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The genus Culicoides is the only group of hematophagous ceratopogonids known from the Park. Although other ceratopogonids such as Atrichopogon spp., Dasyhelea oppressa Thompsen , Forcipomyia edmistoni Wirth and Spinelli , and Parabezzia sp. occur in the Park, none of these are known to feed on vertebrates. Some species of North American Forcipomyia ( Lasiohelea ) and Leptoconops feed on reptile blood and bite humans ( Pechuman & Wirth 1961; Mullens et al. 1997).
Culicoides species are some of the most annoying biters of humans in the Park. These flies tend to feed in the morning and afternoon and inflict painful bites. Some Culicoides species are important vectors of Haemoproteus spp., Mansonella spp., the virus that causes epizootic hemorrhagic disease of deer, and other arbovirues ( Reeves et al. 1970; Blanton & Wirth 1979). Rabinowitz et al. (1985) reported a filarial nematode, M. llewellyni (Price) , that infects raccoons, Procyon lotor (L.), in the Park. Culicoides species are the probable vectors of this filarial worm; other species of Mansonella are transmitted by Ceratopogonidae and Simuliidae ( Bain & Chaubaud 1986) .
Collections: North Carolina, Swain Co., Hazel Creek at campsite 86, 11 August 1999, light trap, coll. J. Cooper; Oconaluftee/Ravensford, 31 May 2001, treehole, larvae; 9 June 2001, treehole, larvae.
Culicoides arboricola developed in waterfilled treeholes with C. guttipennis (Coquillett) and Ochlerotatus triseriatus (Say) . No biting records were established in our study. Culicoides arboricola is primarily ornithophilic and apparently does not bite humans ( Blanton & Wirth 1979; Snow et al. 1957). However, Humphreys & Turner (1973) collected an engorged female from caged rabbits, and Wright & DeFoliart (1970 b) collected a single female from a caged beaver.
Collection: Tennessee, Blount Co., Cades Cove (520 meters, 35.5920 °N, 83.8375 °W), 1986 and 1987, emergence trap, ( Root & Gerhardt 1991).
Root & Gerhardt (1991) studied the seasonal emergence of Culicoides spp. throughout eastern Tennessee and caught C. baueri in the Park. Blanton & Wirth (1979) provided records of this species feeding on humans in Florida.
Collections: Tennessee, Blount Co., Cades Cove, 1984, ex deer and cows, (Gerhardt 1986).
Culicoides bickleyi is mammalophilic and ornithophilic and feeds on deer and cows in the Park ( Blanton & Wirth 1979; Gerhardt 1986). Engorged C. bickleyi have been collected on goats, humans, and turkeys ( Wirth & Hubert 1962; Humphreys & Turner 1973, Blanton & Wirth 1979).
Collection: Tennessee, Blount Co., Cades Cove, 25 May 2001, Shannon trap.
Culicoides biguttatus was active early in the spring when the air temperature fell below freezing at night. Culicoides biguttatus feeds on deer and cows in the Park (Gerhardt 1986). Snow et al. (1957) noted that it feeds on horses. Wright & DeFoliart (1970 b) found C. biguttatus engorged on chickens and Hayes et al. (1984) collected it on cattle. Snow & Pickard (1958) reported being bitten by C. biguttatus during daylight hours in lowland forests, but according to Jamback (1965) it rarely bites humans in New York. Malloch (1915) collected as many as 13 females from humans in one day. Childer & Wingo (1968) collected a single female from a human in Missouri.
A single female of C. chiopterus was trapped in the winter, suggesting that the species is active during the colder months of the year or that adults can overwinter and become active on warm days. This species feeds on humans and birds ( Blanton & Wirth 1979).
Collections: North Carolina, Deep Creek , 13 September 2003, biting human; Tennessee, Blount Co., Cades Cove, 1984, ex deer and cows, (Gerhardt 1986).
Blanton & Wirth (1979) noted that C. debilipalpis feeds on birds in Virginia but bites humans in South America and Asia. Remm (1956) reported that C. debilipalpis is an important pest of humans in Estonia during the spring and fall. Culicoides debilipalpis feeds on deer, cows, and humans in the Park (Gerhardt 1986). Wright & DeFoliart (1970 b) recorded C. debilipalpis from turkeys and chickens.
