ref>

==History==
Wilson and his team at the [[Marine Biological Association]] (MBA), [[University of East Anglia]] and [[Plymouth Marine Laboratory]] (PML), first observed the virus in 1999. Later in the summer of 2005 researchers at the [[Plymouth Marine Laboratory]] (Willie Wilson et al.) and at the [[Sanger Institute]] (Holden et al.) sequenced the genome for the EhV-86 strain finding it to have 472 protein-coding genes making it a "giant-virus", and the largest known marine virus by genome{{Ref|giant}}.

From initial investigation of the ''Coccolithoviruses'' genome, a sequence of genes responsible for production of [[ceramide]] was discovered{{Ref|press}}. Ceramide is a controlling factor in cell death, and it is currently thought that ''Coccolithovirus'' uses this to prolong the life of ''[[Emiliania huxleyi]]'' while it uses the host cell to replicate. This is a unique ability unseen in any other viral genome to date.

==See also==
* ''[[Mimivirus]]'' – largest giant virus on record by genome
* ''[[Mycoplasma genitalium]]'', ''[[Pelagibacter ubique]]'' – some of the smallest known [[bacterium|bacteria]]
* ''[[Nanoarchaeum]]'' – smallest known [[archaeum]]
* [[Smallest organisms]]
* [[Parvovirus]] – smallest known family of [[virus]]es
* ''[[Phycodnaviridae]]'' – algae infecting viruses

==Notes==
{{refbegin}}

# {{note|virology}} [http://www.virologyj.com/content/2/1/52 Giant viruses in the oceans: the 4th Algal Virus Workshop ''Virology Journal'' 2005]
# {{note|sanger}} [http://www.sanger.ac.uk/Projects/EhV/ Sanger institute home for ''Emiliania huxleyi'' virus 86]
# {{note|giant}} [http://www.giantvirus.org/top.html Giantviruses.org top viruses by genome size.]
# {{note|press}} [https://web.archive.org/web/20051101000538/http://www.pml.ac.uk/pml/news/050812.htm Plymouth Marine Laboratory press release.]

{{refend}}

==References==
{{Reflist}}

==Further reading==
* Wilson, W. H., Schroeder, D. C., Allen, M. J., Holden, M. T. G., Parkhill, J., Barrell, B. G., Churcher, C., Hamlin, N., Mungall, K., Norbertczak, H., Quail, M. A., Price, C., Rabbinowitsch, E., Walker, D., Craigon, M., Roy, D. and Ghazal, P. (2005) [http://www.sciencemag.org/cgi/content/abstract/309/5737/1090 Complete Genome Sequence and Lytic Phase Transcription Profile of a Coccolithovirus]. ''Science'' '''5737''', 1090–1092
* Allen, M. J., Schroeder, D. C., Holden, M. T. and Wilson, W. H. (2006) [http://mbe.oxfordjournals.org/cgi/content/abstract/23/1/86 Evolutionary History of the Coccolithoviridae]. ''Mol Biol Evol.'' '''23''', 86–92 ([[Athens access and identity management|Athens]] login required)

==External links==
* [http://viralzone.expasy.org/all_by_species/589.html '''Viralzone''': Coccolithovirus]
* [http://ictvonline.org/virusTaxonomy.asp '''ICTV''']
{{Baltimore classification}}

{{Taxonbar|from=Q962359}}

[[Category:Nucleocytoplasmic large DNA viruses]]
[[Category:Phycodnaviridae]]

Enterobakteriophage MS2

2018-03-23T03:09:45Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q4840020}} (2 sig. taxon IDs); WP:GenFixes using AWB

{{Taxobox
| image = Ms2capsid surface.png
| image_width = 200px
| image_alt = MS2 capsid
| image_caption = Bacteriophage MS2 capsid
| color = violet
| virus_group = iv
| familia = '''[[Leviviridae]]'''
| genus = [[Levivirus]]
| species = '''Bacteriophage MS2'''}}

The '''bacteriophage MS2''' is an icosahedral, positive-sense single-stranded [[RNA]] virus that infects the bacterium ''[[Escherichia coli]]'' and other members of the ''[[Enterobacteriaceae]]''.<ref name="van Duin">{{cite book |last=van Duin |first=J. |last2=Tsareva |first2=N. |chapter=Single-stranded RNA phages. Chapter 15 |pages=175–196 |editor-last=Calendar |editor-first=R. L. |title=The Bacteriophages |edition=Second |publisher=Oxford University Press |year=2006 |isbn=0195148509 }}</ref> MS2 is a member of a family of closely related bacterial viruses that includes [[bacteriophage f2]], [[bacteriophage Qβ]], R17, and GA.<ref>Ni, C. Z. et al. [https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2143325/ Crystal structure of the coat protein from the GA bacteriophage: model of the unassembled dimer]. Protein Sci. 5, 2485–2493 (1996)</ref>

==History==

In 1961, MS2 was isolated by Alvin John Clark and recognized as an RNA-containing phage very similar to [[bacteriophage f2]].<ref>{{cite journal |last=Davis |first=J. E. |last2=Strauss |first2=J. H. |last3=Sinsheimer |first3=R. L. |title=Bacteriophage MS2: another RNA phage |year=1961 |journal=[[Science (journal)|Science]] |volume=134 |issue=3488 |page=1427 |doi=10.1126/science.134.3488.1425 }}</ref>

In 1976, the MS2 genome was the first genome to be completely sequenced.<ref name="Fiers">{{cite journal |last=Fiers |first=W. |last2=Contreras |first2=R. |last3=Duerinck |first3=F |last4=Haegeman |first4=G. |last5=Iserentant |first5=D. |last6=Merregaert |first6=J. |last7=Min Jou |first7=W. |last8=Molemans |first8=F. |last9=Raeymaekers |first9=A. |last10=Van den Berghe |first10=A. |last11=Volckaert |first11=G. |last12=Ysebaert |first12=M. |title=Complete nucleotide sequence of bacteriophage MS2 RNA: primary and secondary structure of the replicase gene |journal=[[Nature (journal)|Nature]] |year=1976 |volume=260 |issue=5551 |pages=500–507 |doi=10.1038/260500a0 |pmid=1264203}}</ref> This was accomplished by [[Walter Fiers]] and his team, building upon their earlier milestone in 1972 of the first gene to be completely sequenced, the MS2 coat protein.<ref>{{cite journal |last=Min Jou |first=W. |last2=Haegeman |first2=G. |last3=Ysebaert |first3=M. |last4=Fiers |first4=W. |title=Nucleotide sequence of the gene coding for the bacteriophage MS2 coat protein |journal=Nature |year=1972 |volume=237 |issue=5350 |pages=82–88 |doi=10.1038/237082a0 |pmid=4555447}}</ref> These sequences were determined at the RNA level, whereas the next landmark achievement, the sequence of the bacteriophage [[Phi X 174|ΦX174]] genome in 1977, was determined using DNA.<ref>{{cite journal |last=Sanger |first=F. F. |year=1977 |title=Nucleotide sequence of bacteriophage φX174 DNA |journal=Nature |volume=265 |issue=5596 |pages=687–695 |doi=10.1038/265687a0 |pmid=870828|display-authors=etal}}</ref>
The first effort at a statistical analysis of the MS2 genome was a search for patterns in the nucleotide sequence. Several non-coding sequences were identified, however at the time of this investigation (1979), the functions of the non-coding patterns were unknown.<ref>A search for patterns in the nucleotide sequence of the MS2 genome, Erickson, JW and Altman, GG; J. Mathematical Biology, April 1979, Volume 7, pp219–230</ref>

==Virology==

===Genome and gene products===
The [https://www.ncbi.nlm.nih.gov/nuccore/176120924?report=genbank MS2 genome] is one of the smallest known, consisting of 3569 nucleotides of single-stranded RNA.<ref name="Fiers" /> It encodes just four proteins: the maturation protein (A-protein), the [[lysis]] protein, the coat protein, and the [[replicase]] protein.<ref name="van Duin" /> The gene encoding lysis protein (''lys'') overlaps both the 3'-end of the upstream gene (''cp'') and the 5'-end of the downstream gene (''rep''), and was one of the first known examples of [[overlapping gene]]s. The positive-stranded RNA genome serves as [[messenger RNA]], and is translated upon viral uncoating within the host cell. Although the four proteins are encoded by the same messenger/viral RNA, they are not all [[Gene expression|expressed]] at the same levels; expression of these proteins is regulated by a complex interplay between [[Translation (biology)|translation]] and [[Nucleic acid secondary structure|RNA secondary structure]].

[[File:MS2_phage_gene_map.svg|frame|500|left|alt=caption|Location of protein-coding genes within bacteriophage MS2 RNA. Note that the ''lys'' gene overlaps segments of both the ''cp'' and ''rep'' genes.|Location of protein-coding genes within bacteriophage MS2 RNA. Note that the ''lys'' gene overlaps segments of both the ''cp'' and ''rep'' genes. Scale is approximate.]]

{| class="wikitable sortable"
|-
! Gene !! Gene length !! Gene product !! Amino acids
|-
| '''''[https://www.ncbi.nlm.nih.gov/gene?cmd=Retrieve&dopt=full_report&list_uids=1260898 mat] '''''(MS2g1) || 1487 nt || [https://www.ncbi.nlm.nih.gov/protein/9626312 maturation protein] || 393
|-
| '''''[https://www.ncbi.nlm.nih.gov/gene?cmd=Retrieve&dopt=full_report&list_uids=1260899 cp]''' ''(MS2g2) || 510 nt || [https://www.ncbi.nlm.nih.gov/protein/9626313 coat protein] || 130
|-
| '''''[https://www.ncbi.nlm.nih.gov/gene?cmd=Retrieve&dopt=full_report&list_uids=1260897 lys] '''''(MS2g3) || 295 nt || [https://www.ncbi.nlm.nih.gov/protein/9626314 lysis protein] || 75
|-
| '''''[https://www.ncbi.nlm.nih.gov/gene?cmd=Retrieve&dopt=full_report&list_uids=1260900 rep]''''' (MS2g4) || 2055 nt || [https://www.ncbi.nlm.nih.gov/protein/9626315 RNA replicase, beta subunit] || 545
|}

===Capsid structure===
[[File:Ms2capsid.png|thumb|right|280px|'''MS2 capsid structure'''. The three quasi-equivalent conformers in the structure are labelled blue (chain a), green (chain b) and magenta (chain c).]]
An MS2 [[virion]] (viral particle) is about 27 nm in diameter, as determined by electron microscopy.<ref>{{cite journal |last=Strauss |first=J. H. |last2=Sinsheimer |first2=R. L. |title=Purification and properties of bacteriophage MS2 and of its ribonucleic acid |journal=[[Journal of Molecular Biology|J Mol Biol]] |year=1963 |volume=7 |issue=1 |pages=43–54 |doi=10.1016/S0022-2836(63)80017-0 }}</ref> It consists of one copy of the maturation protein and 180 copies of the coat protein (organized as 90 dimers) arranged into an [[icosahedral]] shell with triangulation number [[capsid#T-number|T=3]], protecting the genomic RNA inside.<ref>{{cite journal |last=Valegård |first=K. |last2=Lilias |first2=L. |last3=Fridborg |first3=K. |last4=Unge |first4=T. |title=The three-dimensional structure of the bacterial virus MS2 |journal=Nature |year=1990 |volume=345 |issue=6270 |pages=36–41 |doi=10.1038/345036a0 }}</ref> The virion has an [[isoelectric point]] (pI) of 3.9.<ref>{{cite journal |last=Dowd |first=S. E. |title=Delineating the Specific Influence of Virus Isoelectric Point and Size on Virus Adsorption and Transport Through Sandy Soils |journal=[[Applied and Environmental Microbiology|Appl. Environ. Microbiol.]] |volume=64 |issue=2 |year=1998 |pages=405–410 |doi= |url=http://aem.asm.org/content/64/2/405.abstract |display-authors=etal}}</ref>

The structure of the coat protein is a five-stranded [[beta sheet|β-sheet]] with two [[alpha helix|α-helices]] and a [[beta hairpin|hairpin]]. When the [[capsid]] is assembled, the helices and hairpin face the exterior of the particle, while the β-sheet faces the interior.<ref name=Golmohammadi>{{cite journal |last=Golmohammadi |first=R. |last2=Valegård |first2=K. |last3=Fridborg |first3=K. |last4=Liljas |first4=L. |year=1993 |title=The refined structure of bacteriophage MS2 at 2·8 Å resolution |journal=[[Journal of Molecular Biology|J Mol Biol]] |volume=234 |issue=3 |pages=620–639 |doi=10.1006/jmbi.1993.1616 }}</ref>

===Life cycle===
MS2 infects enteric bacteria carrying the [[Fertility factor (bacteria)|fertility (F) factor]], a [[plasmid]] that allows cells to serve as DNA donors in [[bacterial conjugation]]. Genes on the F plasmid lead to the production of an F [[pilus]], which serves as the viral receptor. MS2 attaches to the side of the pilus via its single maturation protein. The precise mechanism by which phage RNA enters the bacterium is unknown.

Once the viral RNA has entered the cell, it begins to function as a [[messenger RNA]] for the production of phage proteins. The gene for the most abundant protein, the coat protein, can be immediately translated. The translation start of the replicase gene is normally hidden within RNA secondary structure, but can be transiently opened as [[ribosome]]s pass through the coat protein gene. Replicase translation is also shut down once large amounts of coat protein have been made; coat protein dimers bind and stabilize the RNA "operator [[stem-loop|hairpin]]", blocking the replicase start. The start of the maturation protein gene is accessible in RNA being replicated but hidden within RNA secondary structure in the completed MS2 RNA; this ensures translation of only a very few copies of maturation protein per RNA. Finally, the lysis protein gene can only be initiated by ribosomes that have completed translation of the coat protein gene and "slip back" to the start of the lysis protein gene, at about a 5% frequency.<ref name="van Duin" />

Replication of the plus-strand MS2 genome requires synthesis of the complementary minus strand RNA, which can then be used as a template for synthesis of a new plus strand RNA. MS2 replication has been much less well studied than replication of the highly related [[bacteriophage Qβ]], partly because the MS2 replicase has been difficult to isolate, but is likely to be similar.<ref name="van Duin" />

The formation of the virion is thought to be initiated by binding of maturation protein to the MS2 RNA; in fact, the complex of maturation protein and RNA is infectious. The assembly of the icosahedral shell or [[capsid]] from coat proteins can occur in the absence of RNA; however, capsid assembly is nucleated by coat protein dimer binding to the operator hairpin, and assembly occurs at much lower concentrations of coat protein when MS2 RNA is present.<ref name="van Duin" />

Bacterial lysis and release of newly formed virions occurs when sufficient lysis protein has accumulated. Lysis protein forms pores in the cytoplasmic membrane, which leads to loss of [[membrane potential]] and breakdown of the [[cell wall#Bacterial cell walls|cell wall]].<ref name="van Duin" />

==Applications==

Since 1998,<ref>Bertrand, E., Chartrand, P., Schaefer, M., Shenoy, S.M., Singer, R.H., and Long, R.M. (1998). Localization of ASH1 mRNA particles in living yeast" ''Mol. Cell'' 2, 437–445.</ref> the MS2 operator hairpin and coat protein have found utility in the detection of RNA in living cells (see [[MS2 tagging]]). MS2 and other viral capsids are also currently under investigation as agents in drug delivery, tumor imaging, and light harvesting applications <ref>Glasgow, J. & Tullman-Ercek, D. [https://link.springer.com/article/10.1007%2Fs00253-014-5787-3 Production and applications of engineered viral capsids.] Appl. Microbiol. Biotechnol. 98, 5847–58 (2014).</ref>

MS-2, due to its structural similarities to noroviruses, its similar optimum proliferation conditions, and non-pathogenicity to humans, has been used as substitute for noroviruses in studies of disease transmission.<ref>{{cite web|url=http://www.today.com/health/viruses-spread-crazy-office-1D80134085|title=Viruses spread 'like crazy' in an office, study finds|author=Maggie Fox}}</ref>

==See also==
{{Portal|Viruses}}
* [[bacteriophage f2]]
* [[bacteriophage Qβ]]
* [[Phi-X174 phage]]
* [[Bacteriophage]]

==References==
{{Reflist|2}}

==External links==
* [https://www.ncbi.nlm.nih.gov/nuccore/EF204940.1 Complete genome] (also isolates [https://www.ncbi.nlm.nih.gov/nuccore/EF108465.1 R17], [https://www.ncbi.nlm.nih.gov/nuccore/EF108464.1 DL16], and [https://www.ncbi.nlm.nih.gov/nuccore/EF204939.1 J20])

{{Taxonbar|from=Q4840020}}

{{DEFAULTSORT:Bacteriophage Ms2}}
[[Category:Leviviridae]]

Pegivirus

2018-03-23T03:06:24Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q16917856}} (1 sig. taxon ID); WP:GenFixe using AWB

{{taxobox
| color = violet
| name = Flaviviruses
| virus_group = iv
| familia = ''[[Flaviviridae]]''
| genus = '''''Pegivirus'''''
| subdivision_ranks = Species
| subdivision =
''[[Pegivirus A]]'' 
''[[Pegivirus B]]'' 
''[[GBV-C|Pegivirus C]] 
''[[Pegivirus D]]'' 
''[[Equine pegivirus|Pegivirus E]]'' 
''[[Pegivirus F]]'' 
''[[Pegivirus G]]'' 
''[[Pegivirus H]]'' 
''[[Pegivirus I]]'' 
''[[Pegivirus J]]'' 
''[[Pegivirus K]]'' 
}}

'''''Pegivirus''''' is the approved name for a [[genus]] of single positive stranded [[RNA virus]]es in the family ''Flaviviridae''.<ref>{{cite journal |doi=10.1099/jgv.0.000672 |pmid=28218572 |pmc=5370391 |title=ICTV Virus Taxonomy Profile: Flaviviridae |journal=Journal of General Virology |volume=98 |issue=1 |pages=2–3 |year=2017 |last1=Simmonds |first1=Peter |last2=Becher |first2=Paul |last3=Bukh |first3=Jens |last4=Gould |first4=Ernest A |last5=Meyers |first5=Gregor |last6=Monath |first6=Tom |last7=Muerhoff |first7=Scott |last8=Pletnev |first8=Alexander |last9=Rico-Hesse |first9=Rebecca |last10=Smith |first10=Donald B |last11=Stapleton |first11=Jack T |last12=Ictv Report |first12=Consortium }}</ref><ref name="ICTVReport">{{cite web|title=ICTV Report Flaviviridae|url=http://www.ictv.global/report/flaviviridae}}</ref><ref name="Stapleton2011a">{{cite journal |doi=10.1099/vir.0.027490-0 |pmid=21084497 |pmc=3081076 |title=The GB viruses: A review and proposed classification of GBV-A, GBV-C (HGV), and GBV-D in genus Pegivirus within the family Flaviviridae |journal=Journal of General Virology |volume=92 |issue=2 |pages=233–46 |year=2010 |last1=Stapleton |first1=J. T |last2=Foung |first2=S |last3=Muerhoff |first3=A. S |last4=Bukh |first4=J |last5=Simmonds |first5=P }}</ref><ref name=v2013>{{cite journal |doi=10.1007/s00705-013-1749-9 |pmid=23836393 |title=Recently agreed changes to the International Code of Virus Classification and Nomenclature |journal=Archives of Virology |volume=158 |issue=12 |pages=2633–9 |year=2013 |last1=Adams |first1=M. J |last2=Lefkowitz |first2=E. J |last3=King |first3=A. M. Q |last4=Carstens |first4=E. B }}</ref> The name is a derived one: "Pe" stands for "persistent" and "g" is a reference to [[GB virus C|Hepatitis G]], a former name of the ''C'' species.

==Taxonomy==

There are eleven named species within the ''Pegivirus'' genus.<ref name=ICTVReport/> Isolates belonging to the species ''Pegivirus C'' are monophyletic and show < 50% nucleotide (55% amino acid) sequence divergence between aligned sequences from the polyprotein from each other. However all differ by >50% nucleotide (>55% amino acid) divergence from other members of this genus. Pegiviruses assigned to this species (Pegivirus A) originate from primate host species (humans, chimpanzees and several New World monkey species). The sequence U22303 has been assigned the type member of the species as this was the first pegivirus to be described for this species. Terminology to describe viruses with different hosts has not been approved by the International Committee on Taxonomy of Viruses (ICTV); however, Pegivirus A viruses have been called HPgV for human pegivirus, SPgV for new world simian pegiviruses, and SPgVcpz for chimpanzee simian virus. This remains an area requiring further clarification.{{citation needed|date=February 2018}}

A second species within the pegiviruses is termed ''Pegivirus B''. Only one virus was included in the naming proposal, which was a complete genome of a virus found in the bat species ''[[Pteropus giganteus]]''. This sequence differs by >50% nucleotide (>55% amino acid) divergence from all proposed members of the primate-derived Pegivirus C species that originate from primate host species (humans, chimpanzees and several New World monkey species). The sequence GU566734 has been assigned the type member of the species as this was the first pegivirus to be described for this species.{{citation needed|date=February 2018}}

The use of deep sequencing technologies has identified additional viruses that differ from ''Pegivirus B'' species by >50% nucleotide (>55% amino acid) and ''Pegivirus C'' in rodents, horses, and in different bat species, and old world monkeys and the number of ''Pegivirus'' species has been expanded to eleven.<ref name=Smith2016/>

===Revised taxonomy===

The species known in 2016 have been classified into 11 species—Pegivirus A–K.<ref name=ICTVReport/><ref name=Smith2016>{{cite journal |doi=10.1099/jgv.0.000612 |pmid=27692039 |pmc=5770844 |title=Proposed update to the taxonomy of the genera Hepacivirus and Pegivirus within the family Flaviviridae |journal=Journal of General Virology |volume=97 |issue=11 |pages=2894–2907 |year=2016 |last1=Smith |first1=Donald B |last2=Becher |first2=Paul |last3=Bukh |first3=Jens |last4=Gould |first4=Ernest A |last5=Meyers |first5=Gregor |last6=Monath |first6=Thomas |last7=Muerhoff |first7=A. Scott |last8=Pletnev |first8=Alexander |last9=Rico-Hesse |first9=Rebecca |last10=Stapleton |first10=Jack T |last11=Simmonds |first11=Peter }}</ref>

* ''Pegivirus A'' includes the virus GBV-A
* ''Pegivirus B'' includes the virus GBV-D
* ''Pegivirus C'' includes the virus [[GBV-C]]
* ''Pegivirus D'' includes the virus Theiler’s disease-associated virus
* ''Pegivirus E'' includes the virus [[Equine pegivirus]]
* ''Pegivirus F'' includes the virus Bat pegivirus
* ''Pegivirus G'' includes the virus Bat pegivirus
* ''Pegivirus H'' includes the virus [[HPgV-2|Human pegivirus 2]]
* ''Pegivirus I'' includes the virus Bat pegivirus
* ''Pegivirus J'' includes the virus Rodent pegivirus
* ''Pegivirus K'' includes the virus Porcine pegivirus

==History==

*In 1967 it was reported that experimental inoculation of serum from a [[surgeon]] (G. Barker) with acute hepatitis into [[tamarin]]s resulted in [[hepatitis]].
*In 1995, two new members of the family ''Flaviviridae'' (GBV-A and GBV-B) were identified in tamarins that developed hepatitis following inoculation with the 11th GB passage. A number of GBV-A variants were later identified in wild [[New World]] monkeys that were captured.
*Subsequently, in 1995 a human virus was identified [GBV-C or hepatitis G virus (HGV)].
*A more distantly related virus (GBV-D) was later discovered in the bat (''[[Pteropus giganteus]]'').<ref name=Epstein2010>{{cite journal |doi=10.1371/journal.ppat.1000972 |pmid=20617167 |pmc=2895649 |title=Identification of GBV-D, a Novel GB-like Flavivirus from Old World Frugivorous Bats (Pteropus giganteus) in Bangladesh |journal=PLoS Pathogens |volume=6 |issue=7 |pages=e1000972 |year=2010 |last1=Epstein |first1=Jonathan H |last2=Quan |first2=Phenix-Lan |last3=Briese |first3=Thomas |last4=Street |first4=Craig |last5=Jabado |first5=Omar |last6=Conlan |first6=Sean |last7=Ali Khan |first7=Shahneaz |last8=Verdugo |first8=Dawn |last9=Hossain |first9=M. Jahangir |last10=Hutchison |first10=Stephen K |last11=Egholm |first11=Michael |last12=Luby |first12=Stephen P |last13=Daszak |first13=Peter |last14=Lipkin |first14=W. Ian }}</ref> Another virus—rodent pegavirus—has been isolated from the white throated woodrat (''[[Neotoma albigula]]'').<ref name=Kapoor2013>{{cite journal |doi=10.1128/mBio.00216-13 |pmid=23572554 |pmc=3622934 |title=Identification of Rodent Homologs of Hepatitis C Virus and Pegiviruses |journal=MBio |volume=4 |issue=2 |pages=e00216–13 |year=2013 |last1=Kapoor |first1=A |last2=Simmonds |first2=P |last3=Scheel |first3=T. K. H |last4=Hjelle |first4=B |last5=Cullen |first5=J. M |last6=Burbelo |first6=P. D |last7=Chauhan |first7=L. V |last8=Duraisamy |first8=R |last9=Sanchez Leon |first9=M |last10=Jain |first10=K |last11=Vandegrift |first11=K. J |last12=Calisher |first12=C. H |last13=Rice |first13=C. M |last14=Lipkin |first14=W. I }}</ref> A pegivirus (equine pegivirus) has also been isolated from a horse.<ref>{{cite journal |doi=10.1128/JVI.00324-13 |pmid=23596285 |pmc=3676142 |title=Identification of a Pegivirus (GB Virus-Like Virus) That Infects Horses |journal=Journal of Virology |volume=87 |issue=12 |pages=7185–90 |year=2013 |last1=Kapoor |first1=A |last2=Simmonds |first2=P |last3=Cullen |first3=J. M |last4=Scheel |first4=T. K. H |last5=Medina |first5=J. L |last6=Giannitti |first6=F |last7=Nishiuchi |first7=E |last8=Brock |first8=K. V |last9=Burbelo |first9=P. D |last10=Rice |first10=C. M |last11=Lipkin |first11=W. I }}</ref>
*The genus ''Pegivirus'' was proposed in 2011.<ref name="Stapleton2011a"/> GBV-B was assigned to the ''[[Hepacivirus]]'' genus,<ref name=ICTVReport/> whereas GBV-A together with GBV-C were assigned to the new ''Pegivirus'' genus.<ref name=ICTVReport/>
*[[Theiler's disease]] — a form of equine hepatitis — also appears to be caused by a pegivirus—[[Theiler disease–associated virus]].<ref name=Chandriani2013>{{cite journal |doi=10.1073/pnas.1219217110 |pmid=23509292 |pmc=3625295 |title=Identification of a previously undescribed divergent virus from the Flaviviridae in an outbreak of equine serum hepatitis |journal=Proceedings of the National Academy of Sciences |volume=110 |issue=15 |pages=E1407–15 |year=2013 |last1=Chandriani |first1=S |last2=Skewes-Cox |first2=P |last3=Zhong |first3=W |last4=Ganem |first4=D. E |last5=Divers |first5=T. J |last6=Van Blaricum |first6=A. J |last7=Tennant |first7=B. C |last8=Kistler |first8=A. L |bibcode=2013PNAS..110E1407C }}</ref>
*Human hepegivirus 1 or Human Pegivirus 2 (HPgV2) is a virus isolated from 2 multiply transfused hemophiliacs and two transfused patients.<ref name=Kapoor2015>{{cite journal |doi=10.1128/mBio.01466-15 |pmid=26396247 |pmc=4600124 |title=Virome Analysis of Transfusion Recipients Reveals a Novel Human Virus That Shares Genomic Features with Hepaciviruses and Pegiviruses |journal=MBio |volume=6 |issue=5 |pages=e01466–15 |year=2015 |last1=Kapoor |first1=Amit |last2=Kumar |first2=Arvind |last3=Simmonds |first3=Peter |last4=Bhuva |first4=Nishit |last5=Singh Chauhan |first5=Lokendra |last6=Lee |first6=Bohyun |last7=Sall |first7=Amadou Alpha |last8=Jin |first8=Zhezhen |last9=Morse |first9=Stephen S |last10=Shaz |first10=Beth |last11=Burbelo |first11=Peter D |last12=Lipkin |first12=W. Ian }}</ref> This virus appears to belong to a new clade in the Pegiviruses.
*A related virus has been isolated from the graceful catshark (''[[Proscyllium habereri]]).<ref name=Shi2015>{{cite journal |doi=10.1128/JVI.02036-15 |pmid=26491167 |pmc=4702705 |title=Divergent Viruses Discovered in Arthropods and Vertebrates Revise the Evolutionary History of the Flaviviridae and Related Viruses |journal=Journal of Virology |volume=90 |issue=2 |pages=659–69 |year=2016 |last1=Shi |first1=Mang |last2=Lin |first2=Xian-Dan |last3=Vasilakis |first3=Nikos |last4=Tian |first4=Jun-Hua |last5=Li |first5=Ci-Xiu |last6=Chen |first6=Liang-Jun |last7=Eastwood |first7=Gillian |last8=Diao |first8=Xiu-Nian |last9=Chen |first9=Ming-Hui |last10=Chen |first10=Xiao |last11=Qin |first11=Xin-Cheng |last12=Widen |first12=Steven G |last13=Wood |first13=Thomas G |last14=Tesh |first14=Robert B |last15=Xu |first15=Jianguo |last16=Holmes |first16=Edward C |last17=Zhang |first17=Yong-Zhen }}</ref>
*The human pegiviruses appear to be related to the non-human primate species.<ref name=Thézé2015>{{cite journal |doi=10.1093/gbe/evv202 |pmid=26494702 |pmc=5635594 |title=Evolutionary and Phylogenetic Analysis of the Hepaciviruses and Pegiviruses |journal=Genome Biology and Evolution |volume=7 |issue=11 |pages=2996–3008 |year=2015 |last1=Thézé |first1=Julien |last2=Lowes |first2=Sophia |last3=Parker |first3=Joe |last4=Pybus |first4=Oliver G }}</ref>
*In 2016 Pegivirus genus was divided into 11 species – pegiviruses A–K with GBV-C classified under pegivirus C species.<ref name=Smith2016/>

==References==
{{Reflist|2}}

{{Taxonbar|from=Q16917856}}

[[Category:Flaviviridae]]
[[Category:Pegiviruses]]

Hepacivirus

2018-03-23T02:32:00Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q2082778}} (1 sig. taxon ID); WP:GenFixes, using AWB

{{Use dmy dates|date=April 2017}}
{{taxobox
| virus_group = iv
| familia = ''[[Flaviviridae]]''
| genus = '''''Hepacivirus'''''
| subdivision_ranks = Species
| subdivision =
''[[Canine hepacivirus|Hepacivirus A]]'' 
''[[Hepacivirus B]]'' 
''[[Hepatitis C virus|Hepacivirus C]]'' 
''[[Hepacivirus D]]'' 
''[[Hepacivirus E]]'' 
''[[Hepacivirus F]]'' 
''[[Hepacivirus G]]'' 
''[[Hepacivirus H]]'' 
''[[Hepacivirus I]]'' 
''[[Hepacivirus J]]'' 
''[[Hepacivirus K]]'' 
''[[Hepacivirus L]]'' 
''[[Hepacivirus M]]'' 
''[[Hepacivirus N]]''
}}

'''''Hepacivirus''''' is a genus of positive-sense, single-stranded RNA [[viruses]] in the family ''[[Flaviviridae]]''.<ref name=ICTVReport>{{cite web|title=Flaviviridae|url=http://www.ictv.global/report/flaviviridae|website=International Committee on Taxonomy of Viruses (ICTV)|language=en}}</ref> The virus [[hepatitis C virus]](HCV), belonging to the species ''Hepacivirus C'', has humans as its only known natural host, and is associated with hepatitis and hepatocellular carcinoma.<ref name=ICTVReport/><ref name=ViralZone>{{cite web|title=Viral Zone|url=http://viralzone.expasy.org/all_by_species/37.html|publisher=ExPASy|accessdate=15 June 2015}}</ref>

==Taxonomy==
<big>'''Group: ssRNA(+)'''</big>
{{Collapsible list|title= <big>Order: Unassigned</big>
|1={{Collapsible list| framestyle=border:none; padding:1.0em;|title=Family: [[Flaviviridae]]
|1={{hidden begin|title=Genus: Hepacivirus}}
*'''''[[Hepatitis C virus]]'''''
{{hidden end}}
}}
}}<ref>{{cite web|title=Taxonomy|url=https://talk.ictvonline.org/taxonomy/|website=International Committee on Taxonomy of Viruses (ICTV)|language=en}}</ref>

===Revised taxonomy===

The viruses belonging to this genus have been classified into 14 species - ''Hepacivirus A-N''.<ref name=pmid27692039>{{cite journal |doi=10.1099/jgv.0.000612 |pmid=27692039 |pmc=5770844 |title=Proposed update to the taxonomy of the genera Hepacivirus and Pegivirus within the Flaviviridae family |journal=Journal of General Virology |volume=97 |issue=11 |pages=2894–2907 |year=2016 |last1=Smith |first1=Donald B |last2=Becher |first2=Paul |last3=Bukh |first3=Jens |last4=Gould |first4=Ernest A |last5=Meyers |first5=Gregor |last6=Monath |first6=Thomas |last7=Muerhoff |first7=A. Scott |last8=Pletnev |first8=Alexander |last9=Rico-Hesse |first9=Rebecca |last10=Stapleton |first10=Jack T |last11=Simmonds |first11=Peter }}</ref><ref name=ICTVReport/>

* The species ''Hepacivirus A'' includes the virus canine hepacivirus/non-primate hepacivirus/equine hepacivirus
* The species ''Hepacivirus B'' includes the virus GBV-B
* The species ''Hepacivirus C'' includes the virus Hepatitis C virus (previously included in the species ''Hepatitis C virus'')
* The species ''Hepacivirus D'' includes the virus Guereza hepacivirus
* The species ''Hepacivirus E'' includes the virus rodent hepacivirus-339
* The species ''Hepacivirus F'' includes the virus rodent hepacivirus-NLR07-oct70
* The species ''Hepacivirus G'' includes the virus Norway rat hepacivirus 1
* The species ''Hepacivirus H'' includes the virus Norway rat hepacivirus 2
* The species ''Hepacivirus I'' includes the virus rodent hepacivirus-SAR-3/RSA/2008
* The species ''Hepacivirus J'' incVan Nguyenludes the virus rodent hepacivirus- RMU10-3382/GER/2010
* The species ''Hepacivirus K'' includes the virus bat hepacivirus-PDB-829
* The species ''Hepacivirus L'' includes the virus bat hepacivirus-PDB-112
* The species ''Hepacivirus M'' includes the virus bat hepacivirus-PDB-491.1
* The species ''Hepacivirus N'' includes the virus bovine hepacivirus

A new virus has been isolated from the bamboo rat (''[[Rhizomys pruinosus]]'') while belonging to this genus does not appear to belong to any previously described species.<ref name=pmid29495551>{{cite journal |doi=10.3390/v10030102 |pmid=29495551 |title=Detection and Characterization of Homologues of Human Hepatitis Viruses and Pegiviruses in Rodents and Bats in Vietnam |journal=Viruses |volume=10 |issue=3 |pages=102 |year=2018 |last1=Van Nguyen |first1=Dung |last2=Van Nguyen |first2=Cuong |last3=Bonsall |first3=David |last4=Ngo |first4=Tue |last5=Carrique-Mas |first5=Juan |last6=Pham |first6=Anh |last7=Bryant |first7=Juliet |last8=Thwaites |first8=Guy |last9=Baker |first9=Stephen |last10=Woolhouse |first10=Mark |last11=Simmonds |first11=Peter }}</ref>

==Structure==
Viruses in the genus ''Hepacivirus'' are enveloped, with spherical geometries. The diameter is around 50 nm. Genomes are linear and non-segmented, around 10kb in length.<ref name=ICTVReport/><ref name=ViralZone />

{| class="wikitable sortable" style="text-align:center"
|-
! Genus !! Structure || Symmetry !! Capsid !! Genomic arrangement !! Genomic segmentation
|-
|Hepacivirus||Icosahedral-like||Pseudo T=3||Enveloped||Linear||Monopartite
|}

==Life cycle==
Entry into the host cell is achieved by attachment of the viral envelope protein E to host receptors, which mediates clathrin-mediated endocytosis. Replication follows the positive stranded RNA virus replication model. Positive stranded rna virus transcription is the method of transcription. Translation takes place by viral initiation. Human serve as the natural host. Transmission routes are sexual, blood, and contact.<ref name=ICTVReport/><ref name=ViralZone />

{| class="wikitable sortable" style="text-align:center"
|-
! Genus !! Host details !! Tissue tropism !! Entry details !! Release details !! Replication site !! Assembly site !! Transmission
|-
|Hepacivirus||Humans||Epithelium: skin; epithelium: kidney; epithelium: intestine; epithelium: testes||Clathrin-mediated endocytosis||Secretion||Cytoplasm||Cytoplasm||Sex; blood
|}

==History==
Hepatitis C virus (HCV), which is the causative agent of [[hepatitis C]] in humans, and a member of the species ''Hepacivirus C'', was discovered in 1989.<ref name=pmid21084497/> Seven genotypes (1-7) and eighty six subtypes (1a, 1b etc.) of hepatitis C virus have been named.<ref>{{cite web|title=HCV Classification|url=http://talk.ictvonline.org/links/hcv/hcv-classification.htm|website=International Committee on Taxonomy of Viruses (ICTV)|language=en}}</ref>

[[GBV-B]] virus (also known as GB virus B) discovered in 1995 is capable of infecting [[New World monkeys]], in particular [[tamarins]]. Like HCV it is transmitted by the blood-borne route and similar to HCV it is associated with the [[viral hepatitis]]. However GBV-B has never been identified in wild animals and its natural host is not known.<ref name=pmid21084497/>

