Simuliidae - Black Flies, Buffalo Gnats

Black flies are common inhabitants of most flowing waters, with adult females often requiring an obligate blood meal for egg development, including man. Up to 54 species have been reported to bite humans (Adler, Currie and Wood 2004), and the prodigous numbers in which they emerge and bite make many species an annoying pest, and in some cases a health concern. In western Africa, black flies are associated with the transmission of a filarial onchocercid nematode that causes onchocerciasis (river blindness). Black flies also attack and affect the health of other mammals and birds, and are associated with the transmission of a variety of parasitic forms (reviewed by Adler, Currie and Wood 2004).

The adult is usually a small, dark fly with a characteristic humped thorax, whose appearance gives them the common name buffalo gnats. Black fly eggs, larvae, and pupae are aquatic, and almost exclusively lotic. Larvae, which orient themselves into the current, use a cephalic structure with fine fans to filter feed on organic matter suspended. Larvae remain attached to a substrate by using a posterior ringed structure that hooks onto a pad of silk built by the larva. Larvae also graze upon periphyton growing upon substrates. When food and competitive conditions are optimal, one can find larval densities that are quite staggering. Black flies pupate in a silken cocoon which is attached to a substrate.

The phylogeny, systematics and cytology, distribution, biology and ecology of the Nearctic species of Simuliidae north of Mexico has been superbly reviewed in a recent work by Adler, Currie and Wood (2004), and their classification scheme is followed. The Holarctic region contains two subfamilies, Parasimuliinae and Simuliinae. Species of the former are found in North America only in the western part of the continent. All species of black flies recorded in Michigan are classified in the subfamily Simuliinae, divided into two tribes: Prosimuliini (Twinnia, Helodon and Prosimulium) and Simuliini (Greniera, Stegopterna, Cnephia, Ectemnia, Metacnephia and Simulium). This arrangement differs from the organization found in Stone et al. (1965), which provided the systematic organization for the previous list.

Cnephia Enderlein, 1921 (Tribe Simuliini)

Cnephia dacotensis (Dyar and Shannon), 1927 Eusimulium - 1
      . Simulium lascivum Twinn, 1936
      • Cnephia 'sp.A' Pruess et al., 2000
Cnephia ornithophilia Davies, Peterson and Wood 1962 - 1
      • Cnephia '(near pecuarum)' Hoicking and Richards, 1952
      • Cnephia (Cnephia) 'U' Bennett, 1960

Ectemnia Enderlein, 1930 (Tribe Simuliini)

Ectemnia invenusta (Walker), 1848 Silmulium - Adler et al. 2004
      • Cnephia 'sp.1' Jamnback, 1951
      • Cnephia (Ectemnia) loisae Stone and Jamnback, 1955
Ectemnia taeniatifrons (Enderlein), 1925 Cnetha - 2

Greniera Doby and David, 1959 (Tribe Simuliini)

Greniera abdita (Peterson), 1962 Cnephia – ON (may be found in e. UP)
Greniera abditoides Wood, 1963 – ON (may be found in UP)
Greniera denaria Davies, Peterson, and Wood, 1962 – ON (may be found in UP)

Helodon Enderlein, 1921 (Tribe Prosimuliini) (=Prosimulium)

Helodon (Parahelodon) decemarticulatus (Twinn), 1936 Simulium (Prosimulium) – ON, WI (Adler et al. 2004, expect in the UP)
Helodon (Parahelodon) gibsoni (Twinn), 1936 Simulium - 1, Adler et al. 2004
Helodon (Parahelodon) vernalis (Shewell), 1952 Prosimulium – ON (se. LP of MI?)

Metacnephia Crosskey, 1969 (Tribe Simuliini)

Metacnephia saskatchewana (Shewell and Fredeen), 1958 Cnephia (widespread boreal, may be found in Isle Royale or the extreme northern UP)
      • Simulium 'sp.' Twinn et al., 1948
      • Cnephia 'N' Fredeen, 1951
      • Cnephia 'species N' Hocking and Pickering, 1954

Prosimulium Roubard, 1906 (Tribe Prosimuliini)

Prosimulium albionense Rothfels, 1956 - Adler et al. 2004
Prosimulium approximatum Peterson, 1970 - Adler et al. 2004
      • Prosimulium mysticum Peterson, 1970
Prosimulium arvum Adler and Kim, 1985 - Adler et al. 2004
      • Prosimulium hirtipes '4' Syme, 1957
Prosimulium canutum Adler, Currie & Wood 2004 - Adler et al. 2004
Prosimulium fontanum Syme and Davies, 1958 - Adler et al. 2004
Prosimulium fuscum Syme and Davies, 1958 - 1
Prosimulium mixtum Syme and Davies, 1958 - 1
Prosimulium multidentatum (Twinn), 1936 Simulium - 1

