title: Filoviruses in bats: current knowledge and future directions.
abstract: Filoviruses, including Ebolavirus and Marburgvirus, pose significant threats to public health and species conservation by causing hemorrhagic fever outbreaks with high mortality rates. Since the first outbreak in 1967, their origins, natural history, and ecology remained elusive until recent studies linked them through molecular, serological, and virological studies to bats. We review the ecology, epidemiology, and natural history of these systems, drawing on examples from other bat-borne zoonoses, and highlight key areas for future research. We compare and contrast results from ecological and virological studies of bats and filoviruses with those of other systems. We also highlight how advanced methods, such as more recent serological assays, can be interlinked with flexible statistical methods and experimental studies to inform the field studies necessary to understand filovirus persistence in wildlife populations and cross-species transmission leading to outbreaks. We highlight the need for a more unified, global surveillance strategy for filoviruses in wildlife, and advocate for more integrated, multi-disciplinary approaches to understand dynamics in bat populations to ultimately mitigate or prevent potentially devastating disease outbreaks.
authors.affiliation: EcoHealth Alliance, 460 W. 34th Street, New York, NY 10001, USA. olival@ecohealthalliance.org.
authors.forename: Kevin J
authors.initials: KJ
authors.lastname: Olival
authors.affiliation: Department of Biology, Colorado State University, Fort Collins, CO 80523, USA. davidtshayman@gmail.com.
authors.forename: David T S
authors.initials: DT
authors.lastname: Hayman
mesh_terms: D000818:Animals; D002685:Chiroptera / Q000821:virology*; D016563:Filoviridae / Q000145:classification / Q000302:isolation & purification*; D018702:Filoviridae Infections / Q000453:epidemiology / Q000635:transmission / Q000662:veterinary* / Q000821:virology; D006801:Humans; D015047:Zoonoses / Q000453:epidemiology* / Q000635:transmission / Q000821:virology*
references.citation: J Infect Dis. 2011 Nov;204 Suppl 3:S776-84
references.pmid: 21987750
references.citation: BMC Infect Dis. 2009 Sep 28;9:159
references.pmid: 19785757
references.citation: PLoS Genet. 2010 Nov 18;6(11):e1001191
references.pmid: 21124940
references.citation: Wkly Epidemiol Rec. 2007 Aug 17;82(33):297-8
references.pmid: 17703537
references.citation: PLoS One. 2012;7(9):e45479
references.pmid: 23029039
references.citation: Vector Borne Zoonotic Dis. 2010 Mar;10(2):183-90
references.pmid: 19402762
references.citation: J Virol Methods. 2013 Nov;193(2):295-303
references.pmid: 23835034
references.citation: PLoS Biol. 2005 Nov;3(11):e371
references.pmid: 16231972
references.citation: J Virol Methods. 2007 Jun;142(1-2):29-40
references.pmid: 17292974
references.citation: Proc Biol Sci. 2013 Feb 01;280(1756):20122753
references.pmid: 23378666
references.citation: Nature. 2008 Feb 21;451(7181):990-3
references.pmid: 18288193
references.citation: Ecohealth. 2010 Dec;7(4):517-25
references.pmid: 21207105
references.citation: PLoS One. 2007 Aug 22;2(8):e764
references.pmid: 17712412
references.citation: Viruses. 2012 Aug;4(8):1318-27
references.pmid: 23012628
references.citation: Lancet. 1982 Apr 10;1(8276):816-20