==Additional Information==
Additional hepaciviruses have been described from bats, rodents including bank voles, horses and dogs.<ref name=pmid23572554>{{cite journal |doi=10.1128/mBio.00216-13 |pmid=23572554 |pmc=3622934 |title=Identification of Rodent Homologs of Hepatitis C Virus and Pegiviruses |journal=M ''Bio'' |volume=4 |issue=2 |pages=e00216–13 |year=2013 |last1=Kapoor |first1=A |last2=Simmonds |first2=P |last3=Scheel |first3=T. K. H |last4=Hjelle |first4=B |last5=Cullen |first5=J. M |last6=Burbelo |first6=P. D |last7=Chauhan |first7=L. V |last8=Duraisamy |first8=R |last9=Sanchez Leon |first9=M |last10=Jain |first10=K |last11=Vandegrift |first11=K. J |last12=Calisher |first12=C. H |last13=Rice |first13=C. M |last14=Lipkin |first14=W. I }}</ref><ref name=pmid23818848>{{cite journal |doi=10.1371/journal.ppat.1003438 |pmid=23818848 |pmc=3688547 |title=Evidence for Novel Hepaciviruses in Rodents |journal=PLoS Pathogens |volume=9 |issue=6 |pages=e1003438 |year=2013 |last1=Drexler |first1=Jan Felix |last2=Corman |first2=Victor Max |last3=Müller |first3=Marcel Alexander |last4=Lukashev |first4=Alexander N |last5=Gmyl |first5=Anatoly |last6=Coutard |first6=Bruno |last7=Adam |first7=Alexander |last8=Ritz |first8=Daniel |last9=Leijten |first9=Lonneke M |last10=Van Riel |first10=Debby |last11=Kallies |first11=Rene |last12=Klose |first12=Stefan M |last13=Gloza-Rausch |first13=Florian |last14=Binger |first14=Tabea |last15=Annan |first15=Augustina |last16=Adu-Sarkodie |first16=Yaw |last17=Oppong |first17=Samuel |last18=Bourgarel |first18=Mathieu |last19=Rupp |first19=Daniel |last20=Hoffmann |first20=Bernd |last21=Schlegel |first21=Mathias |last22=Kümmerer |first22=Beate M |last23=Krüger |first23=Detlev H |last24=Schmidt-Chanasit |first24=Jonas |last25=Setién |first25=Alvaro Aguilar |last26=Cottontail |first26=Veronika M |last27=Hemachudha |first27=Thiravat |last28=Wacharapluesadee |first28=Supaporn |last29=Osterrieder |first29=Klaus |last30=Bartenschlager |first30=Ralf |display-authors=29 }}</ref><ref name=pmid23740998>{{cite journal |doi=10.1128/JVI.00888-13 |pmid=23740998 |pmc=3754081 |title=A Novel Hepacivirus with an Unusually Long and Intrinsically Disordered NS5A Protein in a Wild Old World Primate |journal=Journal of Virology |volume=87 |issue=16 |pages=8971–81 |year=2013 |last1=Lauck |first1=M |last2=Sibley |first2=S. D |last3=Lara |first3=J |last4=Purdy |first4=M. A |last5=Khudyakov |first5=Y |last6=Hyeroba |first6=D |last7=Tumukunde |first7=A |last8=Weny |first8=G |last9=Switzer |first9=W. M |last10=Chapman |first10=C. A |last11=Hughes |first11=A. L |last12=Friedrich |first12=T. C |last13=O'Connor |first13=D. H |last14=Goldberg |first14=T. L }}</ref>

Cattle also appear to be a host for viruses belonging to the species ''Hepacivirus N''.<ref name=pmid25787289>{{cite journal |doi=10.1128/JVI.00393-15 |pmid=25787289 |pmc=4442428 |title=Highly Divergent Hepaciviruses from African Cattle |journal=Journal of Virology |volume=89 |issue=11 |pages=5876–82 |year=2015 |last1=Corman |first1=Victor Max |last2=Grundhoff |first2=Adam |last3=Baechlein |first3=Christine |last4=Fischer |first4=Nicole |last5=Gmyl |first5=Anatoly |last6=Wollny |first6=Robert |last7=Dei |first7=Dickson |last8=Ritz |first8=Daniel |last9=Binger |first9=Tabea |last10=Adankwah |first10=Ernest |last11=Marfo |first11=Kwadwo Sarfo |last12=Annison |first12=Lawrence |last13=Annan |first13=Augustina |last14=Adu-Sarkodie |first14=Yaw |last15=Oppong |first15=Samuel |last16=Becher |first16=Paul |last17=Drosten |first17=Christian |last18=Drexler |first18=Jan Felix }}</ref><ref name=pmid25926652>{{cite journal |doi=10.1128/JVI.00534-15 |pmid=25926652 |pmc=4473572 |title=Identification of a Novel Hepacivirus in Domestic Cattle from Germany |journal=Journal of Virology |volume=89 |issue=14 |pages=7007–15 |year=2015 |last1=Baechlein |first1=Christine |last2=Fischer |first2=Nicole |last3=Grundhoff |first3=Adam |last4=Alawi |first4=Malik |last5=Indenbirken |first5=Daniela |last6=Postel |first6=Alexander |last7=Baron |first7=Anna Lena |last8=Offinger |first8=Jennifer |last9=Becker |first9=Kathrin |last10=Beineke |first10=Andreas |last11=Rehage |first11=Juergen |last12=Becher |first12=Paul }}</ref>

[[Rodent hepacivirus]] is found in the deer mouse (''[[Peromyscus maniculatus]]'').<ref name=pmid21084497>{{cite journal |doi=10.1099/vir.0.027490-0 |pmid=21084497 |pmc=3081076 |title=The GB viruses: A review and proposed classification of GBV-A, GBV-C (HGV), and GBV-D in genus Pegivirus within the family Flaviviridae |journal=Journal of General Virology |volume=92 |issue=2 |pages=233–46 |year=2010 |last1=Stapleton |first1=J. T |last2=Foung |first2=S |last3=Muerhoff |first3=A. S |last4=Bukh |first4=J |last5=Simmonds |first5=P }}</ref>

A virus related to hepaciviruses has been isolated from the graceful catshark (''[[Proscyllium habereri]]'').<ref name=pmid26491167>{{cite journal |doi=10.1128/JVI.02036-15 |pmid=26491167 |pmc=4702705 |title=Divergent Viruses Discovered in Arthropods and Vertebrates Revise the Evolutionary History of the Flaviviridae and Related Viruses |journal=Journal of Virology |volume=90 |issue=2 |pages=659–69 |year=2016 |last1=Shi |first1=Mang |last2=Lin |first2=Xian-Dan |last3=Vasilakis |first3=Nikos |last4=Tian |first4=Jun-Hua |last5=Li |first5=Ci-Xiu |last6=Chen |first6=Liang-Jun |last7=Eastwood |first7=Gillian |last8=Diao |first8=Xiu-Nian |last9=Chen |first9=Ming-Hui |last10=Chen |first10=Xiao |last11=Qin |first11=Xin-Cheng |last12=Widen |first12=Steven G |last13=Wood |first13=Thomas G |last14=Tesh |first14=Robert B |last15=Xu |first15=Jianguo |last16=Holmes |first16=Edward C |last17=Zhang |first17=Yong-Zhen }}</ref>

There are at least two subtypes of equine hepacivirus.<ref name=pmid27882682>{{cite journal |doi=10.1111/tbed.12587 |pmid=27882682 |title=Prevalence of Equine Hepacivirus Infections in France and Evidence for Two Viral Subtypes Circulating Worldwide |journal=Transboundary and Emerging Diseases |volume=64 |issue=6 |pages=1884–1897 |year=2017 |last1=Pronost |first1=S |last2=Hue |first2=E |last3=Fortier |first3=C |last4=Foursin |first4=M |last5=Fortier |first5=G |last6=Desbrosse |first6=F |last7=Rey |first7=F. A |last8=Pitel |first8=P.-H |last9=Richard |first9=E |last10=Saunier |first10=B }}</ref>

The viruses most closely related to hepatitis C virus are the equine hepaciviruses belonging to the species ''Hepacivirus A'' .<ref name=pmid26494702>{{cite journal |doi=10.1093/gbe/evv202 |pmid=26494702 |pmc=5635594 |title=Evolutionary and Phylogenetic Analysis of the Hepaciviruses and Pegiviruses |journal=Genome Biology and Evolution |volume=7 |issue=11 |pages=2996–3008 |year=2015 |last1=Thézé |first1=Julien |last2=Lowes |first2=Sophia |last3=Parker |first3=Joe |last4=Pybus |first4=Oliver G }}</ref>

==References==
{{Reflist}}

==External links==
* [http://www.ictv.global/report/flaviviridae '''ICTV Report''': Flaviviridae]
* [http://viralzone.expasy.org/all_by_species/37.html '''Viralzone''': Hepacivirus]
* [http://www.viprbrc.org/brc/home.do?decorator=flavi Virus Pathogen Database and Analysis Resource (ViPR): Flaviviridae]
{{Baltimore classification}}

{{Taxonbar|from=Q2082778}}

[[Category:Hepaciviruses]]

Heteromeles arbutifolia

2018-03-23T02:03:20Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q2655434}} (6 sig. taxon IDs); WP:GenFixes using AWB

{{Redirect|Toyon|the community in California|Toyon, California}}
{{Taxobox
| image = Heteromeles arbutifolia 1.jpg
| image_caption = Toyon bush in habitat
| regnum = [[Plant]]ae
| unranked_divisio = [[Angiosperms]]
| unranked_classis = [[Eudicots]]
| unranked_ordo = [[Rosids]]
| ordo = [[Rosales]]
| familia = [[Rosaceae]]<ref>Germplasm Resources Information Network, 1910</ref>
| subfamilia = [[Amygdaloideae]]<ref name=Potter>Potter, D., et al. (2007). Phylogeny and classification of Rosaceae. ''Plant Systematics and Evolution''. 266(1–2): 5–43. <nowiki>[Referring to the subfamily by the name "Spiraeoideae"]</nowiki></ref>
| tribus = [[Maleae]]
| subtribus = [[Malinae]]
| genus = '''''Heteromeles'''''
| genus_authority = [[Max Joseph Roemer|M.Roem.]] ''nom. cons.'' 1847
| species = '''''H. arbutifolia'''''
| binomial = ''Heteromeles arbutifolia''
| binomial_authority = ([[John Lindley|Lindl.]]) [[Max Joseph Roemer|M.Roem.]]<ref>[http://ucjeps.berkeley.edu/cgi-bin/get_JM_treatment.pl?Heteromeles+arbutifolia Jepson Flora Project (1993) ''Heteromeles arbutifolia'', University of California, Berkeley]</ref>
| synonyms =
* ''Photinia arbutifolia'' Lindl.
* ''Crataegus arbutifolia'' W.T.Aiton nom. illeg.<ref>{{citation |url=http://www.tropicos.org/Name/50051852 |title=Tropicos.org |accessdate=11 November 2016}}</ref>
* ''Heteromeles fremontiana'' Decaisne
* ''Heteromeles salicifolia'' (C.Presl) Abrams
* ''Photinia salicifolia'' C.Presl
| synonyms_ref = <ref name=PhippsHA>{{citation |chapter-url=http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=250100233 |chapter=''Heteromeles arbutifolia'' (Lindley) M. Roemer, Fam. Nat. Syn. Monogr. 3: 105. 1847 |title=Flora of North America |volume=9 |year=2015 |author=James B. Phipps}}</ref>
|range_map =Heteromeles_arbutifolia_range_map.jpg
|range_map_caption = Natural range
}}

'''''Heteromeles arbutifolia''''' ({{IPAc-en|ˌ|h|ɛ|t|ᵻ|r|oʊ-|ˈ|m|iː|l|iː|z|_|ɑːr|ˌ|b|juː|t|ᵻ|ˈ|f|oʊ|l|i|ə}};<ref>''Sunset Western Garden Book,'' 1995:606–607</ref> more commonly {{IPAc-en|ˌ|h|ɛ|t|ə|ˈ|r|ɒ|m|əl|iː|z}} by Californian botanists), commonly known as '''toyon''', is a common [[Perennial plant|perennial]] [[shrub]] native to extreme southwest [[Oregon]],{{citation needed|date=January 2014}} [[California]], [[Baja California]],<ref name=PhippsHA/> and [[British Columbia]].<ref name=Jepson1/> It is the sole species in the genus '''''Heteromeles'''''.

Toyon is a prominent component of the [[coastal sage scrub]] plant community, and is a part of drought-adapted [[chaparral]] and mixed [[California oak woodland|oak woodland]] [[habitat]]s.<ref>C.M. Hogan, 2008</ref> It is also known by the common names '''Christmas berry'''<ref name=FNA>{{citation |chapter-url=http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=115321 |chapter=''Heteromeles'' M. Roemer, Fam. Nat. Syn. Monogr. 3: 100, 105. 1847. [name conserved] |title=Flora of North America |year=2015 |volume=9 |author=James B. Phipps}}</ref> and '''California holly'''. Accordingly, "the abundance of this species in the hills above [[Los Angeles]] gave rise to the name [[Hollywood]]."<ref>{{cite book|title=Introduction to the Plant Life of Southern California: Coast to Foothills|year=2005|publisher=University of California Press|location=Berkeley and Los Angeles|isbn=0-520-24199-1|page=103|author=Rundel, Philip W|author2=Gustafson, Robert}}</ref>

==Description==
Toyon typically grows from 2–5 m (rarely up to 10 m in shaded conditions) and has a rounded to irregular top. Its [[leaf|leaves]] are [[evergreen]], alternate, sharply toothed, have short [[Petiole (botany)|petiole]]s, and are 5–10 cm in length and 2–4 cm wide. In the early summer it produces small white [[flower]]s 6–10 mm diameter in dense terminal [[corymb]]s. Flowering peaks in June <ref>[https://www.inaturalist.org/taxa/53405-Heteromeles-arbutifolia Heteromeles arbutifolia] at iNaturalist</ref>

The five [[petal]]s are rounded. The [[fruit]] is a small [[pome]],<ref name=Jepson1>{{cite web|title=''Heteromeles arbutifolia'', in Jepson Flora Project|url=http://ucjeps.berkeley.edu/cgi-bin/get_IJM.pl?tid=28072|publisher=Regents of the University of California|accessdate=14 November 2013}}</ref> 5–10 mm across, bright red and berry-like, produced in large quantities, maturing in the fall and persisting well into the winter.

==Cultivation==
Toyon can be grown in domestic gardens in well-drained soil, and is cultivated as an ornamental plant as far north as Southern England. It can survive temperatures as low as -12 °C.{{Citation needed|date=April 2009}} In winter, the bright red pomes (which birds often eat voraciously) are showy.

Like many other genera in the [[Rosaceae]] tribe [[Maleae]], toyon includes some cultivars that are susceptible to [[fireblight]].<ref>[http://www.aces.edu/pubs/docs/A/ANR-0542/ANR-0542.pdf Austin Hagan, Edward Sikora, William Gazaway, Nancy Kokalis- Burelle, 2004. ''Fire Blight on Fruit Trees and Woody Ornamentals'', Alabama A&M and Auburn Universities]</ref> It survives on little water, making it suitable for xeriscape gardening, and is less of a fire hazard than some chaparral plants.<ref>[http://davesgarden.com/guides/pf/go/58417/ Dave's Garden]</ref>

==Wildlife value==
They are visited by [[butterfly|butterflies]], and have a mild, [[Crataegus|hawthorn]]-like scent. The fruit are consumed by [[bird]]s, including [[mockingbird]]s, [[American robin]]s, [[cedar waxwing]]s and [[hermit thrush]]es.<ref>{{cite | title=Ask The Naturalist: How Important Are Red Toyon Berries To the Winter Food Chain?| first1=Alan | last1=Kaplan | first2=Alison | last2=Hawkes | work=Bay Nature | date=December 22, 2016|url=http://baynature.org/article/ask-the-naturalist-how-important-are-all-those-red-berries-we-see-to-the-winter-food-chain/}}</ref> [[Mammal]]s including [[coyote]]s and [[bear]]s also eat and disperse the pomes.

==Traditional use==
The pomes provided food for local [[Native Americans in the United States|Native American]] tribes, such as the [[Chumash (tribe)|Chumash]], [[Tongva people|Tongva]], and [[Tataviam people|Tataviam]]. The pomes also can be made into a jelly. Native Americans also made a tea from the leaves as a stomach remedy. Most were dried and stored, then later cooked into porridge or pancakes. Later settlers added sugar to make [[custard]] and [[wine]].<ref>{{cite web |url=http://www.ethnoherbalist.com/southern-california-native-plants-medicinal/toyon-berries/ |title=Ethnobotany of southern California native plants: Toyon (Heteromeles arbutifolia) |website= EthnoHerbalist}}</ref>

==Toxicity==
Toyon pomes are [[acid]]ic and astringent, and contain a small amount of cyanogenic [[glycoside]]s, which break down into [[hydrocyanic acid]] on digestion. This is removed by mild cooking.{{Citation needed|date=January 2010}}

Some pomes, though mealy, astringent and acid when raw, were eaten fresh, or mashed into water to make a beverage.

==Legislation==
In the 1920s,{{citation needed|date=November 2016}} collecting toyon branches for [[Christmas]] became so popular in Los Angeles that the State of [[California]] passed a law forbidding collecting on public land or on any land not owned by the person picking any plant without the landowner's written permission (CA Penal Code § 384a).<ref>{{cite news|title=California Holly Adds Color to Trail Up Mt. Hollywood|first=JOHN |last=McKINNEY|work=Los Angeles Times |date=December 6, 1986|page=12|url=http://articles.latimes.com/1986-12-06/news/vw-1626_1_hollywood-trail}}</ref><ref>[http://www.leginfo.ca.gov/cgi-bin/displaycode?section=pen&group=00001-01000&file=369a-402c California Penal Code Section 384a] {{webarchive|url=https://web.archive.org/web/20090627103906/http://www.leginfo.ca.gov/cgi-bin/displaycode?section=pen&group=00001-01000&file=369a-402c |date=2009-06-27 }}</ref>

Toyon was adopted as the official native plant of the city of Los Angeles by the LA City Council on April 17, 2012.<ref>{{cite web|title=Item No. (28)|url=http://ens.lacity.org/clk/oldactions/clkcouncilactions276843_04272012.pdf|work=Journal/Council Proceedings|publisher=LA City Council|accessdate=23 November 2013}}</ref>

==Taxonomy==
The genera ''[[Photinia]]'', ''[[Aronia]]'', ''[[Pourthiaea]]'', and ''[[Stranvaesia]]'' have historically been variously combined by different taxonomists.<ref name=Nesom>{{Citation |last1=Nesom |first1=G.L. |last2=Gandhi |first2=K. |author2-link=Kanchi Gandhi |date=2009 |title=(1884–1885) Proposals to conserve the names ''Photinia'', with a conserved type, and ''Heteromeles'' (Rosaceae) |journal=Taxon |volume=58 |issue=1 |pages=310–311 |lastauthoramp=yes }}</ref> The genus ''Heteromeles'' as originally published by [[Max Joseph Roemer]] was [[Monotypic taxon|monospecific]], including ''Photinia arbutifolia'' Lindl. (1820), as ''H. arbutifolia'' (Lindl.) M. Roem, but the name was [[Nomen illegitimum|illegitimate]] (superfluous) because it included the [[type (biology)|type]] of the genus ''Photinia''.<ref name=Nesom/> This has since been corrected by [[conserved name|conservation]],<ref>{{citation |url=http://botany.si.edu/references/codes/props/index.cfm |title=International Code of Nomenclature for algae, fungi, and plants: Appendices II-VIII (Appendix III)}}</ref> and the name is therefore often written as ''Heteromeles'' M. Roem. ''nom. cons.'' (1847).

==See also==
*[[California native plants]]

==References==
{{Reflist}}

==External links==
*{{GRIN | ''Heteromeles salicifolia'' | 414959}}
*[http://globaltwitcher.auderis.se/artspec_information.asp?thingid=84109C. Michael Hogan (2008) ''Toyon: Heteromeles arbutifolia'', GlobalTwitcher.com, ed. N. Stromberg]{{dead link|date=November 2016}}
*[http://www.bahiker.com/plantpages/toyon.html Photos of Toyon in flower and fruit]
*[http://naeb.brit.org/uses/species/1860/ University of Michigan: Dearborn — Native American Ethnobotany (''Heteromeles arbutifolia'')]
*[http://clkrep.lacity.org/onlinedocs/2012/12-0353_CA_04-17-12.pdf Los Angeles City Clerk - Council Files: Toyon]
{{Commons category|Heteromeles arbutifolia}}
*[http://www.calflora.org/cgi-bin/species_query.cgi?where-calrecnum=4140 CalFlora database: ''Heteromeles arbutifolia'']
*[http://www.livingwild.org/winter/toyon/ The Living Wild Project: Toyon]
*[http://plants.usda.gov/java/nameSearch?mode=symbol&keywordquery=HEAR5 USDA Plants Profile for ''Heteromeles arbutifolia'' (toyon)]
*[http://calphotos.berkeley.edu/cgi/img_query?query_src=photos_index&where-taxon=Heteromeles+arbutifolia ''Heteromeles arbutifolia'' — U.C. Photo gallery]

{{Taxonbar|from=Q2655434}}

[[Category:Maleae]]
[[Category:Monotypic Rosaceae genera]]
[[Category:Flora of California]]
[[Category:Flora of Baja California]]
[[Category:Natural history of the California chaparral and woodlands]]
[[Category:Flora of the Sierra Nevada (U.S.)]]
[[Category:Natural history of the California Coast Ranges]]
[[Category:Natural history of the Peninsular Ranges]]
[[Category:Natural history of the San Francisco Bay Area]]
[[Category:Natural history of the Santa Monica Mountains]]
[[Category:Natural history of the Transverse Ranges]]
[[Category:Plants used in traditional Native American medicine]]
[[Category:Bird food plants]]
[[Category:Garden plants of North America]]
[[Category:Drought-tolerant plants]]

Marseilleviridae

2018-03-23T01:41:05Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q6773274}} (1 sig. taxon ID); WP:GenFixes, using AWB

{{Use dmy dates|date=April 2017}}
{{taxobox
| virus_group = i
|ordo =
| familia = '''''Marseilleviridae'''''
| subdivision_ranks = Genera
| subdivision =
*''[[Marseillevirus]]''
}}

'''''Marseilleviridae''''' is a [[Family (biology)|family]] of [[viruses]] first named in 2012.<ref name=pmid23188494>{{cite journal |doi=10.1007/s00705-012-1537-y |pmid=23188494 |title='Marseilleviridae', a new family of giant viruses infecting amoebae |journal=Archives of Virology |volume=158 |issue=4 |pages=915–20 |year=2012 |last1=Colson |first1=Philippe |last2=Pagnier |first2=Isabelle |last3=Yoosuf |first3=Niyaz |last4=Fournous |first4=Ghislain |last5=La Scola |first5=Bernard |last6=Raoult |first6=Didier }}</ref> The [[genome]]s of these viruses are [[double-stranded DNA]]. [[Amoeba]] are often [[Host (biology)|hosts]], but there is evidence that they are found in [[humans]] as well.<ref name=pmid25218687>{{cite journal |doi=10.1016/j.micpath.2014.09.005 |pmid=25218687 |title=Looking at protists as a source of pathogenic viruses |journal=Microbial Pathogenesis |volume=77 |pages=131–5 |year=2014 |last1=La Scola |first1=Bernard }}</ref><ref name=pmid23664726>{{cite journal |doi=10.1016/j.jcv.2013.03.018 |pmid=23664726 |title=Evidence of the megavirome in humans |journal=Journal of Clinical Virology |volume=57 |issue=3 |pages=191–200 |year=2013 |last1=Colson |first1=Philippe |last2=Fancello |first2=Laura |last3=Gimenez |first3=Gregory |last4=Armougom |first4=Fabrice |last5=Desnues |first5=Christelle |last6=Fournous |first6=Ghislain |last7=Yoosuf |first7=Niyaz |last8=Million |first8=Matthieu |last9=La Scola |first9=Bernard |last10=Raoult |first10=Didier }}</ref><ref name=pmid23821720>{{cite journal |doi=10.1093/infdis/jit292 |pmid=23821720 |title=Marseillevirus-Like Virus Recovered from Blood Donated by Asymptomatic Humans |journal=The Journal of Infectious Diseases |volume=208 |issue=7 |pages=1042–50 |year=2013 |last1=Popgeorgiev |first1=Nikolay |last2=Boyer |first2=Mickaël |last3=Fancello |first3=Laura |last4=Monteil |first4=Sonia |last5=Robert |first5=Catherine |last6=Rivet |first6=Romain |last7=Nappez |first7=Claude |last8=Azza |first8=Said |last9=Chiaroni |first9=Jacques |last10=Raoult |first10=Didier |last11=Desnues |first11=Christelle }}</ref><ref name=pmid27502174>{{cite journal |doi=10.1016/S1473-3099(16)30051-2 |pmid=27502174 |title=Marseillevirus in lymphoma: A giant in the lymph node |journal=The Lancet Infectious Diseases |volume=16 |issue=10 |pages=e225–e234 |year=2016 |last1=Aherfi |first1=Sarah |last2=Colson |first2=Philippe |last3=Audoly |first3=Gilles |last4=Nappez |first4=Claude |last5=Xerri |first5=Luc |last6=Valensi |first6=Audrey |last7=Million |first7=Matthieu |last8=Lepidi |first8=Hubert |last9=Costello |first9=Regis |last10=Raoult |first10=Didier }}</ref> As of 2016, the [[International Committee on Taxonomy of Viruses]] recognize four species in this family, divided among 2 genera.<ref name=ViralZone>{{cite web |title=Viral Zone |url=http://viralzone.expasy.org/all_by_species/4740.html |publisher=ExPASy |accessdate=12 August 2015 }}</ref><ref name=ICTV>{{cite web |publisher=ICTV |title=Virus Taxonomy: 2016 Release |url=http://ictvonline.org/virusTaxonomy.asp |accessdate=6 July 2017 }}</ref> It is a member of the [[nucleocytoplasmic large DNA viruses]] clade.

==Taxonomy==
<big>'''Group: dsDNA'''</big>
{{Collapsible list|title= <big>Order: Unassigned</big>
|1={{Collapsible list| framestyle=border:none; padding:1.0em;|title=Family: Marseilleviridae
|1={{hidden begin|title=Genus: [[Marseillevirus]]}}
*'''''[[Marseillevirus marseillevirus]]'''''
*[[Senegalvirus marseillevirus]]
{{hidden end}}
|2={{hidden begin|title=Genus: Unassigned}}
*[[Lausannevirus]]
*[[Tunisvirus]]
{{hidden end}}
}}
}}
<ref name=ICTV />

===Related Viruses===
[[File:MelVfig2j.jpg|thumb|Images of cryo-frozen ''Marseilleviridae'' particles (left and center) and enlarged diagram of structure near a vertex. Black arrows indicate Large Dense Bodies. White arrows indicate lipid bilayer.]]
Additional species have since been recognized.<ref name=pmid23188494/> The first member of this family recognized has been named ''Acanthamoeba polyphaga marseillevirus''. A second member is ''Acanthamoeba castellanii lausannevirus''. Two additional viruses have been isolated but have yet to be named. Another member of this family has been isolated from blood donors.<ref name=pmid23821720/> An isolate from insects—''Insectomime'' virus—has also been reported.<ref name=pmid24157885>{{cite journal |doi=10.1159/000354560 |pmid=24157885 |title=First Isolation of a Marseillevirus in the Diptera Syrphidae ''Eristalis'' tenax |journal=Intervirology |volume=56 |issue=6 |pages=386–94 |year=2013 |last1=Boughalmi |first1=Mondher |last2=Pagnier |first2=Isabelle |last3=Aherfi |first3=Sarah |last4=Colson |first4=Philippe |last5=Raoult |first5=Didier |last6=La Scola |first6=Bernard }}</ref>

The viruses appear to fall into at least 3 lineages: (1) ''Marseillevirus'' and ''Cannes8virus'' (2) ''Insectomime'' and ''Tunisvirus'' and (3) ''Lausannevirus''. A sixth potential member of this family—''Melbournevirus''—appears to be related to the ''Marseillevirus''/''Cannes8virus'' clade.<ref name=pmid25275139>{{cite journal |doi=10.1128/JVI.02414-14 |pmid=25275139 |pmc=4249118 |title=Genome Analysis of the First Marseilleviridae Representative from Australia Indicates that Most of Its Genes Contribute to Virus Fitness |journal=Journal of Virology |volume=88 |issue=24 |pages=14340–9 |year=2014 |last1=Doutre |first1=G |last2=Philippe |first2=N |last3=Abergel |first3=C |last4=Claverie |first4=J.-M }}</ref>

A seventh virus—Brazilian Marseillevirus—has been reported.<ref name=pmid26978387>{{cite journal |doi=10.3390/v8030076 |pmid=26978387 |pmc=4810266 |title=A Brazilian Marseillevirus is the Founding Member of a Lineage in Family Marseilleviridae |journal=Viruses |volume=8 |issue=3 |pages=76 |year=2016 |last1=Dornas |first1=Fábio |last2=Assis |first2=Felipe |last3=Aherfi |first3=Sarah |last4=Arantes |first4=Thalita |last5=Abrahão |first5=Jônatas |last6=Colson |first6=Philippe |last7=La Scola |first7=Bernard }}</ref> This virus appears to belong to a fourth lineage of virus in this family.

Another virus—Tokyovirus—has also been reported.<ref name=pmid27284144>{{cite journal |doi=10.1128/genomeA.00429-16 |pmid=27284144 |pmc=4901213 |title=Draft Genome Sequence of ''Tokyovirus'', a Member of the Family ''Marseilleviridae'' Isolated from the Arakawa River of Tokyo, Japan |journal=Genome Announcements |volume=4 |issue=3 |pages=e00429–16 |year=2016 |last1=Takemura |first1=Masaharu }}</ref>

Another member of this family is Kurlavirus.<ref name=pmid28685284>{{cite journal |doi=10.1007/s00705-017-3469-z |pmid=28685284 |title=Complete genome sequence of Kurlavirus, a novel member of the family Marseilleviridae isolated in Mumbai, India |journal=Archives of Virology |volume=162 |issue=10 |pages=3243 |year=2017 |last1=Chatterjee |first1=Anirvan |last2=Kondabagil |first2=Kiran }}</ref>

==Structure==
Viruses in Marseilleviridae have icosahedral geometries. The diameter is around 250 nm. Genomes are circular, around 372kb in length. The genome has 457 open reading frames.<ref name=ViralZone/>

{| class="wikitable sortable" style="text-align:center"
|-
! Genus !! Structure || Symmetry !! Capsid !! Genomic arrangement !! Genomic segmentation
|-
|Unassigned||Head-Tail||T=16||Non-enveloped||Linear||Monopartite
|-
|Marseillevirus||Icosahedral||||||Circular||
|}

==Life cycle==
Dna templated transcription is the method of transcription. Amoeba serve as the natural host.<ref name=ViralZone/>

{| class="wikitable sortable" style="text-align:center"
|-
! Genus !! Host details !! Tissue tropism !! Entry details !! Release details !! Replication site !! Assembly site !! Transmission
|-
|Marseillevirus||Amoeba||None||Fusion||Lysis||Cytoplasm||Cytoplasm||Diffusion in Water
|}

==Genomics==

A promoter sequence—AAATATTT—has been found associated with 55% of the identified genes in this virus.<ref name=pmid28794030>{{cite journal |doi=10.1128/JVI.01088-17 |pmid=28794030 |pmc=5640848 |title=The Investigation of Promoter Sequences of Marseilleviruses Highlights a Remarkable Abundance of the AAATATTT Motif in Intergenic Regions |journal=Journal of Virology |volume=91 |issue=21 |pages=e01088–17 |year=2017 |last1=Oliveira |first1=Graziele Pereira |last2=Lima |first2=Maurício Teixeira |last3=Arantes |first3=Thalita Souza |last4=Assis |first4=Felipe Lopes |last5=Rodrigues |first5=Rodrigo Araújo Lima |last6=Da Fonseca |first6=Flávio Guimarães |last7=Bonjardim |first7=Cláudio Antônio |last8=Kroon |first8=Erna Geessien |last9=Colson |first9=Philippe |last10=La Scola |first10=Bernard |last11=Abrahão |first11=Jônatas Santos }}</ref> Most of these sequences occur in multiple copies.

==History==

One of the first members of this family was described in 2009.<ref name=pmid20007369>{{cite journal |doi=10.1073/pnas.0911354106 |pmid=20007369 |pmc=2799887 |title=Giant Marseillevirus highlights the role of amoebae as a melting pot in emergence of chimeric microorganisms |journal=Proceedings of the National Academy of Sciences |volume=106 |issue=51 |pages=21848–53 |year=2009 |last1=Boyer |first1=M |last2=Yutin |first2=N |last3=Pagnier |first3=I |last4=Barrassi |first4=L |last5=Fournous |first5=G |last6=Espinosa |first6=L |last7=Robert |first7=C |last8=Azza |first8=S |last9=Sun |first9=S |last10=Rossmann |first10=M. G |last11=Suzan-Monti |first11=M |last12=La Scola |first12=B |last13=Koonin |first13=E. V |last14=Raoult |first14=D |bibcode=2009PNAS..10621848B }}</ref> Other members described around then (2007) and since then have been documented.<ref name=pmid25104553>{{cite journal |doi=10.1016/j.virol.2014.07.014 |pmid=25104553 |title=The expanding family Marseilleviridae |journal=Virology |volume=466-467 |pages=27–37 |year=2014 |last1=Aherfi |first1=Sarah |last2=La Scola |first2=Bernard |last3=Pagnier |first3=Isabelle |last4=Raoult |first4=Didier |last5=Colson |first5=Philippe }}</ref>

==References==
{{Reflist|2}}

==External links==
* [http://viralzone.expasy.org/all_by_species/4740.html '''Viralzone''': Marseilleviridae]
* [http://ictvonline.org/virusTaxonomy.asp '''ICTV''']
{{Baltimore classification}}

{{Taxonbar|from=Q6773274}}

[[Category:Marseilleviridae| ]]
[[Category:Nucleocytoplasmic large DNA viruses]]
[[Category:Virus families]]

Obama (Gattung)

2018-03-23T01:34:36Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q18355807}} (3 sig. taxon IDs); WP:GenFix using AWB

{{Italic title}}
{{Taxobox
| name = ''Obama''
| image = Unknown Brown Geoplanidae from France JL57 - white background.JPG
| image_caption = ''Obama nungara''
| regnum = [[Animal]]ia
| phylum = [[Platyhelminthes]]
| classis = [[Rhabditophora]]
| subclassis =
| superordo =
| ordo = [[Tricladida]]
| subordo = [[Continenticola]]
| infraordo =
| unranked_familia =
| superfamilia =
| familia = [[Geoplanidae]]
| subfamilia = [[Geoplaninae]]
| genus = '''''Obama'''''
| genus_authority = Carbayo et al., 2013
| type_species = ''[[Obama fryi|Geoplana fryi]]''
| type_species_authority = [[Ludwig von Graff|Graff]], 1899
| synonyms =
}}

'''''Obama''''' is a [[genus]] of [[land planarian]]s from [[South America]]. It contains several species adapted to human-disturbed environments,<ref name="Boll2016">{{cite journal | last1 = Boll | first1 = Piter Kehoma | last2=Leal-Zanchet|first2=Ana Maria|title=Preference for different prey allows the coexistence of several land planarians in areas of the Atlantic Forest | journal = Zoology | year = 2016 | issn = 0944-2006 | doi=10.1016/j.zool.2016.04.002 | volume=119 | pages=162–168}}</ref> including the only invasive land planarian native to the [[Neotropical ecozone]], ''[[Obama nungara]]'', which has been accidentally introduced in [[Europe]].<ref name="Carbayo2016a">{{cite journal | last1 = Carbayo | first1 = Fernando | last2 = Álvarez-Presas | first2 = Marta | last3 = Jones | first3 = Hugh D. | last4 = Riutort | first4 = Marta | title = The true identity of ''Obama'' (Platyhelminthes: Geoplanidae) flatworm spreading across Europe|journal=Zoological Journal of the Linnean Society | volume = 177 | issue = 1 | year = 2016 | pages = 5–28 | issn = 0024-4082 | doi = 10.1111/zoj.12358}}</ref>

== Description ==
The genus ''Obama'' is characterized by having a leaf-shaped body. Most species are about {{convert|10|cm|in}} long, but some may reach over {{convert|20|cm|in}}. The hundreds of eyes distributed along the body are of two types: monolobulated, which are simple and circular, and trilobulated, which have three lobes.<ref name="Carbayo2013">{{Cite journal | doi = 10.1111/zsc.12019| title = Molecular phylogeny of Geoplaninae (Platyhelminthes) challenges current classification: Proposal of taxonomic actions| journal = Zoologica Scripta| volume = 42| issue = 5| pages = 508–528| year = 2013| last1 = Carbayo | first1 = F. | last2 = Álvarez-Presas | first2 = M. | last3 = Olivares | first3 = C. U. T. | last4 = Marques | first4 = F. P. L. | last5 = Froehlich | first5 = E. X. M. | last6 = Riutort | first6 = M. }}</ref>

The [[reproductive system of planarians|copulatory apparatus]] of ''Obama'' has a protrusible penis occupying the entire male atrium or most of it. Morphologically the genus can be divided in two subgroups depending on the shape of the penis papilla, which may be symmetrical or asymmetrical. The group with an asymmetrical penis papilla includes 10 species (''O. anthropophilla'', ''O. carbayoi'', ''O. carrierei'', ''O. decidualis'', ''O. josefi'', ''O. ladislavii'', ''O. marmorata'', ''O. nungara'', ''O. otavioi'' and ''O. ruiva'') and seems to form a [[monophyletic]] clade within ''Obama''.<ref name="Carbayo2016">{{cite journal | last1 = Carbayo | first1 = Fernando | last2 = Francoy | first2 = Tiago M.| last3 = Giribet | first3 = Gonzalo | title = Non-destructive imaging to describe a new species of ''Obama'' land planarian (Platyhelminthes, Tricladida) | journal = Zoologica Scripta | year = 2016 | pages = n/a–n/a | issn=0300-3256 | doi = 10.1111/zsc.12175}}</ref> On the other hand, the group with a symmetrical penis papilla is [[paraphyletic]], indicating that this is the ancestral state within the genus.<ref name="Carbayo2016" />

== Etymology ==
The name ''Obama'' is formed by a composition of the [[Tupi language|Tupi]] words ''oba'' (leaf) and ''ma'' (animal), being a reference to the body shape of species in this genus.<ref name="Carbayo2013" />

