midges are abundant hematophagous flies that vector arboviruses of veterinary and medical importance. Dramatic changes in the epidemiology of -borne arboviruses have occurred since 1998, including the emergence of exotic viruses in northern temperate regions, increases in global disease incidence, and enhanced virus diversity in tropical zones. Drivers may include changes in climate, land use, trade, and animal husbandry. New species and new wild reservoir hosts have been implicated in transmission, highlighting the dynamic nature of pathogen-vector-host interactions. Focusing on potential vector species worldwide and key elements of vectorial capacity, we review the sensitivity of life cycles to abiotic and biotic factors. We consider implications for designing control measures and understanding impacts of environmental change in different ecological contexts. Critical geographical, biological, and taxonomic knowledge gaps are prioritized. Recent developments in genomics and mathematical modeling may enhance ecological understanding of these complex arbovirus systems.


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Literature Cited

  1. Acevedo P, Ruiz-Fons F, Estrada R, Márquez AL, Miranda MA. 1.  et al. 2010. A broad assessment of factors determining Culicoides imicola abundance: modelling the present and forecasting its future in climate change scenarios. PLOS ONE 5:e14236 [Google Scholar]
  2. Akey DH, Potter HW, Jones RH. 2.  1978. Effects of rearing temperature and larval density on longevity, size and fecundity in the biting gnat Culicoides variipennis. Ann. Entomol. Soc. Am. 71:411–18 [Google Scholar]
  3. Allingham PG. 3.  1991. Effect of temperature on late immature stages of Culicoides brevitarsis (Diptera: Ceratopogonidae). J. Med. Entomol. 28:878–81 [Google Scholar]
  4. Barnard DR, Jones RH. 4.  1980. Diel and seasonal patterns of flight activity of Ceratopogonidae in northeastern Colorado: Culicoides. Environ. Entomol. 9:446–51 [Google Scholar]
  5. Baylis M, Mellor PS, Meiswinkel R. 5.  1998. Horse sickness and ENSO in South Africa. Nature 397:574 [Google Scholar]
  6. Beer M, Conraths FJ, Van Der Poel WHM. 6.  2013. ‘Schmallenberg virus’—a novel orthobunyavirus emerging in Europe. Epidemiol. Infect. 141:1–8 [Google Scholar]
  7. Bellis G, Dyce A, Gopurenko D, Yanase T, Garros C. 7.  et al. 2014. Revision of the Culicoides (Avaritia) imicola complex Khamala and Kettle (Diptera: Ceratopogonidae) from the Australasian region. Zootaxa 3768:401–27 [Google Scholar]
  8. Bellis GA, Melville LF, Hunt NT, Hearnden MN. 8.  2004. Temporal activity of biting midges (Diptera: Ceratopogonidae) on cattle near Darwin, Northern Territory, Australia. Vet. Ital. 40:324–28 [Google Scholar]
  9. Berry BS, Magori K, Perofsky AC, Stallknecht D, Park AW. 9.  2013. Wetland cover dynamics drive hemorrhagic disease patterns in white-tailed deer in the United States. J. Wildl. Dis. 49:501–9 [Google Scholar]
  10. Bessell PR, Searle KR, Auty HK, Handel IG, Purse BV, Bronsvoort BM. 10.  2013. Epidemic potential of an emerging vector borne disease in a marginal environment: Schmallenberg in Scotland. Nat. Sci. Rep. 3:1178 [Google Scholar]
  11. Bidlingmayer WL. 11.  1961. Field activity studies of adult Culicoides furens. Ann. Entomol. Soc. Am. 54:149–56 [Google Scholar]
  12. Bishop AL, McKenzie HJ, Barchia IM, Harris AM. 12.  1995. The effects of habitat on the distribution of Culicoides brevitarsis Kieffer (Diptera: Certatopogonidae) during its resting phase. Aust. J. Zool. 43:531–39 [Google Scholar]
  13. Bishop AL, McKenzie HJ, Barchia IM, Harris AM. 13.  1996. Effect of temperature regimes on the development, survival and emergence of Culicoides brevitarsis Kieffer (Diptera: Ceratopogonidae) in bovine dung. Aust. J. Entomol. 35:361–68 [Google Scholar]
  14. Bishop AL, McKenzie HJ, Barchia IM, Murison R, Spohr LJ. 14.  1996. Positions of juvenile stages of Culicoides brevitarsis Kieffer (Diptera: Ceratopogonidae) and of four other flies in bovine dung. Aust. J. Entomol. 35:209–12 [Google Scholar]
