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=== As disease reservoirs === [[File:Egyptian fruit bat.jpg|thumb|alt=A yellowish-gray megabat sits atop a skewer of fruit slices, including banana and apple.|The [[Egyptian fruit bat]] (''Rousettus aegyptiacus''), which has tested positive for [[Marburg virus]] and antibodies against the [[Ebola virus]], though not the actual virus]] [[File:Hendra-distribution-map.jpg|upright=1.3|thumb|alt=A map of Madagascar, Asia, and Oceania. Madagascar, Southern and Southeast Asia, and most of Oceania is delimited as flying fox distribution. The northeast coast of Australia shows small red icons that indicate Hendra virus outbreaks. South and Southeast Asia has several blue icons that indicate Nipah virus outbreaks.|[[Henipavirus]] outbreaks overlaid on flying fox distribution map, with [[Nipah virus]] as blue icons and [[Hendra virus]] as red icons]] Megabats are the [[natural reservoir|reservoirs]] of several [[virus]]es that can affect humans and cause disease. They can carry [[filovirus]]es, including the [[Ebola virus]] (EBOV) and ''[[Marburgvirus]]''.<ref name="Hassanin" /> The presence of ''Marburgvirus'', which causes [[Marburg virus disease]], has been confirmed in one species, the Egyptian fruit bat. The disease is rare, but the fatality rate of an outbreak can reach up to 88%.<ref name="Hassanin"/><ref name="who" /> The virus was first recognized after simultaneous outbreaks in the German cities of [[Marburg]] and [[Frankfurt]] as well as [[Belgrade|Belgrade, Serbia]], in 1967,<ref name="who">{{cite web|url=https://www.afro.who.int/health-topics/marburg-haemorrhagic-fever| title=Marburg Haemorrhagic Fever| website=World Health Organization| access-date=30 May 2019}}</ref> where 31 people became ill and seven died.<ref name="cdc" /> The outbreak was traced to [[laboratory]] work with [[vervet monkey]]s from [[Uganda]].<ref name="who"/> The virus can pass from a bat host to a human (who has usually spent a prolonged period in a mine or cave where Egyptian fruit bats live); from there, it can spread person-to-person through contact with infected bodily fluids, including blood and [[semen]].<ref name="who" /> The United States [[Centers for Disease Control and Prevention]] lists a total of 601 confirmed cases of Marburg virus disease from 1967 to 2014, of which 373 people died (62% overall mortality).<ref name="cdc">{{cite web|url=https://www.cdc.gov/vhf/marburg/resources/outbreak-table.html| title=Chronology of Marburg Hemorrhagic Fever Outbreaks| website=Centers for Disease Control and Prevention|date=9 October 2014|access-date=30 May 2019}}</ref> Species that have tested positive for the presence of EBOV include [[Franquet's epauletted fruit bat]] (''Epomops franqueti''), the hammer-headed fruit bat, and the little collared fruit bat. Additionally, [[antibody|antibodies]] against EBOV have been found in the straw-colored fruit bat, [[Gambian epauletted fruit bat]] (''Epomophorus gambianus''), [[Peters's dwarf epauletted fruit bat]] (''Micropteropus pusillus''), [[Veldkamp's dwarf epauletted fruit bat]] (''Nanonycteris veldkampii''), Leschenault's rousette, and the Egyptian fruit bat.<ref name="Hassanin">{{cite journal|doi=10.1016/j.crvi.2016.09.005|pmid=27746072|title=Comparative phylogeography of African fruit bats (Chiroptera, Pteropodidae) provide new insights into the outbreak of Ebola virus disease in West Africa, 2014–2016|journal=[[Comptes Rendus Biologies]]|volume=339|issue=11–12|pages=517–528|year=2016|last1=Hassanin|first1=A. |last2=Nesi|first2=N. |last3=Marin|first3=J. |last4=Kadjo|first4=B. |last5=Pourrut|first5=X. |last6=Leroy|first6=É. |last7=Gembu|first7=G. |last8=Musaba Akawa|first8=P. |last9=Ngoagouni|first9=C. |last10=Nakouné|first10=E. |last11=Ruedi|first11=M. |last12=Tshikung|first12=D. |last13=Pongombo Shongo|first13=C.|last14=Bonillo|first14=C.|s2cid=44412099 |url=https://hal.sorbonne-universite.fr/hal-01382796/document|doi-access=free}}</ref> Much of how humans contract the Ebola virus is unknown. Scientists hypothesize that humans initially become infected through contact with an infected animal such as a megabat or non-human primate.<ref>{{cite web|url=https://www.cdc.gov/vhf/ebola/transmission/index.html| title=Ebola (Ebola Virus Disease): Transmission| access-date=30 May 2019|date=17 May 2019| website=Centers for Disease Control and Prevention}}</ref> Megabats are presumed to be a natural reservoir of the Ebola virus, but this has not been firmly established.