== Species ==
{{commons category|Obama (Geoplanidae)|Obama (genus)}}
There are 38 species assigned to the genus ''Obama'':
{{div col|colwidth=30em}}
*''[[Obama apeva]]'' (Froehlich, 1959)
*''[[Obama applanata]]'' ([[Ludwig von Graff|Graff]], 1899)
*''[[Obama anthropophila]]'' Amaral, Leal-Zanchet & Carbayo, 2015
*''[[Obama argus]]'' ([[Ludwig von Graff|Graff]], 1899)
*''[[Obama assu]]'' (Froehlich, 1959)
*''[[Obama baptistae]]'' (Leal-Zanchet & Oliveira, 2012)
*''[[Obama braunsi]]'' ([[Ludwig von Graff|Graff]], 1899)
*''[[Obama burmeisteri]]'' (Schultze & Müller, 1857)
*''[[Obama carbayoi]]'' (Oliveira & Leal-Zanchet, 2012)
*''[[Obama carinata]]'' (Riester, 1938)
*''[[Obama carrierei]]'' ([[Ludwig von Graff|Graff]], 1897)
*''[[Obama catharina]]'' (Hyman, 1957)
*''[[Obama decidualis]]'' Amaral & Leal-Zanchet, 2015
*''[[Obama dictyonota]]'' (Riester, 1938)
*''[[Obama divae]]'' (Marcus, 1951)
*''[[Obama eudoxiae]]'' (Ogren & Kawakatsu, 1990)
*''[[Obama eudoximariae]]'' (Ogren & Kawakatsu, 1990)
*''[[Obama evelinae]]'' (Marcus, 1951)
*''[[Obama ferussaci]]'' ([[Ludwig von Graff|Graff]], 1897)
*''[[Obama ficki]]'' (Amaral & Leal-Zanchet, 2012)
*''[[Obama fryi]]'' ([[Ludwig von Graff|Graff]], 1899)
*''[[Obama glieschi]]'' (Froehlich, 1959)
*''[[Obama itatiayana]]'' (Schirch, 1929)
*''[[Obama josefi]]'' (Carbayo & Leal-Zanchet, 2001)
*''[[Obama ladislavii]]'' ([[Ludwig von Graff|Graff]], 1899)
*''[[Obama livia]]'' (E. M. Froehlich, 1955)
*''[[Obama maculipunctata]]'' Rossi, Amaral, Ribeiro, Cauduro, Fick, Valiati & Leal-Zanchet, 2015
*''[[Obama marmorata]]'' (Schultze & Müller, 1857)
*''[[Obama metzi]]'' ([[Ludwig von Graff|Graff]], 1899)
*''[[Obama nungara]]'' Carbayo, Álvarez-Presas, Jones & Riutort, 2016 <ref name="Carbayo2016a" />
*''[[Obama otavioi]]'' Carbayo, 2016
*''[[Obama poca]]'' (Froehlich, 1958)
*''[[Obama polyophthalma]]'' ([[Ludwig von Graff|Graff]], 1899)
*''[[Obama riesteri]]'' (Froehlich, 1955)
*''[[Obama rufiventris]]'' (Schultze & Müller, 1857)
*''[[Obama ruiva]]'' (E. M. Froehlich, 1972)
*''[[Obama schubarti]]'' (Froehlich, 1958)
*''[[Obama trigueira]]'' (E. M. Froehlich, 1955)
{{div col end}}

== Phylogeny ==
[[File:Obama three species.jpg|thumb|Three specimens of ''Obama'' of different species in a Petri dish. From top to bottom: ''O. ladislavii'', ''O. anthropophila'' and ''O. nungara'']]
The genus ''Obama'' was erected after a study of [[molecular phylogeny]] with the subfamily Geoplaninae revealed that the genus ''[[Geoplana]]'', originally containing more than a hundred species, was [[polyphyletic]].<ref name="Carbayo2013" /> One of the monophyletic clades revealed by the study was separate from ''Geoplana'' as the new genus ''Obama''. All species within the new genus share a similar morphology, including the leaf-shaped body, the presence of a permanent penis papilla, [[reproductive system of planarians|ovovitelline ducts]] entering the female atrium dorsally and dorsal eyes of two types: mono- and trilobulated.<ref name="Carbayo2013" />

The sister-group of ''Obama'' seems to be the genus ''[[Cratera (genus)|Cratera]]'', which has a very similar appearance but lacks trilobulated eyes.<ref name="Carbayo2013" /><ref name="Rossi2015">{{cite journal | last1 = Rossi | first1 = Ilana | last2 = Amaral | first2 = Silvana Vargas | last3 = Ribeiro | first3 = Giovana Gamino | last4 = Cauduro | first4 = Guilherme Pinto | last5 = Fick | first5 = Israel | last6 = Valiati | first6 = Victor Hugo | last7 = Leal-Zanchet | first7 = Ana Maria | title = Two new Geoplaninae species (Platyhelminthes: Continenticola) from Southern Brazil based on an integrative taxonomic approach | journal = Journal of Natural History | volume = 50 | issue = 13-14 | year = 2015 | pages = 787–815 | issn = 0022-2933 | doi = 10.1080/00222933.2015.1084057}}</ref>

The following phylogenetic tree shows the relationship of several species of ''Obama'' after several molecular studies:<ref name="Carbayo2016a" /><ref name="Carbayo2013" /><ref name="Carbayo2016" /><ref name="Rossi2015" />

{{clade| style=font-size:100%;line-height:80%
|1={{clade
|1={{clade
|1=''Obama baptistae''
|2={{clade
|1=''Obama braunsi''
|2={{clade
|1=''Obama apeva''
|2=''Obama ficki''
}}
}}
}}
|2={{clade
|1=''Obama fryi''
|2={{clade
|1={{clade
|1=''Obama eudoximariae''
|2={{clade
|1=''Obama carinata''
|2=''Obama itatiayana''
|3={{clade
|1=''Obama burmeisteri''
|2=''Obama maculipunctata''
|3={{clade
|1=''Obama applanata''
|2=''Obama argus'' sensu Graff, 1899
}}
}}
}}
}}
|2={{clade
|1=''Obama argus'' sensu Schirch, 1929
|2={{clade
|1=''Obama ladislavii''
|2={{clade
|1=''Obama nungara''
|2=''Obama octavioi''
|3={{clade
|1=''Obama josefi''
|2={{clade
|1=''Obama marmorata''
|2={{clade
|1=''Obama anthropophila''
|2=''Obama decidualis''
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}

== References ==
{{Reflist}}

{{Taxonbar|from=Q18355807}}

[[Category:Geoplanidae]]
[[Category:Rhabditophora genera]]

Enterobakteriophage PhiX174

2018-03-23T01:33:08Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q1063448}} (1 sig. taxon ID); WP:GenFixes, using AWB

{{Taxobox
|image = Bacteriophage_Phi_X_174_Electron_micrograph.gif
|image_caption = Electron micrograph of phage ΦX174
|virus_group = ii
|familia = ''[[Microviridae]]''
|genus = ''[[Microvirus]]''
|species = '''''Enterobacteria phage phiX174'''''
}}

[[File:phiX174.jpg|thumb|right|250 px| Structure of phage ΦX174 capsid]]
[[File:Genome map of the bacteriophage ΦX174 showing overlapping genes.svg|thumb|right|250px|Genome of the bacteriophage ΦX174 showing its 11 genes
<ref name="NCBI NC_001422">Enterobacteria phage phiX174 ''sensu lato'', complete genome. [https://www.ncbi.nlm.nih.gov/nuccore/NC_001422 "Complete genome: accession NC_001422"], ''[[National Center for Biotechnology Information]]''. Retrieved on 30 January 2016.</ref>]]

The '''phi X 174''' (or '''ΦX174''') [[bacteriophage]] is a single-stranded DNA ([[SsDNA virus|ssDNA]]) virus and the first DNA-based [[genome]] to be sequenced. This work was completed by [[Fred Sanger]] and his team in 1977.<ref>{{cite journal |doi=10.1038/265687a0 |title=Nucleotide sequence of bacteriophage ΦX174 DNA |year=1977 |last1=Sanger |first1=F. |last2=Air |first2=G. M. |last3=Barrell |first3=B. G. |last4=Brown |first4=N. L. |last5=Coulson |first5=A. R. |last6=Fiddes |first6=J. C. |last7=Hutchison |first7=C. A. |last8=Slocombe |first8=P. M. |last9=Smith |first9=M. |journal=Nature |volume=265 |issue=5596 |pages=687–95 |pmid=870828|bibcode = 1977Natur.265..687S }}</ref> In 1962, [[Walter Fiers]] and Robert Sinsheimer had already demonstrated the physical, covalently closed circularity of ΦX174 DNA.<ref>{{cite journal |doi=10.1016/S0022-2836(62)80031-X |title=The structure of the DNA of bacteriophage ΦX174 |year=1962 |last1=Fiers |first1=Walter |last2=Sinsheimer |first2=Robert L. |journal=Journal of Molecular Biology |volume=5 |issue=4 |pages=424}}</ref> Nobel prize winner [[Arthur Kornberg]] used ΦX174 as a model to first prove that DNA synthesized in a test tube by purified enzymes could produce all the features of a natural virus, ushering in the age of [[synthetic biology]].<ref>National Library of Medicine Profiles in Science. The Arthur Kornberg Papers. "Creating Life in the Test Tube," 1959-1970. [http://profiles.nlm.nih.gov/ps/retrieve/Narrative/WH/p-nid/209/p-docs/true link]{{psc|date=February 2013}}</ref><ref>{{cite journal |first1=Mehran |last1=Goulian |first2=Arthur |last2=Kornberg |first3=Robert L. |last3=Sinsheimer |title=Enzymatic Synthesis of DNA, XXIV. Synthesis of Infectious Phage ΦX174 DNA |doi=10.1073/pnas.58.6.2321 |jstor=58720 |bibcode=1967PNAS...58.2321G |pmc=223838 |year=1967 |journal=Proceedings of the National Academy of Sciences |volume=58 |issue=6 |pages=2321–2328 |pmid=4873588}}</ref> In 2003, it was reported by [[Craig Venter|Craig Venter's]] group that the genome of ΦX174 was the first to be completely assembled ''in vitro'' from synthesized oligonucleotides.<ref>{{cite journal |first1=Hamilton O. |last1=Smith |first2=Clyde A. |last2=Hutchison |first3=Cynthia |last3=Pfannkoch |first4=J. Craig |last4=Venter |doi=10.1073/pnas.2237126100 |title=Generating a Synthetic Genome by Whole Genome Assembly: ΦX174 Bacteriophage from Synthetic Oligonucleotides |year=2003 |journal=Proceedings of the National Academy of Sciences |volume=100 |issue=26 |pages=15440–5 |bibcode=2003PNAS..10015440S |jstor=3149024 |pmc=307586 |pmid=14657399}}</ref> The ΦX174 virus particle has also been successfully assembled ''in vitro''.<ref>{{cite journal |doi=10.1016/j.jmb.2011.07.070 |title=In Vitro Assembly of the ΦX174 Procapsid from External Scaffolding Protein Oligomers and Early Pentameric Assembly Intermediates |year=2011 |last1=Cherwa |first1=James E. |last2=Organtini |first2=Lindsey J. |last3=Ashley |first3=Robert E. |last4=Hafenstein |first4=Susan L. |last5=Fane |first5=Bentley A. |journal=Journal of Molecular Biology |volume=412 |issue=3 |pages=387–96 |pmid=21840317}}</ref> Recently, it was shown how its highly [[overlapping gene|overlapping genome]] can be fully decompressed and still remain functional.<ref>{{cite journal |doi=10.1016/j.virol.2012.09.020 |title=A fully decompressed synthetic bacteriophage ΦX174 genome assembled and archived in yeast |year=2012 |last1=Jaschke |first1=Paul R. |last2=Lieberman |first2=Erica K. |last3=Rodriguez |first3=Jon |last4=Sierra |first4=Adrian |last5=Endy |first5=Drew |journal=Virology |volume=434 |issue=2 |pages=278–84 |pmid=23079106}}</ref>

==Virology==
This [[bacteriophage]] has a [+] circular single-stranded [[DNA]] genome of 5386 [[nucleotide]]s encoding 11 [[protein]]s.<ref name="NCBI NC_001422" /> Of these 11 genes, only 8 are essential to viral [[morphogenesis]]. The [[GC-content]] is 44% and 95% of nucleotides belong to coding genes.

{| class="wikitable"
|-
! Protein !! Copies !! Function<ref>{{cite book|last1=Fane|first1=Bentley A.|last2=Brentlinger|first2=Karie L.|last3=Burch|first3=April D.|last4=Chen|first4=Min|last5=Hafenstein|first5=Susan|last6=Moore|first6=Erica|last7=Novak|first7=Christopher R.|last8=Uchiyama|first8=Asako|editor1-last=Calender|editor1-first=Richard|title=The Bacteriophages|date=2006|publisher=Oxford Univ. Press|location=New York|isbn=978-0195148503|page=130|edition=2nd|chapter=ɸX174 et al., the ''Microviridae''}}</ref>
|-
| A || -- || Nicks RF DNA to initiate [[rolling circle replication]]; ligates ends of linear phage DNA to form single-stranded circular DNA
|-
| A* || -- || Inhibits host cell DNA replication; blocks superinfecting phage; not essential
|-
| B || 60 in [[procapsid]] || Internal scaffolding protein involved in procapsid assembly
|-
| C || -- || DNA packaging
|-
| D || 240 in procapsid || External scaffolding protein involved in procapsid assembly
|-
| E || -- || Host cell [[lysis]]
|-
| F || 60 in virion || Major capsid protein
|-
| G || 60 in virion || Major spike protein
|-
| H || 12 in virion || DNA pilot protein (or minor spike protein)
|-
| J || 60 in virion || Binds to new single-stranded phage DNA; accompanies phage DNA into procapsid
|-
| K || -- || Optimizes burst size; not essential
|}

Infection begins when G protein binds to [[lipopolysaccharide]]s on the bacterial host cell surface. H protein (or the DNA Pilot Protein) pilots the viral genome through the bacterial membrane of ''[[E.coli]]'' bacteria (Jazwinski ''et al.'' 1975) most likely via a predicted N-terminal [[transmembrane domain]] helix (Tusnady and Simon, 2001). However, it has become apparent that H protein is a multifunctional protein (Cherwa, Young and Fane, 2011). This is the only viral [[capsid]] protein of ΦX174 to lack a crystal structure for a couple of reasons. It has low aromatic content and high [[glycine]] content, making the protein structure very flexible and in addition, individual hydrogen atoms (the R group for glycines) are difficult to detect in protein crystallography. Additionally, H protein induces [[lysis]] of the bacterial host at high concentrations as the predicted N-terminal transmembrane helix easily pokes holes through the bacterial wall. By [[bioinformatics]], this protein contains four predicted [[coiled-coil]] domains which has a significant homology to known transcription factors. Additionally, it was determined by Ruboyianes ''et al.'' (2009) that ''de novo'' H protein was required for optimal synthesis of other viral proteins. Mutations in H protein that prevent viral incorporation, can be overcome when excess amounts of protein B, the internal scaffolding protein, are supplied.

The DNA is ejected through a hydrophilic channel at the 5-fold vertex (McKenna ''et al.'' 1992). It is understood that H protein resides in this area but experimental evidence has not verified its exact location. Once inside the host bacterium, replication of the [+] ssDNA genome proceeds via negative sense DNA]] intermediate. This is done as the phage genome supercoils and the secondary structure formed by such supercoiling attracts a [[primosome]] protein complex. This translocates once around the genome and synthesizes a [-]ssDNA from the positive original genome. [+]ssDNA genomes to package into viruses are created from this by a rolling circle mechanism. This is the mechanism by which the double stranded supercoiled genome is nicked on the negative strand by a virus-encoded A protein, also attracting a bacterial [[DNA polymerase]] (DNAP) to the site of cleavage. DNAP will use the negative strand as a template to make positive sense DNA. As it translocates around the genome it displaces the outer strand of already-synthesised DNA, which is immediately coated by [[SSBP]] proteins. The A protein will cleave the complete genome every time it recognises the origin sequence.

As D protein is the most abundant gene transcript, it is the most protein in the viral procapsid. Similarly, gene transcripts for F, J, and G are more abundant than for H as the [[stoichiometry]] for these structural proteins is 5:5:5:1. The primosomes are protein complexes which attach/bind the enzyme [[helicase]] on the template. Primosomes gives RNA primers for DNA synthesis to strands.

==Notes==
Phi X is regularly used as a [[Scientific control#Positive and negative control|positive control]] in [[DNA sequencing]] due to its relatively small genome size in comparison to other organisms, its relatively balanced nucleotide content — about 23% G, 22% C, 24% A, and 31% T, i.e., 45% G+C and 55% A+T, see the accession NC_001422.1<ref name="NCBI NC_001422"/> for its 5,386 nucleotide long sequence —, and the extensive work that has been done on it.

==See also==
{{Portal|Viruses}}
* [[Bacteriophage MS2]]

==References==
{{Reflist|2}}


==External links==

* {{cite web |url=http://www.rcsb.org/pdb/101/motm.do?momID=2 |title=Bacteriophage phiX174 |date=February 2000 |work=Molecule of the Month |first=David |last=Goodsell |publisher=RCSB-PDB}}

{{Taxonbar|from=Q1063448}}

[[Category:Microviridae]]

Escherichia-Phage T7

2018-03-23T01:32:20Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q7669119}} (2 sig. taxon IDs); WP:GenFixes using AWB

{{copypaste|url=https://books.google.co.jp/books?id=uBCzZSJqdWQC&pg=PA226&lpg=PA226&dq=d’Herelle+T7&source=bl&ots=BCOlDJTdRN&sig=33cr-Fab_K2sgpGDKUTK67q5Qpo&hl=ja&sa=X&ved=0ahUKEwjtt43nkOrWAhVGgbwKHXatDXYQ6AEISTAI#v=onepage&q=d’Herelle%20T7&f=false|date=October 2017}}
{{Taxobox
| name = ''Enterobacteria phage T7''
| virus_group = i
| ordo = ''[[Caudovirales]]''
| familia = ''[[Podoviridae]]''
| genus = ''[[T7likevirus|T7-like viruses]]''
| species = ''T7 phage''
}}

'''Bacteriophage T7''' (or the '''T7 phage''') is a [[bacteriophage]], a virus that infects susceptible [[bacteria]]l cells, that is composed of DNA and infects most strains of ''[[Escherichia coli]]''. Bacteriophage T7 has a lytic life cycle and several properties that make it an ideal phage for experimentation.

==Discovery==
Bacteriophage T7 was identified in 1945 as a member of the seven Type (“T”) phages that grow lytically on ''Escherichia coli'' B,<ref name="Demerec1945">{{Cite journal
| last1 = Demerec | first1 = M.
| last2 = Fano | first2 = U.
| title = Bacteriophage-Resistant Mutants in Escherichia Coli
| journal = Genetics
| volume = 30
| issue = 2
| pages = 119–136
| year = 1945
| pmid = 17247150
| pmc = 1209279
}}</ref> although it is probably identical to phage δ, used earlier by Delbrück. A close relative of T7 was likely studied by d’Herelle in the 1920s.<ref name="d'Herelle1926">d’Herelle, F. (1926). The Bacteriophage and Its Behavior. Baltimore, MD: Williams & Wilkins</ref>

==Hosts==
T7 grows on rough strains of ''E. coli'' (i.e. those without full-length [[Lipopolysaccharide|O-antigen]] polysaccharide on their surface) and some other enteric bacteria, but close relatives also infect smooth and even capsulated strains.<ref name="Molineux2006">Molineux, I. J. (2006). Chapter 20: The T7 group. In: The Bacteriophages (R. Calendar, ed.), pp. 277. Oxford University Press, Oxford.</ref> It infects ''[[Escherichia coli O157:H7|E. coli O157:H7]]'',{{Citation needed|date=May 2015}} a strain of ''E. coli'' that can cause [[foodborne illness]]).

==Virion structure==
The virus has complex structural symmetry, with a [[capsid]] of the phage that is icosahedral with an inner diameter of 55 [[Nanometer|nm]] and a tail 19 nm in diameter and 28.5 nm long attached to the capsid.

==Genome==
The genome of phage T7<ref>{{cite web|title=Genome of bacteriophage T7|url=https://www.ncbi.nlm.nih.gov/nuccore/V01146.1}}</ref> was among the first completely sequenced genomes and was published in 1983.<ref>{{cite journal | pmid = 6864790 | doi = 10.1016/S0022-2836(83)80282-4 | url = http://www.sciencedirect.com/science/article/pii/S0022283683802824 | year = 1983 | author1 = Dunn | first1 = J. J. | title = Complete nucleotide sequence of bacteriophage T7 DNA and the locations of T7 genetic elements | journal = Journal of Molecular Biology | volume = 166 | issue = 4 | pages = 477–535 | last2 = Studier | first2 = F. W. }}</ref> The head of the phage particle contains the roughly 40 [[Base pair#Length measurements|kbp]] [[dsDNA]] genome which encodes 55 proteins.<ref>{{cite web|title=Uniprot: reference proteome of bacteriophage T7|url=http://www.uniprot.org/uniprot/?query=organism:10760+keyword:1185}}</ref>

[[File:T7 phage genome.png|middle|middle|500 px|Schematic view of the phage T7 genome. Boxes are genes, numbers are gene numbers. Colors indicate functional groups as shown. White boxes are genes of unknown function or without annotation. Modified after <ref name="Häuser2012">{{Cite book
| pmid = 22748812
| year = 2012
| author1 = Häuser
| first1 = R
| title = Bacteriophage protein-protein interactions
| volume = 83
| pages = 219–98
| last2 = Blasche
| first2 = S
| last3 = Dokland
| first3 = T
| last4 = Haggård-Ljungquist
| first4 = E
| last5 = von Brunn
| first5 = A
| last6 = Salas
| first6 = M
| last7 = Casjens
| first7 = S
| last8 = Molineux
| first8 = I
| last9 = Uetz
| first9 = P
| doi = 10.1016/B978-0-12-394438-2.00006-2
| chapter = Bacteriophage Protein–Protein Interactions
| series = Advances in Virus Research
| isbn = 9780123944382
| pmc = 3461333
}}</ref> ]]

==Life cycle==
T7 has a short life cycle of 17 min at 37˚C, i.e. the time from infection to the lysis of the host cell when new phage are released. Due to the short latent period most physiological studies are being conducted at 30˚C where infected cells lyse after 30 min. However, high fitness-strains of T7 have been isolated with a latent period of only ~11 min at 37˚C growing under optimal conditions in rich media results. This adapted phage can undergo an effective expansion of its population by more than 1013 in one hour of growth.<ref name="Heinema2007">{{Cite journal
| last1 = Heineman | first1 = R. H.
| last2 = Bull | first2 = J. J.
| doi = 10.1111/j.1558-5646.2007.00132.x
| title = Testing Optimality with Experimental Evolution: Lysis Time in a Bacteriophage
| journal = Evolution
| volume = 61
| issue = 7
| pages = 1695–1709
| year = 2007
| pmid = 17598749
| pmc =1974807
}}</ref>

===Infection of host bacteria===
The T7 phage recognizes receptors on the surface of ''E.coli'' cells. It adheres to the cell surface by the tail fibers. In some strains of T7 the tail fibers are replaced with tailspikes with enzymatic activity that degrades the O- or K-antigens on the cell surface. The adsorbtion and penetration process use such [[lysozyme]]s to create an opening within the peptidoglycan layer of the bacterial cell wall allowing transfer of the viral DNA into the bacterium. The short, stubby tail of the T7-like phages is too short to span the cell envelope and, in order to eject the phage genome into the cell at the initiation of infection, virion proteins must first make a channel from the tip of the tail into the cell cytoplasm.<ref>{{cite journal | pmid = 20036409 | pmc = 2825023 | year = 2010 | author1 = Chang | first1 = C. Y. | title = Gp15 and gp16 cooperate in translocating bacteriophage T7 DNA into the infected cell | journal = Virology | volume = 398 | issue = 2 | pages = 176–86 | last2 = Kemp | first2 = P | last3 = Molineux | first3 = I. J. | doi = 10.1016/j.virol.2009.12.002 }}</ref>
The phage also injects proteins needed to begin replication of the viral genome and cleave the host genome.<ref name="microbemagazine.org">{{cite web| title=New Details about Bacteriophage T7-Host Interactions |url=http://www.microbemagazine.org/index.php/03-2010-home/1428-new-details-about-bacteriophage-t7-host-interactions}}</ref> T7 bacteriophage overcomes several of the host bacteria's defenses including the peptidoglycan cell wall and the CRISPR system.<ref name="microbemagazine.org"/> Once the T7 phage has inserted the viral genome the process of DNA replication of the host genome is halted and replication of viral genome begins.

Under optimal conditions, the T7 phage can complete the lytic process within 25 minutes, leading to the death of the ''E. coli'' host cell. At the time of lysis the virus can produce over 100 progeny.<ref name="microbemagazine.org"/>

===Components===
Gp5 (encoded by gene ''gp5'') is T7 phage's [[DNA polymerase]]. T7 polymerase uses [[E. coli]]'s endogenous [[thioredoxin]] as a sliding clamp during phage [[DNA replication]] (though thioredoxin normally has a different function). The sliding clamp functions to hold the polymerase onto the DNA, which increases the rate of synthesis; initiation, the process by which a polymerase binds to DNA, is time-consuming.{{citation needed|date=November 2016}}

==Applications in molecular biology==

The T7 promoter sequence is used extensively in [[molecular biology]] due to its extremely high affinity for [[T7 RNA polymerase]] and thus high level of expression.

T7 has been used as a model in [[synthetic biology]]. Chan ''et al.'' (2005) "[[refactored]]" the genome of T7, replacing approximately 12 [[Base pair#Length measurements|kbp]] of its genome with engineered DNA.<ref>{{cite journal |author=Leon Y. Chan, Sriram Kosuri, [[Drew Endy]] |year=2005 |title=Refactoring bacteriophage T7 |journal=Molecular Systems Biology |volume=1 |issue=1 |pages=2005.0018 |doi=10.1038/msb4100025 |url= |accessdate= |quote= |pmid=16729053 |pmc=1681472 }}</ref> The engineered DNA was designed to be easier to work with in a number of ways: individual functional elements were separated by [[restriction endonuclease]] sites for simple modification, and overlapping protein coding domains were separated and, where necessary, modified by single base pair [[silent mutations]].
T7 has been tested on human [[osteosarcoma]] to treat tumor cells. Chen ''et al''. (1998)"To test the utility of the system ''[[in vivo]]'' tumor ablation, a T7 cancer gene therapy [[plasmid]] vector, pT7T7/T7TK, was constructed. This nonviral vector contains a T7 autogene, T7T7, and a [[human herpes simplex virus]] [[thymidine kinase]] (HSV-TK) gene driven by a second T7 promoter (T7TK). When co-[[transfected]] with T7 [[RNA polymerase]] ([[T7 RNAP]]) into cultured human osteosarcoma 143B cells, about 10-20% of the cells were found to express HSV-TK, and more than 90% of the cells were killed in the presence of 1 microM [[ganciclovir]] (GCV) within 4 days after DNA transfection."<ref>{{cite journal |author=Xiaozhuo Chen, Li, Xiong,Aizicovici, Xie,Zhu,Sturtz, Shulok,Snodgrass, Wagner,Platika,|date=March 2008 |title=Cancer Gene Therapy by Direct Tumor Injections of a Nonviral T7 Vector Encoding a Thymidine Kinase Gene |journal=Human Gene Therapy |volume=9 |issue=5 |pages=729–736 |doi=10.1089/hum.1998.9.5-729|url=http://online.liebertpub.com/doi/abs/10.1089/hum.1998.9.5-729 |accessdate=2012 |quote= |pmid= 9551620|pmc= }}</ref>

==References==
{{Reflist}}

==External links==
*{{MeshName|T7+Phage}}
*[http://au.expasy.org/viralzone/all_by_species/518.html] ViralZone
*[http://www.microbemagazine.org/index.php?option=com_content&view=article&id=1428:new-details-about-bacteriophage-t7-host-interactions&catid=376:featured&Itemid=531 New Details about T7-host interactions]. Microbe Magazine
*[http://www.uniprot.org/uniprot/?query=organism:10760+keyword:1185 T7 reference proteome] in [[Uniprot]]

{{Taxonbar|from=Q7669119}}

{{DEFAULTSORT:T7 Phage}}
[[Category:Podoviridae]]

Rhizopus microsporus var. oligosporus

2018-03-23T01:27:08Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q4911649}} (2 sig. taxon IDs); WP:GenFixes using AWB

{{Speciesbox
| image = Tempeh_Rhizopus_oligosporus.JPG
| image_caption = ''Rhizopus oligosporus'' on homemade [[tempeh]]
| taxon = Rhizopus oligosporus
| authority = Saito
}}

'''''Rhizopus oligosporus''''' is a [[fungus]] of the [[family]] [[Mucoraceae]] and is a widely used [[starter culture]] for the production of [[tempeh]] at home and industrially. As the mold grows it produces fluffy, white [[Mycelium|mycelia]], binding the beans together to create an edible "cake" of partly [[catabolized]] [[soybeans]]. The domestication of the microbe is thought to have occurred in [[Indonesia]] several centuries ago.<ref>Shurtleff, W. & Aoyagi, A. 2001. The book of tempeh. 2 2. Ten Speed Press. Berkeley, California pp.</ref>

''R. oligosporus'' is the preferred [[starter culture]] for tempeh production for several reasons. It grows effectively at high temperatures (30-40 °C) which are typical of the Indonesian islands, it exhibits strong [[lipolytic]] and [[proteolytic]] activity that create desirable properties in tempeh and it produces [[metabolites]] that allows it to inhibit and thus outcompete other molds and [[gram-positive]] bacteria, including the potentially harmful ''Aspergillus flavus'' and ''[[Staphylococcus aureus]]''.<ref>{{cite journal | last1 = Nout | first1 = M.J.R. | year = 1989 | title = Effect of Rhizopus and Neurospora spp. on growth of Aspergillus flavus and A. parasiticus and accumulation of aflatoxin B1 in groundnut | url = | journal = Mycological Research | volume = 93 | issue = | pages = 518–523 | doi=10.1016/s0953-7562(89)80046-2}}</ref><ref>{{cite journal | last1 = Kobayasi | first1 = Sin-ya | last2 = Naoto | first2 = OKAZAKI | last3 = Takuya | first3 = KosEKI | year = 1992 | title = Purification and Characterization of an Antibiotic Substance Produced from Rhizopus oligosporus IFO 8631 | url = | journal = Biosci. Biotechnol. Biochem. | volume = 56 | issue = 1| pages = 94–98 | doi=10.1271/bbb.56.94 | pmid=1368137}}</ref>

''R. oligosporus'' is at present considered to be a [[domesticated]] form of [[Rhizopus microsporus]] and its proper [[taxonomic]] position is thus ''Rhizopus microsporus var. oligosporus''. ''R. microsporus'' produces several potentially toxic metabolites, [[rhizoxin]] and rhizonins A and B, but it appears the domestication and mutation of the ''R. oligosporus'' [[genome]] has led to the loss of genetic material responsible for [[toxin]] production.<ref>{{cite journal | last1 = Jennessen | first1 = J. | last2 = Nielsen | first2 = K.F. | last3 = Houbraken | first3 = J. | last4 = Lyhne | first4 = E.K. | last5 = Schnürer | first5 = J. | last6 = Frisvad | first6 = J.C. | last7 = Samson | first7 = R.A. | year = 2005 | title = Secondary metabolite and mycotoxin production by the Rhizopus microsporus group | url = | journal = Journal of Agricultural and Food Chemistry | volume = 53 | issue = | pages = 1833–1840 | doi=10.1021/jf048147n | pmid=15740082}}</ref>

==''Rhizopus oligosporus'' properties==
''Rhizopus oligosporus'' is a fungus that belongs to the class [[Zygomycetes]], which is one of two classes in the [[phylum]] [[Zygomycota]].<ref>{{cite journal|last1=Yanai|first1=K|last2=Takaya|first2=N|last3=Kojima|first3=N|last4=Horiuchi|first4=H|last5=Ohta|first5=A|last6=Takagi|first6=M|title=Purification of two chitinases from Rhizopus oligosporus and isolation and sequencing of the encoding genes|journal=American Society for Microbiology|date=1992|page=7398|url=http://jb.asm.org/content/174/22/7398.short|accessdate=29 May 2014}}</ref> ''Rhizopus oligosporus'' belongs to the ''[[Rhizopus microsporus]]'' group. This [[species group|group]] is made of taxa with similar [[Morphology (biology)|morphology]] that are associated with undesired metabolite production, [[pathogenesis]] and food fermentation. Although other varieties in ''Rhizopus microscopus'' may be harmful, ''Rhizopus oligosporus'' is not associated with production of potentially harmful metabolites. It is not found in nature and is frequently used by humans.<ref name="sporangiospores2008">{{cite journal|last1=Jennessen|first1=Jennifer|last2=Schnürer|first2=Johan|last3=Samson|first3=Robert A.|last4=Dijksterhuis|first4=Jan|title=Morphological characteristics of sporangiospores of the tempe fungus Rhizopus oligosporus differentiate it from other taxa of the R. microscopus group|journal=Mycological Research|date=2008|volume=112|pages=547–562|doi=10.1016/j.mycres.2007.11.006|pmid=18400482}}</ref>
''Rhizopus oligosporus'' strains have a large (up to 43 mm) and irregular spores with the most variable sizes. This is, for instance, reflected as high values in the spore volume (96–223 mm3/spore).''Rhizopus oligosporus'' has large, subglobose to globose spores, and high proportion irregular spores (>10 %). ''Rhizopus oligosporus''also has spores with nonparallel valleys and ridges, and plateaus that sometimes are granular<ref name="sporangiospores2008"/>

==''Rhizopus oligosporous'' role in Tempeh fermentation==
A popular Indonesian food, [[Tempeh]], is created by [[fermenting]] [[soybeans]] in combination with ''Rhizopus oligosporus''.<ref name="spores1940">{{cite journal|last1=Hessel Tine|first1=W|last2=Swain|first2=E.W.|last3=Wang|first3=Hwa L.|title=Mass production of Rhizopus oligosporus spores and their application in tempeh fermentation|journal=Journal of Food Science|date=1940|pages=168–170|url=http://onlinelibrary.wiley.com/store/10.1111/j.1365-2621.1975.tb03762.x/asset/j.1365-2621.1975.tb03762.x.pdf|accessdate=29 May 2014|doi=10.1111/j.1365-2621.1975.tb03762.x|volume=40}}</ref> In order to create tempeh, soybeans must first be soaked in water (usually overnight) at a temperature similar to the environment it is placed in. The soybean’s outer covering is then removed and the beans are partially cooked. [[Lactic acid bacteria]], like [[Lactococcus]] and Lb. casei [[species]], play a major role in the fermentation of tempeh.<ref name="microorganisms1999">{{cite journal|last1=Caplice|first1=Elizabeth|last2=Fitzgerald|first2=Gerald F.|title=Food Fermentations: role of microorganisms in food production and preservation|journal=International Journal of Food Microbiology|date=1999|volume=50|page=143|url=http://www.sciencedirect.com/science/article/pii/S0168160599000823#|accessdate=27 May 2014|doi=10.1016/S0168-1605(99)00082-3}}</ref> For the tempeh to ferment, there needs to be a suitable, pure [[Fermentation starter|inoculum]]. Also, [[spores]] with a tendency for fast germinability are needed, as well.<ref name=spores1940 /> In order for the tempeh to attain its characteristic compact ‘cake’ form after fermentation, the soybeans become compressed due to the [[mycelia]] of ''Rhizopus oligosporus''.<ref name=microorganisms1999 /> Rapidly growing mycelia helps speed up the growth of this fungus. Because mycelia are quite sensitive to [[dehydration]] and adverse temperatures, preserving tempeh for extended periods of time can be challenging.<ref name=spores1940 /> When the soybeans are bound together by the white mycelium, the fungus releases [[enzymes]] that make this heavily protein-rich product more digestible for humans.<ref name=sporangiospores2008 /> Tempeh-like foods can also be created from cereal grains such as [[wheat]] and [[rice]]. Many times, a good inoculum for this new fermentation actually comes from tiny pieces of old tempeh that have already been fermented.<ref name=spores1940 />

==Uses of Tempeh==
Tempeh has the potential to be used in many high-protein foods due to its mild flavor when fried in vegetable oil.<ref name=spores1940 /> Containing more than 40% [[protein]], tempeh is often used as a meat-substitute. This product is used in soups or can simply be sliced and seasoned.<ref name=microorganisms1999 />

==Effects of ''Rhizopus oligosporus''==
Even after it is consumed, ''Rhizopus oligosporous'' produces an [[antibiotic]] that limits [[gram-positive bacteria]] like [[Staphylococcus aureus]] (potentially harmful) and [[Bacillus subtilis]] (beneficial). Thus, people who eat tempeh tend to have fewer [[intestinal]] infections.<ref>{{cite journal|last1=Kobayasi|first1=S|last2=Okazaki|first2=N|last3=Koseki|first3=T|title=Purification and characterization of an antibiotic substance produced from Rhizopus oligosporus IFO 8631|journal=Bioscience, Biotechnology, and Biochemistry |volume=56|issue=2|pages=94–98|pmid=1368137|doi=10.1271/bbb.56.94|date=January 1992}}</ref> Tempeh contains [[ergosterol]] ([[provitamin]] D2). Beneficial effects of tempeh include inhibiting [[tumor]] development, lowering [[cholesterol]] and decreasing [[diarrhea]] issues, iron-deficient [[anaemia]], [[lipid oxidation]] and [[hypertension]].<ref>{{cite journal|last1=Chang-Tien|first1=Chang|last2=Hsu|first2=Cheng-Kuang|last3=Chou|first3=Su-Tze|last4=Chen|first4=Ya-Chen|last5=Huang|first5=Feng-Sheng|last6=Chung|first6=Yun-Chin|title=Effect of fermentation time on the antioxidant activities of tempeh prepared from fermented soybean using Rhizopus oligosporous|journal=International Journal of Food Science and Technology|date=2009|volume=44|page=799|url=http://onlinelibrary.wiley.com/doi/10.1111/j.1365-2621.2009.01907.x/full|accessdate=27 May 2014|doi=10.1111/j.1365-2621.2009.01907.x}}</ref> This fungus can also treat waste and wastewater, produce industrial enzymes and ferment other substrates like other legumes and cereals.<ref name=sporangiospores2008 />

==See also==
*''[[Aspergillus oryzae]]''
*[[Medicinal molds]]
*''[[Saccharomyces boulardii]]''
*''[[Saccharomyces cerevisiae]]''

==References==
{{Reflist}}

==External links==
*[http://www.indexfungorum.org/Names/NamesRecord.asp?RecordID=155475 Index Fungorum page (synonyms)]
*[https://books.google.com/books?id=hdKzFlaqWT8C&printsec=frontcover ''The Book of Tempeh: Professional Edition''] - by Shurtleff and Aoyagi (1979).
*[https://books.google.com/books?id=CgA5Z2VaCjkC&printsec=frontcover ''Tempeh production: a craft and technical manual''] - By William Shurtleff, Akiko Aoyagi, Soyfoods Center (Lafayette, Calif.)

{{Taxonbar|from=Q4911649}}

[[Category:Mucoraceae]]
[[Category:Molds used in food production]]

Globisporangium sylvaticum

2018-03-23T00:54:31Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q7263903}} (1 sig. taxon ID); WP:GenFixes, using AWB

{{Italic title}}
{{Taxobox
| name = Globisporangium sylvaticum
| unranked_regnum = [[SAR supergroup|SAR]]
| superphylum = [[Heterokont]]a
| phylum = [[Oomycota]]
| classis = [[Oomycete]]
| ordo = [[Pythiales]]
| familia = [[Pythiaceae]]
| genus = ''[[Globisporangium]]''
| species = ''G. sylvaticum''
| binomial = ''Globisporangium sylvaticum''
| binomial_authority = (W.A. Campb. & F.F. Hendrix) Uzuhashi, Tojo & Kakish., 2010
| synonyms =
''Pythium sylvaticum'' 
}}

'''''Globisporangium sylvaticum''''' is a plant [[pathogen]], an [[oomycete]] known to cause [[root rot]] and [[damping off]] in a multitude of species. These species include [[Apples]], [[Carrot]], [[Cherry Laurel]], [[Garden cress|Cress]], [[Cucumber]], [[Garlic]], [[Lettuce]], [[Pea]], [[Rhododendron]], and [[Spinach]].<ref>"A Desk Study to Review Global Knowledge on Best Practice of Oomycete Root-rot Detection and Control." (n.d.): n. pag. http://horticulture.ahdb.org.uk/sites/default/files/research_papers/CP%20126_Report_Final_2014.pdf. Agriculture and Horticulture Development Board, Apr. 2015. Web. 5 Dec. 2016. <http://horticulture.ahdb.org.uk/sites/default/files/research_papers/CP%20126_Report_Final_2014.pdf>.</ref> Symptoms of infection include [[stunt (botany)|stunt]]ing, [[wilting|wilt]], [[chlorosis]], and browning and eventual [[necrosis]] of roots.<ref>Pythium (Plant Diseases)." Plant Diseases (Penn State Extension). Pennsylvania State University, n.d. Web. 15 Nov. 2016. [http://extension.psu.edu/pests/plant-diseases/all-fact-sheets/pythium].</ref> The pathogen can by identified by the presence of thick, microscopic, round spores within the cells of the root.