Simulium Latrielle, 1802 (Tribe Simuliini)

Simulium (Nevermannia) aestivum Davies, Peterson and Wood 1962 – Adler et al. 2004 (expect in UP)
      • Eusimulium 'Labrador’ Brockhouse, 1984
Simulium (Hellichiella) anatinum Wood, 1963 – Adler et al. 2004
      • Simulium (Eusimulium) ‘H’ Bennett, 1960
      • Cnephia « sp. ‘T’ Anderson & DeFoliart, 1961
Simulium (Boreosimulium) annulus (Lundström), 1911 Melusina – Adler et al. 2004
      • Simulium euryadminiculum Davies, 1949
Simulium (Simulium) appalachiense Adler, Currie and Wood 2004 – Adler et al. 2004
      • Simulium tuberosum ‘CDE sibling’ Landau, 1962
      • Simulium tuberosum ‘CDE-2’ Mason, 1982
      • Simulium ‘CDEM’ tuberosum Mason, 1984
Simulium (Eusimulium) bracteatum – Adler et al. 2004
      • Eusimulium aureum bracteatum Dyar and Shannon, 1927
      • Eusimulium aureum ‘Sibling A’ Dunbar, 1958
Simulium (Nevermannia) burgeri Adler, Currie and Wood 2004 – Adler et al. 2004 (expect in UP)
      • ‘undescribed species near S. fionae’ Currie, 1997
Simulium (Eusimulium) congareenarum (Dyar and Shannon), 1927 Eusimulium - ON, widespread east and Great Slave Lake (NWT)
Simulium (Simulium) conundrum Adler, Currie and Wood 2004 – Adler et al. 2004
      • Simulium tuberosum ‘FGH sibling’ Landau, 1962
Simulium (Nevermannia) craigi Adler and Currie, 1986 – Adler et al. 2004
      • Simulium (Eusimulium) vernum ‘complex’ Adler, 1983
      • Simulium (Eusimulium) vernum ‘cytotype A’ Rothfels and Brockhouse in Okaeme, 1983
      • Eusimulium vernum ‘Cypress Hills’ Brockhouse, 1984
      • Eusimulium vernum ‘Nipigon’ Brockhouse, 1984
      • Simulium ‘X’ Hunter, 1987
Simulium (Nevermannia) croxtoni Nicholson and Mickel, 1950 – Adler et al. 2004
      • Simulium (Eusimulium) ‘sp. E’ Shewell, 1957
Simulium (Simulium) decorum Walker, 1848 – Adler et al. 2004
      • Simulium venustum ‘var. piscicidium’ Johannsen, 1902
      • Simulium decorum katmai Dyar and Shannon, 1927
      • Simulium (Simulium) ottawaense Twinn, 1936
      • Simulium ‘C’ Fredeen, 1956
      • Simulium ‘sp. 1’ Anderson and Dicke, 1960
Simulium (Boreosimulium) emarginatum Davies, Peterson & Wood, 1962 – Adler et al. 2004 (Commencement Creek, MI, coll. Twinn, 1936)
      • Simulium ‘n. sp.’ Bennett, 1960
      • Simulium (Eusimulium) ‘sp. near emarginatum’ Adler, 1983
Simulium (Hellichiella) excisum Davies, Peterson & Wood, 1962 – 1, Adler et al. 2004
      • Simulium (Hellichiella) ‘sp. Near subexcisum’ Shewell, 1957
Simulium (Eusimulium) exulatum Adler, Currie and Wood 2004 – Adler et al. 2004 (expect in boreal UP)
      • Eusimulium aureum ‘Sibling C’ Dunbar, 1958
Simulium (Nevermannia) fionae Adler, 1990 – Adler et al. 2004
      • Eusimulium vernum ‘A-Type’ Brockhouse, 1984
      • Eusimulium vernum ‘Eastmain’ Brockhouse, 1985
      • Simulium (Nevermannia) ‘species near furculatum/croxtoni’ Adler and Kim, 1986
      • Simulium ‘sp.’ Hunter and Connolly, 1986
      • Simulium ‘sp. near furculatum/croxtoni’ Hunter, 1987
Simulium (Schoenbaueria) furculatum (Shewell), 1952 Eusimulium – Adler et al. 2004 (expect in boreal UP)
      • Eusimulium ‘species B’ Twinn et al., 1948
Simulium (Nevermannia) gouldingi – Adler et al. 2004
      • Simulium (Eusimulium) ‘species 1’ DeFoliart, 1951
      • ‘Sp. 27’ Jamnback, 1951
Simulium (Hellichiella) innocens (Shewell), 1952 Eusimulium – Adler et al. 2004