references.pmid: 6122054
references.citation: PLoS One. 2010 Aug 04;5(8):e11978
references.pmid: 20694141
references.citation: Ecohealth. 2009 Dec;6(4):522-39
references.pmid: 20049506
references.citation: Emerg Infect Dis. 2012 Jul;18(7):1207-9
references.pmid: 22710257
references.citation: Microbes Infect. 2005 Jun;7(7-8):1005-14
references.pmid: 16002313
references.citation: Emerg Infect Dis. 2009 Aug;15(8):1171-5
references.pmid: 19751577
references.citation: J Theor Biol. 2004 Jul 7;229(1):119-26
references.pmid: 15178190
references.citation: PLoS Pathog. 2011 Oct;7(10):e1002304
references.pmid: 22039362
references.citation: Mol Biol Evol. 1997 Aug;14(8):800-6
references.pmid: 9254917
references.citation: Adv Virus Res. 2011;79:239-89
references.pmid: 21601050
references.citation: Science. 1968 Sep 27;161(3848):1364
references.pmid: 4970585
references.citation: Virology. 2010 Dec 20;408(2):183-9
references.pmid: 20951400
references.citation: PLoS Pathog. 2008 Nov;4(11):e1000212
references.pmid: 19023410
references.citation: Epidemiol Infect. 2007 May;135(4):610-21
references.pmid: 16999875
references.citation: Emerg Infect Dis. 2004 Jan;10(1):40-7
references.pmid: 15078595
references.citation: Arch Virol Suppl. 1996;11:115-34
references.pmid: 8800793
references.citation: Science. 2004 Jan 16;303(5656):387-90
references.pmid: 14726594
references.citation: Emerg Infect Dis. 2013 Jan;19(1):51-60
references.pmid: 23261015
references.citation: Arch Virol Suppl. 1996;11:101-14
references.pmid: 8800792
references.citation: Nat Commun. 2013;4:2770
references.pmid: 24253424
references.citation: Emerg Infect Dis. 2007 Dec;13(12):1847-51
references.pmid: 18258034
references.citation: Nature. 2005 Dec 1;438(7068):575-6
references.pmid: 16319873
references.citation: Microbes Infect. 2002 Feb;4(2):145-51
references.pmid: 11880045
references.citation: J Gen Virol. 2001 Dec;82(Pt 12):2839-2848
references.pmid: 11714958
references.citation: Zoonoses Public Health. 2013 Feb;60(1):2-21
references.pmid: 22958281
references.citation: Parasit Vectors. 2013 Aug 08;6:231
references.pmid: 23924629
references.citation: J Reprod Fertil. 1992 Nov;96(2):765-73
references.pmid: 1339856
references.citation: PLoS Pathog. 2012;8(10):e1002877
references.pmid: 23055920
references.citation: Lancet. 1995 May 20;345(8960):1271-4
references.pmid: 7746057
references.citation: Vector Borne Zoonotic Dis. 2012 Aug;12(8):666-73
references.pmid: 22607069
references.citation: Curr Top Microbiol Immunol. 2007;315:363-87
references.pmid: 17848072
references.citation: Virology. 2013 Nov;446(1-2):230-7
references.pmid: 24074586
references.citation: Med Trop (Mars). 1999;59(2):201-4
references.pmid: 10546197
references.citation: Emerg Infect Dis. 2002 Dec;8(12):1468-73
references.pmid: 12498665
references.citation: Vector Borne Zoonotic Dis. 2009 Dec;9(6):723-8
references.pmid: 19323614
references.citation: Am J Trop Med Hyg. 1992 Jun;46(6):664-71
references.pmid: 1621890
references.citation: Proc Biol Sci. 2014 Jul 7;281(1786):
references.pmid: 24827436
references.citation: J Infect Dis. 2011 Nov;204 Suppl 3:S804-9
references.pmid: 21987755
references.citation: J Infect Dis. 1999 Feb;179 Suppl 1:S177-87
references.pmid: 9988182
references.citation: PLoS One. 2008 Jul 23;3(7):e2739
