Aquatic Insects of Michigan

by Ethan Bright, Museum of Zoology Insect Division and School of Natural Resources and Environment
University of Michigan

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.

As a consequence of the transmission of disease and their biting habitats, the biology, ecology and taxonomy of Simuliidae are well-studied, and our regional inventory is fairly well documented. The phylogeny, systematics and cytology, distribution, biology and ecology of the Nearctic species of Simuliidae north of Mexico has been superbly reviewed 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 on this website.

Currently, there are 51 species in genera are denoted in 13 genera, with another 16 species and 2 genera that may be encountered based on their known regional distribution. Genera are denoted in green; species denoted in bold blue have been recorded in Michigan; other species denoted in black bold are expected to occur based on their known distribution. You can use the Find function in your web browser to locate the desired taxon or author. Synonomous species (indented) are listed below the current valid species (denoted in bold) with its taxonomic status indicated.

Cnephia Enderlein, 1921: 199 (Tribe Simuliini)
Cnephia dacotensis (Dyar & Shannon, 1927: 20 as Eusimulium) - Merritt et al. 1978
(syn.) Simulium lascivum Twinn, 1936: 127
(syn.) Cnephia 'sp. A' Pruess et al. 2000: 287
Cnephia ornithophilia Davies, Peterson & Wood 1962: 102 - Merritt et al. 1978
(syn.) Cnephia '(near pecuarum)' Hocking & Richards, 1952: 241
(syn.) Cnephia (Cnephia) 'U' Bennett, 1960: 380

Ectemnia Enderlein, 1930: 88 (Tribe Simuliini)
Ectemnia invenusta (Walker, 1848: 112 as Simulium) - Adler et al. 2004
(syn.) '? Nov. sp.' Jamnback, 1951: 3
(syn.) Cnephia 'sp.1' Jamnback, 1953: 43
(syn.) Cnephia (Ectemnia) loisae Stone & Jamnback, 1955: 35
Ectemnia taeniatifrons (Enderlein, 1925: 206 as Cnetha) - Stone 1964

Greniera Doby & David, 1959: 763 (Tribe Simuliini)
Greniera abdita (Peterson, 1962: 96 as Cnephia) - ON (may be found in e. UP)
Greniera abditoides Wood, 1963: 95 - ON (may be found in UP)
Greniera denaria Davies, Peterson, & Wood, 1962: 97 - ON (may be found in UP)
Greniera michiganensis Taber, 2008: 223 - Taber, 2008: 223

Helodon Enderlein, 1921: 122 (Tribe Prosimuliini)
Helodon (Parahelodon) decemarticulatus (Twinn, 1936: 110 as Simulium) - ON, WI (Adler et al. 2004, expect in the UP)
Helodon (Parahelodon) gibsoni (Twinn, 1936: 108 as Simulium) - Merritt et al. 1978, Adler et al. 2004
Helodon (Parahelodon) vernalis (Shewell, 1952: 33 as Prosimulium) - ON (se. LP of MI?)

Metacnephia Crosskey, 1969: 26 (Tribe Simuliini)
Metacnephia saskatchewana (Shewell & Fredeen, 1958: 733 as Cnephia) - widespread boreal, expect on Isle Royale or extreme n. UP)
(syn.) Simulium 'sp.' Twinn et al. 1948: 351
(syn.) Cnephia 'N' Fredeen, 1951: 11
(syn.) Cnephia 'species N' Hocking & Pickering, 1954: 100

Prosimulium Roubard, 1906: 519 (Tribe Prosimuliini)
Prosimulium albionense Rothfels, 1956: 119 - Adler et al. 2004
Prosimulium approximatum Peterson, 1970b: 74 - Adler et al. 2004
(syn.) Prosimulium mysticum Peterson, 1970b:
Prosimulium arvum Adler & Kim, 1985: 41 - Adler et al. 2004
(syn.) Prosimulium hirtipes '4' Syme, 1957: 51
Prosimulium canutum Adler, Currie & Wood 2004: 278 - Adler et al. 2004
Prosimulium fontanum Syme & Davies, 1958: 708 - Adler et al. 2004
(syn.) Prosimulium '16' Rothfels, 1956: 114
(syn.) Prosimulium hirtipes '3' Syme & Davies, 1958: 700
Prosimulium fuscum Syme & Davies, 1958: 702 - Merritt et al. 1978
(syn.) Prosimulium hirtipes '1' Rothfels, 1956: 114
(syn.) Prosimulium (Prosimulium) mysticum Peterson, 1970b: 122
Prosimulium mixtum Syme & Davies, 1958: 706 - Merritt et al. 1978
(nom. nud.) Simulium calceatum Say, in Harris 1835: 595
(syn.) Prosimulium hirtipes '2' Rothfels, 1956: 114

