Reeves, Will K., Adler, Peter H., Grogan, William L. & Super, Paul E., 2004, Hematophagous and Parasitic Diptera (Insecta) in the Great Smoky Mountains National Park, USA, Zootaxa 483, pp. 1-44: 7-12

publication ID

http://doi.org/ 10.5281/zenodo.157582

publication LSID


persistent identifier


treatment provided by


scientific name



Family Ceratopogonidae 

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)  .

Culicoides arboricola Root & Hoffman 

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.

Culicoides baueri Hoffman 

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.

Culicoides bickleyi Wirth & Hubert 

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).

Culicoides biguttatus Coquillett 

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).

Culicoides debilipalpis Lutz 

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.

Culicoides footei Wirth & Blanton 

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.

Culicoides guttipennis (Coquillett) 

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).

Culicoides obsoletus (Meigen) 

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.

Culicoides paraensis (Goeldi) 

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).

Culicoides sanguisuga (Coquillett) 

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.

Culicoides snowi Wirth & Jones 

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).

Culicoides spinosus Root & Hoffman 

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.

Culicoides stellifer Coquillett 

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.

Culicoides venustus Hoffman 

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.

Culicoides villosipennis Root and Hoffman 

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.













Reeves, Will K., Adler, Peter H., Grogan, William L. & Super, Paul E. 2004

Simuliidae (

Bain & Chaubaud 1986