  15. Bishop AL, McKenzie HJ, Spohr LJ. 15.  2008. Attraction of Culicoides brevitarsis Kieffer (Diptera: Ceratopogonidae) and Culex annulirostris Skuse (Diptera: Culicidae) to simulated visual and chemical stimuli from cattle. Aust. J. Entomol. 47:121–27 [Google Scholar]
  16. Bishop AL, Spohr LJ, Barchia IM. 16.  2004. Effects of altitude, distance and waves of movement on the dispersal in Australia of the arbovirus vector, Culicoides brevitarsis Kieffer (Diptera: Ceratopogonidae). Prev. Vet. Med. 65:135–45 [Google Scholar]
  17. Blackwell A, Lock KA, Marshall B, Boag B, Gordon SC. 17.  1999. The spatial distribution of larvae of Culicoides impunctatus biting midges. Med. Vet. Entomol. 13:362–71 [Google Scholar]
  18. Blanton FS, Wirth WW. 18.  1979. The sand flies (Culicoides) of Florida (Diptera: Ceratopogonidae). Bur. Entomol. Rep. 424 Fla. Dep. Agric. Consum. Serv., Div. Plant Ind., Gainesville, Fla. [Google Scholar]
  19. Boinas F, Calistri P, Domingo M, Martínez Avilés M, Martínez López B. 19.  et al. 2009. Scientific review on African horse sickness. Extern. Rep. Quest. No. EFSA-Q-2007-00234 Eur. Food Saf. Auth., Parma, Italy. http://www.efsa.europa.eu/en/supporting/doc/4e.pdf [Google Scholar]
  20. Borkent A. 20.  2014. World species of biting midges (Diptera: Ceratopogonidae) Work. Pap., R. B.C. Mus., Am. Mus. Nat. Hist., Inst. Nac. Biodivers. [Google Scholar]
  21. Braverman Y, Hulley PE. 21.  1979. Relationship between the numbers and distribution of some antennal and palpal sense-organs and host preference in some Culicoides (Diptera: Ceratopogonidae) from southern Africa. J. Med. Entomol. 15:419–24 [Google Scholar]
  22. Brenner RJ, Wargo MJ, Staines GS, Mulla MS. 22.  1984. The dispersal of Culicoides mohave (Diptera: Ceratopogonidae) in the desert of southern California. Mosq. News 44:343–50 [Google Scholar]
  23. Burgin LE, Gloster J, Sanders C, Mellor PS, Gubbins S, Carpenter S. 23.  2013. Investigating incursions of bluetongue virus using a model of long-distance Culicoides biting midge dispersal. Transbound. Emerg. Dis. 60:263–72 [Google Scholar]
  24. Calvo JH, Berzal B, Calvete C, Miranda MA, Estrada R, Lucientes J. 24.  2012. Host feeding patterns of Culicoides species (Diptera: Ceratopogonidae) within the Picos de Europa National Park in northern Spain. Bull. Entomol. Res. 102:692–97 [Google Scholar]
  25. Carpenter S, Lunt HL, Arav D, Venter GJ, Mellor PS. 25.  2006. Oral susceptibility to bluetongue virus of Culicoides (Diptera: Ceratopogonidae) from the United Kingdom. J. Med. Entomol. 43:73–78 [Google Scholar]
  26. Carpenter S, Mellor PS, Torr SJ. 26.  2008. Control techniques for Culicoides biting midges and their application in the U.K. and northwestern Palaearctic. Med. Vet. Entomol. 22:175–87 [Google Scholar]
  27. Carpenter S, Mordue AJ, Mordue W. 27.  2006. Oogenesis and laboratory survival in the Scottish biting midge Culicoides impunctatus. Physiol. Entomol. 31:89–93 [Google Scholar]
  28. Carpenter S, Mordue W. 28.  Mordue (Luntz) J. 2008. Selection of resting areas by emerging Culicoides impunctatus (Diptera: Ceratopogonidae) on downy birch (Betula pubescens). Int. J. Pest Manag. 54:39–42 [Google Scholar]
  29. Carpenter S, Szmaragd C, Barber J, Labuschagne K, Gubbins S, Mellor P. 29.  2008. An assessment of Culicoides surveillance techniques in northern Europe: Have we underestimated a potential bluetongue virus vector?. J. Appl. Ecol. 45:1237–45 [Google Scholar]
  30. Carpenter S, Wilson A, Barber J, Veronesi E, Mellor P. 30.  et al. 2011. Temperature dependence of the extrinsic incubation period of orbiviruses in Culicoides biting midges. PLOS ONE 6:e27987 [Google Scholar]
  31. Carpenter S, Wilson A, Mellor PS. 31.  2009. Culicoides and the emergence of bluetongue virus in northern Europe. Trends Microbiol. 17:172–78 [Google Scholar]