Collection: Tennessee, Sevier Co., Twin Creeks, 17 August – 2 September 1999, funnel trap, coll. Parker.
Larvae of C. footei live in moist treeholes ( Blanton & Wirth 1979) and adults are not known to bite humans.
Collections: North Carolina, Haywood Co., Purchase Knob , 14 May 2002, treehole; Swain Co., Oconaluftee/Ravensford, 9 May 2002, biting horse; 4 July 2002, treehole; Twentymile Creek ranger station, treehole; Tennessee, Blount Co., Cades Cove, 1984, ex deer and cows, (Gerhardt 1986); Cocke Co., Cosby Nature Trail (503 meters, 35.7778 °N, 83.2131 °W), 29 May 2001, treehole.
Culicoides guttipennis was one of the most frequently collected species of Culicoides in treeholes. Larvae of this species coexist with those of C. arboricola , Ochlerotatus triseriatus , and Toxorhynchites rutilus (Coquillett) . A single biting female was taken from a horse at dusk, and Gerhardt (1986) collected this species on deer and cows in the Park. Humphreys & Turner (1973) reported C. guttipennis most commonly on goats, rabbits, and turkeys. Hair & Turner (1968) demonstrated that females prefer humans and small mammals. Snow (1955) reported that C. guttipennis feeds on humans at ground level and in the canopy. However, Jamnback (1965) noted no biting records from humans in New York and suggested that due to the presence of sensilla coeloconica on all flagellomeres, the species might be primarily ornithophilic. Wright & DeFoliart (1970 b) recorded engorged females from chickens.
Culicoides haematopotus is a vector of the blood parasite Haemoproteus meleagridis Levine of wild and domestic turkey ( Mullen 2002). Fallis & Bennett (1961) recorded C. haematopotus feeding on crows and grouse, whereas Hayes et al (1984) collected females from cattle. This species is widely reported to feed on humans (e.g., Malloch 1915).
Collection: North Carolina, Haywood Co., Cataloochee , 19 March 2002, biting elk; Tennessee, Blount Co., Cades Cove, 1984, ex deer and cows, (Gerhardt 1986).
Culicoides obsoletus is a vector of bluetongue virus ( Mullen 2002). Elk were reintroduced into the Park in 2001. The biting record of C. obsoletus on elk could represent a historic host association and is relevant for the epidemiology of epidemic hemorrhagic disease of deer in Cataloochee Cove. This midge also feeds on deer and cows in the Park (Gerhardt 1986) and is an annoying pest of humans at locations up to 2000 meters elevation near Newfound Gap ( Snow et al. 1957). Jamnback (1965) noted that this species feeds on horses and cattle. Wright & DeFoliart (1970 b) collected engorged specimens from rabbits and gray squirrels, and Humphreys & Turner (1973) found a single female on a turkey.
Collections: North Carolina, Haywood, Cataloochee , 22 July 2002, biting humans; Purchase Knob, 22 July 2002, Shannon trap; Swain Co., Oconaluftee/Ravensford, 15 August 2002, biting cat and humans; 11 June 2003, biting horse; Tennessee, Blount Co., Cades Cove, 1984, ex deer and cows, (Gerhardt 1986); Sevier Co., Gatlinburg, 2 September 1954, ( Snow et al. 1957); Rich Mountain Blowhole Cave entrance, 25 August 1999, biting humans.
Culicoides paraensis is a tropical and subtropical biting midge that ranges into the Appalachians during the summer. Culicoides paraensis feeds on deer and cows in the Park (Gerhardt 1986) and can be the most widespread and annoying biter of humans during daylight hours ( Snow et al. 1957). It bites in the afternoon along the edge of woodlands, and feeds on humans in the canopy ( Snow 1955; Snow & Pickard 1958). According to Humphreys & Turner (1973), C. paraensis also feeds on small mammals and birds. We collected C. paraensis feeding on a feral cat in the Park. Because C. paraensis feeds on carnivores, it could serve as a vector of M. llewellyni , a filarial nematode of raccoons. The only laboratory transmission of M. llewellyni involved an apparently misidentified vector. Yates et al. (1982) suggested that C. hollensis (Melander & Brues) was the vector of this nematode, but they used wildcaught flies from Mississippi where C. hollensis does not occur. Culicoides paraensis is a vector of M. ozzardi Manson , a filarial nematode of humans ( Mullen 2002). It also transmits Oropouche virus to humans in South America ( Eldridge et al. 2000).