Simulium (Hemicnethia) innoxium Comstock and Comstock, 1895 – Adler et al. 2004
      • Schoenbaueria aldrichiana Enderlein, 1936
      • Simulium pictipes‘B’ Bedo, 1975
Simulium(Simulium) irritatum Lugger, 1897 – Adler et al. 2004 (expect in boreal UP)
      • Simulium venustum ‘CC9gB)B’ Rothfels, 1979
      • Simulium venustum ‘CC3’ Rothfels, 1981
Simulium (Simulium) jenningsi Malloch, 1914 – Adler et al. 2004
      • Simulium venustum var. a Johannsen, 1902
      • Simulium (Simulium) nigroparvum Twinn, 1936
Simulium (Boreosimulium) johannseni Hart in Forbes, 1912 – IL, MN, ON, WI, Adler et al. 2004 (expect in sw. LP and the UP)
      • Simulium ‘2’ Fredeen, 1956
      • Simulium (Hellichia) johannseni duplex Shewell & Fredeen, 1958
Simulium (Simulium) luggeri Nicholson and Mickel, 1950 – 1 (LP), 2, Adler et al. 2004
      • Simulium (Simulium) ‘L’ Fredeen, 1951
      • ‘species…resembling Simulium luggeri ‘ Davies et al., 1962
      • Simulium ‘n. sp. aff. luggeri’ Pistrang, 1983
Simulium (Byssodon) meridionale Riley, 1887 – WI, Adler et al. 2004 (expect in the w. UP)
      • Simulium occidentale Townsend, 1891
      • Simulium tamaulipense Townsend, 1897
      • Simulium forbesi Malloch, 1914
Simulium (Simulium) minutum Lugger, 1897 – Adler et al. 2004
      • Simulium venustum ‘CC1 (Ca 1 and CTC1)’ Rothfels et al., 1978
Simulium (Nevermannia) modicum Adler, Currie and Wood 2004 – Adler et al. 2004
      • Eusimulium pugetense ‘IL-st’ Choate, 1984
      • Eusimulium pugetense ‘cytotype A’ Hunter and Connolly, 1986
Simulium (Simulium) molestum Harris, 1841 – Adler et al. 2004
      • Simulium ‘A/C venustum’ Rothfels et al., 1978 (proposed synonym by Adler, Currie and Wood 2004)
Simulium (Simulium) murmanum Enderlein, 1935 – 2, Adler et al. 2004
      • Simulium ‘sp. O’ O’Kane, 1926
      • Simulium ‘sp. II’ Puri, 1926
      • Simulium (Simulium) corbis Twinn, 1936
      • Simulium (s. str.) relictum Rubtsov, 1940
      • Gnus forsi Carlsson, 1962
Simulium (Simulium) perissum Dyar and Shannon, 1927 – IL, IN, MN, OH, ON, WI - Adler et al. 2004
      • Simulium tuberosum ‘CKL-type’ Landau, 1962
      • Simulium (Simulium) tuberosum ‘Complex’ Preuss and Peterson 1987
Simulium (Hemicnethia) pictipes Hagen, 1880 – 2, Adler et al. 2004
      • Simulium ‘n. sp. nr. pictipes’ Shewell, 1957
      • Simulium (Hagenomyia) longistylatum Shewell, 1957
      • Simulium pictipes ‘n. sp. nr. pictipes’ Wolfe and Peterson, 1959
Simulium (Eusimulium) pilosum (Knowlton and Rowe), 1934 Eusimulium – Adler et al. 2004
      • Eusimulium utahense Knowlton and Rowe, 1934
      • Eusimulium aureum ‘Sibling B’ Dunbar, 1958
      • Simulium (Eusimulium) aureum ‘Complex’ Pruess and Peterson, 1987
Simulium (Nevermannia) quebecense Twinn, 1936 – Adler et al. 2004
Simulium (Simulium) rostratum (Lundström), 1911 Melusina – Adler et al. 2004
      • Simulium ‘sp.1’ Malloch, 1919
      • Simulium wilhelmii Enderlein, 1922
      • Simulium groenlandicum Enderlein, 1935
      • Simulium (Simulium) sublacustre Davis, 1966
      • Simulium ‘ACD verecundum’ Rothfels et al., 1978
Simulium (Simulium) rugglesi Nicholson and Mickel, 1950 – 2, Adler et al. 2004
      • Simulium (Simulium) ‘M’ Fredeen, 1951
Simulium (Nevermannia) silvestre (Rubtsov), 1956 Eusimulium – Adler et al. 2004
      • Eusimulium latipes ‘St. Rose’ Landau in Dunbar, 1962
      • Simulium (Eusimulium) vernum ‘complex’ Lake, 1980