references.pmid: 18648649
references.citation: J Virol. 2006 Jul;80(13):6497-516
references.pmid: 16775337
references.citation: J Infect Dis. 2011 Nov;204 Suppl 3:S757-60
references.pmid: 21987747
references.citation: Science. 2013 Jan 25;339(6118):456-60
references.pmid: 23258410
references.citation: Science. 2009 Jul 10;325(5937):204-6
references.pmid: 19590002
references.citation: Proc Biol Sci. 2011 Dec 22;278(1725):3703-12
references.pmid: 21561971
references.citation: Sci Rep. 2012;2:811
references.pmid: 23155478
references.citation: Zoonoses Public Health. 2013 Feb;60(1):104-16
references.pmid: 23302292
references.citation: Med Trop (Mars). 2000;60(2 Suppl):23-6
references.pmid: 11100456
references.citation: J Infect Dis. 1999 Feb;179 Suppl 1:S115-9
references.pmid: 9988174
references.citation: Ecohealth. 2013 Sep;10(3):298-313
references.pmid: 23918033
references.citation: Emerg Infect Dis. 2011 Mar;17(3):449-56
references.pmid: 21392436
references.citation: Lancet. 1967 Nov 25;2(7526):1119-21
references.pmid: 4168558
references.citation: Epidemiol Infect. 2011 Oct;139(10):1570-9
references.pmid: 21524339
references.citation: PLoS One. 2010 Nov 10;5(11):e13903
references.pmid: 21085717
references.citation: PLoS Pathog. 2006 Oct;2(10):e90
references.pmid: 17069458
references.citation: Virol J. 2012 Oct 13;9:236
references.pmid: 23062147
references.citation: J Comp Pathol. 2002 Feb-Apr;126(2-3):124-36
references.pmid: 11945001
references.citation: J Infect Dis. 1999 Feb;179 Suppl 1:S108-14
references.pmid: 9988173
references.citation: PLoS Pathog. 2009 Jul;5(7):e1000536
references.pmid: 19649327
references.citation: J Am Vet Med Assoc. 2003 Mar 1;222(5):633-9
references.pmid: 12619845
references.citation: PLoS Pathog. 2010 Jul 29;6(7):e1001030
references.pmid: 20686665
references.citation: PLoS One. 2009 Dec 11;4(12):e8266
references.pmid: 20011515
references.citation: Arch Virol. 2014 May;159(5):1229-37
references.pmid: 24190508
references.citation: N Engl J Med. 2006 Aug 31;355(9):909-19
references.pmid: 16943403
references.citation: Exp Anim. 2002 Apr;51(2):173-9
references.pmid: 12012728
references.citation: Am Nat. 2007 May;169(5):684-9
references.pmid: 17427138
references.citation: Lancet. 2011 Mar 5;377(9768):849-62
references.pmid: 21084112
references.citation: J Infect Dis. 2011 Jul 15;204(2):200-8
references.pmid: 21571728
references.citation: PLoS One. 2013 Aug 30;8(8):e72942
references.pmid: 24023659
references.citation: J Anim Ecol. 2014 Mar;83(2):415-28
references.pmid: 24111634
references.citation: Curr Top Microbiol Immunol. 1999;235:1-21
references.pmid: 9893375
references.citation: Science. 2006 Dec 8;314(5805):1564
references.pmid: 17158318
references.citation: Emerg Infect Dis. 2010 Feb;16(2):352-4
references.pmid: 20113584
references.citation: BMC Evol Biol. 2010 Jun 22;10:193
references.pmid: 20569424
references.citation: Microbes Infect. 1999 Dec;1(14):1193-201
references.pmid: 10580275
references.citation: Nat Rev Immunol. 2003 Aug;3(8):677-85
references.pmid: 12974482
references.citation: Nature. 2003 Apr 10;422(6932):611-4
references.pmid: 12679788
references.citation: Nat Rev Immunol. 2007 Jul;7(7):556-67
references.pmid: 17589545
references.citation: Emerg Infect Dis. 2004 Dec;10(12):2073-81
references.pmid: 15663841