  32. Conte A, Goffredo M, Ippoliti C, Meiswinkel R. 32.  2007. Influence of biotic and abiotic factors on the distribution and abundance of Culicoides imicola and the Obsoletus Complex in Italy. Vet. Parasitol. 150:333–44 [Google Scholar]
  33. Del Rio Lopez R, Miranda MA, Paredes-Esquivel C, Lucientes J, Calvete C. 33.  et al. 2012. Recovery rates of bluetongue virus serotypes 1, 2, 4, and 8 Spanish strains from orally infected Culicoides imicola in South Africa. Med. Vet. Entomol. 26:162–67 [Google Scholar]
  34. Eagles D, Deveson T, Walker PJ, Zaluck MP, Durr P. 34.  2012. Evaluation of long-distance dispersal of Culicoides midges into northern Australia using a migration model. Med. Vet. Entomol. 211:334–40 [Google Scholar]
  35. Elbers ARW, Meiswinkel R, van Weezep E, van Oldruitenborgh-Oosterbaan MMS, Kooi EA. 35.  2013. Schmallenberg virus in Culicoides spp. biting midges, the Netherlands, 2011. Emerg. Infect. Dis. 19:106–9 [Google Scholar]
  36. Gerry AC, Mullens BA. 36.  2000. Seasonal abundance and survivorship of Culicoides sonorensis (Diptera: Ceratopogonidae) at a southern California dairy, with reference to potential bluetongue virus transmission and persistence. J. Med. Entomol. 37:675–88 [Google Scholar]
  37. Gerry AC, Mullens BA, MacLachlan NJ, Mecham OJ. 37.  2001. Seasonal transmission of bluetongue virus by Culicoides sonorensis (Diptera: Ceratopogonidae) at a southern California dairy and evaluation of vectorial capacity as a predictor of bluetongue virus transmission. J. Med. Entomol. 38:197–209 [Google Scholar]
  38. Gerry AC, Sarto i Monteys V, Moreno Vidal JO, Francino O, Mullens BA. 38.  2009. Biting rates of Culicoides midges (Diptera: Ceratopogonidae) on sheep in northeastern Spain in relation to midge capture using UV light and carbon dioxide-baited traps. J. Med. Entomol. 46:615–24 [Google Scholar]
  39. Gibbs EPJ, Greiner EC. 39.  1994. The epidemiology of bluetongue. Comp. Immunol. Microbiol. Infect. Dis. 17:207–20 [Google Scholar]
  40. Glukhova VM. 40.  1979. Larvae of biting midges of the subfamilies Palpomyiinae and Ceratopogoninae of the USSR fauna (Diptera, Ceratopogonidae = Heleidae). Proc. Zool. Inst. Russ. Acad. Sci. 121:1–230 [Google Scholar]
  41. Goffredo M, Romeo G, Monaco F, Di Gennaro A, Savini G. 41.  2004. Laboratory survival and blood feeding response of wild-caught Culicoides obsoletus complex (Diptera: Ceratopogonidae) through natural and artificial membranes. Vet. Ital. 40:282–85 [Google Scholar]
  42. Græsbøll K, Bødker R, Enøe C, Christiansen LE. 42.  2012. Simulating spread of bluetongue virus by flying vectors between hosts on pasture. Nat. Sci. Rep. 2:863 [Google Scholar]
  43. Gubbins S, Carpenter S, Baylis M, Wood JLN, Mellor PS. 43.  2008. Assessing the risk of bluetongue to UK livestock: uncertainty and sensitivity analyses of a temperature-dependent model for the basic reproduction number. J. R. Soc. Interface 5:363–71 [Google Scholar]
  44. Harrup LE, Logan JG, Cook JI, Golding N, Birkett MA. 44.  et al. 2012. Collection of Culicoides (Diptera: Ceratopogonidae) using CO2 and enantiomers of 1-octen-3-ol in the United Kingdom. J. Med. Entomol. 49:112–21 [Google Scholar]
  45. Harrup LE, Purse BV, Golding N, Mellor PS, Carpenter S. 45.  2013. Larval development and emergence sites of farm-associated Culicoides in the United Kingdom. Med. Vet. Entomol. 27:441–49 [Google Scholar]
  46. Hartemink NA, Purse BV, Meiswinkel R, Brown HE, De Koeijer AA. 46.  et al. 2009. Mapping the basic reproduction number (R0) for vector-borne diseases: a case study on bluetongue virus. Epidemics 1:153–61 [Google Scholar]
  47. Hoch AL, Roberts DR, Pinheiro FP. 47.  1986. Breeding sites of Culicoides paraensis and options for control by environmental management. Bull. Pan Am. Health Org. 20:284–93 [Google Scholar]
  48. Hoffmann B, Bauer B, Bauer C, Batza HJ, Beer M. 48.  et al. 2009. Monitoring of putative vectors of bluetongue virus serotype 8, Germany. Emerg. Infect. Dis. 15:1481–84 [Google Scholar]