<ref>{{cite web|url=https://www.cdc.gov/ncezid/stories-features/global-stories/ebola-reservoir-study.html| title=Ebola Reservoir Study| website=Centers for Disease Control and Prevention|access-date=30 May 2019|date=9 July 2018}}</ref> Microbats are also being investigated as the reservoir of the virus, with the [[greater long-fingered bat]] (''Miniopterus inflatus'') once found to harbor a fifth of the virus's genome (though not testing positive for the actual virus) in 2019.<ref>{{cite web|url=https://www.science.org/content/article/bat-species-may-be-source-ebola-epidemic-killed-more-11000-people-west-africa| title=This bat species may be the source of the Ebola epidemic that killed more than 11,000 people in West Africa| first=K.| last= Kupferschmidt|date=24 January 2019| access-date=30 May 2019| website=Science}}</ref> Due to the likely association between Ebola infection and "hunting, butchering and processing meat from infected animals", several West African countries banned bushmeat (including megabats) or issued warnings about it during the [[Western African Ebola virus epidemic|2013–2016 epidemic]]; many bans have since been lifted.<ref>{{cite web| url=https://www.apnews.com/2ff0034f651a4e229c6d9a74b21bc80f| title=Post-Ebola, West Africans flock back to bushmeat, with risk| website=Associated Press| first1=H.| last1=Zon| first2=C.| last2=Petesch| date=21 September 2016| access-date=30 May 2019| archive-date=25 December 2019| archive-url=https://web.archive.org/web/20191225100557/https://apnews.com/2ff0034f651a4e229c6d9a74b21bc80f| url-status=dead}}</ref> Other megabats implicated as disease reservoirs are primarily ''Pteropus'' species. Notably, flying foxes can transmit [[Australian bat lyssavirus]], which, along with the [[rabies virus]], causes [[rabies]]. Australian bat lyssavirus was first identified in 1996; it is very rarely transmitted to humans. Transmission occurs from the bite or scratch of an infected animal but can also occur from getting the infected animal's saliva in a [[mucous membrane]] or an open [[wound]]. Exposure to flying fox blood, urine, or feces cannot cause infections of Australian bat lyssavirus. Since 1994, there have been three records of people becoming infected with it in [[Queensland]]—each case was fatal.<ref name="NSW">{{cite web| url=http://www.health.nsw.gov.au/Infectious/factsheets/Pages/rabies-australian-bat-lyssavirus-infection.aspx| title=Rabies and Australian bat lyssavirus infection fact sheet| date=30 November 2015| website=health.nsw.gov.au| publisher=State of New South Wales NSW Ministry of Health 2015| access-date=14 June 2018}}</ref> Flying foxes are also reservoirs of [[henipavirus]]es such as [[Henipavirus#Hendra virus|Hendra virus]] and [[Henipavirus#Nipah virus|Nipah virus]]. Hendra virus was first identified in 1994; it rarely occurs in humans. From 1994 to 2013, there have been seven reported cases of Hendra virus affecting people, four of which were fatal. The hypothesized primary route of human infection is via contact with [[horse]]s that have come into contact with flying fox [[Urination|urine]].<ref name="CDC">{{cite web| url=https://www.cdc.gov/vhf/hendra/pdf/factsheet.pdf| title=Hendra Virus Disease(HeV)| website=cdc.gov| publisher=U.S. Department of Health & Human Services| access-date=14 June 2018}}</ref> There are no documented instances of direct transmission between flying foxes and humans.<ref name="Sanchez 2016">{{cite journal|doi=10.1371/journal.pntd.0004411|pmid=26829399|title=Disease Risk Perception and Safety Practices: A Survey of Australian Flying Fox Rehabilitators|journal=PLOS Neglected Tropical Diseases|volume=10|issue=2|page=e0004411|year=2016|last1=Sánchez|first1=C. A. |last2=Baker|first2=M. L.|pmc=4734781 |doi-access=free }}</ref> As of 2012, there is a [[vaccine]] available for horses to decrease the likelihood of infection and transmission.<ref name="AVA">{{cite web| url=http://www.ava.com.au/hendra-virus-vaccine| title=The Hendra vaccine| website=ava.com| publisher=The Australian Veterinary Association Ltd (AVA)| date=2018| access-date=14 June 2018| archive-date=14 June 2018| archive-url=https://web.archive.org/web/20180614194613/http://www.ava.com.au/hendra-virus-vaccine| url-status=dead}}</ref> Nipah virus was first identified in 1998 in Malaysia. Since 1998, there have been several Nipah outbreaks in Malaysia, [[Singapore]], India, and Bangladesh, resulting in over 100 casualties. A [[2018 Nipah virus outbreak in Kerala|2018 outbreak]] in [[Kerala|Kerala, India]], resulted in 19 humans becoming infected—17 died.