The species was formerly placed in the genus ''Pythium'', but that genus has been divided into five distinct [[clades]], each characterized by the morphology of the [[sporangium]].<ref>''Globisporangium sylvaticum'' (W. A. Campb. &F. F. Hendrix) Uzuhashi, Tojo & Kakish., comb. nov. Basionym: ''Pythium sylvaticum'' W. A. Campb. & F. F. Hendrix, Mycologia 59: 274, 19 https://www.researchgate.net/publication/225748808_Phylogeny_of_the_genus_Pythium_and_description_of_new_genera</ref>

==Disease cycle==
''Globisporangium sylvaticum'' is known to be [[heterothallic]], whereas other species in the [[basionym]] ''Pythium'' were consistently known to be [[homothallic]].<ref>Papa, K. E. et al. “Sexuality in Pythium Sylvaticum: Heterothallism.” Mycologia, vol. 59, no. 4, 1967, pp. 589–595. www.jstor.org/stable/3757088.</ref> Additionally, [[oospores]] have been shown to occur only in specific pairings of certain sporangial isolates, a finding which reinforces its [[heterothallic]] nature.<ref>Papa, K. E. et al. “Sexuality in Pythium Sylvaticum: Heterothallism.” Mycologia, vol. 59, no. 4, 1967, pp. 589–595. www.jstor.org/stable/3757088.</ref> Since the pathogen is an [[oomycete]], it is known to produce several types of spores including [[sporangia]], [[zoospores]], and [[oospores]].<ref>Link, Virginia Heffer, Mary L. Powelson, and Kenneth B. Johnson. "Oomycetes." Oomycetes. American Pathological Society, n.d. Web. 16 Nov. 2016. [http://www.apsnet.org/EDCENTER/INTROPP/LABEXERCISES/Pages/Oomycetes.aspx].</ref>

The pathogen begins its life cycle in the growth stage by constructing a well developed [[mycelium]] made of hyaline [[hyphae]]. This [[mycelium]] is the actively growing body of the pathogen and is responsible for the infection of the host, the subsequent colonization of the host plant, and the uptake of nutrients from its host.

This hyphal growth can occur in two different ways within ''Globisporangium sylvaticum'', asexually or sexually.

During its aesexual life cycle the pathogen produces sporangium that occur terminally and intercalary, and are globose and thin walled in shape as well as sub-globose and lemon shaped, respectively.<ref>Campbell, W. A., and F. F. Hendrix. “A New Heterothallic Pythium from Southern United States.” Mycologia, vol. 59, no. 2, 1967, pp. 274–278. www.jstor.org/stable/3756800.</ref> After a period of time biflagellate [[zoospores]] begin to develop within the protoplasm of the [[sporangium]] and are subsequently released during the proper conditions.<ref>Globisporangium sylvaticum (W. A. Campb. &F. F. Hendrix) Uzuhashi, Tojo & Kakish., comb. nov. Basionym: Pythium sylvaticum W. A. Campb. & F. F. Hendrix, Mycologia 59: 274, 19 https://www.researchgate.net/publication/225748808_Phylogeny_of_the_genus_Pythium_and_description_of_new_genera</ref> The motile [[zoospores]] proliferate through the vector medium until they reach the infection court of its host, in this case the seed or the roots. Once at the infection court, the [[zoospores]] encyst themselves on the root tips or seed of the host thereby infecting it and resulting in the occurrence of more mycelial growth and the eventual damping off/ root rot of the host plant.

However, during the sexual lifecycle the pathogen, as previously mentioned, needs two different compatible isolates to form a [[diploid]] structure called an [[oospore]]. The [[oospore]] is a survival structure for the pathogen which only germinates during non-adverse conditions.<ref>"A Desk Study to Review Global Knowledge on Best Practice of Oomycete Root-rot Detection and Control." (n.d.): n. pag. http://horticulture.ahdb.org.uk/sites/default/files/research_papers/CP%20126_Report_Final_2014.pdf. Agriculture and Horticulture Development Board, Apr. 2015. Web. 5 Dec. 2016. <http://horticulture.ahdb.org.uk/sites/default/files/research_papers/CP%20126_Report_Final_2014.pdf>.</ref> In the case of ''Globisporangium sylvaticum'', the [[oospore]] is globose, [[aplerotic]](space between oospore wall and oogonium wall) and thick walled.<ref>Globisporangium sylvaticum (W. A. Campb. &F. F. Hendrix) Uzuhashi, Tojo & Kakish., comb. nov. Basionym: Pythium sylvaticum W. A. Campb. & F. F. Hendrix, Mycologia 59: 274, 19 https://www.researchgate.net/publication/225748808_Phylogeny_of_the_genus_Pythium_and_description_of_new_genera</ref><ref>Campbell, W. A., and F. F. Hendrix. “A New Heterothallic Pythium from Southern United States.” Mycologia, vol. 59, no. 2, 1967, pp. 274–278. www.jstor.org/stable/3756800.</ref> The formation of the [[oospore]] is caused by the fusion of an [[antheridia]] and an [[oogonium]], the male and female reproductive parts, respectively. The antheridia in ''Globisporangium sylvaticum'' is characterized as being [[diclinous]] and branched while the [[oogonia]] are [[Intercalation (biochemistry)|intercalary]] and sub-globose.<ref>Globisporangium sylvaticum (W. A. Campb. &F. F. Hendrix) Uzuhashi, Tojo & Kakish., comb. nov. Basionym: Pythium sylvaticum W. A. Campb. & F. F. Hendrix, Mycologia 59: 274, 19 https://www.researchgate.net/publication/225748808_Phylogeny_of_the_genus_Pythium_and_description_of_new_genera</ref><ref>Campbell, W. A., and F. F. Hendrix. “A New Heterothallic Pythium from Southern United States.” Mycologia, vol. 59, no. 2, 1967, pp. 274–278. www.jstor.org/stable/3756800.</ref> After an [[oospore]] is formed as a result of the fusion between the aforementioned reproductive structures the pathogen continues to undergo cell division giving rise to the [[mycelium]] where both cycles repeat under the proper conditions.

==Environment==
A moist environment is required for the propagation and dispersal of ''Globisporangium sylvaticum''. The pathogen produces highly motile [[zoospores]], which use [[Flagellum|flagella]] for locomotion. In the presence of water, the zoospore will use its flagella to propel itself through the surrounding water, in the direction of a prospective [[Glossary of phytopathology|infection court]].<ref>Raftoyannis, Yannis, and Michael W. Dick. "Zoospore Encystment and Pathogenicity of Phytophthora and Pythium Species on Plant Roots." Zoospore Encystment and Pathogenicity of Phytophthora and Pythium Species on Plant Roots. N.p., 1 June 2005. Web. 16 Nov. 2016. [http://www.sciencedirect.com/science/article/pii/S0944501305000479].</ref> In adverse environmental conditions, the pathogen exists in a structure called an [[oospore]]. This structure protects the pathogen for extended periods until conditions for its propagation are satisfied.<ref>"Pythium." Brill’s New Pauly (n.d.): n. pag. Michigan State University. http://www.ipm.msu.edu/uploads/files/herbperennials/pythium.pdf</ref> Additionally, during ideal temperature conditions, it has been observed that optimal radial growth for ''Globisporangium sylvaticum'' occurs between 28 °C and 30 °C. During this temperature range the pathogen has been observed to grow 2.7 cm over a 24-hour period.<ref>Campbell, W. A., and F. F. Hendrix. “A New Heterothallic Pythium from Southern United States.” Mycologia, vol. 59, no. 2, 1967, pp. 274–278. www.jstor.org/stable/3756800.Copy</ref>

== External links ==
* [http://www.speciesfungorum.org/Names/Names.asp Index Fungorum]
* [https://www.webcitation.org/5QK694Uju?url=http://nt.ars-grin.gov/fungaldatabases USDA ARS Fungal Database]

== References ==
<references />

{{Taxonbar|from=Q7263903}}

[[Category:Water mould plant pathogens and diseases]]
[[Category:Carrot diseases]]
[[Category:Leaf vegetable diseases]]
[[Category:Peronosporales]]
[[Category:Eukaryotes described in 1967]]

Orthobunyavirus

2018-03-23T00:30:20Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q2580182}} (1 sig. taxon ID); WP:GenFixes, using AWB

{{Taxobox

| name = ''Orthobunyavirus''
| virus_group = v
| ordo = ''[[Bunyavirales]]''
| familia = ''[[Peribunyaviridae]]''
| genus = '''''Orthobunyavirus'''''
| type_species = ''Bunyamwera virus''
| subdivision_ranks = Species
| subdivision =''[[Acara virus]]'' 
''[[Akabane virus]]'' 
''[[Alajuela virus]]'' 
''[[Anopheles A virus]]'' 
''[[Anopheles B virus]]'' 
''[[Bakau virus]]'' 
''[[Batai virus]]'' 
''[[Batama virus]]'' 
''[[Bellavista virus]]'' 
''[[Benevides virus]]'' 
''[[Bertioga virus]]'' 
''[[Bimiti virus]]'' 
''[[Botambi virus]]'' 
''[[Bunyamwera virus]]'' 
''[[Bushbush virus]]'' 
''[[Bwamba virus]]'' 
''[[California encephalitis virus]]'' 
''[[Capim virus]]'' 
''[[Caraparu virus]]'' 
''[[Catu virus]]'' 
''[[Estero Real virus]]'' 
''[[Gamboa virus]]'' 
''[[Guajara virus]]'' 
''[[Guama virus]]'' 
''[[Guaroa virus]]'' 
''[[Jatobal virus]]'' 
''[[Kaeng Khoi virus]]'' 
''[[Kairi virus]]'' 
''[[Koongol virus]]'' 
''[[Kowanyama virus]]'' 
''[[La Crosse virus]]'' 
''[[M'Poko virus]]'' 
''[[Madrid virus]]'' 
''[[Maguari virus]]'' 
''[[Main Drain virus]]'' 
''[[Manzanilla virus]]'' 
''[[Marituba virus]]'' 
''[[Minatitlan virus]]'' 
''[[Nyando virus]]'' 
''[[Olifantsvlei virus]]'' 
''[[Oriboca virus]]'' 
''[[Oropouche virus]]'' 
''[[Patois virus]]'' 
''[[Sathuperi virus]]'' 
''[[Schmallenberg virus]]"''<ref>[http://www.n-tv.de/wissen/Unbekanntes-Virus-entdeckt-article4825956.html Provisional name, discovered in 2011]</ref> 
''[[Shamonda virus]]'' 
''[[Shuni virus]]'' 
''[[Simbu virus]]'' 
''[[South River virus]]'' 
''[[Tacaiuma virus]]'' 
''[[Tahyna virus]]'' 
''[[Tensaw virus]]'' 
''[[Tete virus]]'' 
''[[Thimiri virus]]'' 
''[[Timboteua virus]]'' 
''[[Turlock virus]]'' 
''[[Wyeomyia virus]]'' 
''[[Zegla virus]]''
}}

'''''Orthobunyavirus''''' is a [[genus]] of the [[Peribunyaviridae]] [[family (taxonomy)|family]] in the order [[Bunyavirales]]. There are currently ~170 viruses recognised in this genus. These have been assembled into 48 species and 19 serogroups.

The [[type species]] is [[Bunyamwera virus]].

==Epidemiology==

The genus is most diverse in [[Africa]], [[Australia]] and [[Oceania]], but occurs almost worldwide. Most orthobunyavirus [[species]] are transmitted by [[gnat]]s and cause diseases of [[cattle]].{{citation needed|date=September 2015}} The ''[[California encephalitis virus]]'', the ''[[La Crosse virus]]'' and the ''[[Jamestown Canyon virus]]''  are [[North America]]n species that cause [[encephalitis]] in humans.

==Virology==

*The type species is the Bunyamwera virus.<ref>[http://ictvdb.bio-mirror.cn/Ictv/fs_bunya.htm International Committee on Taxonomy of Viruses, Bunyaviridae]</ref>
*The virus is spherical, diameter 80 nm to 120 nm, and comprises three single stranded RNA molecules encapsulated in a ribonucleocapsid.<ref>[http://viralzone.expasy.org/all_by_species/250.html Orthobunyavirus by ViralZone]</ref>
*The three RNAs are described as S, M and L (for Small, Medium and Large) and are circa 1kb (kilobases), 4.5kb and 6.5kb in length<ref>[https://dx.doi.org/10.1016/0042-6822(77)90030-7 Richard J. Kascsak & Michael J. Lyons, (1971). Bunyamwera virus I. The molecular complexity of the virion RNA, Virology, Volume 82, Issue 1, 1 October 1977, Pages 37–47]</ref><ref>[http://viralzone.expasy.org/all_by_species/250.html Orthobunyavirus by ViralZone]</ref>
*The S RNA encodes the Nucleocapsid protein (N protein) and a non structural protein (NS Protein).<ref>Genbank: [https://www.ncbi.nlm.nih.gov/nuccore/17225332?report=genbank Bunyamwera virus segment S, complete sequence]</ref>
*The M RNA encodes a polyprotein which is cleaved by host protease into Gn, NSm and Gc proteins.<ref>[http://viralzone.expasy.org/all_by_species/250.html Orthobunyavirus by ViralZone]</ref>
*The L RNA encodes the viral RNA dependent RNA Polymerase or L Protein<ref>Genbank: [https://www.ncbi.nlm.nih.gov/nuccore/9630656?report=genbank Bunyamwera virus L segment, complete sequence]</ref>

==Life cycle==

{{Empty section|date=September 2012}}

==Serogroups==

The taxonomy remains somewhat fluid as relatively few viral genomes in this genus have been sequenced. Several of the viruses listed have been shown to be recombinants of other viruses and may be reclassified.

18 serogroups have been recognized on the basis of the results of cross-hemagglutination inhibition and antibody neutralization relationships. Another - Wyeomyia - has since been recognised. Several viruses have not yet been classified into one of the serogroups.

The Simbu serogroup is the largest and contains at least 25 members. There are at least 13 members in the Group C serogroup.

Medically important viruses belong to the Bwamba, Bunyamwera, California, Group C and Simbu serogroups.

===Anopheles A serogroup===

Anopheles A virus 
Tacaiuma virus 
Virgin River virus

Trombetas complex
*Arumateua virus
*Caraipé virus
*Trombetas virus
*Tucuruí virus

===Anopheles B serogroup===

Anopheles B virus 
Boraceia virus

===Bakau serogroup===

Bakau virus 
Nola virus

===Bunyamwera serogroup===

Birao virus 
Bozo virus 
[[Bunyamwera virus]] 
Cache Valley virus 
Fort Sherman virus 
Germiston virus 
Guaroa virus 
Ilesha virus 
Kairi virus 
[[Maguari virus]] 
Main Drain virus 
Northway virus 
Playas virus 
Potosi virus 
Shokwe virus 
Stanfield virus 
[[Tensaw virus]] 
Xingu virus

Batai complex
*Batai virus
*Čalovo virus
*Chittoor virus

Ngari complex
*Garissa virus
*KV-141 virus
*Ngari virus

===Bwamba serogroup===

Bwamba virus 
Pongola virus

===California serogroup===

[[California encephalitis virus]] 
Chatanga virus 
Inkoo virus 
[[Jamestown Canyon virus]] 
Jerry Slough virus 
Keystone virus 
Khatanga virus 
[[La Crosse virus]] 
Lumbo virus 
Melao virus 
Morro Bay virus 
San Angelo virus 
Serra do Navio virus 
Snowshoe hare virus 
South River virus 
[[Tahyna virus]] 
Trivittatus virus

===Capim serogroup===

Acara virus 
Benevides virus 
Capim virus

===Gamboa serogroup===

Alajuela virus 
Gamboa virus 
Pueblo Viejo virus 
San Juan virus

===Group C serogroup===

Bruconha virus 
Ossa virus

Caraparu complex
*Apeu virus
*Bruconha virus
*Caraparu virus
*Vinces virus

Madrid complex
*Madrid virus
*Marituba complex
*Gumbo limbo virus
*Marituba virus
*Murutucu virus
*Nepuyo virus
*Restan virus

Oriboca complex
*Itaqui virus
*Oriboca virus

===Guama serogroup===

Ananindeua virus 
Bertioga virus 
Bimiti virus 
Cananeia virus 
Catu virus 
Guama virus 
Guaratuba virus 
Itimirim virus 
Mahogany hammock virus 
Mirim virus 
Timboteua virus

===Koongol serogroup===

Koongol virus 
Wongal virus

===Mapputta serogroup===

Buffalo Creek virus 
Mapputta virus 
Maprik virus 
Murrumbidgee virus 
Salt Ash virus

===Minatitlan serogroup===

Minatitlan virus 
Palestina virus

===Nyando serogroup===

Eretmapodites virus 
Nyamdo virus

===Olifanstlei serogroup===

Botambi virus 
Olifanstlei virus

===Patois serogroup===

Abras virus 
Babahoyo virus 
Pahayokee virus 
Patois virus 
Shark River virus

===Simbu serogroup===

Iquitos virus 
Jatobal virus 
Leanyer virus 
Oropouche serocomplex
*Madre de Dios virus (MDDV)
*Oropouche virus
Oya virus 
Thimiri virus

Akabane serocomplex
*Akabane virus
*Tinaroo virus

Sathuperi serocomplex
*Douglas virus
*Sathuperi virus

Shuni serocomplex
*Aino virus
*Shuni virus

Shamonda serocomplex
*Peaton virus
*Shamonda virus

Simbu complex
*Schmallenberg virus
*Simbu virus

===Tete serogroup===

Batama virus 
[[Tete virus]]

===Turlock serogroup===

M'Poko virus 
Turlock virus 
Umbre virus

===Wyeomyia serogroup===

[[Anhembi virus]] 
Cachoeira Porteira virus 
Iaco virus 
Macaua virus 
Sororoca virus 
Taiassui virus 
Tucunduba virus 
Wyeomyia virus

===Unclassified===

Batama virus 
Bellavista virus 
Belmont virus 
Enseada virus 
Estero Real virus 
Jurona virus 
Kaeng Khei virus 
Kowanyama virus 
Mojuí dos Campos virus 
Wolkberg virus

==References==
{{Reflist}}

==External links==
* [http://www.expasy.org/viralzone/all_by_species/250.html '''Viralzone''': Orthobunyavirus]

{{Taxonbar|from=Q2580182}}

[[Category:Orthobunyaviruses]]

Virgaviridae

2018-03-23T00:16:57Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q3560593}} (1 sig. taxon ID); WP:GenFixes, using AWB

{{Use dmy dates|date=April 2017}}
{{taxobox
| familia = '''''Virgaviridae'''''
| virus_group = iv
| subdivision_ranks = Genera
| subdivision =
*''[[Furovirus]]''
*''[[Goravirus]]''
*''[[Hordeivirus]]''
*''[[Pecluvirus]]''
*''[[Pomovirus]]''
*''[[Tobamovirus]]''
*''[[Tobravirus]]''
}}

'''''Virgaviridae''''' is a family of positive sense, single-stranded [[RNA virus]]es. Plants serve as natural hosts.<ref>{{cite journal|last1=Adams|first1=Michael J.|last2=Adkins|first2=Scott|last3=Bragard|first3=Claude|last4=Gilmer|first4=David|last5=Li|first5=Dawei|last6=MacFarlane|first6=Stuart A.|last7=Wong|first7=Sek-Man|last8=Melcher|first8=Ulrich|last9=Ratti|first9=Claudio|last10=Ryu|first10=Ki Hyun|title=ICTV Virus Taxonomy Profile: Virgaviridae|journal=Journal of General Virology|date=1 August 2017|volume=98|issue=8|pages=1999–2000|doi=10.1099/jgv.0.000884}}</ref><ref name="ICTVReport">{{cite web|title=ICTV Report Virgaviridae|url=http://www.ictv.global/report/virgaviridae}}</ref><ref name=ViralZone /><ref name=Adams2009>Adams MJ, Antoniw JF, Kreuze J (2009) Virgaviridae: a new family of rod-shaped plant viruses. Arch Virol 154(12):1967–1972</ref> There are currently 59 species in this family, divided among 7 genera.<ref name=ICTVReport /><ref name=ViralZone>{{cite web|title=Viral Zone|url=http://viralzone.expasy.org/all_by_species/734.html|publisher=ExPASy|accessdate=15 June 2015}}</ref><ref name="ICTV">{{cite web|url=http://ictvonline.org/virusTaxonomy.asp|title=Virus Taxonomy: 2015 Release|last1=ICTV|accessdate=4 June 2016}}</ref> The name of the family is derived from the Latin word ''virga'' (rod), as all viruses in this family are rod-shaped.

==Taxonomy==
Viruses include in the family ''Virgaviridae'' are characterized by unique alpha-like replication proteins.

<big>'''Group: ssRNA(+)'''</big>
{{Collapsible list|title= <big>Order: Unassigned</big>
|1={{Collapsible list| framestyle=border:none; padding:1.0em;|title=Family: Virgaviridae
|1={{hidden begin|title=Genus: [[Furovirus]]}}
*[[Chinese wheat mosaic virus]]
*[[Japanese soil-borne wheat mosaic virus]]
*[[Oat golden stripe virus]]
*[[Soil-borne cereal mosaic virus]]
*'''''[[Soil-borne wheat mosaic virus]]'''''
*[[Sorghum chlorotic spot virus]]
{{hidden end}}
|2={{hidden begin|title=Genus: [[Hordeivirus]]}}
*[[Anthoxanthum latent blanching virus]]
*'''''[[Barley stripe mosaic virus]]'''''
*[[Lychnis ringspot virus]]
*[[Poa semilatent virus]]
{{hidden end}}
|3={{hidden begin|title=Genus: [[Pecluvirus]]}}
*[[Indian peanut clump virus]]
*'''''[[Peanut clump virus]]'''''
{{hidden end}}
|4={{hidden begin|title=Genus: [[Pomovirus]]}}
*[[Beet soil-borne virus]]
*[[Beet virus Q]]
*[[Broad bean necrosis virus]]
*'''''[[Potato mop-top virus]]'''''
{{hidden end}}
|5={{hidden begin|title=Genus: [[Tobamovirus]]}}
*[[Bell pepper mottle virus]]
*[[Brugmansia mild mottle virus]]
*[[Cactus mild mottle virus]]
*[[Clitoria yellow mottle virus]]
*[[Cucumber fruit mottle mosaic virus]]
*[[Cucumber green mottle mosaic virus]]
*[[Cucumber mottle virus]]
*[[Frangipani mosaic virus]]
*[[Hibiscus latent Fort Pierce virus]]
*[[Hibiscus latent Singapore virus]]
*[[Kyuri green mottle mosaic virus]]
*[[Maracuja mosaic virus]]
*[[Obuda pepper virus]]
*[[Odontoglossum ringspot virus]]
*[[Paprika mild mottle virus]]
*[[Passion fruit mosaic virus]]
*[[Pepper mild mottle virus]]
*[[Rattail cactus necrosis-associated virus]]
*[[Rehmannia mosaic virus]]
*[[Ribgrass mosaic virus]]
*[[Sammons's Opuntia virus]]
*[[Streptocarpus flower break virus]]
*[[Sunn-hemp mosaic virus]]
*[[Tobacco latent virus]]
*[[Tobacco mild green mosaic virus]]
*'''''[[Tobacco mosaic virus]]'''''
*[[Tomato mosaic virus]]
*[[Tomato mottle mosaic virus]]
*[[Tropical soda apple mosaic virus]]
*[[Turnip vein-clearing virus]]
*[[Ullucus mild mottle virus]]
*[[Wasabi mottle virus]]
*[[Yellow tailflower mild mottle virus]]
*[[Youcai mosaic virus]]
*[[Zucchini green mottle mosaic virus]]
{{hidden end}}
|6={{hidden begin|title=Genus: [[Tobravirus]]}}
*[[Pea early-browning virus]]
*[[Pepper ringspot virus]]
*'''''[[Tobacco rattle virus]]'''''
{{hidden end}}
|7={{hidden begin|title=Genus: [[Goravirus]]}}
*[[Drakaea virus A]]
*'''''[[Gentian ovary ringspot virus]]'''''
{{hidden end}}
}}
}}<ref name=ICTVReport />

==Structure==
Viruses in Virgaviridae are non-enveloped, with rigid helical rod geometries, and helical symmetry. The diameter is around 20-25 nm,<ref name=ICTVReport /><ref name=ViralZone /> and virions have a central "canal." Genomes are linear, single-stranded, positive sense RNA<ref name=ICTVReport /><ref name=ViralZone /> with a 3'-tRNA like structure and no polyA tail. They may be in one, two, or three segments, depending on the genus. Coat proteins are about 19–24 [[Dalton (unit)|kiloDalton]]s.

{| class="wikitable sortable" style="text-align:center"
|-
! Genus !! Structure || Symmetry !! Capsid !! Genomic arrangement !! Genomic segmentation
|-
|''Tobamovirus''||Rod-shaped||Helical||Non-enveloped||Linear||Non-Segmented
|-
|''Furovirus''||Rod-shaped||Helical||Non-enveloped||Linear||Segmented
|-
|''Pecluvirus''||Rod-shaped||Helical||Non-enveloped||Linear||Segmented
|-
|''Hordeivirus''||Rod-shaped||Helical||Non-enveloped||Linear||Segmented
|-
|''Tobravirus''||Rod-shaped||Helical||Non-enveloped||Linear||Segmented
|-
|''Pomovirus''||Rod-shaped||Helical||Non-enveloped||Linear||Segmented
|}

==Life cycle==
Viral replication is cytoplasmic. Entry into the host cell is achieved by penetration into the host cell. Replication follows the positive stranded RNA virus replication model. Positive stranded RNA virus transcription is the method of transcription. Translation takes place by leaky scanning, and suppression of termination. The virus exits the host cell by tripartite non-tubule guided viral movement, and monopartite non-tubule guided viral movement.
Plants serve as the natural host.<ref name=ICTVReport /><ref name=ViralZone />

{| class="wikitable sortable" style="text-align:center"
|-
! Genus !! Host details !! Tissue tropism !! Entry details !! Release details !! Replication site !! Assembly site !! Transmission
|-
|''Tobamovirus''||Plants||None||Unknown||Viral movement||Cytoplasm||Cytoplasm||Mechanical
|-
|''Furovirus''||Plants||None||Unknown||Viral movement||Cytoplasm||Cytoplasm||Mechanical inoculation: fungus
|-
|''Pecluvirus''||Plants||None||Unknown||Viral movement||Cytoplasm||Cytoplasm||Mechanical inoculation: fungus. Mechanical contact; seed
|-
|''Hordeivirus''||Plants||None||Unknown||Viral movement||Cytoplasm||Cytoplasm||Mechanical: contact; seed
|-
|''Tobravirus''||Plants||None||Unknown||Viral movement||Cytoplasm||Cytoplasm||Mechanical inoculation: nematodes
|-
|''Pomovirus''||Plants||None||Unknown||Viral movement||Cytoplasm||Cytoplasm||Mechanical inoculation: fungus
|}

==Notes==
The genus ''[[Benyvirus]]'', although its members are rod shaped and infect plants, is not included in this family as its proteins appear to be only very distantly related, but is instead included in the family ''Benyviridae''.<ref>{{cite web|title=ICTV Report Benyviridae|url=http://www.ictv.global/report/benyviridae}}</ref>

==References==
{{Reflist}}

==External links==
* [http://www.ictv.global/report/virgaviridae '''ICTV Online Report''': ''Virgaviridae'']
* [http://viralzone.expasy.org/all_by_species/734.html '''Viralzone''': Virgaviridae]
{{Baltimore classification}}

{{Taxonbar|from=Q3560593}}

[[Category:Virgaviridae]]
[[Category:Viral plant pathogens and diseases]]
[[Category:Virus families]]

Trichoderma reesei

2018-03-23T00:03:17Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q2749932}} (3 sig. taxon IDs); WP:GenFixes using AWB

{{Italic title}}
{{Taxobox
| image =Trichoderma.reesei.jpg
| image_caption =
| name = ''Trichoderma reesei''
| regnum = [[Fungi]]
| divisio = [[Ascomycota]]
| subdivisio = [[Pezizomycotina]]
| classis = [[Sordariomycetes]]
| ordo = [[Hypocreales]]
| familia = [[Hypocreaceae]]
| genus = ''[[Trichoderma]]''
| species = '''''T. reesei'''''
| binomial = ''Trichoderma reesei''
}}

'''''Trichoderma reesei''''' is a [[mesophilic]] and [[Hypha|filamentous]] [[fungus]]. It is an [[anamorph]] of the fungus '''''Hypocrea jecorina'''''.
''T. reesei'' has the capacity to secrete large amounts of [[cellulolytic]] [[enzyme]]s ([[cellulase]]s and [[hemicellulase]]s). Microbial [[cellulase]]s have industrial application in the conversion of [[cellulose]], a major component of plant [[biomass]], into [[glucose]].<ref>{{cite journal |vauthors=Kumar R, Singh S, Singh OV |title=Bioconversion of lignocellulosic biomass: biochemical and molecular perspectives |journal=J. Ind. Microbiol. Biotechnol. |volume=35 |issue=5 |pages=377–91 |date=May 2008 |pmid=18338189 |doi=10.1007/s10295-008-0327-8 }}</ref>

''T. reesei'' isolate QM6a was originally isolated from the [[Solomon Islands]] during [[World War II]] because of its degradation of canvas and garments of the [[US army]].<ref name=Seidl09>{{cite journal |vauthors=Seidl V, Seibel C, Kubicek CP, Schmoll M |title=Sexual development in the industrial workhorse Trichoderma reesei |journal=PNAS |volume=106 |issue=33 |pages=13909–13914 |year=2009 |doi=10.1073/pnas.0904936106 |pmid=19667182 |pmc=2728994}}</ref> All strains currently used in biotechnology and basic research were derived from this one isolate.<ref name=Seidl09 />

Recent advances in the [[biochemistry]] of cellulase [[enzymology]], the mechanism of cellulose [[hydrolysis]] ([[cellulolysis]]), [[Strain (biology)|strain]] improvement, molecular cloning and process engineering are bringing ''T. reesei'' cellulases closer to being a commercially viable route to cellulose hydrolysis.<ref>{{cite journal |vauthors=Viikari L, Alapuranen M, Puranen T, Vehmaanperä J, Siika-Aho M |title=Thermostable enzymes in lignocellulose hydrolysis |journal=Adv. Biochem. Eng. Biotechnol. |volume=108 |issue= |pages=121–45 |year=2007 |pmid=17589813 |doi=10.1007/10_2007_065 }}</ref> Several industrially useful strains have been developed and characterised, e.g. Rut-C30,<ref>{{cite journal |vauthors=Seidl V, Gamauf C, Druzhinina IS, Seiboth B, Hartl L, Kubicek CP |title=The ''Hypocrea jecorina'' (''Trichoderma reesei'') hypercellulolytic mutant RUT C30 lacks an 85 kb (29 gene-encoding) region of the wild-type genome |journal=BMC Genomics |volume=9|pages=327 |year=2008 |pmid=18620557 |pmc=2483294 |doi=10.1186/1471-2164-9-327 |url=http://www.biomedcentral.com/1471-2164/9/327}}</ref> RL-P37 and MCG-80. The genome of this organism was released in 2008.<ref>{{cite journal |vauthors=Martinez D, Berka RM, Henrissat B, etal |title=Genome sequencing and analysis of the biomass-degrading fungus Trichoderma reesei (syn. Hypocrea jecorina) |journal=Nat. Biotechnol. |volume=26 |issue=5 |pages=553–60 |date=May 2008 |pmid=18454138 |doi=10.1038/nbt1403 }}</ref> This organism also has a mating type dependent characterised sexual cycle.<ref name=Seidl09 />

==Sexual development==

''T. reesei'' QM6a has a ''MAT1-2'' mating type locus. The opposite mating type ''MAT1-1'', was recently found, thus proving that ''T. reesei'' is a heterothallic species.<ref name=Seidl09 /> After being regarded as asexual since its discovery more than 50 years ago, sexual reproduction can now be induced in ''T. reesei'' QM6a leading to formation of fertilized stromata and mature ascospores.<ref name=Seidl09 />

==Use in industry==

''T. reesei'' is an important commercial and industrial micro-organism due to its cellulase production ability. Many strains of ''T. reesei'' have been developed since its discovery, with heavy emphasis on increasing cellulase production. These "improvement programs" originally consisted of classical (ionising-radiation-based and chemical-based) mutagenesis, which led to strains capable of producing 20 times as much cellulase as the QM6a isolate.<ref name=Biofuelbook>{{cite book
| last1=Seiboth
| first1=Bernhard
| last2=Ivanova
| first2=Christa
| last3=Seidl-Seiboth
| first3= Verena
| editor-last=Dos Santos Bernardes
| editor-first=Marco Aurélio
| title=Biofuel Production-Recent Developments and Prospects
| publisher=InTech
| date=September 15, 2011
| pages=321
| chapter=Chapter 13: Trichoderma reesei: A Fungal Enzyme Producer for Cellulosic Biofuels
| isbn=978-953-307-478-8
| doi=10.5772/16848 }}</ref> The ultimate aim in the creation of hypercellulolytic strains was to obtain a [[Catabolite repression|carbon catabolite derepressed]] strain. This derepression would allow the ''T. reesei'' strain to produce cellulases under any set of growth conditions, even in the presence of glucose.

However, with the advent of modern genetic engineering tools such as targeted deletion, targeted knockout, and more, a new generation of strains dubbed "hyperproducers" has emerged. Some of the highest performing industrial strains produce up to 100 grams of cellulases per litre, more than 3 times as much as the RUT-C30 strain (which itself produces twice as much as the parent strain NG14 from which it was derived).<ref name=Biofuelbook />

== See also ==
{{Portal|Fungi}}
*[[Cellobiohydrolase (disambiguation)|Cellobiohydrolase]] (CBH)
*[[Cellulosic ethanol]]
*[[Endoglucanase]] (EG)

==References==
{{Reflist}}

==External links==
* Risk Assessment Summary, CEPA 1999. [https://web.archive.org/web/20131120135831/http://www.bch.gc.ca/default.asp?lang=En&n=3CC8DDA3-1&xsl=bchdescriptor,showfull&xml=664AFEB2-AE08-454F-ADD8-38C5A94C1DA4 ''Trichoderma reesei'' 1391A]
* Risk Assessment Summary, CEPA 1999. [http://www.ec.gc.ca/subsnouvelles-newsubs/default.asp?lang=En&n=AFDD052F-1 ''Trichoderma reesei'' P59G]
* Risk Assessment Summary, CEPA 1999. [http://www.ec.gc.ca/subsnouvelles-newsubs/default.asp?lang=En&n=894BCDC6-1 ''Trichoderma reesei'' P210A]
* Risk Assessment Summary, CEPA 1999. [http://www.ec.gc.ca/subsnouvelles-newsubs/default.asp?lang=En&n=71588E9F-1 ''Trichoderma reesei'' P345A]
* https://web.archive.org/web/20061006145854/http://www.eere.energy.gov/cleancities/progs/afdc/vwbs2.cgi?200
* [https://web.archive.org/web/20070319220040/http://www.nysaes.cornell.edu/ent/biocontrol/pathogens/trichoderma.html Trichoderma spp].
* [https://web.archive.org/web/20061015220341/http://www.genencor.com/wt/gcor/pr_1059584144 from genencor DNA sequence] and [https://web.archive.org/web/20070405105320/http://trichoderma.iq.usp.br/ Trichoderma reesei EST Database and Mitochondrial Genome].
*{{cite journal |vauthors=Nidetzky B, Steiner W, Claeyssens M |title=Cellulose hydrolysis by the cellulases from ''Trichoderma reesei'': adsorptions of two cellobiohydrolases, two endocellulases and their core proteins on filter paper and their relation to hydrolysis |journal=Biochem. J. |volume=303 |issue=Pt 3 |pages=817–23 |date=November 1994 |pmid=7980450 |pmc=1137620 }}
* {{cite journal |vauthors=Miettinen-Oinonen A, Suominen P |title=Enhanced production of Trichoderma reesei endoglucanases and use of the new cellulase preparations in producing the stonewashed effect on denim fabric |journal=Appl. Environ. Microbiol. |volume=68 |issue=8 |pages=3956–64 |date=August 2002 |pmid=12147496 |pmc=124001 |url=http://aem.asm.org/cgi/content/full/68/8/3956 |doi=10.1128/AEM.68.8.3956-3964.2002}}
* [https://www.megazyme.com/downloads/en/data/E-CBHI.pdf CBH I from Trichoderma sp.].{{dead link|date=July 2015}}
* [http://www.megazyme.com/booklets/ECBHII.pdf CBH I from Trichoderma sp.].{{dead link|date=July 2015}}
* {{cite journal |vauthors=Medve J, Ståhlberg J, Tjerneld F |title=Adsorption and synergism of cellobiohydrolase I and II of Trichoderma reesei during hydrolysis of microcrystalline cellulose |journal=Biotechnol. Bioeng. |volume=44 |issue=9 |pages=1064–73 |date=November 1994 |pmid=18623023 |doi=10.1002/bit.260440907 }}

{{DoE}}
{{Taxonbar|from=Q2749932}}

{{DEFAULTSORT:Trichoderma Reesei}}
[[Category:Trichoderma]]
[[Category:Ethanol]]

{{Hypocreales-stub}}

Orthonairovirus

2018-03-23T00:00:23Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q3067948}} (1 sig. taxon ID); WP:GenFixes, using AWB

{{Taxobox

| virus_group = v
| ordo = ''[[Bunyavirales]]''
| familia = ''[[Nairoviridae]]''
| genus = '''''Orthonairovirus'''''
| type_species = ''[[Dugbe virus]]''
| subdivision_ranks = Species
| subdivision =
[[Abu Hammad virus]] 
[[Abu Mina virus]] 
[[Artashat virus]] 
[[Avalon virus]] 
[[Bandia virus]] 
[[Burana virus]] 
[[Caspiy virus]] 
[[Chim virus]] 
[[Clo Mor virus]] 
[[Crimean-Congo hemorrhagic fever virus]] 
[[Dera Ghazi Khan virus]] 
[[Dugbe virus]] 
[[Erve virus]] 
[[Farallon virus]] 
[[Ganjam virus]] 
[[Geran virus]] 
[[Gossas virus]] 
[[Hazara virus]] 
[[Hughes virus]] 
[[Issyk-Kul virus]] 
[[Kasokero virus]] 
[[Keterah virus]] 
[[Kupe virus]] 
[[Leopards Hill virus]] 
[[Nairobi sheep disease virus]] 
[[Paramushir virus]] 
[[Puffin Island virus]] 
[[Punte Salinas virus]] 
[[Qalyub orthonairovirus|Qalyub virus]] 
[[Raza virus]] 
[[Sakhalin virus]] 
[[Soldado virus]] 
[[Tamdy virus]] 
[[Thiafora virus]] 
[[Tillamook virus]] 
[[Tofla virus]] 
[[Yogue virus]] 
[[Zirqa virus]]
}}

'''''Orthonairovirus''''' is a genus in the family ''[[Nairoviridae]]'' of the order [[Bunyavirales]] that include viruses with circular, [[RNA virus#Group V - negative-sense ssRNA viruses|negative-sense single stranded RNA]]. It got its name from the Nairobi sheep disease that affects the gastrointestinal tracts of sheep and goats. The vast majority, and perhaps all viruses in this genus are [[arbovirus|tick-borne virus]]es that can have human or other vertebrate hosts.<ref name=crabtree>Crabtree, Mary B., Rosemary Sang, and Barry R. Miller. "Kupe Virus, a New Virus in the Family Bunyaviridae, Genus Nairovirus, Kenya." Emerging Infectious Diseases 15 (2009): 147–54.</ref>

==Taxonomy==

The Nairobi sheep disease virus (NSD) is the prototype virus of the genus.<ref name=crabtree/><ref>Clerx, John PM, Jordi Casals, and David HL Bishop. "Structural Characteristics of Nairoviruses (Genus Nairovirus, Bunyaviridae)." Journal of General Virology 55 (1981): 165–78.</ref> NSD is found in East and Central Africa, and causes acute hemorrhagic gastroenteritis in sheep and goats.