  49. Hunt GJ, Tabachnick WJ. 49.  1995. Cold-storage effects on egg hatch in laboratory-reared Culicoides variipennis sonorensis (Diptera: Ceratopogonidae). J. Am. Mosq. Control Assoc. 11:335–38 [Google Scholar]
  50. Isberg E, Hillbur Y, Ignell R. 50.  2013. Comparative study of antennal and maxillary palp olfactory sensilla of female biting midges (Diptera: Ceratopogonidae: Culicoides) in the context of host preference and phylogeny. J. Med. Entomol. 50:485–92 [Google Scholar]
  51. Jamnbach H. 51.  1961. Observations on Culicoides obsoletus (Meigen) in the laboratory (Diperta: Ceratopogonidae). Mosq. News 21:48–53 [Google Scholar]
  52. Jennings M, Mellor PS. 52.  1989. Culicoides: biological vectors of Akabane virus. Vet. Microbiol. 21:125–31 [Google Scholar]
  53. Johansen CA, Farrow RA, Morrisen A, Foley P, Bellis G. 53.  et al. 2003. Collection of wind-borne haematophagous insects in the Torres Strait, Australia. Med. Vet. Entomol. 17:102–9 [Google Scholar]
  54. Kang CH, Yu HS. 54.  1991. Seasonal abundance and host blood meal sources of the genus Culicoides (Diptera: Ceratopogonidae) from cattle and poultry farms in Kyeonggi Province, Korea. Korean J. Entomol. 21:29–36 [Google Scholar]
  55. Kettle DS. 55.  1962. The bionomics and control of Culicoides and Leptoconops (Diptera, Ceratopogonidae = Heleidae). Annu. Rev. Entomol. 7:401–18 [Google Scholar]
  56. Kettle DS. 56.  1969. The biting habits of Culicoides furens (Poey) and C. barbosai Wirth & Blanton. I. The 24-h cycle, with a note on differences between collectors. Bull. Entomol. Res. 59:21–31 [Google Scholar]
  57. Kettle DS. 57.  1977. Biology and bionomics of blood sucking ceratopogonids. Annu. Rev. Entomol. 22:33–51 [Google Scholar]
  58. Kitaoka S. 58.  1982. Effects of rearing temperature on length of larval period and size of adults in Culicoides arakawae and Culicoides maculatus (Dipteria: Ceratopogonidae). Natl. Inst. Anim. Health Q. 22:159–62 [Google Scholar]
  59. Lassen SB, Nielsen SA, Kristensen M. 59.  2012. Identity and diversity of blood meal hosts of biting midges (Diptera: Ceratopogonidae: Culicoides Latreille) in Denmark. Parasites Vectors 5:143 [Google Scholar]
  60. LeDuc JW, Pinheiro FP. 60.  1989. Oropouche fever. The Arboviruses: Epidemiology and Ecology TP Monath 41–14 Boca Raton, FL: CRC Press [Google Scholar]
  61. Lillie TH, Marquardt WC, Jones RH. 61.  1981. The flight range of Culicoides variipennis (Diptera: Ceratopogonidae).. Can. Entomol. 113:419–26 [Google Scholar]
  62. Linden A, Desmecht D, Volpe R, Wirtgen M, Gregoire F. 62.  et al. 2012. Epizootic spread of Schmallenberg virus among wild cervids, Belgium, fall 2011. Emerg. Infect. Dis. 18:12 Dec.; doi: 10.3201/eid1812.121067 [Google Scholar]
  63. Linley JR. 63.  1966. The ovarian cycle in Culicoides barbosai Wirth & Blanton and C. furens (Poey) (Diptera: Ceratopogonidae). Bull. Entomol. Res. 57:1–17 [Google Scholar]
  64. Linley JR. 64.  1983. Autogeny in the Ceratopogonidae: literature and notes. Fla. Entomol. 66:228–34 [Google Scholar]
  65. Linthicum KJ, Davies FG. 65.  1984. Aerial collection of Culicoides schultzei group (Diptera: Ceratopogonidae) in Kenya. Mosq. News 44:601–3 [Google Scholar]
  66. Lo Iacono G, Robin CA, Newton JR, Gubbins S, Wood JLN. 66.  2013. Where are the horses? With the sheep or cows? Uncertain host location, vector-feeding preferences and the risk of African horse sickness transmission in Great Britain. J. R. Soc. Interface 10:20130194 [Google Scholar]
  67. Logan JG, Cook JI, Mordue (Luntz) AJ, Kline DL. 67.  2010. Understanding and exploiting olfaction for the surveillance and control of Culicoides biting midges. Ecology and Control of Vector-Borne Diseases: Olfaction in Vector-Host Interactions W Takken, BGJ Knols 2217–46 Wageningen, The Neth: Wageningen Acad. [Google Scholar]
  68. Lord CC, Woolhouse MEJ, Heesterbeek JAP, Mellor PS. 68.  1996. Vector-borne diseases and the basic reproduction number: a case study of African horse sickness. Med. Vet. Entomol. 10:19–28 [Google Scholar]