Prosimulium multidentatum (Twinn, 1936: 106 as Simulium) - Merritt et al. 1978

Simulium Latrielle, 1802: 426 (Tribe Simuliini)
Simulium (Nevermannia) aestivum Davies, Peterson & Wood 1962: 104 - Adler et al. 2004 (expect in UP)
(syn.) Eusimulium 'Labrador' Brockhouse, 1984: 42
Simulium (Hellichiella) anatinum Wood, 1963: 96 - Adler et al. 2004: 307
(syn.) Simulium (Eusimulium) ‘H’ Bennett, 1960: 379
(syn.) Cnephia sp. ‘T’ Anderson & DeFoliart, 1961: 719
Simulium (Boreosimulium) annulus (Lundström, 1911: 17 as Melusina)- Adler et al. 2004: 314
(syn.) Simulium euryadminiculum Davies, 1949: 45
Simulium (Simulium) appalachiense Adler, Currie & Wood 2004: 413 - Adler et al. 2004
(syn.) Simulium tuberosum ‘CDE sibling’ Landau, 1962: 928
(syn.) Simulium tuberosum ‘CDE-2’ Mason, 1982: 293
(syn.) Simulium ‘CDEM’ tuberosum Mason, 1984: 654
Simulium (Eusimulium) bracteatum Coquillett, 1898: 69 - Adler et al. 2004: 328
(comb.) Eusimulium aureum bracteatum Dyar & Shannon, 1927: 14
(syn.) Eusimulium aureum ‘Sibling A’ Dunbar, 1958: 23
Simulium (Nevermannia) burgeri Adler, Currie & Wood 2004: 333 - Adler et al. 2004 (expect in UP)
(syn.) ‘undescribed species near S. fionae’ Currie, 1997: 579
Simulium (Hellichiella) congareenarum (Dyar & Shannon, 1927: 20 as Eusimulium) - ON (one record from Bruce Peninsula), east to Great Slave Lake (NWT); Adler et al. 2004: 308
(syn.) Eusimulium congareenarum 'b' Dunbar 1967: 387
Simulium (Simulium) conundrum Adler, Currie & Wood 2004: 415 - Adler et al. 2004
(syn.) Simulium tuberosum ‘FGH sibling’ Landau, 1962: 928
Simulium (Nevermannia) craigi Adler & Currie, 1986: 1208 - Adler et al. 2004
(syn.) Simulium (Eusimulium) vernum ‘complex’ Adler, 1983: 203
(syn.) Simulium (Eusimulium) vernum‘cytotype A’ Rothfels & Brockhouse, in Okaeme, 1983: 57
(syn.) Eusimulium vernum ‘Cypress Hills’ Brockhouse, 1984: 28
(syn.) Eusimulium vernum ‘Nipigon’ Brockhouse, 1984: 33
(syn.) Simulium ‘X’ Hunter, 1987: 64
Simulium (Nevermannia) croxtoni Nicholson & Mickel, 1950: 41- Adler et al. 2004
(syn.) Simulium (Eusimulium) ‘sp. E’ Shewell, 1957: 2
Simulium (Simulium) decorum Walker, 1848: 112 - Adler et al. 2004
(syn.) Simulium venustum ‘var. piscicidium’ Johannsen, 1902: 48
(syn.) Simulium decorum katmai Dyar & Shannon, 1927: 31
(syn.) Simulium (Simulium) ottawaense Twinn, 1936: 146
(syn.) Simulium ‘C’ Fredeen, 1956: 4
(syn.) Simulium ‘sp. 1’ Anderson & Dicke, 1960: 397
Simulium (Boreosimulium) emarginatum Davies, Peterson & Wood, 1962: 110 - Adler et al. 2004: 317 (Commencement Creek, MI, coll. Twinn, 1936)
(syn.) Simulium ‘n. sp.’ Bennett, 1960: 380
(syn.) Simulium (Eusimulium) ‘sp. near emarginatum’ Adler, 1983: 205
Simulium (Hellichiella) excisum Davies, Peterson & Wood, 1962: 113 - Merritt et al. 1978, Adler et al. 2004: 312
(syn.) Simulium (Hellichiella) ‘sp. Near subexcisum’ Shewell, 1957: 2
Simulium (Eusimulium) exulatum Adler, Currie & Wood 2004: 329- Adler et al. 2004 (expect in boreal UP)
(syn.) Eusimulium aureum ‘Sibling C’ Dunbar, 1958: 23, 38
Simulium (Nevermannia) fionae Adler, 1990: 431 - Adler et al. 2004
(syn.) Eusimulium vernum ‘A-Type’ Brockhouse, 1984: 40
(syn.) Eusimulium vernum ‘Eastmain’ Brockhouse, 1985: 2148
(syn.) Simulium (Nevermannia) ‘species near furculatum/croxtoni’ Adler & Kim, 1986: 29
(syn.) Simulium ‘sp.’ Hunter & Connolly, 1986: 300
(syn.) Simulium ‘sp. near furculatum/croxtoni’ Hunter, 1987: 52
Simulium (Schoenbaueria) furculatum (Shewell, 1952: 40 as Eusimulium) - Adler et al. 2004 (expect in boreal UP)