      • Simulium (Eusimulium) vernum ‘cytotype B’ Rothfels and Brockhouse in Okaeme, 1983
      • Eusimulium vernum ‘Caledon’ Brockhouse, 1984
      • Simulium caledonense Adler and Currie, 1986
Simulium (Psilozia) tribulatum Lugger, 1897 – Adler et al. 2004
      • Simulium glaucum Coquillett, 1902
      • Simulium venustoides Hart in Forbes, 1912
      • Simulium vittatum ‘IIIL-1’ Rothfels and Featherston, 1981
      • Simulium ‘sp.’ Osborne et al., 1985
      • Simulium vittatum ‘cytotype IIIL-2’ Sutcliffe and McIver, 1987
      • ?Simulium ‘spp.’ Mullens and Dada, 1992 (in part)
Simulium (Simulium) truncatum (Lundström), 1911 Melusina – Adler et al. 2004
      • Simulium venustum ‘EFG/C (CEFG, CEF)’ Rothfels et al., 1978
      • ‘species near Simulium truncatum’ Gordon and Cupp, 1980
Simulium (Simulium) tuberosum s.s., (‘cytotype AB’) (Lundström), 1911 Melusina – 1, Adler et al. 2004
      • Simulium (Simulium) tuberosum ‘A sibling’ Wong, 1976
      • Simulium tuberosum ‘Y 2’ Mason, 1982
Simulium (Simulium) vandalicum Dyar and Shannon, 1927 – Adler et al. 2004
      • Simulium (Simulium) turmale Twinn, 1938
      • Eusimulium ‘species’ Frost, 1949
      • Simulium tuberosum ‘FG sibling’ Landau, 1962
      • Simulium (Simulium) tuberosum ‘B sibling’ Wong, 1976
      • Simulium tuberosum ‘s.l.’ Tang et al., 1996
Simulium (Simulium) venustum s.s. Say, 1823 – 1, Adler et al. 2004
      • Simulium sp. Forbes, 1912
      • Simulium ‘CC venustum’ Rothfels et al., 1987
Simulium (Simulium) verecundrum s.s., Stone and Jamnback, 1955 – Adler et al. 2004
      • Simulium ‘(probably venustum)’ Davies, 1971
      • Simulium ‘AA venustum (Transposition Sibling)’ Rothfels et al., 1978
      • Simulium ‘A/C verecundum (PS3)’ Rothfels et al., 1978
      • Simulium ‘AA-AC verecundum’ Gordon and Cupp, 1980
      • Simulium ‘CC ver.’ Rothfels., 1981
      • Simulium ‘verecundum ‘AA-A/C-CC’ Ciborowski and Adler, 1990
Simulium violator Adler, Currie and Wood 2004 – Adler et al. 2004
      • Eusimulium aureum ‘Sibling D’ Dunbar, 1958
Simulium (Psilozia) vittatum Zetterstedt, 1838 – Adler et al. 2004
      • Simulium vittata Zetterstedt, 1837 (nomen nudum)
      • Simulium nasale Gistel, 1848
      • Psilozia groenlandica Enderlein, 1936
      • Simulium asakakae Smart, 1944
      • ‘simuliid’ Craig, 1968
      • Simulium vittatum ‘IS-7’ Rothfels and Featherston, 1981
      • ?Simulium ‘sp.’ Grant, 1991

Stegopterna Enderlein, 1930 (Tribe Simuliini)

Stegopterna diplomutata Currie and Hunter, 2003 - ON, WI (expect in Dickinson Co. environs by border, also rest of UP)
      • Cnephia mutata '2n' Basrur, 1957
      • Cnephia mutata 'diploid' Basrur and Rothfels, 1959
Stegopterna emergens (Stone), 1952 Cnephia - Adler et al. 2004
      • Cnephia 'species #2' Anderson and Dicke, 1960
      • Stegopterna tshukotensis Rubtsov, 1971
Stegopterna mutata (Malloch), 1914 Prosimulium
      • Simulium 'species undescribed' Strickland, 1911
      • Simulium 'Undescribed species No.2' Ritcher, 1931
      • Simulium 'sp.A' Johannsen, 1934
      • Cnephia mutata 'triploid' Basrur and Rothfels, 1959
      • Stegopterna mutata '(IIIL-1 sibling)' Crosskey, 1990

Twinnia Stone and Jamnback, 1955 (Tribe Prosimuliini)

Twinnia tibblesi Stone and Jamnback, 1955 - northeastern

Page created: November 29, 2002 - Last updated: June 4, 2011 (EB)