  69. Lysyk TJ, Danyk T. 69.  2007. Effect of temperature on life history parameters of adult Culicoides sonorensis (Diptera: Ceratopogonidae) in relation to geographic origin and vectorial capacity for bluetongue virus. J. Med. Entomol. 44:741–51 [Google Scholar]
  70. Maan S, Maan NS, Nomikou K, Anthony SJ, Ross-Smith N. 70.  et al. 2009. Molecular epidemiology studies of bluetongue virus. Bluetongue P Mellor, M Baylis, PPC Mertens 135–66 New York: Academic [Google Scholar]
  71. Maclachlan NJ, Drew CP, Darpel KE, Worwa G. 71.  2009. The pathology and pathogenesis of bluetongue. J. Comp. Pathol. 141:1–16 [Google Scholar]
  72. Maclachlan NJ, Mayo CE. 72.  2013. Potential strategies for control of bluetongue, a globally emerging, Culicoides-transmitted viral disease of ruminant livestock and wildlife. Antivir. Res. 99:79–90 [Google Scholar]
  73. Maclachlan NJ, Wilson WC, Crossley BM, Mayo CE, Jasperson DE. 73.  et al. 2013. Novel serotype of Bluetongue virus, western North America [letter]. Emerg. Infect. Dis. 19:4 [Google Scholar]
  74. Mayo CE, Mullens BA, Gerry AC, Barker CM, Mertens PPC. 74.  et al. 2012. The combination of abundance and infection rates of Culicoides sonorensis estimates risk of subsequent bluetongue virus infection of sentinel cattle on California dairy farms. Vet. Parasitol. 187:295–301 [Google Scholar]
  75. Meiswinkel R, Paweska JT. 75.  2003. Evidence for a new field Culicoides vector of African horse sickness in South Africa. Prev. Vet. Med. 60:243–53 [Google Scholar]
  76. Mellor PS, Boorman J, Baylis M. 76.  2000. Culicoides biting midges: their role as arbovirus vectors. Annu. Rev. Entomol. 45:307–40 [Google Scholar]
  77. Mellor PS, Carpenter S, White DM. 77.  2009. Bluetongue in the insect host. Bluetongue Monograph PS Mellor, M Baylis, PPC Mertens 295–321 London: Elsevier, 1st ed.. [Google Scholar]
  78. Melville LF, Hunt NT, Bellis G, Pinch D. 78.  2004. An assessment of insecticides to minimize the transmission of arboviruses in cattle. Arbovirus Res. Aust. 8:249–55 [Google Scholar]
  79. Mercer DR, Spinelli GR, Watts DM, Tesh RB. 79.  2003. Biting rates and developmental substrates for biting midges (Diptera: Ceratopogonidae) in Iquitos, Peru. J. Med. Entomol. 40:807–12 [Google Scholar]
  80. Mullen GR, Hribar LJ. 80.  1988. Biology and feeding behavior of ceratopogonid larvae (Diptera: Ceratopogonidae) in North America. Bull. Soc. Vector Ecol. 13:60–81 [Google Scholar]
  81. Mullens BA, Gerry AC. 81.  1998. Comparison of bait cattle and carbon dioxide-baited suction traps for collecting Culicoides variipennis sonorensis (Diptera: Ceratopogonidae) and Culex quinquefasciatus (Diptera: Culicidae). J. Med. Entomol. 35:245–50 [Google Scholar]
  82. Mullens BA, Gerry AC, Lysyk TJ, Schmidtmann ET. 82.  2004. Environmental effects on vector competence and virogenesis of bluetongue virus in Culicoides: interpreting laboratory data in a field context. Vet. Ital. 40:160–66 [Google Scholar]
  83. Mullens BA, Gerry AC, Velten RK. 83.  2001. Failure of a permethrin treatment regime to protect against bluetongue virus. J. Vet. Entomol. 38:760–62 [Google Scholar]
  84. Mullens BA, Rodriguez JL. 84.  1985. Effect of experimental habitat shading on the distribution of Culicoides variipennis (Diptera: Ceratopogonidae) larvae. Environ. Entomol. 14:749–54 [Google Scholar]
  85. Mullens BA, Rodriguez JL. 85.  1988. Colonization and response of Culicoides variipennis (Diptera: Ceratopogonidae) to pollution levels in experimental dairy wastewater ponds. J. Med. Entomol. 25:441–51 [Google Scholar]
  86. Mullens BA, Rodriguez JL. 86.  1992. Survival and vertical distribution of larvae of Culicoides variipennis (Diptera: Ceratopogonidae) in drying mud habitats. J. Med. Entomol. 29:745–49 [Google Scholar]
  87. Mullens BA, Rutz DA. 87.  1983. Development of immature Culicoides variipennis (Diptera: Ceratopogonidae) at constant laboratory temperatures. Ann. Entomol. Soc. Am. 76:747–51 [Google Scholar]