Collections: North Carolina, Haywood Co., Big Creek Campground , 19 July 1999, light trap, coll. J. Cooper; Cataloochee, 22 July 2002, biting humans; Purchase Knob, 4 August 2003, biting human; Swain Co., Clingman's Dome (1924 meters, 35.5561 °N, 83.4961 °W), 22 July 2001, sewage treatment plant, biting human; Oconaluftee/Ravensford, 9 May 2002, biting human; Straight Fork Motor Trail (959 meters, 35.6215 °N, 83.2110 °W), 19 July 2001, biting human during rainstorm; Tennessee, Blount Co., Abrams Creek sewage treatment plant, 19 June 2002, 2 females, feeding on little brown bat, coll. J. Bellwood; Tremont Institute, 2 August 1999, black light trap, coll. J. Ciegler.
This species, along with C. paraensis , feeds on the exposed arms and legs of Park visitors during daylight hours and at dusk. Two engorged females of C. sanguisuga were removed from the wing of a little brown bat that was trapped as it escaped a maintenance building, representing the first record of C. sanguisuga feeding on bats. This species also feeds on deer and cows in the Park (Gerhardt 1986). Jamnback (1965) recorded this species feeding on horses, rabbits, hares, ducks, grouse, blue jays, and sparrows. Humphreys & Turner (1973) collected over 22,000 flies from goats, 3400 from turkeys, and more than 2000 from rabbits.
Collection: Tennessee, Sevier Co, W. tributary of Porters Creek, 27 July 1999, Malaise trap, coll. J. Cooper.
The single collection of C. snowi in a Malaise trap provides no biological information other than the summer collection date. Culicoides snowi is attracted to humans ( Wirth & Hubert 1962; Blanton & Wirth 1979).
Collections: Tennessee, Blount Co., Cades Cove, 1984, ex deer and cows, (Gerhardt 1986).
Culicoides spinosus bites humans, deer, and cows ( Snow et al. 1957; Blanton & Wirth 1979; Hayes et al. 1984; Gerhardt 1986). Humphreys & Turner (1973) recorded collections of C. spinosus from goats, rabbits, turkeys, and chickens.
Collection: Tennessee, Blount Co., Tremont Institute (524 meters, Thunderhead Mountain 35.621 ºN 83.678 ºW), 22 August 2001, biting human, coll. P. Super; Cades Cove, 1984, ex deer and cows, (Gerhardt 1986).
Culicoides stellifer was a humanbiting species collected during the later summer. Blanton & Wirth (1979) noted that there were no confirmed records of C. stellifer biting humans, as earlier records ( Snow et al 1957; Jamnback 1965) probably referred to C. paraensis . Our specimens verify the humanbiting habits of this species. Culicoides stellifer also feeds on deer and cows in the Park (Gerhardt 1986). Humphreys & Turner (1973) recorded the species from goats and turkey, and Hayes et al. (1984) recorded it from cattle.
Collection: Tennessee, Blount Co., Cades Cove, 1984, ex deer and cows, (Gerhardt 1986).
Culicoides venustus feeds on deer and cows in the Park (Gerhardt 1986). This species apparently has never been collected biting humans. Jamnback (1965) suggested that C. venustus is strictly mammalophilic due to the small number of sensilla coeloconica on its flagellum.
Collections: North Carolina, Deep Creek , 20 July 1999, light trap, coll. C. Parker and J. Cooper; Tennessee, Sevier Co., LeConte Creek at Twin Creeks, 12 July 1999, hand collection below dam, coll. J. Cooper.
A single specimen was collected in the Park while hovering around a human. Culicoides villosipennis feeds on humans and birds ( Hair & Turner 1968; Wright & DeFoliart 1970 b; Blanton & Wirth 1979). Jamnback (1965) suggests that this species is primarily ornithophilic because of the greater number of flagellar sensilla coeloconica.
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