(syn.) Eusimulium ‘species B’ Twinn et al., 1948: 353
Simulium (Nevermannia) gouldingi Stone, 1952: 90 - Adler et al. 2004
(syn.) Simulium (Eusimulium) ‘species 1’ DeFoliart, 1951: 67
(syn.) ‘Sp. 27’ Jamnback, 1951: 3
Simulium (Hellichiella) innocens (Shewell, 1952: 38 as Eusimulium) - Adler et al. 2004: 308
Simulium (Hemicnethia) innoxium Comstock & Comstock, 1895: 452 - Adler et al. 2004
(syn.) Schoenbaueria aldrichiana Enderlein, 1936: 120
(syn.) Simulium pictipes‘B’ Bedo, 1975: 1147
(syn.) Simulium "sp." Riley, 1887: 592
(syn.) Simulium innoxium Williston, in Phillips 1890: 21 [unavail., unpubl. thesis name]
Simulium (Simulium) irritatum Lugger, 1897: 203- Adler et al. 2004 (expect in boreal UP)
(syn.) Simulium venustum ‘CC9gB)B’ Rothfels, 1979: 526
(syn.) Simulium venustum ‘CC3’ Rothfels, 1981: 27
Simulium (Simulium) jenningsi Malloch, 1914: 41 - Adler et al. 2004
(syn.) Simulium venustum var. a Johannsen, 1903: 381
(syn.) Simulium (Simulium) nigroparvum Twinn, 1936: 142
Simulium (Boreosimulium) johannseni Hart, in Forbes, 1912: 31 - IL, MN, ON, WI, Adler et al. 2004: 321 (expect in sw. LP and the UP)
(syn.) Simulium ‘2’ Fredeen, 1956: 4
(syn.) Simulium (Hellichia) johannseni duplex Shewell & Fredeen, 1958: 734
Simulium (Simulium) luggeri Nicholson & Mickel, 1950: 54 - Merritt et al. 1978 (LP), 2, Adler et al. 2004
(syn.) Simulium (Simulium) ‘L’ Fredeen, 1951: 12, 25
(syn.) ‘species…resembling Simulium luggeri‘ Davies et al., 1962: 81
(syn.) Simulium ‘n. sp. aff. luggeri’ Pistrang, 1983: 33
Simulium (Byssodon) meridionale Riley, 1887: 513 - WI, Adler et al. 2004: 324 (expect in the w. UP)
(syn.) Simulium occidentale Townsend, 1891: 107
(syn.) Simulium tamaulipense Townsend, 1897: 171
(syn.) Simulium forbesi Malloch, 1914: 63
Simulium (Simulium) minutum Lugger, 1897: 201 - Adler et al. 2004
(syn.) Simulium venustum ‘CC1 (Ca 1 and CTC1)’ Rothfels et al. 1978: 1118
Simulium (Nevermannia) modicum Adler, Currie & Wood 2004: 342 - Adler et al. 2004
(syn.) Eusimulium pugetense ‘IL-st’ Choate, 1984: 60
(syn.) Eusimulium pugetense ‘cytotype A’ Hunter & Connolly, 1986: 302
Simulium (Simulium) molestum Harris, 1841: 405 - Adler et al. 2004
(syn.) Simulium ‘A/C venustum’ Rothfels et al. 1978: 1119
Simulium (Simulium) murmanum Enderlein, 1935: 363 - Stone 1964, Adler et al. 2004
(syn.) Simulium ‘sp. O’ O'Kane, 1926: 21
(syn.) Simulium ‘sp. II’ Puri, 1926: 164
(syn.) Simulium (Simulium) corbis Twinn, 1936: 147
(syn.) Simulium (s. str.) relictum Rubtsov, 1940: 425
(syn.) Gnus forsi Carlsson, 1962: 90
Simulium (Simulium) perissum Dyar & Shannon, 1927: 43 - IL, IN, MN, OH, ON, WI - Adler et al. 2004
(syn.) Simulium tuberosum ‘CKL-type’ Landau, 1962: 928
(syn.) Simulium (Simulium) tuberosum ‘Complex’ Preuss & Peterson, 1987: 532
Simulium (Hemicnethia) pictipes Hagen, 1880: 305 - Stone 1964, Adler et al. 2004
(syn.) Simulium ‘n. sp. nr. pictipes’ Shewell, 1957: 2
(syn.) Simulium (Hagenomyia) longistylatum Shewell, 1959: 84
(syn.) Simulium pictipes ‘n. sp. nr. pictipes’ Wolfe & Peterson, 1959: 148
Simulium (Eusimulium) pilosum (Knowlton & Rowe, 1934: 580 as Eusimulium) - Adler et al. 2004
(syn.) Eusimulium utahense Knowlton & Rowe, 1934: 582
(syn.) Eusimulium aureum ‘Sibling B’ Dunbar, 1958: 23
(syn.) Simulium (Eusimulium) aureum ‘Complex’ Pruess & Peterson, 1987: 529
Simulium (Nevermannia) quebecense Twinn, 1936: 117 - Adler et al. 2004