===Serogroups===

The genus is divided into at least nine [[serogroup]]s. The Hughes and Sakhalin serogroups appear to be sister groups.

====Serogroups====

Crimean-Congo hemorrhagic fever 
Dera Ghazi Khan 
Hughes 
Kasokero 
Nairobi sheep disease 
Qalyub 
Sakhalin 
Tamdy 
Thiafora

Within each serogroup are a number of related viruses:

'''Crimean-Congo hemorrhagic fever serogroup''':
:[[Crimean-Congo hemorrhagic fever virus]]
:[[Hazara virus]]
:[[Tofla virus]]

'''Dera Ghazi Khan serogroup''':
:[[Abu Hammad virus]]
:[[Abu Mina virus]]

'''Hughes serogroup''':
:[[Farallon virus]]
:[[Hughes virus]]
:[[Puffin Island virus]]
:[[Punte Salinas virus]]
:[[Raza virus]]
:[[Soldado virus]]
:[[Zirqa virus]]

'''Kasokero serogroup''':
:[[Kasokero virus]]
:[[Yogue virus]]

'''Nairobi sheep disease serogroup''':
:[[Dugbe virus]]
:[[Ganjam virus]]
:[[Kupe virus]]
:[[Nairobi sheep disease virus]]

'''Qalyub serogroup''':
:[[Bandia virus]]
:[[Chim virus]]
:[[Geran virus]]
:[[Qalyub orthonairovirus|Qalyub virus]]

'''Sakhalin serogroup''':
:[[Avalon virus]]
:[[Paramushir virus]]
:[[Sakhalin virus]]
:[[Tillamook virus]]

'''Tamdy serogroup''':
:[[Burana virus]]
:[[Tamdy virus]]

'''Thiafora serogroup''':
:[[Erve virus]]
:[[Thiafora virus]]

== Genome ==

Nairovirus genomes are [[Antisense RNA|negative sense, single-stranded RNA]]. The complete [[genome]] is about 17,100–22,800 [[nucleotide]]s long, and is divided into three segments: large, medium, and small.<ref name=crabtree/> The large segment is about 11000–14400 nucleotides long (11–14.4 kb), and it encodes the viral polymerase.<ref name=osmond>Büchen-Osmond, Cornelia. "00.011.0.03. Nairovirus." ICTVdb Virus Descriptions. 25 Apr. 2006. International Committee on Taxonomy of Viruses. 17 Apr. 2009 <http://phene.cpmc.columbia.edu/ICTVdB/00.011.0.03.htm>.</ref><ref name=nairo>"Nairovirus." Nairovirus. Viral Zone. <http://www.expasy.ch/viralzone/all_by_species/251.html>.</ref> The medium segment is about 4,400–6,300 nucleotides long (4.4–6.3 kb), and it encodes for [[glycoprotein]]s G¬n and Gc.<ref name=osmond/><ref name=nairo/> The small segment is about 1,700–2,100 nucleotides long (1.7–2.1 kb), and it encodes the nucleocapsid protein.<ref name=crabtree/><ref name=osmond/><ref name=nairo/>

The genome has terminally redundant sequences, with the sequences being repeated at both ends. The terminal nucleotides are base-paired forming, non-covalently closed, [[circular RNA]].<ref name=osmond/> Both the 5’ and 3’ ends have conserved regions, 9 nucleotides in length. The sequences are, 5’end: UCUCAAAGA, and 3’end: AGAGUUUCU.<ref name=osmond/>

== Virion ==

The [[Virus#Structure|virions]] for viruses in this genus have a spherical shape.<ref name=nairo/> They range in size from about 80–120 nm in diameter, with 50% of their weight attributed to proteins and 20–30% of their weight attributed to lipids.<ref name=osmond/> The ribonucleocapsid is filamentous, having a length of about 200-300 nm and a width of about 2–2.5 nm.<ref name=osmond/>

These [[virus#Structure|nucleocapsids]] are surrounded by a single envelope that has projections made of glycoproteins protruding from its surface. These projections evenly cover the surface of the virion, and are about 5–10 nm long.<ref name=osmond/> They aid in attachment to the host receptor in [[DNA replication|replication]].

== Replication ==

Nairoviruses attach to the host receptor by their Gn-Gc glycoprotein dimer.<ref name=nairo/> The virus is then [[endocytosis|endocytosed]] into the host cell via a [[vesicle (biology)|vesicle]]. The ribonucleocapsid segments are released into the [[cytoplasm]], commencing [[transcription (genetics)|transcription]].<ref name=nairo/> Transcription and replication occur within the cell, and the newly synthesized virions are released by [[budding]].

== Transmission & Distribution ==

Members of this viral genus infect many different [[vertebrate]] [[host (biology)|hosts]], and are transmitted via ticks.<ref name=osmond/>

Members of the genus Nairovirus may be found the world over, wherever their [[arthropod]] vectors and vertebrate hosts are found together.<ref name=nairo/>

==Evolution==

Phylogenetic analysis has shown that these viruses fall into two major monophyletic groups, the hard (''[[Ixodidae]]'') and soft (''[[Argasidae]]'') tick-vectored groups.<ref name=Honig2004>Honig JE, Osborne JC, Nichol ST (2004) The high genetic variation of viruses of the genus Nairovirus reflects the diversity of their predominant tick hosts. Virology 318(1):10-16</ref> Fossil and phylogenetic data places the hard tick-soft tick divergence between {{Ma|120}} and {{Ma|92}}. This suggests that the Nairoviruses have been associated with these ticks for over 100 million years.

Additionally, nairoviruses vectored by ticks of the genera ''[[Argas]]'', ''Carios'' and ''[[Ornithodoros]]'' form three separate monophyletic lineages, again supporting the suggestion of host-virus cospeciation.

The hard bodied tick serogroups are

* Crimean-Congo hemorrhagic fever
* Nairobi sheep disease
* Sakhalin
* Tamdy

The soft bodied tick serogroups are

* Hughes
* Dera Ghazi Khan
* Qalyub

The tick vectors for the Kasokero and Thiafora serogroups are not currently known.

==Clinical importance==

Only four viruses in this genus have, to date, been recognised as human pathogens:

* Crimean-Congo hemorrhagic fever virus
* Dugbe virus
* Nairobi sheep disease virus
* Kasokero virus.

A fifth— Erve virus —may also be pathogenic for humans.

==Notes and references==
{{Reflist}}

==External links==
* [http://www.expasy.org/viralzone/all_by_species/251.html '''Viralzone''': Nairovirus]

{{Taxonbar|from=Q3067948}}

[[Category:Nairoviridae]]

Teufelsfarn

2018-03-22T23:48:48Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q2748623}} (11 sig. taxon IDs); WP:GenFixe using AWB

{{Italic title}}
{{Taxobox
| name = ''Osmunda claytoniana'' Interrupted fern
| image = Osmunda claytoniana JSG.jpg
| image_width = 240px
| image_caption = Typical fertile fronds
| regnum = [[Plant]]ae
| divisio = [[Pteridophyta]]
| classis = [[Polypodiopsida]] / &emsp;Pteridopsida {{au|(disputed)}}
| ordo = [[Osmundales]]
| familia = [[Osmundaceae]]
| genus = ''[[Osmunda]]''
| sectio = ''[[Claytosmunda]]''
| species = '''''O. claytoniana'''''
| binomial = ''Osmunda claytoniana''
| binomial_authority = [[Carl Linnaeus|L.]]
}}

'''''Osmunda claytoniana''''', the '''interrupted fern''', is a [[fern]] native to [[East Asia|Eastern Asia]] and eastern [[North America]], in the [[Eastern United States]] and [[Eastern Canada]].

The [[botanical name|specific epithet]] is named after the English-born Virginian botanist [[John Clayton (botanist)|John Clayton]].<ref>Fernald's "Gray's Manual of Botany" (1950)</ref> "Interrupted" describes the gap in middle of the blade left by the fertile portions after they wither and eventually fall off.<ref>University of Wisconsin-Madison Arboretum, [http://uwarboretum.org/foa/plant_sale/Interrupted%20Fern.pdf Interrupted fern profile]</ref>

The plant is known from [[fossil]]s to have grown in [[Europe]], showing a previous [[circumboreal]] distribution. Fragmentary foliage resembling ''Osmunda claytoniana'' has been found in the fossil record as far back as the [[Triassic]], and is known as †''[[Osmunda claytoniites]]''. ''O. claytoniana'' is a paramount example of evolutionary stasis. Paleontological evidence indicates it has remained unchanged, even at the level of fossilized nuclei and chromosomes, for at least 180 million years.<ref name="pmid24653037">{{cite journal |vauthors=Bomfleur B, McLoughlin S, Vajda V |title=Fossilized nuclei and chromosomes reveal 180 million years of genomic stasis in royal ferns |journal=Science |volume=343 |issue=6177 |pages=1376–7 | date=March 2014 |pmid=24653037 |doi=10.1126/science.1249884 |url=}}</ref>

==Distribution==
[[File:OFH-018 Osmunda claytoniana fertile frond.png|thumb|upright|The fertile middle pinnae give the frond an "interrupted" gap]]

===North America===
In eastern North America it occurs in: the [[Great Lakes region]]; eastern Canada – in southern [[Manitoba]], [[Ontario]], [[Quebec]] (north to [[tree line]]); and east to [[Newfoundland and Labrador|Newfoundland]]; eastern United States – upper [[New England]] south through the [[Appalachian Mountains]] and Atlantic seaboard, into the [[Southeastern United States]] in [[Georgia (U.S. state)|Georgia]] and [[Alabama]]; and west across the [[Southern United States]] to [[Mississippi River]], and back up the [[Mississippi embayment]] through the [[Midwestern United States]] to the Great Lakes.

===Asia===
In eastern Asia, the fern is found in the [[subtropical]] and [[temperate]] Asia in: the [[Eastern Himalaya]], [[South Central China]] and [[East China|Eastern China]], [[Taiwan]], the [[Korean Peninsula]], the [[Ryukyu Islands]], and [[Japan]].

===Ecology===
''Osmunda claytoniana'' is found in humid zones, mostly in forests, but also in more open [[habitat]]s and [[biome]]s, although rarely in [[bog]]s. The interrupted fern is often found alongside [[Ostrich fern|ostrich]], [[Osmundastrum cinnamomeum|cinnamon]] and [[sensitive fern]]s.

==Description==
''Osmunda claytoniana'' [[frond]]s are [[bipinnate]], {{convert|40|–|100|cm|in|abbr=on|0}} tall and {{convert|20|–|30|cm|in|abbr=on|0}} broad, the blade formed of alternate segments forming an arching blade tightening to a pointed end. The lower end is also slightly thinner than the rest of the frond because the first segments are shorter. Three to seven short, cinnamon-colored fertile segments are inserted in the middle of the length, giving the plant its name.

In their absence, the plant in all its stages appears similar to ''[[Osmundastrum cinnamomeum]]'' (cinnamon fern). The base of the segments distinguishes the two species: where ''O. cinnamomeum'' has typical felt-like [[trichome|hairs]], the few hairs present on ''O. claytoniana'' are extremely short, usually requiring a [[magnifying glass]] to see well.

Like other species in the family Osmundaceae, it grows a very large [[rhizome]], with persistent stipe bases from previous years. It forms small, dense [[clonal colony|colonies]], spreading locally through its rhizome, and often forming [[fairy ring]]s.

[[File:Interruptedfern-evening.jpg|left|thumb|Interrupted fern in evening light]]

===Hybrids===
''[[Osmunda × ruggii]]'', is a hybrid between ''O. claytoniana'' and ''[[Osmunda spectabilis|O. spectabilis]]'' (American royal fern). The hybrid is considered important because it suggests a closer genetic relationship between ''O. claytoniana'' and ''O. spectabilis'' than between ''O. claytoniana'' and ''O. cinnamomeum'' (a fact which has led to moving ''O. cinnamomeum'' out of ''Osmunda'' and into its own genus ''Osmundastrum''). ''Osmunda'' × ''ruggii'' is sterile and is known from only about two natural populations, despite the many areas in which both ''O. claytoniana'' and ''O. spectabilis'' are found.<ref>{{Cite web | url = http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500836 | title = 4. Osmunda ruggii R. M. Tryon | postscript = . }}</ref>

==Uses==
===Medicinal===
The [[Iroquois]] used the plant as [[medicinal plant|treatment]] for blood disorders and venereal diseases.<ref>[http://www.maquah.net/BritBrn/ethnobotanical/Osmundaceae.htm Univ. Mich.-Dearborn College of Arts, Sciences, and Letters: Native American Ethnobotany: ''Osmunda'' species] (scroll for ''O. claytoniana'') . accessed 12.1.2011</ref>

===Culinary===
Unlike those of the ostrich fern, the interrupted fern's [[fiddlehead]]s are not readily edible, due to their bitter taste and a tendency to cause diarrhea. The base of the stipe and very young buds are edible. Overuse may kill the crown.{{citation needed|date=February 2013}}

===Cultivation===
''Osmunda claytoniana'' is cultivated as an [[ornamental plant]] for use in traditional, [[native plant]], and [[wildlife garden]]s; for woodlands and [[natural landscaping]]; and for [[habitat]] [[ecological restoration|restoration]] projects. Their spreading [[Colony (biology)|colonizing]] habit can be used for some slope stabilization and erosion control measures.{{citation needed|date=February 2013}}

==References==
{{commons category|Osmunda claytoniana|''Osmunda claytoniana''}}
{{Reflist}}
{{refbegin}}
*[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=200002994 Flora of North America: ''Osmunda claytoniana''] [http://www.efloras.org/object_page.aspx?object_id=5111&flora_id=1 RangeMap:]
*[http://tai2.ntu.edu.tw/udth/bin/fot1.exe/browse?bid=1&page=82 Flora of Taiwan: ''Osmunda claytoniana'']
*{{cite book | author=Lamoureux, Gisèle and al. | title=Fougères, prêles et lycopodes | publisher=Fleurbec | year=1993 | isbn=2-920174-13-4}}
* Phipps, C. J., Taylor, T. N., Taylor, E. L., Cuneo, N. R., Boucher, L. D., and Yao, X. (1998). ''Osmunda'' (Osmundaceae) from the Triassic of Antarctica: An example of evolutionary stasis. American Journal of Botany, 85: 888–895
{{refend}}

{{Taxonbar|from=Q2748623}}

[[Category:Osmundaceae]]
[[Category:Fern species]]
[[Category:Pteridophyta of the Americas]]
[[Category:Pteridophyta of Asia]]
[[Category:Flora of China]]
[[Category:Flora of Japan]]
[[Category:Flora of Korea]]
[[Category:Flora of Taiwan]]
[[Category:Flora of the Northeastern United States]]
[[Category:Flora of the North-Central United States]]
[[Category:Flora of the Southeastern United States]]
[[Category:Flora of Eastern Canada]]
[[Category:Flora of Manitoba]]
[[Category:Edible plants]]
[[Category:Plants used in traditional Native American medicine]]
[[Category:Plants described in 1753]]

Acanthamoeba castellanii mamavirus

2018-03-22T23:45:58Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q1769257}} (1 sig. taxon ID); WP:GenFixes, using AWB

{{Taxobox
| name =
| image =
| image_caption =
| virus_group = i
| familia = [[Mimiviridae]]
| genus =
| subdivision_ranks = Species
| subdivision = Mamavirus
}}

'''Mamavirus''' is a large and complex [[viruses|virus]] in the [[DNA virus#Group I: dsDNA viruses|Group I]] family [[mimiviridae]].<ref name=Etten2010>{{cite journal|last=Van Etten|first=James L.|title=DNA Viruses: The Really Big Ones (Giruses)|pmc=2936810|publisher=Annual Reviewof Microbiology|author2=Leslie C. Lane |author3=David D. Dunigan |pages=83–99|date=13 October 2010|pmid=20690825|doi=10.1146/annurev.micro.112408.134338|journal=Annual Review of Microbiology|volume=64}}</ref> The virus is exceptionally large, and larger than many [[bacteria]]. Mamavirus and other mimiviridae belong to [[nucleocytoplasmic large DNA virus]] (NCLDVs) family.<ref name=Claverie2009>{{cite web|last=Claverie|first=Jean-Michel|title=Mimivirus and its Virophage|url=http://www.annualreviews.org/doi/abs/10.1146/annurev-genet-102108-134255|publisher=Annual Review of Genetics|author2=Chantal Abergel|pages=49–66|year=2009}}</ref> Mamavirus can be compared to the similar complex virus [[mimivirus]]; mamavirus was so named because it is similar to but larger than mimivirus.

==Discovery==

Mamavirus was first reported in September 2008. Like mimivirus, mamavirus was isolated from an [[amoeba]] in a cooling tower. The mimiviridae were not discovered until recently because of their size; when filtered the mimiviridae stay with the bacteria which led scientists to believe they were also bacteria. Mimivirus was first isolated in 1992 when scientists were looking for the cause of a pneumonia outbreak in Bradford UK. Due to its size it was named “Bradford coccus” and put in a freezer with scientists thinking it was a bacterium.<ref name="Claverie2009"/> A decade later, Jean-Michel Claverie and Didier Raoult discovered “Bradford coccus” was no bacterium when they tried to digest the cell wall with no success. Deciding to take a different route they looked at it under an [[electron microscope]]. They were surprised to find that it looked like a giant '''[[Iridoviridae|iridovirus]]''', which are icosahedral viruses that infect insects, fish, and frogs. Knowing this paved the way for the discovery of mamavirus because scientists knew to look for other giant viruses.<ref name=Ehrenberg2009>{{cite journal|last=Ehrenberg|first=Rachel|title=Enter the Virosphere: As evidence of the influence of viruses escalates, appreciation of these master manipulators grows|doi=10.1002/scin.5591760820|publisher=Science News|pages=22–25|date=10 October 2009|journal=Science News|volume=176|issue=8}}</ref>

It was originally isolated from ''[[Acanthamoeba]] [[Acanthamoeba polyphaga|polyphaga]]'', but subsequent work has involved ''[[Acanthamoeba castellanii]]''.<ref name="pmid21705471">{{cite journal |vauthors=Colson P, Yutin N, Shabalina SA, etal |title=Viruses with more than 1000 genes: Mamavirus, a new Acanthamoeba castellanii mimivirus strain, and reannotation of mimivirus genes |journal=Genome Biol Evol |volume= 3|issue= |pages= 737–42|date=June 2011 |pmid=21705471 |doi=10.1093/gbe/evr048 |url=http://gbe.oxfordjournals.org/cgi/pmidlookup?view=long&pmid=21705471 |pmc=3163472}}</ref>

==Structure and genome==

Mamavirus, like other mimiviridae, is icosahedral with a core [[capsid]] and a peripheral fiber layer. It contains a linear double-stranded DNA [[genome]] which has a very high coding density that is characteristic of NCLDVs. The mimiviridae contain very similar genomes due to gene duplications, and a fair piece of the genome is associated with functions not previously found in a virus.<ref name="Etten2010"/>

==Replication ==

Mamavirus possesses its own [[Transcription (genetics)|transcription]] machinery, and it packages transcription proteins in its particles. Transcription is believed to occur in the core particles. The core releases viral DNA and forms a cytoplasmic replication factory where DNA replication begins and transcription of late genes occurs. The replication factory forms around the viral core and expands until it occupies a large fraction of the amoeba cell volume. Later stages of the replication cycle involve partially assembled procapsids undergoing DNA packaging.<ref name="Etten2010"/>

==Sputnik virophage==

While the mimiviridae were a surprise themselves, mamavirus contained an even bigger surprise. While looking at mamavirus under the electron microscope, Raoult discovered a second, small virus closely associated with mamavirus which was named [[Sputnik virophage]], a satellite virus.<ref name=Pearson2008>{{cite web|last=Pearson|first=Helen|title='Virophage' suggests viruses are alive|url=http://www.igs.cnrs-mrs.fr/SpipInternet/IMG/pdf/PearsonNatureNews.pdf|publisher=Nature|page=677|date=7 August 2008}}</ref> Sputnik contains 21 genes and is tiny compared to mamavirus; however, it is quite powerful in its effects on mamavirus. Sputnik cannot replicate in acanthamoeba cells without a simultaneous infection by mamavirus (or mimivirus) so it infects the viral factory that mamavirus creates and hijacks it to replicate its own genome.<ref name=Scola2008>{{cite web|last=La Scola|first=Bernard|title=The virophage as a unique parasite of the giant mimivirus|url=http://jwbrown.mbio.ncsu.edu/MJC/old/20082009/Kristen_paper.pdf|publisher=Nature |author2=Christelle Desnues |author3=Isabelle Pagnier |author4=Catherine Robert |author5=Lina Barrassi |author6=Ghislain Fournous |author7=Michele Merchat |author8=Marie Suzan-Monti |author9=Patrick Forterre |author10=Eugene Koonin |author11=Didier Raoult|pages=100–105|date=4 September 2008}}</ref> This causes mamavirus to produce fewer viruses that are often deformed and less effective; there is also evidence of a partial thickening of the capsid. The fact that Sputnik can do this suggests that it is a viral parasite, and thus, was named the first virophage. A virophage is like [[bacteriophage]] viruses, which infects and sickens bacteria, but virophages infect viruses. Sputnik contains a circular double-stranded DNA of 18,343 base pairs, and is icosahedral in shape.<ref name=Desnues2010>{{cite web|last=Desnues|first=C|title=Inside the Lifestyle of the Virophage|url=http://content.karger.com/ProdukteDB/produkte.asp?Doi=312914|publisher=Intervirology|author2=D. Raoult|pages=293–303|date=15 June 2010}}</ref> Of the 21 genes it contains, eight encode proteins that have homologues. Of these eight, three are thought to be derived from mamavirus or mimivirus.<ref name=Sun2010>{{cite journal|last=Sun|first=Siyang|title=Structural Studies of the Sputnik Virophage|pmc=2798384|publisher=Journal of Virology|author2=Bernard La Scola |author3=Valorie D. Bownam |author4=Christopher M. Ryan |author5=Julian P. Whitelegge |author6=Didier Raoult |author7=Michael G. Rossmann |pages=894–897| date=January 2010 |pmid=19889775|volume=84|issue=2|doi=10.1128/JVI.01957-09|journal=Journal of Virology}}</ref> This indicates that Sputnik can participate in gene-transfer processes and mediate lateral gene transfer between giant viruses.<ref name=Smallridge2008>{{cite web|last=Smallridge|first=Rachel|title=Virology: A Virus gets a Virus|url=http://www.nature.com/nrmicro/journal/v6/n10/full/nrmicro2002.html|publisher=Nature Reviews|year=2008}}</ref>

==Implications ==

Mamavirus has caused scientists to review the criteria of life; to start questioning whether viruses are alive, revive the debate about the origin of DNA viruses and their possible role in the emergence of the eukaryotic nucleus.<ref name="Pearson2008"/>

== See also==
*[[Mimivirus]]
*[[Sputnik virophage]]
*[[Marseillevirus]]

==References==
{{Reflist}}

==External links==
*[http://www.radiolab.org/story/shrink/ Radiolab: Shrink] (podcast episode on "giant viruses" such as Mimivirus, Mamavirus and Megavirus)

{{Taxonbar|from=Q1769257}}

[[Category:Mimiviridae]]

Westlicher Knopfbusch

2018-03-22T23:22:17Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q4992358}} (11 sig. taxon IDs); WP:GenFixe using AWB

{{Italic title}}
{{taxobox
|image = Cephalanthus occidentalis.jpg
|image_caption = ''C. o.'' var. ''occidentalis''
|regnum = [[Plant]]ae
|unranked_divisio = [[Flowering plant|Angiosperms]]
|unranked_classis = [[Eudicots]]
|name=Buttonbush
|unranked_ordo = [[Asterids]]
|ordo = [[Gentianales]]
|familia = [[Rubiaceae]]
|subfamilia = [[Cinchonoideae]]
|tribus = [[Naucleeae]]
|genus = ''[[Cephalanthus]]''
|species = '''''C. occidentalis'''''
|binomial = ''Cephalanthus occidentalis''
|binomial_authority = [[Carl Linnaeus|L.]], 1753<ref name="GRIN">{{GRIN | accessdate=2010-01-04}}</ref>
|subdivision_ranks = [[Variety (botany)|Varieties]]
|subdivision = ''C. o.'' var. ''californicus'' 
''C. o.'' var.'' occidentalis''
|range_map = Cephalanthus occidentalis range map 1.png
|range_map_caption = Natural range of ''Cephalanthus occidentalis''
}}

'''''Cephalanthus occidentalis''''' is a species of [[flowering plant]] in the [[Coffea|coffee]] family, [[Rubiaceae]], that is native to eastern and southern [[North America]]. Common names include '''buttonbush''', '''common buttonbush''', '''button-willow''' and '''honey-bells'''.

==Description==
''C. occidentalis'' is a [[deciduous]] [[shrub]] or small [[tree]] that averages {{convert|1|-|3|m|ft|abbr=on}} in height, but can reach {{convert|6|m|ft|abbr=on}}. The [[leaf|leaves]] are opposite or in whorls of three, elliptic to ovate, {{convert|7|-|18|cm|in|abbr=on}} long and {{convert|4|-|10|cm|in|abbr=on}} broad, with a smooth edge and a short [[Petiole (botany)|petiole]]. The [[flower]]s are arranged in a dense spherical [[inflorescence]] {{convert|2|-|3.5|cm|in|abbr=on}} in diameter on a short [[Peduncle (botany)|peduncle]]. Each flower has a fused white to pale yellow four-lobed [[Corolla (flower)|corolla]] forming a long slender tube connecting to the [[sepal]]s. The [[gynoecium|stigma]] protrudes slightly from the corolla. The [[fruit]] is a spherical cluster of [[achene]]s (nutlets).<ref>{{cite web |url=http://www.fs.fed.us/rm/pubs_other/iitf_gtr026.pdf |title=''Cephalanthus occidentalis'' L. buttonbush |format=PDF |work=Wildland Shrubs of the United States and its Territories: Thamnic Descriptions |publisher=United States Forest Service |accessdate=2009-09-14}}</ref>

==Taxonomy==

There are two [[variety (biology)|varieties]], not considered distinct by all authorities:
*''Cephalanthus occidentalis'' var. ''occidentalis'' (syn. var. ''pubescens'') – common buttonbush. Eastern North America from [[Nova Scotia]] west to [[Minnesota]] and south to [[Florida]] and eastern [[Texas]].
*''Cephalanthus occidentalis'' var. ''californicus'' – California button-willow. Southwestern North America, from western Texas west to [[California]] ([[Sierra Nevada (U.S.)|Sierra Nevada]] foothills, [[San Joaquin Valley]], [[Sacramento Valley]], and the [[Inner North Coast Ranges]]) and south to [[Mexico]] and [[Central America]].

==Habitat==
Buttonbush is a common shrub of many [[wetland]] habitats in its range, including [[swamp]]s, [[floodplain]]s, [[mangrove]], [[pocosin]], [[riparian zone]]s, and moist forest [[understory]].<ref name=fs>{{cite web |url=http://www.fs.fed.us/database/feis/plants/shrub/cepocc/all.html |title=Cephalanthus occidentalis |work=Fire Effects Information System |publisher=United States Forest Service}}</ref> It is a member of the flora in the [[Everglades]].<ref name=fs/>

==Ecology==
[[Waterfowl]] and other [[bird]]s eat the seeds. [[Wood duck]]s utilize the plant as [[Bird nest|nest]] protection. [[Deer]] [[Browsing (predation)|browse]] the foliage. [[Insect]]s and [[hummingbird]]s take the [[nectar]], with [[honeybee|bees]] using it to make [[honey]].<ref name=fs/><ref name=usda/>

==Distribution==
The species occurs in eastern North America with disjunct populations occurring in the west. In [[Canada]], it occurs from southern [[Ontario]] and [[Quebec]] east to [[New Brunswick]]. Besides the eastern United States, and eastern regions of the [[Midwestern United States|Midwest]], notable areas range into [[Arizona]], the [[Mogollon Rim]], and other mountain ranges; in California, the entire [[San Joaquin Valley]]<ref>Little. ''Atlas of United States Trees, Volume 3, Minor Western Hardwoods'', Little, Elbert L, 1976, US Government Printing Office. Library of Congress No. 79-653298. Map 34-NW, Map 34-SW, ''Cephalanthus occidentalis''.</ref> West of the Great Plains and the Rocky Mountains, only western Texas, Arizona, and California find ''C. occidentalis''.
[[File:Buttonbush -- Cephalanthus occidentalis.jpg|thumb|400|Buttonbush -- Cephalanthus occidentalis from East Texas]]

==Uses==

===Medicinal===
''C. occidentalis'' has a number of historical [[Medicinal plant|medicinal]] uses, but it is also [[Toxin|toxic]] due to the presence of [[cephalathin]].<ref name=fs/><ref name=usda>{{cite web |url=http://plants.usda.gov/plantguide/pdf/pg_ceoc2.pdf |format=PDF |title=Common Buttonbush ''Cephalanthus occidentalis'' L. |work=Natural Resources Conservation Service Plant Guide |publisher=United States Department of Agriculture}}</ref>

===Cultivation===
Buttonbush is cultivated as an [[ornamental plant]] for a nectar source or '[[List of honey plants|honey plant]]' and for aesthetics in [[garden]]s and [[native plant]] landscapes, and is planted on slopes to help control [[erosion]].<ref>{{cite book |url=https://books.google.com/books?id=VSZmm76pVw8C& |title=The Homeowner's Complete Tree & Shrub Handbook: The Essential Guide to Choosing, Planting, and Maintaining Perfect Landscape Plants |first=Penelope |last=O'Sullivan |publisher=Storey Publishing |year=2007 |isbn=978-1-58017-571-5 |page=197}}</ref> Buttonbush is a suitable shrub for butterfly gardens.

==San Joaquin Valley landmark tree==
The town of [[Buttonwillow, California]] was named for the buttonbush (''Cephalanthus occidentalis''). A lone buttonbush served as a landmark on an old trans-San Joaquin Valley trail, and was used by ancient [[Yokut people|Yokut]] Indians as a meeting place. It later became the site of settlers' stock [[rodeo]]s. This buttonbush tree is listed as [[California Historical Landmark]] No. 492, and is now known as the "Buttonwillow Tree."

==References==
{{Reflist}}

==External links==
{{Wikispecies-inline}} 
{{commonscat-inline|Cephalanthus occidentalis|''Cephalanthus occidentalis''}}
*{{cite web |url=http://esp.cr.usgs.gov/data/atlas/little/cephocci.pdf |format=PDF |title=''Cephalanthus occidentalis'' |work=Digital Representations of Tree Species Range Maps from "Atlas of United States Trees" by Elbert L. Little, Jr. (and other publications) |publisher=United States Geological Survey |accessdate=}}
*[http://plants.usda.gov/java/profile?symbol=CEOC2 USDA Plants Profile: ''Cephalanthus occidentalis'']
*[http://www.missouriplants.com/Whiteopp/Cephalanthus_occidentalis_page.html Missouriplants: ''Cephalanthus occidentalis'' var. ''occidentalis'']
*[http://ucjeps.berkeley.edu/cgi-bin/get_JM_treatment.pl?Cephalanthus+occidentalis Jepson Flora Project: ''Cephalanthus occidentalis'' var. ''californicus'']
*[http://calphotos.berkeley.edu/cgi/img_query?query_src=photos_index&where-taxon=Cephalanthus+occidentalis Photo gallery]

{{Taxonbar|from=Q4992358}}

[[Category:Cephalanthus|occidentalis]]
[[Category:Medicinal plants of North America]]
[[Category:Garden plants of North America]]
[[Category:Flora of Mexico]]
[[Category:Flora of Canada]]
[[Category:Flora of Guatemala]]
[[Category:Flora of Belize]]
[[Category:Flora of Honduras]]
[[Category:California Historical Landmarks]]
[[Category:Flora of the United States]]
[[Category:Plants described in 1753]]

Lysimachia asperulifolia

2018-03-22T23:07:19Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q6710172}} (7 sig. taxon IDs); WP:GenFixes using AWB

{{taxobox
|image = Lysimachia asperulifolia.jpg
|status = G3
|status_system = TNC
|regnum = [[Plantae]]
|unranked_divisio = [[Angiosperms]]
|unranked_classis = [[Eudicots]]
|unranked_ordo = [[Asterids]]
|ordo = [[Ericales]]
|familia = [[Primulaceae]]
|genus = ''[[Lysimachia]]''
|species = '''''L. asperulifolia'''''
|binomial = ''Lysimachia asperulifolia''
|binomial_authority = [[Poir.]]
|}}

'''''Lysimachia asperulifolia''''' ([[orth. var.]] ''L. asperulaefolia'') is a rare species of flowering plant in the [[Primulaceae]] known by the common name '''rough-leaved loosestrife''' and '''roughleaf yellow loosestrife'''. It is [[Endemism|endemic]] to the [[Atlantic coastal plain]] in [[North Carolina]] and northern [[South Carolina]] in the United States, where there are 64 known populations.<ref name=tnc>[http://www.natureserve.org/explorer/servlet/NatureServe?searchName=Lysimachia+asperulifolia ''Lysimachia asperulifolia''.] [[The Nature Conservancy]].</ref> It is a federally listed [[endangered species]] of the United States.