<ref name="Gulland 2018">{{cite web| url=https://www.telegraph.co.uk/news/2018/06/12/nipah-virus-control-india-britain-world-must-alert-signs-infected/ |archive-url=https://ghostarchive.org/archive/20220112/https://www.telegraph.co.uk/news/2018/06/12/nipah-virus-control-india-britain-world-must-alert-signs-infected/ |archive-date=12 January 2022 |url-access=subscription |url-status=live| title=Nipah virus 'under control' in India – but Britain and the world must be alert for signs of infected travellers| last=Gulland| first=A.| date=12 June 2018| website=The Telegraph| publisher=Telegraph Media Group Limited 2018| access-date=14 June 2018}}{{cbignore}}</ref> The overall fatality rate is 40–75%. Humans can contract Nipah virus from direct contact with flying foxes or their fluids, through exposure to an intermediate [[host (biology)|host]] such as [[domestic pig]]s, or from contact with an infected person.<ref name="WHO">{{cite web| url=https://www.who.int/news-room/fact-sheets/detail/nipah-virus| title=Nipah virus| website=World Health Organization| publisher=WHO| date=30 May 2018| access-date=14 June 2018}}</ref> A 2014 study of the Indian flying fox and Nipah virus found that while Nipah virus outbreaks are more likely in areas preferred by flying foxes, "the presence of bats in and of itself is not considered a risk factor for Nipah virus infection." Rather, the consumption of [[date palm#Sap|date palm sap]] is a significant route of transmission. The practice of date palm sap collection involves placing collecting pots at date palm trees. Indian flying foxes have been observed licking the sap as it flows into the pots, as well as defecating and urinating in proximity to the pots. In this way, humans who drink palm wine can be exposed to henipaviruses. The use of bamboo skirts on collecting pots lowers the risk of contamination from bat urine.<ref name="Hahn 2014">{{cite journal| last1=Hahn| first1= M. B.| last2=Epstein |first2= J. H.| last3= Gurley| first3= E. S.| last4= Islam| first4= M. S.| last5= Luby| first5= S. P.| last6= Daszak| first6= P.| last7=Patz| first7= J. A.| date=2014| title= Roosting behaviour and habitat selection of Pteropus giganteus reveal potential links to Nipah virus epidemiology| journal=[[Journal of Applied Ecology]]| volume= 51| issue=2| pages=376–387| doi=10.1111/1365-2664.12212| pmid= 24778457| pmc=4000083| bibcode= 2014JApEc..51..376H}}</ref> Flying foxes can transmit several non-lethal diseases as well, such as [[Menangle virus]]<ref name="Bowden 2001">{{cite journal|doi=10.1006/viro.2001.0893|pmid=11336561|title=Molecular Characterization of Menangle Virus, a Novel Paramyxovirus which Infects Pigs, Fruit Bats, and Humans|journal=[[Virology (journal)|Virology]]|volume=283|issue=2|pages=358–73|year=2001|last1=Bowden|first1=T. R. |last2=Westenberg|first2=M.|last3=Wang|first3=L.|last4=Eaton|first4=B. T.|last5=Boyle|first5=D. B.|doi-access=free}}</ref> and [[Nelson Bay virus]].<ref name="Yamanaka 2014">{{cite journal|doi=10.1371/journal.pone.0092777|pmid=24667794|title=Imported Case of Acute Respiratory Tract Infection Associated with a Member of Species Nelson Bay Orthoreovirus|journal=PLOS ONE|volume=9|issue=3 |page=e92777|year=2014 |last1=Yamanaka|first1=A. |last2=Iwakiri|first2=A. |last3=Yoshikawa|first3=T. |last4=Sakai|first4=K. |last5=Singh|first5=H. |last6=Himeji|first6=D. |last7=Kikuchi|first7=I. |last8=Ueda|first8=A. |last9=Yamamoto|first9=S. |last10=Miura|first10=M. |last11=Shioyama|first11=Y. |last12=Kawano|first12=K. |last13=Nagaishi|first13=T. |last14=Saito|first14=M. |last15=Minomo|first15=M. |last16=Iwamoto|first16=N. |last17=Hidaka|first17=Y. |last18=Sohma|first18=H. |last19=Kobayashi|first19=T. |last20=Kanai|first20=Y. |last21=Kawagishi|first21=T. |last22=Nagata|first22=N. |last23=Fukushi|first23=S. |last24=Mizutani|first24=T. |last25=Tani|first25=H. |last26=Taniguchi|first26=S. |last27=Fukuma|first27=A. |last28=Shimojima|first28=M. |last29=Kurane|first29=I. |last30=Kageyama|first30=T.|display-authors=29|bibcode=2014PLoSO...992777Y|pmc=3965453|doi-access=free}}</ref> These viruses rarely affect humans, and few cases have been reported.<ref name="Bowden 2001" /><ref name="Yamanaka 2014" /> Megabats are not suspected to be vectors of [[coronaviruses]].<ref name="Smith 2013">{{Cite journal|doi=10.1016/j.coviro.2012.11.006|pmid=23265969 | title=Bats and their virome: An important source of emerging viruses capable of infecting humans|journal=[[Current Opinion in Virology]] | volume=3 | issue=1 | pages=84–91 | year=2013 | last1=Smith | first1=I. | last2=Wang | first2=L.|pmc=7102720 }}</ref>
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