Simulium (Simulium) rostratum (Lundström, 1911: 13 as Melusina) - Adler et al. 2004
(syn.) Simulium ‘sp.1’ Malloch, 1919: 42
(syn.) Simulium wilhelmii Enderlein, 1922: 74
(syn.) Simulium groenlandicum Enderlein, 1935: 363
(syn.) Simulium (Simulium) sublacustre Davis, 1966: 491
(syn.) Simulium ‘ACD verecundum’ Rothfels et al. 1978: 1122
Simulium (Simulium) rugglesi Nicholson & Mickel, 1950: 60 - Stone 1964, Adler et al. 2004
(syn.) Simulium (Simulium) ‘M’ Fredeen, 1951: 11, 25
Simulium (Nevermannia) silvestre (Rubtsov, 1956: 433 as Eusimulium) - Adler et al. 2004
(syn.) Eusimulium latipes ‘St. Rose’ Landau, in Dunbar, 1962: 31
(syn.) Simulium (Eusimulium) vernum ‘complex’ Lake, 1980: 37
(syn.) Simulium (Eusimulium) vernum ‘cytotype B’ Rothfels & Brockhouse, in Okaeme, 1983: 57
(syn.) Eusimulium vernum ‘Caledon’ Brockhouse, 1984: 35
(syn.) Simulium caledonense Adler & Currie, 1986: 1212
Simulium (Psilozia) tribulatum Lugger, 1897: 205 - Adler et al. 2004
(syn.) Simulium glaucum Coquillett, 1902: 97
(syn.) Simulium venustoides Hart in Forbes, 1912: 42
(syn.) Simulium vittatum ‘IIIL-1’ Rothfels & Featherston, 1981: 1852
(syn.) Simulium ‘sp.’ Osborne et al., 1985: 158
(syn.) Simulium vittatum ‘cytotype IIIL-2’ Sutcliffe & McIver, 1987: 13
(syn.) ?Simulium ‘spp.’ Mullens & Dada, 1992: 478 (in part)
Simulium (Simulium) truncatum (Lundström, 1911: 13 as Melusina) - Adler et al. 2004
(syn.) Simulium venustum ‘EFG/C (CEFG, CEF)’ Rothfels et al. 1978: 117
(syn.) ‘species near Simulium truncatum’ Gordon & Cupp, 1980: 975
Simulium (Simulium) tuberosum s.s., (‘cytotype AB’) (Lundström, 1911: 14 as Melusina) - Merritt et al. 1978, Adler et al. 2004
(syn.) Simulium '(s. str) sp. indet. Edwards 1933: 614
(syn.) ?Simulium 'species 2' Jenkins, 1948: 149
(syn.) Simulium (Simulium) tuberosum ‘A sibling’ Wong, 1976: 40
(syn.) Simulium tuberosum ‘A sibling’ Mason, 1982: 293
(syn.) Simulium tuberosum ‘Y2’ Mason, 1982: 295
Simulium (Simulium) vandalicum Dyar & Shannon, 1927: 44 - Adler et al. 2004
(syn.) Simulium (Simulium) turmale Twinn, 1938: 51
(syn.) Eusimulium ‘species’ Frost, 1949: 129
(syn.) Simulium tuberosum ‘FG sibling’ Landau, 1962: 928
(syn.) Simulium (Simulium) tuberosum ‘B sibling’ Wong, 1976: 41
(syn.) Simulium tuberosum ‘s.l.’ Tang et al. 1996: 40
Simulium (Simulium) venustum s.s. Say, 1823: 28 - Merritt et al. 1978, Adler et al. 2004
(syn.) Simulium sp. Forbes, 1912: 23
(syn.) Simulium ‘CC venustum’ Rothfels et al., 1987: 1118
Simulium (Simulium) verecundrum s.s., Stone & Jamnback, 1955: 83 - Adler et al. 2004
(syn.) Simulium ‘(probably venustum)’ Davis, 1971: 333
(syn.) Simulium ‘AA venustum (Transposition Sibling)’ Rothfels et al. 1978: 1121
(syn.) Simulium ‘A/C verecundum (PS3)’ Rothfels et al. 1978: 1122
(syn.) Simulium ‘AA-AC verecundum’ Gordon & Cupp, 1980: 975
(syn.) Simulium ‘CC ver.’ Rothfels 1981: 26
(syn.) Simulium ‘verecundum ‘AA-A/C-CC’ Ciborowski & Adler, 1990: 2115
Simulium (Eusimulium) violator Adler, Currie & Wood 2004: 330 - Adler et al. 2004
(syn.) Eusimulium aureum ‘Sibling D’ Dunbar, 1958: 23, 38
Simulium (Psilozia) vittatum Zetterstedt, 1838: 803 - Adler et al. 2004
(nom. nud.) Simulium vittata Zetterstedt, 1837: 58
(syn.) Simulium nasale Gistel, 1848: 151
(syn.) Psilozia groenlandica Enderlein, 1936: 114
(syn.) Simulium asakakae Smart, 1944: 131
(syn.) ‘simuliid’ Craig, 1968: 31
(syn.) Simulium vittatum ‘IS-7’ Rothfels & Featherston, 1981: 1858
(syn.) ?Simulium ‘sp.’ Grant, 1991: 436