==Description==

This plant is a [[rhizome|rhizomatous]] perennial herb growing erect to a maximum height around 60 to 70 centimeters. The lower stem is pinkish in color and ribbed, and the upper stem is yellowish and lacks ribs. The stem in the [[inflorescence]] is covered in reddish glands. The leaves are green, lance-shaped, and up to 5 centimeters long by 2 wide. They are borne in whorls of three or four around the stem, or sometimes in opposite pairs. The leaves are not rough in texture as the common name would suggest.<ref name=cpc>[http://www.centerforplantconservation.org/collection/cpc_viewprofile.asp?CPCNum=2750 ''Lysimachia asperulifolia''.] {{webarchive|url=https://web.archive.org/web/20101215100124/http://centerforplantconservation.org/Collection/CPC_ViewProfile.asp?CPCNum=2750 |date=2010-12-15 }} Center for Plant Conservation.</ref> Smaller, tougher, brown-colored leaves are opposite or borne in whorls of up to 7 near the stem base. The top of the stem is occupied by the inflorescence, which is a [[raceme]] of star-shaped yellow flowers interspersed with leaflike green [[bract]]s. Each flower has 4 to 7, but usually five, yellow petals with wide bases and pointed, ragged tips. The petals and green [[sepal]]s are dotted with red glands and streaked with reddish resin canals. The fruit is a red-mottled straw-colored capsule a few millimeters in length.<ref name=tnc/><ref name=fna>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=250092252 ''Lysimachia asperulifolia''.] Flora of North America.</ref>

==Habitat==

This herb grows in a number of [[plant community|plant communities]] on the coastal plains of southern North Carolina and northern South Carolina, including [[pocosin]]s, [[sandhill]]s, [[Flatwoods|pine flatwoods]] and [[Eastern savannas of the United States|pine savannas]].<ref name=cpc/> It is most commonly found in open areas in the [[ecotone]] between [[Pinus palustris|longleaf pine]] [[Upland and lowland (freshwater ecology)|uplands]] and [[Pinus serotina|pond pine]] pocosins.<ref name=tnc/> The soil is seasonally wet to waterlogged or submerged, low in nutrients, rich in [[peat]] over sandy substrates.<ref name=tnc/> The plant is found in sections of these habitat types which, in their pristine state, are openings in a dense layer of [[shrubs]] which are kept open by severe periodic [[wildfire]]s. Fire prevents [[ecological succession]], maintaining a type of [[ecosystem]] in which the taller vegetation is kept small and sparse, allowing the herb layer to flourish in the sun.<ref name=tnc/> Trees, shrubs, and ferns in the area include ''[[Aronia arbutifolia]]'' (chokeberry), ''[[Clethra alnifolia]]'' (summersweet), ''[[Cyrilla|Cyrilla racemiflora]]'' (titi), ''[[Fothergilla gardenii]]'' (dwarf fothergilla), ''[[Ilex glabra]]'' (Appalachian tea), ''[[Magnolia virginiana]]'' (sweetbay), ''[[Osmunda cinnamomea]]'' (cinnamon fern), ''[[Persea palustris]]'' (swampbay), ''[[Symplocos tinctoria]]'' (yellowwood), and ''[[Vaccinium]]'' species (wild blueberries). Herbs, grasses, and [[moss]]es associated with this ecosystem include ''[[Andropogon glomeratus]]'' (bushy bluestem), ''[[Aristida stricta]]'' (wiregrass), ''[[Drosera intermedia]]'' (oblong-leaved sundew), ''[[Drosera capillaris]]'' (pink sundew), ''[[Lachnanthes caroliniana]]'' (redroot), ''[[Peltandra sagittifolia]]'' (spoon flower), ''[[Sarracenia flava]]'' (yellow pitcher plant), and ''[[Sphagnum]]'' species (sphagnum mosses).<ref name=tnc/>

In the 1980s the only known viable populations of the plant were located in [[Green Swamp (North Carolina)|Green Swamp Nature Preserve]] and the [[Croatan National Forest]], and at [[Military Ocean Terminal Sunny Point]].<ref name=rec>USFWS. [http://ecos.fws.gov/docs/recovery_plan/950419b.pdf ''Lysimachia asperulifolia'' Recovery Plan.] April 19, 1995.</ref> Another was located on [[Fort Bragg]].<ref name=rec/> A program of [[prescribed fire]] was instituted on military base territory in the region to preserve habitat for the [[red-cockaded woodpecker]] (''Picoides borealis''). This habitat was appropriate for the plant as well, and as a result some additional populations appeared where it had been crowded out by vegetation overgrowth before. One such population is located at [[Camp Lejeune]], and more occurrences grew at Fort Bragg.<ref name=rec/> The South Carolina occurrence is located on [[Fort Jackson (South Carolina)|Fort Jackson]]. In 1995 there were 64 populations known.<ref name=cpc/> Because the plant usually [[vegetative reproduction|reproduces vegetatively]] by sprouting from its rhizome, creating clones, what appears to be a large population may actually be relatively few genetic individuals with many cloned stems above the ground.<ref name=cpc/>

==Environmental Issues==

The main threat to the species is the loss and degradation of its habitat.<ref name=cpc/> Much of the land in the region has been developed, the wetlands drained and dried to create land for residential, industrial, and recreational purposes.<ref name=tnc/> Habitat that remains is improperly managed, becoming degraded as the natural [[fire regime]] is prevented. As [[fire suppression]] practices have been implemented, the pocosins, sandhills, and swamps have overgrown with brush and woody vegetation. Shrubs are not kept back and they grow to large sizes and shade out the smaller plants in the herb layer. Even where fire is prescribed or allowed to take place, pocosins do not burn thoroughly enough to remove the heavy organic layers that accumulate there.<ref name=tnc/> Activities involved with fire may also be detrimental, as [[firebreak]]s are often plowed in the very ecotone openings where the plant most often grows.<ref name=tnc/><ref name=rec/> [[Agriculture]], including pine plantations, and other types of operations alter the [[hydrology]] of the wetlands, making conditions unsuitable for this and other native plants.<ref name=tnc/><ref name=rec/> Many of the populations are located on military bases and are vulnerable to destruction during military operations.<ref name=rec/>

==References==
{{Reflist}}

==External links==
*[http://plants.usda.gov/java/profile?symbol=LYAS2 USDA Plants Profile for ''Lysimachia asperulifolia'']

{{Taxonbar|from=Q6710172}}

[[Category:Lysimachia|asperulifolia]]
[[Category:Flora of North Carolina]]
[[Category:Flora of South Carolina]]
[[Category:Endemic flora of the United States]]
[[Category:Vulnerable flora of the United States]]

Amerikanische rote Himbeere

2018-03-22T22:45:32Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q210156}} (5 sig. taxon IDs); WP:GenFixes, using AWB

{{taxobox
|image = Rubus_strigosus_8782.JPG
|image_caption = ''Rubus strigosus'' near [[Matanuska Glacier]], [[Alaska]]
|regnum = [[Plant]]ae
|unranked_divisio = [[Angiosperms]]
|unranked_classis = [[Eudicots]]
|unranked_ordo = [[Rosids]]
|ordo = [[Rosales]]
|familia = [[Rosaceae]]
|genus = ''[[Rubus]]''
|subgenus = ''[[Rubus#Scientific classification|Idaeobatus]]''
|species = '''''R. strigosus'''''
|binomial = ''Rubus strigosus''
|binomial_authority = [[André Michaux|Michx.]]
|}}
[[File:Raspberry Leaves.JPG|thumb|''Rubus strigosus'': foliage showing the large leaf of a first-year shoot, and the smaller leaves of a second-year shoot]]

'''''Rubus strigosus''''', the '''American red raspberry''' or '''American raspberry''', is a species of ''[[Rubus]]'' native to much of [[North America]]. It has often been treated as a [[variety (biology)|variety]] or [[subspecies]] of the closely related Eurasian ''[[Rubus idaeus]]'' (raspberry or European raspberry),<ref name=usda>USDA Plants Profile: [http://plants.usda.gov/java/profile?symbol=RUIDS2 ''Rubus idaeus'' subsp. ''strigosus'']</ref><ref name=pbc>Plants of British Columbia: [http://linnet.geog.ubc.ca/Atlas/Atlas.aspx?sciname=Rubus+idaeus ''Rubus idaeus'' subsp. ''strigosus'']</ref> but currently is more commonly treated as a distinct species.<ref name=grin>{{GRIN | accessdate = 14 January 2018}}</ref><ref name=roland>Roland, A. E., & Smith, E. C. (1969, reprinted 1983). ''The Flora of Nova Scotia''. Halifax: Nova Scotia Museum.</ref><ref name=grignon>Grignon, T. (1992). The Dynamics of ''Rubus strigosus'' (Michx.) in Post-Clearcut Mixedwood and Softwood Forests of Nova Scotia. Thesis.</ref> Many of the commercial raspberry [[cultivar]]s grown for their fruit derive from [[Hybrid (biology)|hybrids]] between ''R. strigosus'' and ''R. idaeus''; see [[Raspberry]] for more details.

==Classification==
Botanists have long debated the taxonomic treatment of the Eurasian and American red raspberries, with some viewing all of these plants as members of a single, [[circumboreal]] [[species]] ''Rubus idaeus,'' and others recognizing two (or more) species within this group.<ref name=lhb>{{Cite journal | last1 = Bailey | first1 = L. H. | year = 1945 | title = Species Batorum. The genus ''Rubus'' in North America X. | url = | journal = Gentes Herbarum | volume = 5 | issue = | pages = 859–918 }}</ref><ref name=fernald>{{Cite journal | last1 = Fernald | first1 = M. L. | year = 1900 | title = Rubus idaeus'' and its variety anomalus in America | url = | journal = Rhodora | volume = 22 | issue = | pages = 195–200 }}</ref><ref name=fernald2>{{Cite journal | last1 = Fernald | first1 = M. L. | year = 1919 | title = Rubus idaeus'' and some of its variations in North America | url = | journal = Rhodora | volume = 21 | issue = | pages = 89–98 }}</ref><ref name=hodgdon>{{Cite journal | last1 = Hodgdon | first1 = A. R. | last2 = Pike | first2 = R. B. | year = 1964 | title = Flora of the Wolf Islands, New Brunswick. Part 2. Some phytogeographic considerations | url = | journal = Rhodora | volume = 66 | issue = | page = 140 }}</ref><ref name=whitney>Whitney, G. G. (1978). A demographic analysis of ''Rubus idaeus'' L. and ''Rubus pubescens'' Raf.: the reproductive traits and population dynamics of two temporally isolated members of the genus ''Rubus''. Ph.D. thesis, Yale University. 139 pp.</ref><ref>Fernald doubted this distinction, but Bailey and many other authors rely on it.</ref><ref name=grignon/><ref name=nickerson>{{Cite journal | last1 = Nickerson | first1 = N. L. | last2 = Hall | first2 = I. V. | year = 1978 | title = Large-flowered Trillium, ''Trillium grandiflorum'', in Nova Scotia | url = | journal = Canad. Field-Naturalist | volume = 92 | issue = 3| page = 291 }}</ref><ref name=freedman>Freedman, B. (1989). ''Environmental Ecology: The Impacts of Pollution and Other Stresses on Ecosystem Structure and Function''. Academic Press, Inc., San Diego.</ref> The two species share many similarities, and probably recently diverged from a common ancestor, leading to differences in taxonomic interpretation, particularly regarding the more intermediate eastern Asian plants. A common current treatment, followed here, is to classify the North American red raspberries as ''Rubus strigosus,'' and include only the Eurasian plants in ''Rubus idaeus.'' When the species are combined, as done in some recent publications, the Eurasian plants are ''Rubus idaeus'' ssp. ''idaeus'' (or ''Rubus idaeus'' var. ''idaeus),'' and the American plants ''R. idaeus'' ssp. ''strigosus'' (or ''R. idaeus'' var. ''strigosus).'' Different interpretations are also sometimes made regarding placement of various eastern Asian populations of this group, by some considered to represent additional varieties or subspecies, if not different species altogether. The most distinctive physical difference among these plants is usual presence of [[gland]]-tipped hairs on first-year canes, petioles, pedicels, and calyces of ''R. strigosus,'' lacking in ''R. idaeus.''

==Distribution==
''Rubus strigosus,'' as treated here, is widely distributed in North America, particularly the more boreal regions. Some authors also treat various raspberries in eastern Asia, east from the Aerhtal Shan (Altai) Mountain Range in Mongolia to Dongbei (Manchuria) and Japan in this taxon (where it is suggested to have originated along with a great deal of the North American flora),<ref name=fernald/> but others include all Asian raspberries in ''R. idaeus''.<ref name=grin2>{{GRIN | ''Rubus idaeus'' | 32348 | accessdate = 14 January 2018}}</ref> with the Eurasian plants being ''Rubus idaeus'' ssp. (or var.) ''idaeus''.

==Description==

''R. strigosus'' is a [[perennial plant]] which bears [[biennial plant|biennial]] stems ("canes") from the perennial root system. In its first year, a new stem grows vigorously to its full height of 0.5–2 m, unbranched, and bearing large pinnate [[leaf|leaves]] with three or five (rarely seven) leaflets; normally it does not produce any flowers. In its second year, the stem does not grow taller, but produces several side shoots, which bear smaller leaves with three leaflets.

The [[flower]]s are produced in late spring on short [[raceme]]s on the tips of these side shoots, each flower with five white [[petal]]s 4–7 mm long. The [[fruit]] is 1–1.2 cm diameter, red, edible, sweet but tart-flavored, produced in summer or early autumn; in [[botany|botanical]] terminology, it is not a [[Berry (botany)|berry]] at all, but an [[fruit#Aggregate fruit|aggregate fruit]] of numerous [[drupe]]lets around a central core.<ref name=pbc/><ref name=pfaf>Plants For A Future: [http://www.pfaf.org/database/plants.php?Rubus+strigosus ''Rubus strigosus'']</ref>

== References ==
{{Commons}}
{{Reflist|2}}

{{Taxonbar|from=Q210156}}

[[Category:Rubus|strigosus]]
[[Category:Berries]]
[[Category:Flora of Western Canada]]
[[Category:Flora of the Western United States]]
[[Category:Flora of the Rocky Mountains]]
[[Category:Flora of the North-Central United States]]
[[Category:Flora of the Great Lakes region (North America)]]
[[Category:Flora of Eastern Canada]]
[[Category:Flora of the Northeastern United States]]
[[Category:Flora of Idaho]]
[[Category:Flora of California]]
[[Category:Garden plants of North America]]
[[Category:Vines]]
[[Category:Flora of North America]]

Phyllodoce caerulea

2018-03-22T22:43:58Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q478437}} (11 sig. taxon IDs); WP:GenFixes, using AWB

{{Italic title}}
{{taxobox
| name = ''Phyllodoce caerulea''
| image = Phyllodoce caerulea LC0329.jpg|Phyllodoce caerulea001.jpg
| regnum = [[Plant]]ae
| unranked_divisio = [[Angiosperms]]
| unranked_classis = [[Eudicots]]
| unranked_ordo = [[Asterids]]
| ordo = [[Ericales]]
| familia = [[Ericaceae]]
| genus = ''[[Phyllodoce (plant)|Phyllodoce]]''
| species = '''''P. caerulea'''''
| binomial = ''Phyllodoce caerulea''
| binomial_authority = ([[Carl Linnaeus|L.]]) [[Cardale Babington|Bab.]]
| synonyms =
*''Andromeda caerulea'' L.
*''Menziesia caerulea'' (L.) Sw.
*''Bryanthus caeruleus'' (L.) Dippel
| synonyms_ref =  <ref name="Coker"/><ref name="Iberica">{{cite book |author=L. Villar |year=2003 |chapter=''Phyllodoce'' Salisb. |page=512 |series=Flora Iberica |volume=4 |title=Cruciferae–Monotropaceae |editor=S. Castroviejo |publisher=[[Consejo Superior de Investigaciones Científicas]] |isbn=9788400073855 |url=http://www.floraiberica.es/floraiberica/texto/pdfs/04_074_05_Phyllodoce.pdf |format=[[Portable Document Format|PDF]]}}</ref>
}}

'''''Phyllodoce caerulea''''', known as '''blue heath''' in [[British English]]<ref>{{cite book |author=[[Clive A. Stace]] |year=2010 |chapter=''Phyllodoce'' Salisb. – Blue Heath |page=526 |title=New Flora of the British Isles |edition=3rd |publisher=[[Cambridge University Press]] |isbn=978-0-521-70772-5}}</ref> and '''purple mountain heather'''<ref name="FNA"/> or '''blue mountainheath'''<ref>{{PLANTS|Id=PHCA10|taxon=Phyllodoce caerulea|accessdate=30 September 2015}}</ref> in [[American English]], is an evergreen species of [[dwarf shrub]] that grows up to around {{convert|15|cm|0|abbr=on}} tall, and bears clusters of 2–6 purple flowers. It is native to [[Subarctic climate|boreal]] regions around the [[Northern Hemisphere]], but with large gaps in its distribution.

==Description==
''Phyllodoce caerulea'' is a low [[shrub]], typically growing {{convert|5|-|15|cm|0}} high, and exceptionally reaching {{convert|25|cm|0|abbr=on}}.<ref name="Coker"/> Its evergreen leaves are {{convert|4|-|10|mm|abbr=on}} long and {{convert|1.7|-|3.6|mm|2|abbr=on}} wide, and are borne on {{convert|1|mm|2|adj=on}} long [[petiole (botany)|petioles]]; they are arranged alternately.<ref name="Coker"/>

The flowers are borne in clusters of 2–6; each flower is {{convert|8|-|12|mm|1|abbr=on}} long, with a [[corolla (flower)|corolla]] composed of five fused [[petal]]s that begin purple, but fade to a bluish pink.<ref name="Coker"/> These are surrounded by five [[sepal]]s, and themselves surround the 8–10 free [[stamen]]s and a [[superior ovary]] that produces [[nectar]] at its base.<ref name="Coker"/>

==Distribution==
''Phyllodoce caerulea'' has a patchy [[circumboreal]] distribution, with gaps between [[110th meridian west|110° W]] and [[155th meridian west|155° W]] and between [[70th meridian east|70° E]] and [[125th meridian east|125° E]].<ref name="Coker">{{cite journal |author1=P. D. Coker |author2=A. M. Coker |lastauthoramp=yes |year=1973 |title=''Phyllodoce caerulea'' (L.) Bab. |journal=[[Journal of Ecology]] |volume=61 |issue=3 |pages=901–913 |jstor=2258657}}</ref>

[[File:The Sow of Atholl from the A9 road.jpg|thumb|[[The Sow of Atholl]] from the north, including the site where ''P. caerulea'' was first discovered in the British Isles, in 1810.]]
In Europe, ''P. caerulea'' is found from [[Iceland]] to the [[Kanin Peninsula]].<ref name="Coker"/> Its Icelandic distribution is also disjunct, comprising the area around [[Eyjafjörður]] and a site near Desjarmyri.<ref name="Coker"/> In the [[British Isles]], ''P. caerulea'' is confined to a few sites in the [[Scottish Highlands]]. It was first discovered around a spring at an altitude of {{convert|740|m}} on the slopes of the [[Sow of Atholl]], but has since been found at a few sites in the [[Ben Alder]] forest.<ref name="Coker"/> It became a protected species in the UK in 1975 under the Conservation of Wild Creatures and Wild Plants Act.<ref>http://www.caithness.org/caithnessfieldclub/bulletins/1975/october/conservation.htm</ref> There are reports of the plant's occurrence in the [[Swiss Alps]], but no herbarium specimens have been found to confirm this.<ref name="Coker"/> The species has not been observed on the [[Faroe Islands]], [[Jan Mayen]], [[Bear Island (Norway)|Bjørnøya]], [[Svalbard]] or [[Franz Josef Land]].<ref name="Coker"/>

In Asia, ''Phyllodoce caerulea'' occurs in the [[Ural Mountains]], around [[Lake Baikal]] and in the [[Mongolia]]n [[Khangai Mountains|Khangai]] and [[Khentii Mountains|Kentii]] mountains, but is absent from most of central [[Siberia]]. It occurs on [[Hokkaido]], [[Sakhalin]], the [[Kamchatka Peninsula]] and in [[Beringia]].<ref name="Coker"/>

In North America, ''P. caerulea'' is found in coastal [[Alaska]], the [[Northwest Territories]], [[Quebec]] and [[Labrador]], as well as scattered sites in the [[Gaspé Peninsula]] and the [[White Mountains (New Hampshire)|White Mountains]] of [[New Hampshire]] and [[Vermont]].<ref name="Coker"/> It is widespread and common in [[Greenland]].<ref name="Coker"/> Its absence from the [[Yukon]] has been described as "surprising".<ref name="Coker"/>

==Taxonomy==
''Phyllodoce caerulea'' was first described by [[Carl Linnaeus]] in his 1753 ''{{lang|la|[[Species Plantarum]]}}'', as a species in the genus ''[[Andromeda (genus)|Andromeda]]''. It was transferred to the genus ''[[Phyllodoce (plant)|Phyllodoce]]'' by [[Charles Cardale Babington]] in his 1843 ''Manual of British Botany''.<ref name="FNA">{{cite book |author1=John G. Packer |author2=A. Joyce Gould |lastauthoramp=yes |year= |chapter=''Phyllodoce'' Salisbury, Parad. Lond. 1: plate 36. 1806 |title=Magnoliophyta: Paeoniaceae to Ericaceae |series=[[Flora of North America]] |volume=8 |publisher=[[Oxford University Press]] |isbn=978-0-19-534026-6 |url=http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=200016320}}</ref> In Japan, ''P. caerulea'' [[hybrid (biology)|hybridises]] with the pale yellowish-flowering species ''[[Phyllodoce (plant)|P. aleutica]]'' to produce [[F1 hybrid|F1 offspring]] with flowers that are pink, orange or striped in pink and yellowish white.<ref>{{cite journal |author1=Y. Kameyama, T. Kasagi |author2=G. Kudo |lastauthoramp=yes |year=2008 |title=A hybrid zone dominated by fertile F1s of two alpine shrub species, ''Phyllodoce caerulea'' and ''Phyllodoce aleutica'', along a snowmelt gradient |journal=[[Journal of Evolutionary Biology]] |volume=21 |issue=2 |pages=588–597 |doi=10.1111/j.1420-9101.2007.01476.x}}</ref>

==References==
{{Reflist|32em}}

==Further reading==
*{{cite journal |author=E. C. Nelson |year=1977 |title=The discovery in 1810 and subsequent history of ''Phyllodoce caerulea'' (L.) Bab. in Scotland |journal=Western Naturalist |volume=6 |pages=45–72}}

==External links==
*{{Commons category-inline|Phyllodoce caerulea|''Phyllodoce caerulea''}}
*{{Wikispecies-inline|Phyllodoce caerulea|''Phyllodoce caerulea''}}

{{Taxonbar|from=Q478437}}

[[Category:Ericoideae]]
[[Category:Arctic flora]]
[[Category:Flora of Europe]]
[[Category:Flora of North America]]
[[Category:Flora of temperate Asia]]
[[Category:Plants described in 1753]]

Metriacanthosaurus

2018-03-22T21:45:30Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q1957207}} (1 sig. taxon ID); WP:GenFixes, using AWB

{{Italic title}}
{{automatic taxobox
| name= ''Metriacanthosaurus''
| fossil_range = [[Middle Jurassic]], {{fossilrange|160}}
| image = Metriacanthosaurus.jpg
| image_width = 250px
| image_caption = Life restoration
| type_species = {{extinct}}''Megalosaurus parkeri''
| type_species_authority = [[Friedrich von Huene|von Huene]], 1923
| authority = [[Alick Walker|Walker]], 1964
| subdivision_ranks = [[Species]]
| subdivision =
{{extinct}}'''''Metriacanthosaurus parkeri''''' (von Huene, 1923)
| synonyms =
''[[Megalosaurus]] parkeri'' von Huene, 1926 
''[[Altispinax]] parkeri'' (von Huene, 1926)
}}

'''''Metriacanthosaurus''''' (meaning "moderately-spined lizard") is a [[genus]] of [[Metriacanthosauridae|metriacanthosaurid]] [[dinosaur]] from the upper [[Oxford Clay]] of [[England]], dating to the mid-[[Jurassic Period]], about 160 million years ago (lower [[Oxfordian (stage)|Oxfordian]]).

==History of discovery==
[[File:Metriacanthosaurus ilium.png|thumb|left|Reconstruction of the [[ilium bone|ilium]]]]
In 1923, German paleontologist [[Friedrich von Huene]] wrote a paper on [[Jurassic]] and [[Cretaceous]] European carnivorous dinosaurs. In this paper, he examined a specimen, OUM J.12144, including an incomplete hip, a leg bone, and part of a backbone, assigning it to a new species of ''[[Megalosaurus]]'': ''Megalosaurus parkeri''. The [[specific name (zoology)|specific name]] honours W. Parker who in the nineteenth century had collected the fossils near Jordan's Cliff at [[Weymouth, Dorset|Weymouth]].<ref>{{cite journal | first = F. | last = von Huene | authorlink = Friedrich von Huene | date = 1923 | title = Carnivorous Saurischia in Europe since the Triassic | journal = Bulletin of the Geological Society of America | volume = 34 | pages = 449–458 |doi=10.1130/GSAB-34-449}}</ref> These bones were from the [[Oxford Clay Formation]], which is from the [[Upper Jurassic]].<ref name="naish&martill07"/>

In 1932, however, von Huene concluded it was species of ''[[Altispinax]]'', ''A. parkeri''.<ref>{{cite journal | first = F. | last = von Huene | authorlink = Friedrich von Huene | date = 1932 | title = Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte | journal = Monographien zur Geologie und Palaeontologie | volume = 1 | issue = 4 | pages = 361 }}</ref>

In 1964, scientist [[Alick Walker]] decided these fossils were too different from ''Altispinax'', as it lacked the long vertebral spines, and named a new genus, ''Metriacanthosaurus''.<ref>{{cite journal | first = Alick D. | last = Walker | authorlink = Alick Walker | date = 1964 | title = Triassic reptiles from the Elgin area: ''Ornithosuchus'' and the origin of carnosaurs | journal = Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences | volume = 248 | pages = 53–134 | doi=10.1098/rstb.1964.0009}}</ref> The generic name is derived from Greek ''metrikos'', "moderate", and ''akantha'', "spine". ''Metriacanthosaurus'' thus gets its name from its [[vertebrae]], which are taller than typical carnosaurs, like ''[[Allosaurus]]'', but lower than other high-spined dinosaurs like ''[[Acrocanthosaurus]]''.

==Description==
''Metriacanthosaurus'' was a medium-sized theropod with a femur length of eighty centimetres. [[Gregory S. Paul]] in 1988 estimated its weight at a tonne.<ref name="paul88">{{cite book|last=Paul|first=Gregory S.|authorlink = Gregory S. Paul|date=1988|title=Predatory Dinosaurs of the World|publisher=Simon & Schuster|location=New York}}</ref> ''Metriacanthosaurus'' was named for the height of its [[neural spines]], which are actually not overly tall for theropods.<ref name="naish&martill07">{{cite journal|last=Naish|first=Darren|authorlink1 = Darren Naish|last2=Martill|first2=David M.|authorlink2 = David Martill|date=2007|title=Dinosaurs of Great Britain and the role of the Geological Society of London in their discovery: basal Dinosauria and Saurischia|journal=Quarterly Journal of the Geological Society|volume=164|pages=493–510|citeseerx = 10.1.1.394.9849|doi=10.1144/0016-76492006-032}}</ref> They are similar to other theropods such as ''Megalosaurus'', ''[[Sinraptor]]'', and ''[[Ceratosaurus]]'' in being 1.5 times the height of the [[Body of vertebra|centrum]].<ref name=benson&radley10>{{cite journal|last1=Benson|first1=R. B. J.|last2=Radley|first2=J. D.|date=2010|title=A New Large-Bodied Theropod Dinosaur from the Middle Jurassic of Warwickshire, United Kingdom|journal=Acta Palaeontologica Polonica|volume=55|issue=1|pages=35–42|doi=10.4202/app.2009.0083|url=http://www.bioone.org/doi/full/10.4202/app.2009.0083}}</ref>

==Classification==
Originally named as a species of ''Megalosaurus'' in [[Megalosauridae]], ''Metriacanthosaurus'' was more likely a member of [[Metriacanthosauridae]]. It is thought to be related to genera such as ''[[Yangchuanosaurus]]'', and in 1988 Paul synonymized the two genera. However, a 2007 review of British dinosaurs by [[Darren Naish]] and [[David Martill]] found that they were distinct. ''Metriacanthosaurus'' is the first genus of sinraptorid from Europe, being named before the other possibly member ''[[Lourinhanosaurus]]''.<ref name="naish&martill07"/> ''Metriacanthosaurus'' is likely a member of the subfamily [[Metriacanthosaurinae]].<ref name=tetanurans2012/>

Below is a simplified cladogram of [[Tetanurae]] by [[Matthew Carrano]] ''et al.'' (2012).<ref name=tetanurans2012>{{Cite journal | last1 = Carrano | first1 = M. T. | last2 = Benson | first2 = R. B. J. | last3 = Sampson | first3 = S. D. | doi = 10.1080/14772019.2011.630927 | title = The phylogeny of Tetanurae (Dinosauria: Theropoda) | journal = Journal of Systematic Palaeontology | volume = 10 | issue = 2 | pages = 211–300| year = 2012 | pmid = | pmc = }}</ref>

{{clade| style=font-size:85%; line-height:85%
|label1=[[Metriacanthosauridae]]
|1={{clade
|1={{clade
|1=''[[Yangchuanosaurus zigongensis]]''
|2={{clade
|1=CV 00214
|2=''[[Yangchuanosaurus shangyouensis]]'' }} }}
|label2=[[Metriacanthosaurinae]]
|2={{clade
|1=''[[Shidaisaurus]]''
|2={{clade
|1='''''Metriacanthosaurus'''''
|2={{clade
|1=''[[Sinraptor|"Sinraptor" hepingensis]]''
|2={{clade
|1=''[[Sinraptor dongi]]''
|2=''[[Siamotyrannus]]'' }} }} }} }} }} }}

==References==
{{Reflist}}

{{Portal|Dinosaurs}}
{{Allosauroidea}}
{{Taxonbar|from=Q1957207}}

[[Category:Metriacanthosaurids]]
[[Category:Middle Jurassic dinosaurs of Europe]]
[[Category:Fossil taxa described in 1964]]
[[Category:Taxa named by Alick Walker]]
[[Category:Paleontology in England]]

Schottischer Liebstöckel

2018-03-22T21:31:46Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q3257294}} (9 sig. taxon IDs); WP:GenFixes using AWB

{{Italic title}}
{{taxobox
| name = ''Ligusticum scoticum''
| image = Ligusticum scoticum.jpg
| regnum = [[Plant]]ae
| unranked_divisio = [[Angiosperms]]
| unranked_classis = [[Eudicots]]
| unranked_ordo = [[Asterids]]
| ordo = [[Apiales]]
| familia = [[Apiaceae]]
| genus = ''[[Ligusticum]]''
| species = '''''L. scoticum'''''
| binomial = ''Ligusticum scoticum''
| binomial_authority = [[Carl Linnaeus|L.]]
| synonyms_ref =  <ref name="Palin"/>
| synonyms =
*''Ligusticum scothicum'' L.
*''Haloscias scoticum'' (L.) Fr.
}}

'''''Ligusticum scoticum''''', known as '''Scots lovage''',<ref name="Hackney"/> or '''Scottish licorice-root''',<ref>{{PLANTS|id=LISC3|taxon=Ligusticum scoticum|accessdate=2 June 2015}}</ref> is a [[perennial plant]] of the family [[Apiaceae|Umbelliferae]] (Apiaceae) found near the coasts of northern Europe and north-eastern North America. It grows up to {{convert|60|cm}} tall and is found in rock crevices and cliff-top grassland. It is closely related to, and possibly conspecific with, ''[[Ligusticum hultenii]]'' from the coast of the northern Pacific Ocean. The plant is edible, with a flavour resembling [[parsley]] or [[celery]].

==Description==
''Ligusticum scoticum'' is a [[herbaceous plant|herbaceous]] [[perennial plant]] which typically grows {{convert|15|–|60|cm|0}} tall.<ref name="Palin">{{cite journal |author=M. Anne Palin |year=1988 |title=''Ligusticum scoticum'' L. (''Haloscias scoticum'' (L.) Fr.) |journal=[[Journal of Ecology]] |volume=76 |issue=3 |pages=889–902 |jstor=2260580}}</ref> It has triangular, twice-[[Glossary of leaf morphology|ternate]] leaves, {{convert|5|-|20|cm|abbr=on}} long, with each lobe {{convert|2|–|5|cm|1|abbr=on}} long. The edges of the leaves may be toothed, lobed or serrated, and are typically either a paler green or magenta.<ref name="Palin"/> The [[Plant stem|stem]] branches infrequently, and bears 2–5 [[inflorescence]]s, each of which is a compound [[umbel]] {{convert|4|–|6|cm|abbr=on}} in diameter.<ref name="Palin"/> There are typically 8–12 rays in both the primary and secondary umbels. Each individual flower is around {{convert|2|mm|2|abbr=on}} in diameter and greenish-white in colour.<ref name="Palin"/> The [[fruit]] are {{convert|4|–|6|mm|abbr=on}} long, with five prominent ridges on each [[carpel]].<ref name="Palin"/>

''Ligusticum scoticum'' tastes and smells like [[parsley]]<ref name="Palin"/> or [[celery]],<ref name="Watts">{{cite book |author=D. C. Watts |year=2007 |title=Dictionary of Plant Lore |publisher=[[Academic Press]] |isbn=9780080546025 |chapter=Lovage |page=231 |url=https://books.google.com/books?id=WAagnZNb0cAC&pg=PA231}}</ref> and was formerly widely eaten in western Britain, both for nutrition and to combat [[scurvy]].<ref name="Watts"/>

==Distribution==
''Ligusticum scoticum'' is primarily an [[Arctic]] plant, with a [[disjunct distribution|disjunct range]] extending from [[northern Norway]] to the more northerly shores of the [[British Isles]], and from western [[Greenland]] to [[New England]].<ref>{{cite book |author=R. M. M. Crawford |year=2008 |title=Plants at the Margin: Ecological Limits and Climate Change |publisher=[[Cambridge University Press]] |isbn=9781139469296 |pages=73, 235 |url=https://books.google.com/books?id=fvvIxkPjTFkC&pg=PA235}}</ref> A related species, ''[[Ligusticum hultenii]]'', which was described by [[Merritt Lyndon Fernald]] in 1930<ref>{{cite journal |author=[[Merritt Lyndon Fernald|M. L. Fernald]] |year=1930 |title=''Ligusticum scothicum'' of the North Atlantic and of the North Pacific |journal=[[Contributions from the Gray Herbarium of Harvard University]] |volume=87 |pages=7–9 |jstor=41764456}}</ref> and may be better treated as a [[subspecies]] of ''L. scoticum'', occurs around the northern Pacific Ocean, from [[Japan]] to [[Alaska]].<ref name="Hackney">{{cite web |url=http://www.habitas.org.uk/priority/species.asp?item=3702 |title=''Ligusticum scoticum'' – scots lovage |work=Northern Ireland Priority Species |accessdate=October 18, 2013 |author=Paul Hackney |publisher=[[National Museums Northern Ireland]]}}</ref> The southernmost occurrence of ''L. scoticum'' is at [[Ballyhalbert]] in [[Northern Ireland]].<ref name="Hackney"/>

==Ecology==
Within the British Isles, ''Ligusticum scoticum'' is only found on coasts where the mean annual [[temperature]] is below {{convert|15|C|F}}, and this bound is likely to also apply in other parts of the species' range.<ref name="Palin"/> Towards the southern end of its range, the plant performs poorly on south-facing sites.<ref name="Palin"/> It grows in fissures in rocks, where it may be the only vascular plant, and also in cliff-top grassland communities dominated by ''[[Festuca rubra]]'' and ''[[Plantago maritima]]''.<ref name="Palin"/>

''Ligusticum scoticum'' cannot tolerate [[grazing]], and is harmed by the actions of nesting [[seabird]]s; it is therefore rarely found on bird cliffs, or where grazing sheep and rabbits are found.<ref name="Palin"/> It is, however, tolerant of [[salt spray]], and its growth has been shown to improve when given dilute [[sea water]].<ref name="Palin"/> The leaves of ''L. scoticum'' are frost-tolerant, and die back each winter, but regrow very rapidly the following spring. In the British Isles, [[flowering]] occurs from June to August, and the seeds are ripe in October or November; the timing is expected to be later at higher latitudes.<ref name="Palin"/> The flowers of ''L. scoticum'' are visited by generalist [[pollinator]]s, mostly [[fly|flies]].<ref name="Palin"/>

==Taxonomy==
''Ligusticum scoticum'' was first described by [[Carl Linnaeus]] in his 1753 work ''{{lang|la|[[Species Plantarum]]}}''.<ref>{{cite web |url=http://www.ipni.org/ipni/idPlantNameSearch.do?id=844602-1 |title= Plant Name Details−Apiaceae ''Ligusticum scoticum'' L. |publisher=[[International Plant Names Index]] |accessdate=7 November 2012}}</ref> Linnaeus originally used the [[specific name (botany)|epithet]] ''{{lang|la|scothicum}}'', and this is used by many authors in North America; in Europe, the amended spelling ''{{lang|la|scoticum}}'' is used.<ref name="Palin"/>

==References==
{{Reflist|32em}}

==External links==
*{{Commons category-inline|Ligusticum scoticum|''Ligusticum scoticum''}}

{{Edible Apiaceae}}
{{Taxonbar|from=Q3257294}}

[[Category:Ligusticum|scoticum]]
[[Category:Flora of Northern Europe]]
[[Category:Flora of Subarctic America]]
[[Category:Flora of Eastern Canada]]
[[Category:Flora of the Northeastern United States]]
[[Category:Edible Apiaceae]]
[[Category:Plants described in 1753]]
[[Category:Flora of Saint Pierre and Miquelon]]

Hypomyces

2018-03-22T21:11:07Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q5951106}} (3 sig. taxon IDs); WP:GenFixes using AWB

{{expand Japanese|date=November 2011|ヒポミケス属}}
{{Taxobox
| image = Lobster mushrooms.jpg
| image_caption = ''[[Hypomyces lactifluorum|H. lactifluorum]]''
| regnum = [[Fungi]]
| divisio = [[Ascomycota]]
| classis = [[Sordariomycetes]]
| ordo = [[Hypocreales]]
| familia = [[Hypocreaceae]]
| genus = '''''Hypomyces'''''
| genus_authority = ([[Elias Magnus Fries|Fr.]]) [[Louis René Tulasne|Tul.]] & [[Charles Tulasne|C. Tul.]] (1860)
| type_species = ''Hypomyces lactifluorum''
| type_species_authority = ([[Lewis David de Schweinitz|Schwein.]]) Tul. & C. Tul.
}}

'''''Hypomyces''''' is a [[genus]] of parasitic [[ascomycete]] fungi found in Europe, North America, Australia, and parts of China. The genus contains 53 species.<ref name=Kirk2008>{{cite book |vauthors=Kirk PM, Cannon PF, Minter DW, Stalpers JA |title=Dictionary of the Fungi. |edition=10th |publisher=CAB International |location=Wallingford, UK |year=2008|page=373|isbn=978-0-85199-826-8}}</ref> Better known species include the [[lobster mushroom]] (''Hypomyces lactifluorum'') and the [[bolete eater]] (''Hypomyces chrysospermus'').