  88. Mullens BA, Sarto i Monteys V, Przhiboro AA. 88.  2008. Mermithid parasitism in Ceratopogonidae: a literature review and critical assessment of host impact and potential for biological control of Culicoides. Russ. Entomol. J. 17:87–113 [Google Scholar]
  89. Mullens BA, Tabachnick WJ, Holbrook FR, Thompson LH. 89.  1995. Effects of temperature on virogenesis of bluetongue virus serotype 11 in Culicoides variipennis sonorensis. Med. Vet. Entomol. 9:71–76 [Google Scholar]
  90. Muller MJ. 90.  1985. Experimental infection of Culicoides brevitarsis from southeast Queensland with three serotypes of bluetongue virus. Aust. J. Biol. Sci. 38:73–77 [Google Scholar]
  91. Murray MD. 91.  1991. The seasonal abundance of female biting midges, Culicoides brevitarsis Kieffer (Diptera: Ceratopogonidae), in coastal southeastern Australia. Aust. J. Zool. 39:333–42 [Google Scholar]
  92. Napp S, García-Bocanegra I, Pagès N, Allepuz A, Alba A, Casal J. 92.  2013. Assessment of the risk of a bluetongue outbreak in Europe caused by Culicoides midges introduced through intracontinental transport and trade networks. Med. Vet. Entomol. 27:19–28 [Google Scholar]
  93. Nayduch D, Cohnstaedt LW, Saski C, Lawson D, Kersey P. 93.  et al. 2014. Studying Culicoides vectors of BTV in the post-genomic era: resources, bottlenecks to progress and future directions. Virus Res. 182:43–49 [Google Scholar]
  94. Nevill EM. 94.  1967. Biological studies on some South African Culicoides species (Diptera: Ceratopogonidae) and the morphology of their immature stages MSc thesis. Univ. Pretoria [Google Scholar]
  95. Ng TFF, Willner DL, Lim YW, Schmieder R, Chau B. 95.  et al. 2011. Broad surveys of DNA viral diversity obtained through viral metagenomics of mosquitoes. PLOS ONE 6:e20579 [Google Scholar]
  96. Nie W, Li J, Li D, Gratz N. 96.  2005. A preliminary report of introduced living biting midges (Diptera: Ceratopogonidae) found on entry ships arriving at Qinhuangdao Port, China. Med. Entomol. Zool. 56:359–61 [Google Scholar]
  97. Nunamaker RA, Sieburth PJ, Dean VC, Wigington JG, Nunamaker CE, Mecham JO. 97.  1990. Absence of transovarial transmission of bluetongue virus in Culicoides variipennis: immunogold labelling of bluetongue virus in developing oocytes from Culicoides variipennis. Comp. Biochem. Physiol. A 96:19–31 [Google Scholar]
  98. Parker A. 98.  1950. Studies on the eggs of certain biting midges (Culicoides Latreille) occurring in Scotland. Proc. R. Entomol. Soc. Lond. 25:43–52 [Google Scholar]
  99. Paweska JT, Venter GJ, Hamblin C. 99.  2005. A comparison of the susceptibility of Culicoides imicola and C. bolitinos to oral infection with eight serotypes of epizootic haemorrhagic disease virus. Med. Vet. Entomol. 19:200–7 [Google Scholar]
  100. Pinheiro FP, Darosa A, Darosa J, Ishak R, Freitas RB. 100.  et al. 1981. Oropouche virus. I. A review of clinical, epidemiological and ecological findings. Am. J. Trop. Med. Hyg. 30:149–60 [Google Scholar]
  101. Pioz M, Guis H, Calavas D, Durand B, Abrial D, Ducrot C. 101.  2011. Estimating front-wave velocity of infectious diseases: a simple, efficient method applied to bluetongue. Vet. Res. 42:60 [Google Scholar]
  102. Pioz M, Guis H, Crespin L, Gay E, Calavas D. 102.  et al. 2012. Why did bluetongue spread the way it did? Environmental factors influencing the velocity of bluetongue virus serotype 8 epizootic wave in France. PLOS ONE 7:e43360 [Google Scholar]
  103. Prasad G, Sreenivasulu D, Singh KP, Mertens P, Maan S. 103.  2009. Bluetongue in the Indian subcontinent. Bluetongue PS Mellor, M Baylis, PPC Mertens 167–96 London: Elsevier Acad. [Google Scholar]
  104. Purse BV, Brown HE, Harrup L, Mertens PPC, Rogers DJ. 104.  2008. Invasion of bluetongue and other orbivirus infections into Europe: the role of biological and climatic processes. Rev. Sci. Tech. 27:427–42 [Google Scholar]
  105. Purse BV, Falconer D, Sullivan MJ, Carpenter S, Mellor PS. 105.  et al. 2012. Impacts of climate, host and landscape factors on Culicoides species in Scotland. Med. Vet. Entomol. 26:168–77 [Google Scholar]