references.citation: Emerg Infect Dis. 2013 May;19(5):743-7
references.pmid: 23647732
references.citation: mBio. 2013 Sep 03;4(5):e00598-13
references.pmid: 24003179
references.citation: J Infect Dis. 1999 Feb;179 Suppl 1:S54-9
references.pmid: 9988165
references.citation: Arch Virol. 2010 Dec;155(12):2083-103
references.pmid: 21046175
references.citation: Am J Trop Med Hyg. 1996 Jul;55(1):89-90
references.pmid: 8702028
references.citation: Lancet. 1991 Feb 16;337(8738):425-6
references.pmid: 1671441
references.citation: Viruses. 2012 Oct 01;4(10):1878-927
references.pmid: 23202446
references.citation: Emerg Infect Dis. 2011 Aug;17(8):1559-60
references.pmid: 21801651
references.citation: Viruses. 2012 Sep;4(9):1425-37
references.pmid: 23170166
references.citation: J Infect Dis. 2007 Nov 15;196 Suppl 2:S400-3
references.pmid: 17940976
references.citation: J R Soc Interface. 2012 Jan 7;9(66):89-101
references.pmid: 21632614
references.citation: J Virol. 2013 Mar;87(5):2608-16
references.pmid: 23255795
references.citation: Philos Trans R Soc Lond B Biol Sci. 2012 Oct 19;367(1604):2881-92
references.pmid: 22966143
references.citation: J Virol. 2014 Jan;88(1):99-109
references.pmid: 24131711
references.citation: J Infect Dis. 1999 Feb;179 Suppl 1:S155-63
references.pmid: 9988179
references.citation: Virus Res. 2008 Mar;132(1-2):140-4
references.pmid: 18166242
references.citation: J Mammal. 2012 Oct;93(5):1393-1404
references.pmid: 23525358
references.citation: PLoS One. 2012;7(7):e40740
references.pmid: 22815803
references.citation: BMC Evol Biol. 2011 Nov 17;11:336
references.pmid: 22093762
references.citation: Emerg Infect Dis. 2003 Dec;9(12):1531-7
references.pmid: 14720391
references.citation: J Infect Dis. 1999 Feb;179 Suppl 1:S120-6
references.pmid: 9988175
references.citation: Am J Trop Med Hyg. 2006 Jul;75(1):9-15
references.pmid: 16837700
references.citation: PLoS One. 2010 Feb 09;5(2):e9126
references.pmid: 20161740
references.citation: Proc Natl Acad Sci U S A. 2007 Oct 23;104(43):17123-7
references.pmid: 17942693
references.citation: Lancet. 1990 Mar 3;335(8688):502-5
references.pmid: 1968529
references.citation: Aust Vet J. 1998 Dec;76(12):813-8
references.pmid: 9972433
references.citation: Biometrics. 2006 Dec;62(4):1170-7
references.pmid: 17156292
references.citation: Wkly Epidemiol Rec. 1992 May 8;67(19):142-3
references.pmid: 1581179
references.citation: Nature. 1979 Aug 2;280(5721):361-7
references.pmid: 460412
references.citation: MMWR Morb Mortal Wkly Rep. 1990 Apr 6;39(13):221
references.pmid: 2107388
references.citation: J Infect Dis. 2007 Nov 15;196 Suppl 2:S176-83
references.pmid: 17940947
references.citation: Intervirology. 1982;18(1-2):24-32
references.pmid: 7118520
references.citation: Emerg Infect Dis. 1996 Oct-Dec;2(4):321-5
references.pmid: 8969248
references.citation: Emerg Infect Dis. 2010 Dec;16(12):1990-3
references.pmid: 21122240
references.citation: Viruses. 2012 May;4(5):889-900
references.pmid: 22754654
references.citation: Emerg Infect Dis. 2013 Feb;19(2):270-3
references.pmid: 23343532
references.citation: Virology. 2013 Aug 1;442(2):97-100
references.pmid: 23711383
references.citation: PLoS One. 2013 Nov 21;8(11):e79665
references.pmid: 24278154
references.citation: J Infect Dis. 2007 Nov 15;196 Suppl 2:S148-53
references.pmid: 17940943