==List of noteworthy species==
* ''[[Hypomyces cervinigenus|H. cervinigenus]]'' - on ''[[Helvella lacunosa]]''.<ref name="arora86">{{cite book |last=Arora |first=David|authorlink=David Arora |year=1986 |title=Mushrooms demystified: a comprehensive guide to the fleshy fungi |edition=2nd |location=Berkeley |publisher=Ten Speed Press |isbn=0-89815-169-4| pages=815–16}}</ref>
* ''[[Hypomyces chrysospermus|H. chrysospermus]]'' - Bolete Eater, Cask fungus (Eurasia, Western Australia, North America)
* ''[[Hypomyces hyalinus|H. hyalinus]]'' - Amanita "mold" (North America)
* ''[[Hypomyces lactifluorum|H. lactifluorum]]'' - Lobster mushroom (North America)
* ''[[Hypomyces luteovirens|H. luteovirens]]'' - Yellow-green Russula "mold" (North America)
* ''[[Hypomyces transformans|H. transformans]]'' - Ramaria Eater (North America)

==Other Species==
{{div col|cols=4}}
*''[[Hypomyces agaricola]]''
*''[[Hypomyces albidus]]''
*''[[Hypomyces albus]]''
*''[[Hypomyces amaurodermatis]]''
*''[[Hypomyces apiculatus]]''
*''[[Hypomyces apiosporus]]''
*''[[Hypomyces arachnoideus]]''
*''[[Hypomyces arecae]]''
*''[[Hypomyces arenaceus]]''
*''[[Hypomyces armeniacus]]''
*''[[Hypomyces asclepiadis]]''
*''[[Hypomyces ater]]''
*''[[Hypomyces aurantiicolor]]''
*''[[Hypomyces aurantius]]''
*''[[Hypomyces auriculariicola]]''
*''[[Hypomyces australbidus]]''
*''[[Hypomyces australiensis]]''
*''[[Hypomyces australis]]''
*''[[Hypomyces badius]]''
*''[[Hypomyces banningiae]]''
*''[[Hypomyces batavus]]''
*''[[Hypomyces biasolettianus]]''
*''[[Hypomyces boleticola]]''
*''[[Hypomyces boletinus]]''
*''[[Hypomyces boletiphagus]]''
*''[[Hypomyces bombacinus]]''
*''[[Hypomyces bresadolae]]''
*''[[Hypomyces bresadolanus]]''
*''[[Hypomyces camphorati]]''
*''[[Hypomyces caulicola]]''
*''[[Hypomyces cervinigenus]]''
*''[[Hypomyces cervinus]]''
*''[[Hypomyces cesatii]]''
*''[[Hypomyces chlorinigenus]]''
*''[[Hypomyces chlorinus]]''
*''[[Hypomyces chromaticus]]''
*''[[Hypomyces chrysospermus]]''
*''[[Hypomyces completus]]''
*''[[Hypomyces conviva]]''
*''[[Hypomyces corticiicola]]''
*''[[Hypomyces dactylarioides]]''
*''[[Hypomyces deformans]]''
*''[[Hypomyces destruens-equi]]''
*''[[Hypomyces ekmanii]]''
*''[[Hypomyces epimyces]]''
*''[[Hypomyces favoli]]''
*''[[Hypomyces flavescens]]''
*''[[Hypomyces flavolanatus]]''
*''[[Hypomyces floccosus]]''
*''[[Hypomyces fulgens]]''
*''[[Hypomyces fusisporus]]''
*''[[Hypomyces galericola]]''
*''[[Hypomyces goroshankianus]]''
*''[[Hypomyces hrubyanus]]''
*''[[Hypomyces hyacinthi]]''
*''[[Hypomyces hyalinus]]''
*''[[Hypomyces inaequalis]]''
*''[[Hypomyces insignis]]''
*''[[Hypomyces javanicus]]''
*''[[Hypomyces khaoyaiensis]]''
*''[[Hypomyces lactifluorum]]''
*''[[Hypomyces laeticolor]]''
*''[[Hypomyces lateritius]]''
*''[[Hypomyces leotiarum]]''
*''[[Hypomyces leotiicola]]''
*''[[Hypomyces linearis]]''
*''[[Hypomyces linkii]]''
*''[[Hypomyces lithuanicus]]''
*''[[Hypomyces macrosporus]]''
*''[[Hypomyces melanocarpus]]''
*''[[Hypomyces melanochlorus]]''
*''[[Hypomyces melanostigma]]''
*''[[Hypomyces microspermus]]''
*''[[Hypomyces miliarius]]''
*''[[Hypomyces mycogones]]''
*''[[Hypomyces mycophilus]]''
*''[[Hypomyces niveus]]''
*''[[Hypomyces novae-zelandiae]]''
*''[[Hypomyces ochraceus]]''
*''[[Hypomyces odoratus]]''
*''[[Hypomyces orthosporus]]''
*''[[Hypomyces paeonius]]''
*''[[Hypomyces pallidus]]''
*''[[Hypomyces pannosus]]''
*''[[Hypomyces papulasporae]]''
*''[[Hypomyces papyraceus]]''
*''[[Hypomyces parvisporus]]''
*''[[Hypomyces parvus]]''
*''[[Hypomyces penicillatus]]''
*''[[Hypomyces pergamenus]]''
*''[[Hypomyces perniciosus]]''
*''[[Hypomyces petchii]]''
*''[[Hypomyces pezizae]]''
*''[[Hypomyces polyporinus]]''
*''[[Hypomyces porphyreus]]''
*''[[Hypomyces pseudocorticiicola]]''
*''[[Hypomyces pseudopolyporinus]]''
*''[[Hypomyces psiloti]]''
*''[[Hypomyces puertoricensis]]''
*''[[Hypomyces purpureus]]''
*''[[Hypomyces robledoi]]''
*''[[Hypomyces rosellus]]''
*''[[Hypomyces rostratus]]''
*''[[Hypomyces rubi]]''
*''[[Hypomyces semitranslucens]]''
*''[[Hypomyces sepulchralis]]''
*''[[Hypomyces sepultariae]]''
*''[[Hypomyces siamensis]]''
*''[[Hypomyces sibirinae]]''
*''[[Hypomyces spadiceus]]''
*''[[Hypomyces stephanomatis]]''
*''[[Hypomyces stereicola]]''
*''[[Hypomyces stuhlmannii]]''
*''[[Hypomyces subaurantius]]''
*''[[Hypomyces subiculosus]]''
*''[[Hypomyces succineus]]''
*''[[Hypomyces sulphureus]]''
*''[[Hypomyces sympodiophorus]]''
*''[[Hypomyces tegillum]]''
*''[[Hypomyces terrestris]]''
*''[[Hypomyces thailandicus]]''
*''[[Hypomyces thiryanus]]''
*''[[Hypomyces tomentosus]]''
*''[[Hypomyces torminosus]]''
*''[[Hypomyces transformans]]''
*''[[Hypomyces trichoderma]]''
*''[[Hypomyces triseptatus]]''
*''[[Hypomyces tubericola]]''
*''[[Hypomyces tuberosus]]''
*''[[Hypomyces tulasneanus]]''
*''[[Hypomyces vanbruntianus]]''
*''[[Hypomyces vandae]]''
*''[[Hypomyces villosus]]''
*''[[Hypomyces viridigriseus]]''
*''[[Hypomyces viridis]]''
*''[[Hypomyces volemi]]''
*''[[Hypomyces vuilleminianus]]''
*''[[Hypomyces xyloboli]]''
*''[[Hypomyces xylophilus]]''
{{div col end}}

==References==
{{Reflist}}

==External links==
{{commons category}}
*{{IndexFungorum|2446}}

{{Taxonbar|from=Q5951106}}

[[Category:Sordariomycetes genera]]
[[Category:Parasitic fungi]]
[[Category:Hypocreaceae]]
[[Category:Science articles needing translation from Japanese Wikipedia]]

{{Hypocreales-stub}}
{{parasite-stub}}

Benutzer:Wenz-s/Trichopoda pennipes

2018-03-22T21:02:11Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q1211203}} (5 sig. taxon IDs); WP:GenFixes using AWB

{{Taxobox
| image = Feather-legged_Fly_(Trichopoda_pennipes)_2.jpg
| image_caption = ''Trichopoda pennipes''
| regnum = [[Animal]]ia
| phylum = [[Arthropod]]a
| classis = [[Insect]]a
| ordo = [[fly|Diptera]]
| familia = [[Tachinidae]]
| subfamilia = [[Phasiinae]]
| tribus = [[Trichopodini]]
| genus = ''[[Trichopoda]]''
| subgenus = ''[[Trichopoda (subgenus)|Trichopoda]]''
| species = '''''T. pennipes'''''
| binomial = '''''Trichopoda pennipes'''''
| binomial_authority = ([[Johan Christian Fabricius|Fabricius]], 1781)
|synonyms={{collapsible list|bullets = true
| ''Phasia jugatoria'' Say, 1829
| ''Thereva hirtipes'' Fabricius, 1805
| ''Thereva pennipes'' Fabricius, 1805
| ''Trichopoda cilipes'' Wiedemann, 1830
| ''Trichopoda flavicornis'' Robineau-Desvoidy, 1830 }}
}}

'''''Trichopoda pennipes''''', common name '''feather-legged fly''', is a [[Diptera|fly]] in the family [[Tachinidae]].

==Distribution==
This species is native to [[North America]] ([[U.S.A. ]], [[Mexico]]),<ref>Bisby F.A., Roskov Y.R., Orrell T.M., Nicolson D., Paglinawan L.E., Bailly N., Kirk P.M., Bourgoin T., Baillargeon G., Ouvrard D. [http://www.catalogueoflife.org/annual-checklist/2011/details/species/id/8648091 Catalogue of life]</ref> in [[Hawaiian Islands]] and in [[South America]]<ref name="WISC">{{cite web | url = http://www.entomology.wisc.edu/mbcn/kyf605.html | title = ''Trichopodes pennipes'', Parasitoid of True Bugs | first = Mahr | last = Susan | publisher = University of Wisconsin }}</ref> and has been introduced into southern [[Europe]] ([[France]], [[Italy]] and [[Spain]]).<ref>[https://fauna-eu.org/cdm_dataportal/taxon/3cc92fd7-2842-4998-a066-ef46d3b54838 Fauna europaea]</ref><ref name="BG">{{cite web | url = http://bugguide.net/node/view/13284 | title = Species Trichopoda pennipes | first = Leung | last = Richard | date = 2005-03-20 | publisher = Iowa State University Entomology. }}</ref>

==Habitat==
This species inhabit grasslands, hedge rows and crops where its key hosts are present.<ref name="CU"/>

==Morphology==
[[File: Trichopoda pennipes.jpg |thumb|300px|left|''Trichopoda pennipes'' showing the comb-like fringe of flattened hairs]]
''Trichopoda pennipes'' can reach a length of {{convert|10.5|mm}},<ref>[http://eol.org/pages/763440/overview#161 Encyclopedia of life]</ref> about the size of a large [[housefly]]. These medium-sized flies have a velvety black head. The velvety black or brown [[Thorax (insect anatomy)|thorax]] shows a few yellow stripes. Eyes are large brown with yellow between. The color of the slender [[abdomen]] varies from bright orange to completely black. Females usually have a dark-tipped abdomen, while males have a dark orange apex. Wings are transparent smoky with prominent veins. In the females they are evenly dusky, with the posterior margin sub-hyaline, while male flies have a ferrugineous marking on the wings. This fly has huge [[halteres]]. Legs are black, with yellow feet. The hind legs bear a prominent feather-like fringe of flattened hairs.<ref name="WISC"/><ref name="CU">Pickett, Charles H. [https://biocontrol.entomology.cornell.edu/parasitoids/trichopoda.php "Trichopoda pennipes (Diptera: Tachinidae)"] College of Agriculture and Life Sciences - Cornell University.</ref>

[[File:Nezara viridula f. torquata & Trichopoda pennipes MHNT.jpg|thumb|Egg on ''[[Nezara viridula]]'' head.]]

==Life cycle==
''Trichopoda pennipes'' first appears in the late spring or early summer and feeds on nectar sucked from flowers such as [[Daucus carota|Queen Anne's lace]] and [[Spiraea|meadowsweet]]. It may be seen hovering over other plants in search of suitable bugs on which to lay its eggs, most commonly squash bugs and southern green stinkbugs. The female fly lays several small, pale-coloured, oval [[Egg (biology)|eggs]] on a large [[Nymph (biology)|nymph]] or an adult bug. In fact the larvae are [[parasitoids]] of several [[Hemiptera|true bugs]], particularly [[squash bug]]s and leaf-footed bugs in the family [[Coreidae]] (including the large-sized ''[[Leptoglossus occidentalis]]''), stinkbugs in the family [[Pentatomidae]] and other pentatomorph bugs ([[Largidae]] and [[Scutelleridae]] species).<ref name="BG"/>

When the eggs hatch, the larvae burrow into the bug. If there are several larvae in one host, only one survives. After feeding on the bug's tissues, the cream-coloured larva emerges and falls to the ground where it pupates in a reddish-brown [[pupa]]rium formed from the last larval skin. The bug meanwhile dies. After about two weeks, an adult fly emerges from the pupa. After mating, a female fly may lay several hundred eggs in total. There are up to three generations of the fly each year and the parasitoid overwinters as a second [[instar]] larva within the body of the overwintering [[Host (biology)|host]].<ref name="WISC"/><ref name="CE">Hoffmann, M.P. and Frodsham, A.C. ''Natural Enemies of Vegetable Insect Pests''. (1993) Cooperative Extension, Cornell University, Ithaca, NY.</ref>

==Use in biological control==
''Trichopoda pennipes'' is used as a biological control agent for agricultural [[Pest (organism)|pest]]s. One of the principal host species for ''T. pennipes'' is the southern green stinkbug, ''[[Nezara viridula]]''. This is a crop pest that originated in [[Ethiopia]] but now has a worldwide distribution and is named for the foul-smelling defensive secretion it exudes from a gland on its thorax. This exudate appears to be fairly effective in preventing predation by birds but is well tolerated by ''T. pennipes''<ref name="EES">Eisner, T., Eisner, M. and Siegler, M. ''Secret Weapons: Defenses of Insects, Spiders, Scorpions, and Other Many-Legged Creatures.'' (2005) Belknap Press</ref> which seems to be highly attracted by an aggregation [[pheromone]] produced by the male bugs. This results in a higher proportion of males being parasitised than females.<ref name="WISC"/>

There seems to be different biotypes across the [[United States]], preying on different hosts in different regions. In northern [[California]], a population of the fly parasitised the bordered plant bug, ''[[Largus succinctus]]'', but did not attack the squash bug, ''[[Anasa tristis]]''. In an experiment, ''T. pennipes'' was collected from squash fields in [[New York (state)|New York State]] and released near [[Squash (plant)|squash]] fields in California. The insects targeted the squash bugs and established permanent populations. Their eggs are now found deposited on nearly 50% of the squash bugs in the area but it is unclear how effective this is in controlling the crop pests.<ref name="EES"/> On the southern green stink bug, the rate of parasitism can be as high as 93% and up to 80% on the squash bug. However, ''T. pennipes'' does not prevent all crop damage as the bugs continue to feed and reproduce after being parasitised, though the reproductive organs begin to atrophy when the parasitoid reaches the second instar stage. Control of the pest is more effective when nymphs are parasitised since half of these die before becoming adults and any that overwinter will die before laying eggs.<ref name="WISC"/>

==Gallery==
<gallery widths="240" heights="180">
File: Fly August 2008-1.jpg|Male, with ferrugineous marking on the wings
File: Tachinidae - Trichopoda pennipes.JPG|Female with black abdomen and evenly dusky wings, with the posterior margin sub-hyaline
File: Feather-legged Fly - Trichopoda pennipes, Julie Metz Wetlands, Woodbridge, Virginia.jpg| Female with orange black-tipped abdomen
Feather-legged Fly (Trichopoda pennipes).jpg|Front view
</gallery>

==References==
{{Wikispecies}}
{{Commons}}
{{Reflist}}

{{Taxonbar|from=Q1211203}}

[[Category:Phasiinae]]
[[Category:Insects acting as insect pest control agents]]
[[Category:Diptera of Europe]]
[[Category:Diptera of North America]]
[[Category:Diptera of South America]]
[[Category:Insects described in 1781]]

{{Fly-stub}}

Flueggea

2018-03-22T20:57:20Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q41427}} (11 sig. taxon IDs); WP:GenFixes, using AWB

{{Italic title}}
{{taxobox
|name = Bushweeds
|image = Flueggea leucopyrus Bra54.png
|image_caption = ''[[Flueggea leucopyrus]]''<ref>1874 illustration from tab. 54 of D. Brandis, Illustrations of the Forest Flora of North-West and Central India, 1874</ref>
|regnum = [[Plant]]ae
|unranked_divisio = [[Angiosperms]]
|unranked_classis = [[Eudicots]]
|unranked_ordo = [[Rosids]]
|ordo = [[Malpighiales]]
|familia = [[Phyllanthaceae]]
|tribus = [[Phyllantheae]]
|subtribus = [[Flueggeinae]]
|genus = '''''Flueggea'''''
|genus_authority = [[Carl Ludwig Willdenow|Willd.]] 1806 not Rich. 1807 (Asparagaceae)
|synonyms_ref=<ref name=w/>
|synonyms=*''Bessera'' [[Spreng.]], illegitimate homonym, not Schult. 1809 nor Schult. f. 1829 nor Vell. 1825
*''Colmeiroa'' [[Reut.]]
*''Coilmeroa'' Endl.
*''Geblera'' [[Friedrich Ernst Ludwig von Fischer|Fisch.]] & [[C.A.Mey.]]
*''Neowawraea'' [[Joseph Rock|Rock]]
*''Pleiostemon'' [[Otto Wilhelm Sonder|Sond.]]
*''Villanova '' [[Pourr.]] ex [[Cutanda]] 1861, illegitimate homonym, not Ortega 1797 nor Lag. 1816 (both of the latter Asteraceae)
*''Acidoton'' P.Browne 1856, rejected name, now Sw. 1788
*''Fluggea'' Willd. spelling variant
|}}

'''''Flueggea''''', the '''bushweeds''', is a genus of shrubs and trees in the [[family]] [[Phyllanthaceae]] first described as a genus in 1806.<ref>[https://www.biodiversitylibrary.org/page/566364#page/6/mode/1up Willdenow, Carl Ludwig von. 1806. Species Plantarum. Editio quarta 4(2): 637]</ref><ref>[https://www.biodiversitylibrary.org/page/566364#page/126/mode/1up Willdenow, Carl Ludwig von. 1806. Species Plantarum. Editio quarta 4(2): 757–758]</ref><ref>[http://www.tropicos.org/Name/40009982 Tropicos, genus ''Flueggea'' Willd.]</ref> It is widespread across much of [[Asia]], [[Africa]], and various oceanic islands, with a few species in [[South America]] in on the [[Iberian Peninsula]].<ref name=w>[http://apps.kew.org/wcsp/namedetail.do?name_id=84577 Kew World Checklist of Selected Plant Families]</ref><ref>Govaerts, R., Frodin, D.G. & Radcliffe-Smith, A. (2000). World Checklist and Bibliography of Euphorbiaceae (and Pandaceae) 1-4: 1-1622. The Board of Trustees of the Royal Botanic Gardens, Kew.</ref><ref>Barker, C. & van Welzen, P.C. (2010). ''Flueggea'' (Euphorbiaceae s. l. or Phyllanthaceae) in Malesia. Systematic Botany 35: 541-551.</ref><ref name=s>[http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=112889 Flora of China Vol. 11 Page 177 <big>白饭树属</big> bai fan shu shu ''Flueggea'' Willdenow, Sp. Pl. 4: 637, 757. 1805. ]</ref><ref>Schatz, G. E., S. Andriambololonera, Andrianarivelo, M. W. Callmander, Faranirina, P. P. Lowry, P. B. Phillipson, Rabarimanarivo, J. I. Raharilala, Rajaonary, Rakotonirina, R. H. Ramananjanahary, B. Ramandimbisoa, A. Randrianasolo, N. Ravololomanana, Z.S. Rogers, C.M. Taylor & G. A. Wahlert. 2011. Catalogue of the Vascular Plants of Madagascar. Monographs in systematic botany from the Missouri Botanical Garden</ref><ref>Nasir, E. & S. I. Ali (eds). 1980-2005. Flora of Pakistan University. of Karachi, Karachi.</ref>

The genus is named after [[Johannes Flüggé|John Fluegge]], a [[Germany|German]] cryptogamic [[botanist]].

Members of this genus all have entire ovate leaves and minute green [[flower]]s that form at the [[Leaf|leaf axils]] in the form of [[fascicle (botany)|fascicle]]s or [[inflorescence|cyme]]s. The fruits are [[Berry (botany)|berries]], of the size of [[pea]]s.<ref name=s/>

Many members of the genus were formerly classified under the genus ''[[Securinega]]''.

;Species<ref name=w/>
{{columns-list|2|
# ''[[Flueggea acicularis]]'' - S China
# ''[[Flueggea acidoton]]'' - West Indies
# ''[[Flueggea anatolica]]'' - S Turkey
# ''[[Flueggea elliptica]]'' - Ecuador
# ''[[Flueggea flexuosa]]'' - Philippines, [[Maluku (province)|Maluku]], New Guinea, SW Pacific
# ''[[Flueggea gracilis]]'' - [[Hainan]], [[Palawan]], Borneo, Malaysia, S Thailand
# ''[[Flueggea jullienii]]'' - Cambodia, Laos, Vietnam
# ''[[Flueggea leucopyrus]]'' - [[Socotra]], [[Sajid Island]], S [[India]], [[Sri Lanka]], [[Sichuan]], [[Yunnan]]
# ''[[Flueggea monticola]]'' - [[Sichuan]], [[Yunnan]]
# ''[[Flueggea neowawraea]]'' - [[Hawaii]]
# ''[[Flueggea schuechiana]]'' - [[Pernambuco]]
# ''[[Flueggea spirei]]'' - [[Laos]]
# ''[[Flueggea suffruticosa]]'' - Siberia, Mongolia, China, Korea, Japan
# ''[[Flueggea tinctoria]]'' - Spain, Portugal
# ''[[Flueggea verrucosa]]'' - [[Cape Province]], [[KwaZulu-Natal]]
# ''[[Flueggea virosa]]'' - Africa, Madagascar, SW + S + SE Asia, [[Mascarenes]], N Australia }}

;formerly included<ref name=w/>
moved to other genera ''([[Leptopus]] [[Margaritaria]] [[Meineckia]] [[Ophiopogon]] )''
{{columns-list|2|
# ''F. anceps'' - ''[[Ophiopogon japonicus]]''
# ''F. angulata'' Raf. 1838 not (Schumach. & Thonn.) Schrank 1828 - ''[[Ophiopogon japonicus]]''
# ''F. bailloniana - [[Margaritaria discoidea]] var. triplosphaera''
# ''F. capillipes - [[Leptopus chinensis]]''
# ''F. dracaenoides - [[Ophiopogon dracaenoides]]''
# ''F. dubia - [[Ophiopogon intermedius]]''
# ''F. eglandulosa - [[Margaritaria anomala]]''
# ''F. fagifolia - [[Margaritaria discoidea]] var. fagifolia''
# ''F. griffithii - Ophiopogon intermedius''
# ''F. hilariana - [[Meineckia neogranatensis]] ''subsp''. hilariana''
# ''F. intermedia - [[Ophiopogon intermedius]]''
# ''F. jaburan - [[Ophiopogon jaburan]]''
# ''F. jacquemontiana - [[Ophiopogon intermedius]]''
# ''F. japonica'' (Thunb.) Rich. not 1807 (Miq.) Pax 1890 - ''[[Ophiopogon japonicus]]''
# ''F. major - [[Margaritaria anomala]]''
# ''F. meineckia - [[Meineckia phyllanthoides]]''
# ''F. nitida - [[Margaritaria discoidea]]'' var. ''nitida''
# ''F. obovata'' Baill. 1861 not (Willd.) Wall. ex Fern.-Vill. 1880 - ''[[Margaritaria discoidea]]'' var. ''triplosphaera''
# ''F. prolifera'' - ''[[Ophiopogon caulescens]]''
# ''F. trichogynis - [[Meineckia trichogynis]]''
# ''F. wallichiana'' Kunth 1825 not Baill. 1858 - ''[[Ophiopogon intermedius]]''
}}

==References==
{{Reflist}}

{{Taxonbar|from=Q41427}}

[[Category:Flueggea| ]]
[[Category:Phyllanthaceae]]
[[Category:Malpighiales genera]]
[[Category:Medicinal plants]]
[[Category:Taxa named by Carl Ludwig Willdenow]]

Endeavouria septemlineata

2018-03-22T20:41:43Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q18591209}} (3 sig. taxon IDs); WP:GenFix using AWB

{{Italic title}}
{{Taxobox
| name = ''Endeavouria''
| image = Endeavouria_septemlineata.JPG
| image_caption = ''Endeavouria septemlineata''
| regnum = [[Animal]]ia
| phylum = [[Platyhelminthes]]
| classis = [[Rhabditophora]]
| subclassis =
| superordo =
| ordo = [[Tricladida]]
| subordo = [[Continenticola]]
| infraordo =
| unranked_familia =
| superfamilia =
| familia = [[Geoplanidae]]
| subfamilia = [[Rhynchodeminae]]
| tribus = [[Caenoplanini]]
| genus = '''''Endeavouria'''''
| genus_authority = [[Robert E. Ogren|Ogren]] & Kawakatsu 1991
| species = '''''E. septemlineata'''''
| binomial = ''Endeavouria septemlineata''
| binomial_authority = ([[Libbie Hyman|Hyman]], 1939)
| range_map = Endeavouria septemlineata distribution.png
| range_map_caption = Known distribution of ''E. septemlineata'' in dark blue
| synonyms = ''Geoplana septemlineata'' Hyman, 1939
}}

'''''Endeavouria''''' is a [[monotypic taxon|monotypic]] [[genus]] of [[land planarian]]s from the Pacific region. It contains a single species, ''Endeavouria septemlineata''.

==Description==
[[File:Endeavouria septemlineata ventral view.jpg|thumb|left|Ventral view of ''Endeavouria septemlineata'']]

The genus ''Endeavouria'' is similar to other closely related genera, such as ''[[Caenoplana]]'' and ''[[Kontikia]]'', but can be differentiated by the narrower creeping sole and a thick layer of longitudinal muscle fibers beneath the ventral nerve plate.<ref name="Ogren91">{{cite journal | last1 = Ogren | first1 = Robert E. | last2 = Kawakatsu | first2 = Masaharu | year = 1991 | title = Index to the species of the family Geoplanidae (Turbellaria, Tricladida, Terricola) Part II: Caenoplaninae and Pelmatoplaninae | journal = Bulletin of Fujis Women's College | volume = 29 | pages = 35–59 | url = http://ci.nii.ac.jp/naid/110008426083/en}} {{open access}}.</ref>

''Endeavouria septemlineata'' is a small land planarian, reaching about {{convert|30|mm|in}} in length. The dorsal side may appear completely black at first sight, but a closer look reveals that it has a light-brown background. There are seven dark longitudinal stripes running along the body: one narrown dark-brown stripe, two lateral broad black stripes with diffuse margins, two paramarginal stripes and two marginal stripes. The dorsal color is visible between the median and lateral stripes and between the lateral and paramarginal stripes. The ventral side and the region between the paramarginal and marginal stripes is pale grey. The creeping sole is marked by numerous small dark spots.<ref name="Hyman1939">{{cite journal | last = Hyman | first = Libbie H. | title = Land Planarians from the Hawaiian Islands | year = 1939 | journal = Archives de Zoologie Expérimentale et Genérale | volume = 80 | pages = 116–124}}</ref>

The several eyes are arranged in a single row along the body margins, being located over the marginal stripes. They are closer to each other in the anterior region and posteriorly they gradually become more spaced.<ref name="Hyman1939" />

The [[reproductive system of planarians|copulatory apparatus]] of ''E. septemlineata'' has a big ventral fold and a small [[reproductive system of planarians#Male part of the reproductive system|intra-antral penis papilla]].

==Etymology==
The name ''Endeavouria'' honors the flagship [[HMS Endeavour]] commanded by [[James Cook]] in his [[First voyage of James Cook|first voyage of discovery]] to [[Australia]] and [[New Zealand]].<ref name="Ogren91" /> The specific epithet ''septemlineata'' ([[Latin]] for "seven-striped") refers to the seven dark longitudinal stripes on the dorsum.<ref name="Hyman1939" />

==Origin and Distribution==
The native place of ''Endeavouria septemlineata'' is still unknown. It was first found in [[Hawaii]] and later in several locations in [[Brazil]].<ref name="CarbayoPedroni2007">{{cite journal | last1 = Carbayo | first1 = Fernando | last2 = Pedroni | first2 = Júlio | last3 = Froehlich | first3 = Eudóxia Maria | title = Colonization and extinction of land planarians (Platyhelminthes, Tricladida) in a Brazilian Atlantic Forest regrowth remnant | journal = Biological Invasions | volume = 10 | issue = 7 | year = 2007 | pages = 1131–1134 | issn = 1387-3547 | doi = 10.1007/s10530-007-9190-1 | url = https://link.springer.com/article/10.1007/s10530-007-9190-1}}</ref> As all land planarians in the tribe [[Caenoplanini]], it is most likely originally from somewhere in the Pacific region.

==Ecology and behavior==
[[File:Endeavouria septemlineata feeding on Rhinocricus millipede.png|thumb|left|''Endeavouria septemlineata'' feeding on ''Rhinocricus'' millipede]]
[[File:Tumbling behavior of Endeavouria septemlineata.png|thumb|Tumbling behavior of ''Endeavouria septemlineata'']]
Specimens of ''E. septemlineata'' show a gregarious behavior, gathering in groups of tenths of individuals that remain hidden in the leaf litter and under rocks or fallen logs during the day. They feed on several invertebrate groups, such as [[arthropod]]s and [[mollusk]]s, which they may hunt, but most of the time they feed on dead animals, thus being mainly scavengers.<ref name="BollRossi2015">{{cite journal|last1 = Boll | first1 = Piter K. | last2 = Rossi | first2 = Ilana | last3 = Amaral | first3 = Silvana V. | last4 = Leal-Zanchet | first4=Ana | title = A taste for exotic food: Neotropical land planarians feeding on an invasive flatworm | journal = PeerJ | volume = 3 |year = 2015 | pages = e1307 | issn = 2167-8359 | doi=10.7717/peerj.1307 | url = https://peerj.com/articles/1307/ | pmid=26500817 | pmc=4614845}} {{open access}}</ref>

It has been demonstrated that in Brazil several native land planarians feed on the introduced ''E. septemlineata'', thus possibly controlling its spread as an [[invasive species]].<ref name="BollRossi2015" />

When attacked by a predator, individuals of ''E. septemlineata'' quickly move away. The most frequent strategy to escape is by tumbling, a behavior where the planarian lifts its posterior end and bends it forward until it touchs the substrate ahead of the anterior end. If part of the body is trapped by the predator, it may escape by performing [[autotomy]].<ref name="BollRossi2015" />

==References==
{{Reflist}}

{{Taxonbar|from=Q18591209}}

[[Category:Geoplanidae]]

Cephalanthus

2018-03-22T18:25:04Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q3245386}} (10 sig. taxon IDs); WP:GenFixe using AWB

{{automatic taxobox
|image = Buttonbush in the Everglades.jpg
|image_caption = ''[[Cephalanthus occidentalis]]''
|display_parents = 2
|taxon = Cephalanthus
|authority = [[Carl Linnaeus|L.]]
|type_species = ''[[Cephalanthus occidentalis]]''
|type_species_authority = [[Carl Linnaeus|L.]]
|synonyms = *''Acrodryon'' [[Curt Polycarp Joachim Sprengel|Spreng.]]
*''Axolus'' [[Constantine Samuel Rafinesque|Raf.]]
*''Eresimus'' [[Constantine Samuel Rafinesque|Raf.]]
}}
{{commons category}}{{Wikispecies}}

'''''Cephalanthus''''' is a [[genus]] of [[flowering plant]]s in the [[Rubiaceae]] [[Family (biology)|family]]. There are about six species that are commonly known as '''buttonbush'''.<ref name="pl">[http://www.theplantlist.org/browse/A/Rubiaceae/Cephalanthus The Plant List, ''Cephalanthus'']</ref><ref name=china>[http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=106077 Flora of China, ''Cephalanthus'']</ref>

==Description==
They are [[shrub]]s or small [[tree]]s growing to {{convert|5|-|15|m|ft|abbr=on}} tall. The [[leaf|leaves]] are simple, arranged in opposite pairs or whorls of three. The [[flower]]s form a dense globular [[inflorescence]].

==Distribution and habitat==
''Cephalanthus occidentalis'' is [[Indigenous (ecology)|native]] to the eastern [[United States]] and [[Canada]]. The others [[Range (biology)|occur]] in [[Tropics|tropical]] regions of the [[Americas]], [[Africa]] and [[Asia]].<ref name="Mabberley2008">{{cite book|author=Mabberley DJ|year=2008|title=Mabberley's Plant Book|edition=3|publisher=Cambridge University Press|ISBN=978-0-521-82071-4}}</ref> Two species are known in [[Gardening|cultivation]].<ref name="Huxley1992">Huxley AJ et al. (eds.) ''The New Royal Horticultural Society Dictionary of Gardening.'' The Macmillan Press Limited, London; The Stockton Press, New York. 1992. {{ISBN|978-0-333-47494-5}}</ref>

==Systematics==
''Cephalanthus'' was [[Botanical name|named]] by [[Carl Linnaeus|Linnaeus]] in ''[[Species Plantarum]]'' in 1753.<ref name="linnaeus1753">Linnaeus, C. [https://www.biodiversitylibrary.org/page/26068066 ''Cephalanthus'']. ''Species Plantarum''. 1753. 1: 95</ref> The [[Genus#Generic name|generic name]] is [[Etymology|derived]] from the [[Ancient Greek]] words κέφαλη (''kephale''), meaning "head", and ἄνθος (''anthos''), meaning "flower".<ref name="Quattrocchi2000">{{cite book|author=Quattrocchi, U.|year=2000|title=CRC World Dictionary of Plant Names|volume=1|pages=476|url=https://books.google.nl/books?id=A68qyOyhOdkC&pg=PA476&lpg=PA476&dq=Cephalanthus&hl=nl#v=onepage&q=Cephalanthus&f=true|publisher=CRC Press|location=Boca Raton, New York, Washington DC, London|ISBN=978-0-8493-2675-2}}</ref>

===Taxonomy===
''Cephalanthus'' is the most [[Basal (phylogenetics)|basal]] genus in the [[Tribe (biology)|tribe]] [[Naucleeae]].<ref name="manns2010">Manns, U. and B. Bremer. 2010. [http://www.bergianska.se/polopoly_fs/1.68533.1325518850!/menu/standard/file/Manns_Bremer_2010.pdf Towards a better understanding of intertribal relationships and stable tribal delimitations within Cinchonoideae s.s. (Rubiaceae).] ''Molecular Phylogenetics and Evolution'' 56(1), 21-39. {{doi|10.1016/j.ympev.2010.04.002}}</ref> Some authors have [[Segregate (taxonomy)|segregated]] it into its own [[Monotypic taxon|monotypic]] tribe.<ref name="ridsdale1976">{{cite journal|author=Ridsdale CE|year=1976|title=A revision of the tribe Cephalantheae (Rubiaceae)|journal=Blumea|volume=23|issue=1|pages=177–88}}</ref> The [[type species]] is ''[[Cephalanthus occidentalis]]''.<ref>[http://www.botany.si.edu/ing/INGsearch.cfm?searchword=Cephalanthus ''Cephalanthus''.] Index Nominum Genericorum. Smithsonian National Museum of Natural History.</ref>

===Species===
*''[[Cephalanthus angustifolius]]'' [[João de Loureiro|Lour.]] - [[Laos]], [[Cambodia]], [[Vietnam]]
*''[[Cephalanthus glabratus]]'' ([[Curt Polycarp Joachim Sprengel|Spreng.]]) [[Karl Moritz Schumann|K.Schum.]] - ''sarandí'' - [[Brazil]], [[Argentina]], [[Paraguay]], [[Uruguay]]
*''[[Cephalanthus natalensis]]'' [[Daniel Oliver|Oliv.]] - [[Tanzania]], [[Malawi]], [[Zambia]], [[Lesotho]], [[Swaziland]], [[South Africa]]
*''[[Cephalanthus occidentalis]]'' [[Carl Linnaeus|L.]] - button-willow, common buttonbush, honey-bells - [[Cuba]], eastern [[Canada]], eastern, central and southern [[United States]], [[California]], [[Arizona]], [[New Mexico]]
*''[[Cephalanthus salicifolius]]'' [[Alexander von Humboldt|Humb.]] & [[Aimé Bonpland|Bonpl.]] - Mexican buttonbush, willowleaf buttonbush - [[Mexico]], [[Honduras]], extreme southern tip of [[Texas]]
*''[[Cephalanthus tetrandra]]'' ([[William Roxburgh|Roxb.]]) [[Colin Ernest Ridsdale|Ridsdale]] & [[Reinier Cornelis Bakhuizen van den Brink, Jr.|Bakh.f.]] - [[tropical Asia]] from [[India]] to [[China]] and [[Thailand]]

==References==
{{Reflist}}

==External links==
*[http://apps.kew.org/wcsp/namedetail.do?name_id=36697 Kew World Checklist of Selected Plant Families, ''Cephalanthus'']
*[http://plants.usda.gov/java/profile?symbol=CEPHA USDA PLANTS, ''Cephalanthus'']
*[http://tai2.ntu.edu.tw/udth/bin/fot1.exe/browse?BID=4&page=248 Flora of Taiwan, ''Cephalanthus'']

{{Taxonbar|from=Q3245386}}

[[Category:Cephalanthus| ]]
[[Category:Rubiaceae genera]]
[[Category:Flora of the United States]]
[[Category:Flora of Canada]]
[[Category:Flora of Mexico]]
[[Category:Flora of Honduras]]
[[Category:Flora of India]]
[[Category:Flora of China]]
[[Category:Flora of Assam (region)]]
[[Category:Flora of Bangladesh]]
[[Category:Flora of Indo-China]]
[[Category:Flora of Cuba]]
[[Category:Flora of Brazil]]
[[Category:Flora of Argentina]]
[[Category:Flora of Paraguay]]
[[Category:Flora of Uruguay]]
[[Category:Flora of Tanzania]]
[[Category:Flora of South Africa]]

Cephalanthus salicifolius

2018-03-22T17:53:08Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q5063253}} (7 sig. taxon IDs); WP:GenFixes using AWB

{{Italic title}}
{{taxobox
|name =Mexican buttonbush
|image =
|image_caption =
|regnum = [[Plant]]ae
|unranked_divisio = [[Flowering plant|Angiosperms]]
|unranked_classis = [[Eudicots]]
|unranked_ordo = [[Asterids]]
|ordo = [[Gentianales]]
|familia = [[Rubiaceae]]
|genus = ''[[Cephalanthus]]''
|species = '''''C. salicifolius'''''
|binomial = ''Cephalanthus salicifolius''
|binomial_authority = [[Alexander von Humboldt|Humb.]] & [[Aimé Bonpland|Bonpl.]],<ref name="ITIS"/> 1809<ref>{{cite web |url=http://www.tropicos.org/name/27903092 |title=''Cephalanthus salicifolius'' Bonpl. |work=TROPICOS |publisher=Missouri Botanical Garden |accessdate=2009-10-10}}</ref>
|synonyms_ref=<ref>[http://apps.kew.org/wcsp/synonomy.do?name_id=36739 Kew World Checklist of Selected Plant Families, synonymy, ''Cephalanthus salicifolius'']</ref>
|synonyms=*''Cephalanthus occidentalis'' var. ''salicifolius'' (Humb. & Bonpl.) A.Gray
*''Cephalanthus occidentalis'' subsp. ''salicifolius'' (Humb. & Bonpl.) Borhidi & Diego
*''Cephalanthus peroblongus'' Wernham
|}}

'''''Cephalanthus salicifolius''''' is a species of [[flowering plant]] in the [[cinchona]] family, [[Rubiaceae]].<ref>[https://www.biodiversitylibrary.org/page/279219#page/142/mode/1up Humboldt, Friedrich Wilhelm Heinrich Alexander von & Bonpland, Aimé Jacques Alexandre. 1809. Plantae Aequinoctiales 2: 63–64, pl. 98. ''Cephalanthus salicifolius]</ref><ref>Davidse, G., M. Sousa Sánchez, S. Knapp & F. Chiang Cabrera. 2012. Rubiaceae a Verbenaceae. 4(2): i–xvi, 1–533. In G. Davidse, M. Sousa Sánchez, S. Knapp & F. Chiang Cabrera (eds.) Flora Mesoamericana. Missouri Botanical Garden Press, St. Louis.</ref><ref name="ITIS">{{cite web |url=https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=34787 |title=''Cephalanthus salicifolius'' Humb. & Bonpl. |work=ITIS Standard Reports |publisher=Integrated Taxonomic Information System |accessdate=2009-10-10}}</ref> Common names include '''Mexican buttonbush''', '''mimbre''', '''botoncillo''', and '''Jazmin blanco'''.<ref name="TexasNative">{{cite web |url=http://aggie-horticulture.tamu.edu/ornamentals/nativeshrubs/cephalanthussalicifol.htm |title=Mexican Buttonbush, Mimbre, Botoncillo, Jazmin Blanco ''Cephalanthus salicifolius'' |work=Texas Native Shrubs |publisher=Texas A&M University |accessdate=2009-10-10}}</ref> Its native range extends from the banks of the [[Rio Grande Valley|southernmost stretch]] of the [[Rio Grande]] in [[Cameron County, Texas|Cameron]] and [[Hidalgo County, Texas|Hidalgo Counties]] of [[Texas]]<ref>{{cite paper |url=http://nativeplantproject.com/SABALS/wetland_shrubs_FINAL.pdf |title=Wonderful and Woody Shrubs of the Water’s Edge...and Beyond |first=Christina |last=Mild |page=2 |publisher=Native Plant Project |format=PDF |accessdate=2009-10-10}}</ref> through much of [[Mexico]] from [[Coahuila]] to [[Oaxaca]]; a [[Disjunct distribution|disjunct population]] exists in [[Honduras]].<ref name="Correll">{{cite book |url=https://books.google.com/books?id=yZSrAAAAIAAJ&source=gbs_navlinks_s |title=Aquatic and Wetland Plants of Southwestern United States |volume=Volume 2 |first=Donovan Stewart |last=Correll |author2=Helen B. Correll |publisher=Stanford University Press |year=1975 |isbn=978-0-8047-0866-1 |page=1551}}</ref><ref>[http://bonap.net/MapGallery/County/Cephalanthus%20salicifolius.png Biota of North America Program, distribution map, ''Cephalanthus salicifolius'']</ref>

Like other species in its genus, Mexican Buttonbush grows in the wet soils of [[riparian zone]]s, [[swamp]]s, and pond margins.<ref>{{cite paper |url=http://nativeplantproject.tripod.com/PondandWetland.pdf |title=Native Pond and Wetland Plants of the Rio Grande Valley, Texas: Landscape Uses and Identification |publisher=Native Plant Project |format=PDF |page=32 |accessdate =2009-10-10}}</ref> It is a [[deciduous]] [[shrub]] or small [[tree]], reaching a height of {{convert|8|-|18|ft|m|abbr=on}} and a width of {{convert|4|-|10|ft|m|abbr=on}}.<ref name="TexasNative"/> The oblong [[Leaf|leaves]] reach {{convert|12|cm|in|abbr=on}} in length and {{convert|23|mm|in|abbr=on}} in width.<ref name="Correll"/>
The white [[flower]]s are produced from March to July; the [[fruit]] is a collection of brown [[Pyrena|nutlets]].<ref>{{cite book |url=https://books.google.com/books?id=8cEq7weUPYYC&source=gbs_navlinks_s |title=Trees, Shrubs, and Cacti of South Texas
|first=J. H. |last=Everitt |author2=Dale Lynn Drawe |author3=Robert I. Lonard |publisher=Texas Tech University Press |year=2002 |isbn=978-0-89672-473-0 |page=175}}</ref>

==References==
{{Reflist}}

==External links==
*[https://aggie-horticulture.tamu.edu/ornamentals/nativeshrubs/cephalanthussalicifol.htm Texas A&M University, Texas Native Plants Database, Mexican buttonbush]
*[https://web.archive.org/web/20140501235511/http://it.gardening.eu/arc/piante/Alberature/Cephalanthus-salicifolius-Humb.-Bonpl/13868/ Gardening Europe, ''Cephalanthus salicifolius'']
*[http://fm1.fieldmuseum.org/vrrc/index.php?language=br&page=view&id=47530&PHPSESSID=6f92cce9194c815acd5cb0e8cb60cb88 Field Museum in Chicago, photo of herbarium specimen of ''Cephalanthus salicifolius'' collected in Honduras]

{{Commons|Cephalanthus salicifolius}}

{{Taxonbar|from=Q5063253}}

[[Category:Cephalanthus|salicifolius]]
[[Category:Plants described in 1809]]
[[Category:Flora of Honduras]]
[[Category:Flora of Mexico]]
[[Category:Flora of Texas]]
[[Category:Flora of the U.S. Rio Grande Valleys]]
[[Category:Taxa named by Aimé Bonpland]]
[[Category:Taxa named by Alexander von Humboldt]]

{{rubiaceae-stub}}

Lomatium roseanum

2018-03-22T17:38:55Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q6669268}} (5 sig. taxon IDs); WP:GenFixes using AWB

{{taxobox
|name =
|image = Lomatium roseanum -21235 (14677976582).jpg
|status = G2
|status_system = TNC
|regnum = [[Plantae]]
|unranked_divisio = [[Angiosperms]]
|unranked_classis = [[Eudicots]]
|unranked_ordo = [[Asterids]]
|ordo = [[Apiales]]
|familia = [[Apiaceae]]
|genus = ''[[Lomatium]]''
|species = '''''L. roseanum'''''
|binomial = ''Lomatium roseanum''
|binomial_authority = [[Arthur Cronquist|Cronquist]] 1992
|synonyms_ref=<ref>[http://www.tropicos.org/Name/50098810 Tropicos, ''Lomatium roseanum'' Cronquist ]</ref>
|synonyms=*''Leptotaenia leibergii'' J.M.Coult. & Rose 1900, not ''Lomatium leibergii'' J.M. Coult. & Rose 1900
|}}
'''Adobe parsley''' ('''''Lomatium roseanum'''''), also known as '''adobe lomatium''' and '''rose-flowered desert-parsley''', is a very [[rare species|rare plant]] of the Western U.S., known only from northwestern [[Nevada]] and southeastern [[Oregon]], and which may also occur in northeastern [[California]]. The largest populations occur on the [[Sheldon National Wildlife Refuge]].<ref name="Morefield 2001">
{{cite web
| last = Morefield
| first = J.D.
| title = ''Lomatium roseanum''
| work = Nevada Rare Plant Atlas
| publisher = Nevada Natural Heritage Program
| date = 25 June 2001
| url = http://heritage.nv.gov/atlas/atlas.html
| accessdate = 2007-03-28}}</ref> It is a member of the celery family, the [[Apiaceae|Umbelliferae]], and has yellow flowers.