  106. Purse BV, McCormick BJJ, Mellor PS, Baylis M, Boorman J. 106.  et al. 2007. Incriminating bluetongue virus vectors with climate envelope models. J. Appl. Ecol. 44:1231–42 [Google Scholar]
  107. Purse BV, Mellor PS, Rogers DJ, Samuel AR, Mertens PPC, Baylis M. 107.  2005. Climate change and the recent emergence of bluetongue in Europe. Nat. Rev. Microbiol. 3:171–81 [Google Scholar]
  108. Rigot T, Vercauteren Drubbel M, Delecolle J, Gilbert M. 108.  2013. Farms, pastures and woodlands: the fine-scale distribution of Palearctic Culicoides spp. biting midges along an agro-ecological gradient. Med. Vet. Entomol. 27:29–38 [Google Scholar]
  109. Roberts DR, Hoch AL, Dixon KE, Llewellyn CH. 109.  1981. Oropouche virus. III. Entomological observations from three epidemics in Para, Brazil, 1975. Am. J. Trop. Med. Hyg. 30:165–71 [Google Scholar]
  110. Roberts DR, Pinheiro FdP, Hoch AL, LeDuc JW, Peterson NE. 110.  et al. 1977. Vectors and natural reservoirs of Oropouche virus in the Amazon region. Tech. Rep. NTIS 7815 US Dep. Commer., Washington, DC; access. no. AD-A053 168/1 [Google Scholar]
  111. Ruder M, Howerth E, Stallknecht D, Allison A, Carter D. 111.  et al. 2012. Vector competence of Culicoides sonorensis (Diptera: Ceratopogonidae) to epizootic hemorrhagic disease virus serotype 7. Parasites Vectors 5:236 [Google Scholar]
  112. Sanders CJ, Gubbins S, Mellor PS, Barber J, Golding N. 112.  et al. 2012. Investigation of diel activity of Culicoides biting midges (Diptera: Ceratopogonidae) in the United Kingdom by using a vehicle-mounted trap. J. Med. Entomol. 49:757–65 [Google Scholar]
  113. Sanders CJ, Selby R, Carpenter S, Reynolds DR. 113.  2011. High-altitude flight of Culicoides biting midges. Vet. Rec. 169:208 [Google Scholar]
  114. Santiago-Alarcon D, Havelka P, Schaefer HM, Segelbacher G. 114.  2012. Bloodmeal analysis reveals avian Plasmodium infections and broad host preferences of Culicoides (Diptera: Ceratopogonidae) vectors. PLOS ONE 7:e31098 [Google Scholar]
  115. Savini G, Afonso A, Mellor PS, Aradaib I, Yadin H. 115.  et al. 2011. Epizootic hemorrhagic disease. Res. Vet. Sci. 91:1–17 [Google Scholar]
  116. Schmidtmann ET, Bobian RJ, Belden RP. 116.  2000. Soil chemistries define aquatic habitats with immature populations of the Culicoides variipennis complex (Diptera: Ceratopogonidae). J. Med. Entomol. 37:58–64 [Google Scholar]
  117. Schmidtmann ET, Jones CJ, Gollands B. 117.  1980. Comparative host-seeking activity of Culicoides (Diptera: Ceratopogonidae) attracted to pastured livestock in central New York state, USA. J. Med. Entomol. 17:221–31 [Google Scholar]
  118. Sedda L, Brown HE, Purse BV, Burgin L, Gloster J, Rogers DJ. 118.  2012. A new algorithm quantifies the roles of wind and midge flight activity in the bluetongue epizootic in northwest Europe. Proc. R. Soc. B 279:2354–62 [Google Scholar]
  119. Severson DW, Behura SK. 119.  2012. Mosquito genomics: progress and challenges. Annu. Rev. Entomol. 57:143–66 [Google Scholar]
  120. Sinclair M, Bührmann G, Gummow B. 120.  2006. An epidemiological investigation of the African horse sickness outbreak in the Western Cape province of South Africa in 2004 and its relevance to the current equine export protocol. J. S. Afr. Vet. Assoc. 77:191–96 [Google Scholar]
  121. Tabachnik WJ. 121.  1996. Culicoides variipennis and bluetongue virus epidemiology in the United States. Annu. Rev. Entomol. 41:23–43 [Google Scholar]
  122. Tabachnik WJ. 122.  2004. Culicoides and the global epidemiology of bluetongue virus infection. Vet. Ital. 40:145–50 [Google Scholar]
  123. Takken W, Verhulst NO. 123.  2013. Host preferences of blood-feeding mosquitoes. Annu. Rev. Entomol. 58:433–53 [Google Scholar]
  124. Turner J, Bowers RG, Baylis M. 124.  2013. Two-host, two-vector basic reproduction ratio (R0) for bluetongue. PLOS ONE 8:e53128 [Google Scholar]
  125. Uslu U, Dik B. 125.  2006. Vertical distribution of Culicoides larvae and pupae. Med. Vet. Entomol. 20:350–52 [Google Scholar]
