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Quaranjavirus is a of , one of seven genera in the family . The is , negative-sense segmented , generally with six segments. The genus contains six species. Quaranjaviruses predominantly infect and ; Quaranfil quaranjavirus is the only member of the genus to have been shown to infect humans. The Quaranfil and Johnston Atoll viruses are transmitted between by , resembling members of , another genus of Orthomyxoviridae.


History
was first isolated from humans in in 1953. Johnston Atoll virus and Lake Chad virus were first isolated from birds in 1964 and 1969, respectively. In 1989, based on the appearance of the virus particles under the electron microscope, H.G. Zeller and colleagues suggested that they should be classified as , but this was not accepted by the International Committee on Taxonomy of Viruses (ICTV). In 2009, based on sequence data and the structure of the virus particles, Rachel Presti and colleagues suggested that the three viruses should be classified as a new genus of , originally named "Quarjavirus". Multiple other viruses have subsequently been suggested as genus members. The genus was formally proposed to the ICTV in 2012, under the name '"Quaranjavirus", and formally approved by that body in 2013.


Nomenclature
Quaranfil virus is named for , one of the villages near from which the virus was isolated. The Johnston Atoll virus is named for in the , also where the virus was first isolated. The genus name combines "Quaran", with "ja" for Johnston Atoll.


Virology
The virus particle is and spherical, ovoid or variable in shape, with a diameter generally in the range 80–120 . The virus particle contains around ten -like granules, a feature of arenaviruses. It bears surface projections variously described as 4–5 nm long and 10–14 nm in length and 4–6 nm in diameter.

The , – is linear and segmented, generally with six segments of 0.9–2.4 kb and a total size of around 11.5 kb. Wellfleet Bay virus has a seventh segment of 519 . The genome encodes six or (in Wellfleet Bay virus) seven proteins. The PA, PB1 and PB2 subunits of the trimeric enzyme are encoded by the three largest segments (1–3), as in other orthomyxoviruses. Segment 5 encodes the envelope (GP). Proteins of unknown function are encoded by segments 4 and 6, which have recently been tentatively assigned to the viral and , respectively. Segment 7 of Wellfleet Bay virus encodes an additional protein of unknown function. The quaranjavirus glycoprotein shows no similarity with the glycoproteins (haemagglutinin and neuraminidase), and instead shows some similarities with the gp64 glycoprotein of , which infect insects, as well as the glycoprotein of , a genus of tick-transmitted orthomyxoviruses.

The replication cycle of Quaranfil virus takes around 24–36 hours, which is comparable with thogotoviruses and slower than influenza viruses.


Host range and disease
Quaranjaviruses infect both and hosts. The most common arthropod hosts are species of soft-bodied ( family) . Most members cannot infect cell lines in the laboratory. In 2015, multiple new members of the genus were proposed based on RNA sequences obtained from mosquitos, , other , and the spider.

Aquatic birds that nest in colonies are the most common vertebrate hosts, including , and . Cygnet River and Wellfleet Bay viruses have been associated with an often-fatal disease in farmed and wild species, with symptoms including and . Most genus members tested can infect mice under laboratory conditions; they cause severe pathology and are frequently lethal. Quaranfil virus is the only member of the genus to have been shown to infect humans; infection generally appears to be asymptomatic and has occasionally been reported to be associated with mild fever.


Species and strains
The genus contains the following species, listed by scientific name and followed by the exemplar virus of the species:

  • Quaranjavirus araguariense,
  • Quaranjavirus chadense, Lake Chad virus
  • Quaranjavirus johnstonense, Johnston Atoll virus
  • Quaranjavirus quaranfilense,
  • Quaranjavirus tyulekense,
  • Quaranjavirus wellfleetense, Wellfleet Bay virus

Many other species or strains have been identified from a range of based on RNA data.

Cygnet River Australia
Johnston Atoll 165–200 ticks, Australasia, Pacific
Lake Chad , vitelline masked weaver birdAfrica
Quaranfil≥6100 or 137–156 ticks, human, Africa, Asia
Tyulek6 ticks Asia
Wellfleet Bay≥790–110 North America


Quaranfil virus and related strains
Quaranfil virus was initially isolated in 1953 from two children with mild fever in the villages of Quaranfil and , near Cairo in Egypt. to the virus have been demonstrated in around 8% of people sampled in the area, although no further cases of symptomatic disease have been reported. The geographical range of the virus is wide, also including and in Africa, and , , , and in the Middle East and Asia. The virus has also been isolated from seabirds and from ticks, such as the . The virus has been shown to be transmitted between vertebrates by ticks. In the laboratory, it causes severe disease in mice.


Cygnet River and Wellfleet Bay virus
Cygnet River virus was isolated in 2010 from embryonated eggs of the ( Cairina moschate) from , in Australia. The virus was associated with an outbreak of severe disease in farmed ducks.

Wellfleet Bay virus was isolated from wild ( Somateria mollissima) overwintering at Jeremy Point, Wellfleet Bay in , , USA. The virus was associated with an outbreak of severe disease in 2010, and is hypothesised to have been a factor in a series of similar outbreaks in 1998–2013.

The two viruses are closely related and might be strains of the same virus; they are also related to Quaranfil virus.


Lake Chad virus
Lake Chad virus was isolated in 1969 from a vitelline masked weaver bird ( Ploceus vitellinus) from in , and has yet to be isolated elsewhere. The virus causes severe disease in mice under laboratory conditions. It is most closely related to Quaranfil virus.


Johnston Atoll virus
Johnston Atoll virus was first isolated in 1964 from soft-bodied ticks infesting the ( Anous stolidus) from Sand Island, , in the Pacific. Its range also includes , and . The virus has been shown to be transmitted between vertebrates by ticks. In the laboratory, it causes severe disease in mice and chicks.


Tyulek virus
Tyulek virus was isolated from soft-bodied ticks infesting birds by the Aksu River near the village of in . The virus is closely related to both Quaranfil and 'ohnston Atoll viruses.


Other putative strains
Thirteen RNA sequences with 34–40% sequence identity to Johnston Atoll virus have been identified in and in . Putative hosts include (spider), Atherigona orientalis (pepper fruit fly), Chrysomya megacephala (oriental latrine fly), (flesh flies), (housefly), Culex tritaeniorhynchus, C. quinquefasciatus, Anopheles sinensis and Armigeres subalbatus (mosquitoes), (moth fly), (louse fly), as well as unidentified species in the (horse-fly), (water strider) and (soldier fly) groups. Nearly 50 viruses associated with were identified in a broad variety of insect hosts, after computational virus screening of the largest insect transcriptome collection sampled worldwide. -related viruses were found for the first time in insect hosts, such as , , , , , , and . These findings place Quaranjavirus in a paraphyletic position to the other viruses of with implications for taxonomic evaluations and virus evolution.

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