==Technical description==
*'''[[Plant]]''' perennial, long-lived, usually over 10 cm tall, glabrous, somewhat caulescent at the base.
*'''[[Root]]''' tuberous, thick.
*'''[[Leaf|Leaves]]''' green, shiny, ternate, with ternate-pinnately dissected leaflets, the ultimate segments numerous and small (<1 cm); petiole dilated (flared out) and sheathing at the base.
*'''Scapes''' (peduncles) ascending, arcuate (curved), 15–20 cm long.
*'''Involucels''' with slender bractlets.
*'''[[Flower]]s''' yellow, aging to whitish.
*'''[[Fruit]]''' only very narrowly laterally winged; dorsal ribs wingless.<ref name="Morefield 2001"/><ref name="Cronquist 1997">
{{cite book
| last = Cronquist
| first = Arthur |author2=Noel H. Holmgren |author3=Patricia K. Holmgren
| year = 1997
| chapter = Apiaceae
| title = Intermountain Flora: Vascular Plants of the Intermountain West, U.S.A. Volume Three, Part A: Subclass Rosidae (except Fabales)
| pages = 412–413}}</ref>

==Distribution, habitat, and ecology==
The range of adobe parsley is extremely limited: it is known only from [[Washoe County, Nevada|Washoe County]] in Nevada and nearby places in southeastern Oregon, where it has possibly already been extirpated. It is also suspected to exist in [[Humboldt County, Nevada]]. Fewer than 20 known populations exist, though these can be large.<ref name="Morefield 2001"/>

Adobe parsley lives in loose, rocky habitat. Specifically, it prefers dry [[basalt]] talus scree overlying [[clay]] soils. It associates with the [[low sagebrush]] community, and specific associates include ''[[Artemisia arbuscula]]'', ''[[Poa secunda]]'', ''[[Elymus elymoides]]'', ''[[Arenaria aculeata]]'', ''[[Phlox]]'' spp., ''[[Erigeron linearis]]'', etc.<ref name="Morefield 2001"/>

The [[fire ecology]] is unknown for members of this [[genus]]. The [[low sagebrush]] community type within which adobe parsley lives generally lacks enough [[fuel]]s to carry a fire. When it does burn, these non-fire adapted shrubs are usually killed and replaced by [[Taeniatherum caput-medusae|medusahead]] and [[cheatgrass]], and re-establish low sagebrush vegetation through time (2 to 5 years) via [[seed]]s.<ref name="Steinberg 2002">
{{cite web
| last = Steinberg
| first = Peter D.
| title = ''Artemisia arbuscula''
| work = Fire Effects Information System
| publisher = USDA FS RMRS Fire Sciences Laboratory
| date = 2002
| url = http://www.fs.fed.us/database/feis/
| accessdate = 2007-03-28}}</ref>

==Conservation status and threats==
*[[United States Forest Service|U.S. Forest Service]]: Pacific Southwest Region Sensitive Species
*[[California Native Plant Society]]: Not Listed (not known from California)
*[[Nevada Natural Heritage Program]]: Sensitive Plant
*[[NatureServe]] Nevada State Rank: S2S3; Global Rank: G2

==Field identification==
Adobe parsley occurs between {{convert|5750|ft|m}} and {{convert|6175|ft|m}} above sea level. It flowers and may be most easily recognized in early spring, i.e., between April and June. It stands out because of its tuberous root, yellow flowers, and green shiny leaves.<ref name="Morefield 2001"/> It resembles ''[[Lomatium hendersonii|L. hendersonii]]'', which is found farther north, but is more robust (larger) and somewhat caulescent.<ref name="Cronquist 1997"/>

==References==
{{Reflist}}

==External links==
{{Wikispecies-inline}}
*[http://plants.usda.gov/java/profile?symbol=LORO7&mapType=nativity&photoID= USDA Plants Profile of ''Lomatium roseanum'']
*[http://www.calflora.org/cgi-bin/species_query.cgi?where-calrecnum=9475 Calflora Database: ''Lomatium roseanum'' (Biscuitroot)]
* [http://www.natureserve.org/explorer/index.htm official NatureServe website]
{{commonscat|position=left}}

{{Taxonbar|from=Q6669268}}

[[Category:Lomatium|roseanum]]
[[Category:Flora of California]]
[[Category:Flora of Nevada]]
[[Category:Flora of Oregon]]
[[Category:Flora of the Great Basin]]
[[Category:Endemic flora of the United States]]
[[Category:Endangered flora of the United States]]
[[Category:Endangered flora of California]]
[[Category:Plants described in 1900]]
[[Category:Taxa named by John Merle Coulter]]

Globitermes sulphureus

2018-03-22T17:27:01Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q4039502}} (1 sig. taxon ID); WP:GenFixes, using AWB

{{Taxobox
| image =
| image_caption =
| regnum = [[Animal]]ia
| phylum = [[Arthropod]]a
| classis = [[Insect]]a
| ordo = [[Isoptera]]
| familia = [[Termitidae]]
| genus = ''[[Globitermes]]''
| species = '''''G. sulphureus'''''
| binomial = ''Globitermes sulphureus''
| binomial_authority = [[George Darby Haviland|Haviland]]<ref name="Bordereau" />
}}

'''''Globitermes sulphureus''''' is a species of [[termite]] that is very common in Central and Southern [[Vietnam]]<ref name="Bordereau">{{cite journal|doi=10.1007/s000400050049|last=Bordereau|first=C. |author2=A. Robert |author3=V. Van Tuyen |author4=A. Peppuy|date=August 1997|title=Suicidal defensive behaviour by frontal gland dehiscence in Globitermes sulphureus Haviland soldiers (Isoptera)|journal=Insectes Sociaux|publisher=Birkhäuser Basel|volume=44|issue=3|pages=289–297|issn=1420-9098|url=http://www.springerlink.com/content/m727aywa4mdf04ln/}}</ref> and also present in other areas of South East Asia, including [[Malaysia]].<ref name="malaysia">{{cite journal|last=Majid|first=Abdul Hafiz Ab.|author2=Abu Hassan Ahmad|author3=Rashid M Z A|author4=Che Salmah Md Rawi|year=2007|title=Preliminary field efficacy of imidacloprid on Globitermes sulphureus (Isoptera: Termitidae) (Subterranean termite) in Penang|journal=Jurnal Biosains|volume=18|issue=2|pages=109–114|url=http://myais.fsktm.um.edu.my/6666/1/18-02-Art9SC.pdf|deadurl=yes|archiveurl=https://web.archive.org/web/20110722233232/http://myais.fsktm.um.edu.my/6666/1/18-02-Art9SC.pdf|archivedate=2011-07-22|df=}}</ref> They live in nests made of earth that can be up to 1.5 m tall and can contain tens of thousands of individuals. Between five and 10 per cent of the population are soldier termites which can be recognised by their yellow [[abdomen]] and two large, curved mandibles. The termites use [[autothysis]] as a [[defense mechanism]].<ref name="Bordereau" />

==Defense==
[[File:Globitermes sulphureus soldier.svg|thumb|left|A drawing of the head of a soldier termite showing the head (H), antennae (A) and the mandibles (M). Scale bar = 0.3mm]]
When the nest is damaged, the soldier termites defend the nest and workers rapidly repair any damage to the nest walls. The soldiers stand on alert on their legs and scan the area with their antennae. If the nest is attacked by [[ant]]s, for example ''[[Oecophylla smaragdina]]'', the workers immediately escape by retreating into the nest, while the soldiers remain outside to defend the nest. They will first attempt to catch and pierce the attackers with their large mandibles. If this does not deter the attackers, some soldiers secrete a yellow liquid from a large gland that occupies a large proportion of their body. The liquid is forced out of the gland by contractions of the mandibular muscles which compress the walls of the gland. This liquid rapidly hardens on contact with air, producing a sticky substance that traps ants and other termites.<ref name="Bordereau" /> The secretion also contains a [[pheromone]] which attracts more soldiers to fight the attackers. In some cases, the contractions are so violent that the termites rupture themselves. This form of suicidal [[altruism in animals|altruism]] is known as [[autothysis]].<ref name="Bordereau" />

==Nitrogen fixation==
''G. sulphureus'' is able to [[nitrogen fixation|fix nitrogen]]. An experiment in [[Thailand]] found that they fix around 250 grams of nitrogen per [[hectare]] per year. Although this contribution is only between 7% and 22% of the total nitrogen inputs in the [[ecosystem]] it is thought to be relatively important as termites add it to dead wood, thereby accelerating its [[decomposition]].<ref name="nitrogen">{{cite journal|doi=10.1007/s10021-005-0024-7|last=Yamada|first=Akinori |author2=Tetsushi Inoue |author3=Decha Wiwatwitaya |author4=Moriya Ohkuma |author5=Toshiaki Kudo |author6=Atsuko Sugimoto|date=February 2006|title= Nitrogen Fixation by Termites in Tropical Forests, Thailand |journal=Ecosystems|publisher=Springer New York|volume=9|issue=1|pages=75–83|issn=1435-0629|url=http://www.springerlink.com/content/5n7m7w55g3627v2l/}}</ref>

==Foraging==
''G. sulphureus'' forages areas of up to 62m2 and ventures up to 16m away from its nest.<ref name="lee">{{cite journal|last=Lee|first=Chow-Yang|author2=Jocelyn Yap|author3=Peng-Soon Ngee|author4=Zairi Jaal|year=2003|title=Foraging Colonies of a Higher Mound-building Subterranean Termite, Globitermes sulphureus (Haviland) in Malaysia|journal=Japanese Journal of Environmental Entomological Zoology|volume=14|issue=2|pages=105–112|url=http://idisk.mac.com/chowyang/Public/054.pdf|deadurl=yes|archiveurl=https://web.archive.org/web/20110718182511/http://idisk.mac.com/chowyang/Public/054.pdf|archivedate=2011-07-18|df=}}</ref>

==Control==
The species is a major pest in areas it inhabits as it attacks wooden structures<ref name="Bordereau" /> and can damage [[coconut]] and [[oil palm]] plantations.<ref name="lee" /> Experiments have shown that it can be controlled using the insecticide [[imidacloprid]].<ref name="malaysia" />

==See also==
''[[Camponotus saundersi]]'' is an ant with a similar defensive mechanism.

==References==
{{Reflist}}

==External links==
* [http://www.jcc2u.com/book_termite/images/07.jpg Photograph of a nest and a soldier]

{{Taxonbar|from=Q4039502}}

[[Category:Termites]]
[[Category:Exploding animals]]
[[Category:Insects of Vietnam]]

Chlorovirus

2018-03-22T17:26:02Z

Tom.Bot: Task 3: +{{Taxonbar|from=Q18822090}} (1 sig. taxon ID); WP:GenFixe using AWB

{{Use dmy dates|date=April 2017}}
{{taxobox
| virus_group = i
| familia = ''[[Phycodnaviridae]]''
| genus = '''''Chlorovirus'''''
| subdivision_ranks = Type Species
| subdivision =
*''[[Paramecium bursaria Chlorella virus 1]]''
}}

'''''Chlorovirus''''' is a genus of giant double-stranded [[DNA virus]], in the family [[Phycodnaviridae]]. Alga serve as natural hosts. There are currently 19 species in this genus including the type species [[Paramecium bursaria Chlorella virus 1]].<ref name=ViralZone>{{cite web|title=Viral Zone|url=http://viralzone.expasy.org/all_by_species/588.html|publisher=ExPASy|access-date=15 June 2015}}</ref><ref name=ICTV>{{cite web|last1=ICTV|title=Virus Taxonomy: 2014 Release|url=http://ictvonline.org/virusTaxonomy.asp|access-date=15 June 2015}}</ref> One species, [[Chlorovirus ATCV-1]], commonly found in freshwater lakes, has been found to infect humans.<ref name=":4">{{cite journal | vauthors = Yolken RH, Jones-Brando L, Dunigan DD, Kannan G, Dickerson F, Severance E, Sabunciyan S, Talbot CC, Prandovszky E, Gurnon JR, Agarkova IV, Leister F, Gressitt KL, Chen O, Deuber B, Ma F, Pletnikov MV, Van Etten JL | title = Chlorovirus ATCV-1 is part of the human oropharyngeal virome and is associated with changes in cognitive functions in humans and mice | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 111 | issue = 45 | pages = 16106–11 | date = November 2014 | pmid = 25349393 | pmc = 4234575 | doi = 10.1073/pnas.1418895111 | url = http://www.pnas.org/content/111/45/16106.abstract }}</ref>

==Taxonomy==

Chlorovirus belongs to '''Group 1: dsDNA viruses''', and is a genus of giant [[double stranded DNA]], in the family of '''[[Phycodnaviridae]]'''.

== Structure ==
Viruses in ''Chlorovirus'' are enveloped, with icosahedral and spherical geometries, and T=169 symmetry. The diameter is around 100-220 nm. Genomes are linear, around 330 kb in length. The genome has 700 open reading frames.<ref name=ViralZone /> Chloroviruses are large and contain dsDNA (double stranded DNA). As a group chloroviruses encode from 632 protein families, however, each individual virus only has 330 to 416 protein encoding genes. As part of the DNA modification systems, chloroviruses have [[DNA methylation|methylated bases]] in specific sections of their DNA sequence. Some chloroviruses also contain [[introns]] and inteins, though this is rare within the genus.<ref name="journals.plos.org">{{cite journal | vauthors = Van Etten JL, Dunigan DD | title = Giant Chloroviruses: Five Easy Questions | journal = PLoS Pathogens | volume = 12 | issue = 8 | pages = e1005751 | date = August 2016 | pmid = 27536965 | doi = 10.1371/journal.ppat.1005751 | url = http://journals.plos.org/plospathogens/article?id=10.1371/journal.ppat.1005751 }}</ref>

The type species PBCV-1 (Paramecium bursaria Chlorella virus 1) have a 190 nm diameter <ref name="journals.plos.org"/> and a fivefold axis.<ref name="Cristian F 2016">{{cite journal|last1=Quispe|first1=Cristian F.|last2=Esmael|first2=Ahmed|last3=Sonderman|first3=Olivia|last4=McQuinn|first4=Michelle|last5=Agarkova|first5=Irina|last6=Battah|first6=Mohammed|last7=Duncan|first7=Garry A.|last8=Dunigan|first8=David D.|last9=Smith|first9=Timothy P.L.|last10=De Castro|first10=Cristina|last11=Speciale|first11=Immacolata|last12=Ma|first12=Fangrui|last13=Van Etten|first13=James L.|title=Characterization of a new chlorovirus type with permissive and non-permissive features on phylogenetically related algal strains|journal=Virology|date=January 2017|volume=500|pages=103–113|doi=10.1016/j.virol.2016.10.013}}</ref> One faces juncture has a protruding spike, which is the first part of the virus to contact a host.<ref name="James L 2011">{{cite journal|last1=Van Etten|first1=James L.|last2=Dunigan|first2=David D.|title=Chloroviruses: not your everyday plant virus|journal=Trends in Plant Science|date=January 2012|volume=17|issue=1|pages=1–8|doi=10.1016/j.tplants.2011.10.005}}</ref> The outer [[capsid]] covers a single lipid bilayer membrane, which is obtained from the hosts endoplasmic reticulum.<ref name="Cristian F 2016"/> Some capsomers on the external shell have fibres extending away from the virus to aid in host attachment.<ref name="ReferenceA">{{cite journal|last1=Van Etten|first1=James L.|last2=Dunigan|first2=David D.|last3=Condit|first3=Richard C.|title=Giant Chloroviruses: Five Easy Questions|journal=PLOS Pathogens|date=18 August 2016|volume=12|issue=8|pages=e1005751|doi=10.1371/journal.ppat.1005751}}</ref><ref name="James L 2011"/> The dsDNA is non-sequenced and about 331 kb and is closed with a hairpin structure terminus. Generally, the DNA is only found with a single copy. PBCV-1 has 148 viral proteins and a minimum of one host encoded protein. This type has 416 protein encoding genes.<ref name="journals.plos.org"/>

{| class="wikitable sortable" style="text-align:center"
|-
! Genus !! Structure || Symmetry !! Capsid !! Genomic arrangement !! Genomic segmentation
|-
|Chlorovirus||Icosahedral||T=169||Enveloped||Linear||Monopartite
|}

==Hosts==
Chloroviruses infect certain unicellular, eukaryotic, exsymbiontic chlorella-like green algae, called [[zoochlorellae]]. Chloroviruses are very species and even strain specific. Zoochlorellae are associated with the protozoan ''[[Paramecium bursaria]]'', the coelenterate ''[[Hydra viridis]]'', the heliozoon ''[[Acanthocystis turfacea]]'' and other freshwater and marine invertebrates and protozoans. Chloroviruses are common in inland waters throughout the world with titers as high as thousands of plaque-forming units (PFUs) per milliliter of indigenous water, although titers are typically 1-100 PFUs/mL. The viruses cannot infect zoochlorellae when they are in their symbiotic phase and there is no evidence that zoochlorellae grow free of their hosts in indigenous waters.<ref>{{cite journal | vauthors = Van Etten JL, Dunigan DD | title = Chloroviruses: not your everyday plant virus | journal = Trends in Plant Science | volume = 17 | issue = 1 | pages = 1–8 | date = January 2012 | pmid = 22100667 | pmc = 3259250 | doi = 10.1016/j.tplants.2011.10.005 }}</ref>

==Life cycle==
Viral replication is nucleo-cytoplasmic. Replication follows the DNA strand displacement model, and DNA templated transcription is the method of transcription. The virus exits the host cell by lysis via lytic phospholipids, with passive diffusion being the mechanism behind transmission routes.

In three dimensional recreations of PBCV-1 ([[''Paramecium bursaria chlorella virus'']]), a prototype of chlorovirus, it is seen that the spike first contacts the host’s cell wall<ref>Zhang X, Xiang Y, Dunigan DD, Klose T, Chipman PR, Van Etten JL, Rossmann MG: Three dimensional structure and function of the Paramecium bursaria chlorella virus capsid. Proc. Natl. Acad. Sci. USA 2011, 108:14837–14842. doi: 10.1073/pnas.1107847108 {{PMID|21873222}}</ref> and is aided by fibres in order to secure the virus to the host. The attachment of PBCV-1 to it’s receptor is very specific, and a major source of limitation of PBCV-1 applicable range of hosts. Virus-associated enzymes allow the host cell wall to degrade, and the viral internal membrane fuses with the host membrane. This fusion allows the transfer of viral DNA and virion-associated proteins into the host cell and also triggers depolarization of the host membrane. This is presumably occurring due to a virus encoded K+ channel. Studies predict this channel is within the virus, acting as an internal membrane, and releasing K+ from the cell which may assist in the ejection of viral DNA and proteins from the viral cell to it’s host. The [[depolarization]] of the host’s cell membrane is also though to prevent secondary infection from another virus or secondary transporters.

[[File:Chlorella cells and chlorovirus Paramecium bursaria chlorella virus (PBCV-1).png|thumb|'''Chlorella cells and chlorovirus Paramecium bursaria chlorella virus (PBCV-1)''' (A) PBCV-1 and its symbiotic chlorella cells. (B) Plaques formed as a result of PBCV-1 on Chlorella variabilis. ( C) 5 times averaged electron micrograph of PBCV-1 displayins a long narrow spike at one of it’s verticies with fibres extending. (D) PBCV-1 attached to the cell wall. (E) Surface view of PBCV-1 spike/fibres. (F) Initial attachment of PBCV-1 to a C.variabilis cell. (G) Digestion of the cell wall once PBCV-1 has attached (1-3 minutes postinfection). (H) Virion particles assembling within the cytoplasm, marking virus assembly centers approximately 4 hours post infection. (I) Depiction of PBCV-1 assembling into infectious particles. (J) Localized lysis of cell wall/plasma membrane, and release of progeny viruses approximately 8 hours postinfection.]]<ref name="Van Etten 2012">Van Etten, James L.; Dunigan, David D. (January 2012). "Chloroviruses: not your everyday plant virus". Trends in Plant Science. 17 (1): 1–8. doi:10.1016/j.tplants.2011.10.005.</ref>

Because PBCV-1 does not have an [[RNA polymerase]] gene, its DNA and viral-associated proteins move to the nucleus where transcription begins 5–10 minutes post infection. This rapid transcription is attributed to some component facilitating this transfer or viral DNA to the nucleus. This component is assumed to be a product of the PBCV-a443r gene, which obtains structures resembling proteins involved in nuclear trafficking in mammalian cells.

Host transcription rates decrease in this early phase of infection and host transcription facilitators are reprogrammed to transcribe the new viral DNA. Minutes after infection, host chromosomal DNA degradation begins. This is presumed to occur through PBCV-1 encoded and packaged DNA restriction endonucleases. Degradation of the host chromosomal DNA inhibits host transcription. This results in 33-55% of the polyadenylated [[mRNAs]] in the infected cell being of viral origin by 20 minutes after initial infection.<ref>Blanc G, Mozar M, Agarkova IV, Gurnon JR, Yanai Balser G, Rowe JM, Xia Y, Riethoven JJ, Dunigan DD, Van Etten JL: Deep RNA sequencing reveals hidden features and dynamics of early gene transcription in Paramecium bursaria chlorella virus 1. PLoS ONE 2014, 9:e90989. doi: 10.1371/journal. pone.0090989 {{PMID|24608750}}</ref>

Viral DNA replication initiates after 60 to 90 minutes, which is then followed by the transcription of late genes within the host cell. Roughly 2–3 hours post infection, the assembly of virus capsids begins. This occurs within localized regions of the cytoplasm, with the virus capsids becoming prominent 3–4 hours after initial infection.
5–6 hours after PBCV-1 infection, the cytoplasm of the host cell fills with infectious progeny virus particles. A short amount of time later, (6–8 hours post infection) localized [[lysis]] of the host cell releases progeny. ~1000 particles are released from each infected cell, with ~30% of which forming plaques.<ref name="Van Etten 2012"/>

== Effects of Infection ==
In algae infected with Cloroviruses the result is lysis, and thus death. As such, Chloroviruses are an important mechanism to the termination of algal blooms and play a vital role in the supply of nutrients to the water column.<ref name="ReferenceA"/>

Humans found to be infected with ATCV-1 showed a decreased visual processing ability and reduced visual motor speed. This led to an overall decline in the ability to perform tasks based on vision and spatial reasoning.<ref name=":4" />

Recent studies conducted on ACTV-1 infected mice show changes in the [[Cyclin-dependent kinase 5|Cdk5 pathway]], which aides with learning and memory formation, as well as alterations in gene expression in the dopamine pathway.<ref name=":4" /> Further, mice infected were found to be less social - interacting less with newly introduced companion mice than the control group. Infected mice also spent an increased time in a light exposed portion of a test chamber, where the control mice tended to prefer the dark side and avoided the light. This indicates a decrease in anxiety with ACTV-1 infection. The test mice were also less able to recognize an object had been moved from its previous location, showing a decrease in spatial reference memory.<ref name="Marilyn S 2016">{{cite journal|last1=Petro|first1=Marilyn S.|last2=Agarkova|first2=Irina V.|last3=Petro|first3=Thomas M.|title=Effect of Chlorovirus ATCV-1 infection on behavior of C57Bl/6 mice|journal=Journal of Neuroimmunology|date=August 2016|volume=297|pages=46–55|doi=10.1016/j.jneuroim.2016.05.009}}</ref> As in humans, there is a decrease in vision spatial task ability. Within the hippocampus changes in gene expression occur and infection presents a change in the pathways of immune cell functioning and antigen processing. This indicates an immune system response to the ACTV-1 virus causing inflammation which may be the cause for the cognitive impairments.<ref name=":4" /> The symptoms presented suggest hippocampus and medial prefrontal cortex interference from ACTV-1 infection.<ref name="Marilyn S 2016"/>

== Ecology ==
Chloroviruses are widespread in [[freshwater]] environments in all parts of the globe and have been isolated from freshwater sources in [[Europe]], [[Asia]], [[Australia]], as well as [[North America|North]] and [[South America]].<ref name=":0">{{cite journal | vauthors = Quispe CF, Sonderman O, Seng A, Rasmussen B, Weber G, Mueller C, Dunigan DD, Van Etten JL | title = Three-year survey of abundance, prevalence and genetic diversity of chlorovirus populations in a small urban lake | journal = Archives of Virology | volume = 161 | issue = 7 | pages = 1839–47 | date = July 2016 | pmid = 27068168 | doi = 10.1007/s00705-016-2853-4 }}</ref><ref>{{cite journal | vauthors = Short SM | title = The ecology of viruses that infect eukaryotic algae | journal = Environmental Microbiology | volume = 14 | issue = 9 | pages = 2253–71 | date = September 2012 | pmid = 22360532 | doi = 10.1111/j.1462-2920.2012.02706.x | url = http://onlinelibrary.wiley.com/doi/10.1111/j.1462-2920.2012.02706.x/abstract }}</ref> Natural hosts of chloroviruses include various types of unicellular eukaryotic ''Chlorella''-like algae, with individual virus species typically infecting only within a distinct strain. These algal hosts are known to establish [[Endosymbiont|endosymbiotic]] relationships with larger protists, such as ''[[Paramecium bursaria]]'' (a member of the [[ciliate]]s), ''Acanthocystis turfacea'' (a [[Centrohelid|centroheliozoan]]) and ''Hydra viridis'' (member of the [[hydrozoa]]).<ref name="journals.plos.org"/> While an individual protist can harbour up to several hundred algal cells at any given time, free-floating algae are highly susceptible to chloroviruses, indicating that such endosymbiosis serves to provide resistance from infection.<ref name=":1">{{cite journal | vauthors = DeLong JP, Al-Ameeli Z, Duncan G, Van Etten JL, Dunigan DD | title = Predators catalyze an increase in chloroviruses by foraging on the symbiotic hosts of zoochlorellae | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 113 | issue = 48 | pages = 13780–13784 | date = November 2016 | pmid = 27821770 | doi = 10.1073/pnas.1613843113 | url = http://www.pnas.org/content/113/48/13780 }}</ref>

Chlorovirus titers are variable by season and location, but typically fluctuate between 1 and 100 PFU/mL, although high abundances of up to 100,000 PFU/mL may occur in some environments. Due to the rich genetic diversity and high specialization of individual species with respect to infectious range, variations in their ecology are not unusual, resulting in unique spatio-temporal patterns, which ultimately depend on lifestyle and nature of the host. As such, previous survey data highlighted two prominent seasonal abundance peaks for both ''Chlorella variabilis'' NC64A and ''Chlorella variabilis'' Syngen viruses — one in late fall, and the other in late spring to mid-summer — which is likely attributed to the fact that they share a host species. Conversely, ''Chlorella heliozoae'' SAG viruses peaked at different times of the year and generally exhibited more variability in titers, as compared to the NC64A and Syngen viruses.<ref name=":0" /> Additionally, studies revealed that chloroviruses demonstrate some resilience in response to decreased temperatures observed during the winter season, characterized by presence of infectious particles under ice layers in a stormwater management pond in [[Ontario|Ontario, Canada]].<ref>{{cite journal | vauthors = Long AM, Short SM | title = Seasonal determinations of algal virus decay rates reveal overwintering in a temperate freshwater pond | language = En | journal = The ISME Journal | volume = 10 | issue = 7 | pages = 1602–12 | date = July 2016 | pmid = 26943625 | doi = 10.1038/ismej.2015.240 | url = http://www.nature.com/articles/ismej2015240 }}</ref> Further, DeLong et al. (2016) suggest that predation by small crustaceans can play an indirect role in titer fluctuations, as degradation of protist cells passing through the digestive tract results in liberation of large numbers of unicellular algae that become susceptible to viral infection due to disruption of endosymbiosis.<ref name=":1" /> Overall, seasonal abundance of chloroviruses depends not only on the host species, but also on the abundance of other microorganisms, general nutrient status and ecological conditions.<ref name = "Yanai_2009" />

Collectively, chloroviruses are able to mediate global [[biogeochemical cycle]]s through [[phytoplankton]] turnover. ''Chlorella'', in co-occurrence with other types of microscopic algae like ''[[Microcystis aeruginosa]]'', are known to cause toxic [[algal bloom]]s that typically last from February to June in the Northern hemisphere, resulting in oxygen depletion and deaths of larger organisms in freshwater habitats.<ref>{{cite journal | vauthors = Song H, Lavoie M, Fan X, Tan H, Liu G, Xu P, Fu Z, Paerl HW, Qian H | title = Allelopathic interactions of linoleic acid and nitric oxide increase the competitive ability of Microcystis aeruginosa | language = En | journal = The ISME Journal | volume = 11 | issue = 8 | pages = 1865–1876 | date = August 2017 | pmid = 28398349 | doi = 10.1038/ismej.2017.45 | url = http://www.nature.com/doifinder/10.1038/ismej.2017.45 }}</ref><ref>{{cite journal | vauthors = Rieper M |date=1976-03-01|title=Investigations on the relationships between algal blooms and bacterial populations in the Schlei Fjord (western Baltic Sea)|url=https://link.springer.com/article/10.1007/BF01610792|journal=Helgoländer wissenschaftliche Meeresuntersuchungen|language=en|volume=28|issue=1|pages=1–18|doi=10.1007/bf01610792|issn=0017-9957}}</ref> Lytic infection of unicellular algae by chloroviruses results in termination of algal blooms and the subsequent release of carbon, nitrogen and phosphorus trapped in the cells, transporting them to lower [[trophic level]]s and, ultimately, fueling the food chain.<ref name = "Yanai_2009">{{cite thesis | first = Giane M. | last = Yanai | name-list-format = vanc | publisher = University of Nebraska at Lincoln | date = 2009 | title = Transcription analysis of the chlorovirus Paramecium bursaria chlorella virus-1 | url = https://digitalcommons.unl.edu/bioscidiss/5/ | type = PhD }}</ref>

== Evolution ==
Chloroviruses, as well as the remaining members of the Phycodnaviridae family, are considered part of the broader group of microbes called Nucleocytoplasmic Large DNA Viruses (NCLDVs). Although Phycodnaviruses are diverse genetically and infect different hosts, they display high levels of similarity on the structural level to each other and other NCLDVs. [[Phylogenetics|Phylogenetic]] analysis of the major capsid protein within the group indicates great likelihood of close relatedness, as well as prior divergence from a single common ancestor, which is believed to be a small DNA virus.<ref>{{cite journal | vauthors = Yutin N, Wolf YI, Koonin EV | title = Origin of giant viruses from smaller DNA viruses not from a fourth domain of cellular life | journal = Virology | volume = 466-467 | pages = 38–52 | date = October 2014 | pmid = 25042053 | doi = 10.1016/j.virol.2014.06.032 | url = http://linkinghub.elsevier.com/retrieve/pii/S0042682214003006 }}</ref><ref name=":2">{{cite journal | vauthors = Dunigan DD, Fitzgerald LA, Van Etten JL | title = Phycodnaviruses: a peek at genetic diversity | journal = Virus Research | volume = 117 | issue = 1 | pages = 119–32 | date = April 2006 | pmid = 16516998 | doi = 10.1016/j.virusres.2006.01.024 }}</ref> Additionally, studies suggest that genome gigantism, characteristic of all chloroviruses, is a property which evolved early on in the history of NCLDVs, and subsequent adaptations towards respective hosts and particular habitats resulted in mutations and gene loss events, which ultimately shaped all currently existing chlorovirus species.<ref name=":2" />

[[Whole genome sequencing|Genome sequencing]] and functional screening of proteins from PBCV-1 and [[Acanthocystis turfacea chlorella virus 1|ATCV-1]] revealed large number of [[Horizontal gene transfer|horizontally transferred genes]], which indicates a long history of co-evolution with the unicellular host and lateral gene transfer with other seemingly unrelated organisms.<ref name=":2" /> Further, both viruses were found to encode several so-called "progenitor enzymes", which are smaller, but less specialized than their modern-day analogues. For example, one of the sugar-manipulating enzymes in PBCV-1 ([[GDP-mannose 4,6-dehydratase|GDP-d-mannose 4,6 dehydratase or GMD]]) was shown to mediate catalysis of not only the dehydration of [[GDP-D-mannose|GDP-d-mannose]], but also reduction of the sugar molecule produced in the initially predicted process. Such dual functionality is uncommon among the currently existing sugar-manipulating enzymes, and possibly suggests the ancient nature of the PBCV-1 GMD.<ref name=":3">{{cite journal | vauthors = Van Etten JL, Agarkova I, Dunigan DD, Tonetti M, De Castro C, Duncan GA | title = Chloroviruses Have a Sweet Tooth | journal = Viruses | volume = 9 | issue = 4 | date = April 2017 | pmid = 28441734 | pmc = 5408694 | doi = 10.3390/v9040088 }}</ref>

In addition, infection cycle studies in PBCV-1 revealed that the virus relies on a unique capsid glycosylation process independent of the host's [[Endoplasmic reticulum|ER]] or [[Golgi apparatus|Golgi]] machinery. This feature has not yet been observed in any other virus currently known to science and potentially represents an ancient and conserved pathway, which could have evolved before [[Eukaryote|eukaryogenesis]], which was estimated to occur around 2.0-2.7 billion years ago.<ref name=":3" />

Lastly, recent discovery regarding presence of DNA sequences homologous to ATCV-1 in the human oropharyngeal virome, as well as the subsequent studies demonstrating successful infection of mammalian animal model by ATCV-1, also point to the likelihood of ancient evolutionary history of chloroviruses, which possess structural features and utilize molecular mechanisms that potentially allow for replication within diverse animal hosts.<ref name=":4" /><ref>{{cite journal | vauthors = Petro TM, Agarkova IV, Zhou Y, Yolken RH, Van Etten JL, Dunigan DD | title = Response of Mammalian Macrophages to Challenge with the Chlorovirus Acanthocystis turfacea Chlorella Virus 1 | journal = Journal of Virology | volume = 89 | issue = 23 | pages = 12096–107 | date = December 2015 | pmid = 26401040 | pmc = 4645302 | doi = 10.1128/JVI.01254-15 }}</ref><ref>{{cite journal | vauthors = Petro MS, Agarkova IV, Petro TM | title = Effect of Chlorovirus ATCV-1 infection on behavior of C57Bl/6 mice | journal = Journal of Neuroimmunology | volume = 297 | pages = 46–55 | date = August 2016 | pmid = 27397075 | doi = 10.1016/j.jneuroim.2016.05.009 }}</ref>

== See also ==
* ''[[Phycodnaviridae]]'' - algae infecting viruses

== References ==
{{Reflist}}

== External links ==
* [http://viralzone.expasy.org/all_by_species/588.html '''Viralzone''': Chlorovirus]
* [http://ictvonline.org/virusTaxonomy.asp '''ICTV''']
{{Baltimore classification}}

{{Taxonbar|from=Q18822090}}

[[Category:Phycodnaviridae]]
[[Category:Nucleocytoplasmic large DNA viruses]]