  126. Vasconcelos HB, Nunes MRT, Casseb LMN, Carvalho VL, Pinto da Silva EV. 126.  et al. 2011. Molecular epidemiology of Oropouche virus, Brazil. Emerg. Infect. Dis. 17:5 May; doi: 10.3201/eid1705.101333 [Google Scholar]
  127. Vaughan JA, Turner EC. 127.  1987. Development of immature Culicoides variipennis (Diptera: Ceratopogonidae) from Saltville, Virginia, at constant laboratory temperatures. J. Med. Entomol. 24:390–95 [Google Scholar]
  128. Velthuis AG, Saatkamp HW, Mourits MC, de Koeijer AA, Elbers A. 128.  2010. Financial consequences of the Dutch bluetongue serotype 8 epidemics of 2006 and 2007. Prevent. Vet. Med. 93:294–304 [Google Scholar]
  129. Venter GJ, Paweska JT, Van Dijk AA, Mellor PS, Tabachnick WJ. 129.  1998. Vector competence of Culicoides bolitinos and C. imicola for South African bluetongue virus serotypes 1, 3 and 4. Med. Vet. Entomol. 12:378–85 [Google Scholar]
  130. Veronesi E, Venter GJ, Labuschagne K, Mellor PS, Carpenter S. 130.  2009. Life-history parameters of Culicoides (Avaritia) imicola Kieffer in the laboratory at different rearing temperatures. Vet. Parasitol. 163:370–73 [Google Scholar]
  131. Viennet E, Garros C, Gardès L, Rakotoarivony I, Allène X. 131.  et al. 2012. Host preferences of Palaearctic Culicoides biting midges: implications for transmission of orbiviruses. Med. Vet. Entomol. 27:255–66 [Google Scholar]
  132. Viennet E, Garros C, Lancelot R, Allène X, Gardès L. 132.  et al. 2011. Assessment of vector/host contact: comparison of animal-baited traps and UV-light/suction trap for collecting Culicoides biting midges (Diptera: Ceratopogonidae), vectors of Orbiviruses. Parasites Vectors 4:119 [Google Scholar]
  133. Walker AR. 133.  1977. Seasonal fluctuations of the Culicoides species (Diptera: Ceratopogonidae) in Kenya. Bull. Entomol. Res. 67:217–33 [Google Scholar]
  134. Wang J, Wu Y, Yang G, Aksoy S. 134.  2009. Interactions between mutualist Wigglesworthia and tsetse peptidoglycan recognition protein (PGRP-LB) influence trypanosome transmission. Proc. Natl. Acad. Sci. USA 106:12133–38 [Google Scholar]
  135. Wellby MP, Baylis M, Rawlings P, Mellor PS. 135.  1996. Effect of temperature on survival and rate of virogenesis of African horse sickness virus in Culicoides variipennis sonorensis (Diptera: Ceratopogonidae) and its significance in relation to the epidemiology of the disease. Bull. Entomol. Res. 86:715–20 [Google Scholar]
  136. White DM, Wilson WC, Blair CD, Beaty BJ. 136.  2005. Studies on overwintering of bluetongue viruses in insects. J. Gen. Virol. 86:453–62 [Google Scholar]
  137. Wittmann EJ. 137.  2000. Temperature and the transmission of arboviruses by Culicoides biting midges PhD thesis. Univ. Bristol, City of Bristol [Google Scholar]
  138. Wittmann EJ, Mellor PS, Baylis M. 138.  2002. Effect of temperature on the transmission of orbiviruses by the biting midge, Culicoides sonorensis. Med. Vet. Entomol. 16:147–56 [Google Scholar]
  139. Yoshioka M, Couret J, Kim F, McMillan J, Burkot TR. 139.  et al. 2012. Diet and density dependent competition affect larval performance and oviposition site selection in the mosquito species Aedes albopictus (Diptera: Culicidae). Parasites Vectors 5:225 [Google Scholar]
  140. Zimmer J, Saegerman C, Losson B, Haubruge E. 140.  2008. Breeding sites of bluetongue virus vectors, Belgium. Emerg. Infect. Dis. 16:575–76 [Google Scholar]
  141. Zimmer JY, Saegerman C, Losson B, Beckers Y, Haubruge E, Francis F. 141.  2013. Chemical composition of silage residues sustaining the larval development of the Culicoides obsoletus/Culicoides scoticus species (Diptera: Ceratopogonidae). Vet. Parasitol. 191:197–201 [Google Scholar]
  142. Zimmerman RH, Turner EC. 142.  1984. Dispersal and gonotrophic age of Culicoides variipennis (Diptera: Certaopogonidae) at an isolated site in southwestern Virginia, USA. J. Med. Entomol. 21:527–35 [Google Scholar]

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