Stegopterna Enderlein, 1930: 89 (Tribe Simuliini)
Stegopterna diplomutata Currie & Hunter, 2003: 1 - ON, WI (expect in Dickinson Co. environs by border, also rest of UP)
(syn.) Cnephia mutata '2n' Basrur, 1957: 4
(syn.) Cnephia mutata 'diploid' Basrur & Rothfels, 1959: 571
Stegopterna emergens (Stone, 1952: 80 as Cnephia) - Adler et al. 2004
(syn.) Cnephia 'species #2' Anderson & Dicke, 1960: 395
(syn.) Stegopterna tshukotensis Rubtsov, 1971: 174
Stegopterna mutata (Malloch, 1914: 20 as Prosimulium) - Adler et al. 2004
(syn.) Simulium 'species undescribed' Strickland, 1911: 321
(syn.) Simulium 'Undescribed species No.2' Ritcher, 1931: 242
(syn.) Simulium 'sp.A' Johannsen, 1934: 64
(syn.) Cnephia mutata 'triploid' Basrur & Rothfels, 1959: 571
(syn.) Stegopterna mutata '(IIIL-1 sibling)' Crosskey, 1990: 288

Twinnia Stone & Jamnback, 1955: 18 (Tribe Prosimuliini)
Twinnia tibblesi Stone & Jamnback, 1955: 19 - northeastern

References - Distribution

Adler PH, Currie DC, Wood DW. 2004. The black flies (Simuliidae) of North America. Cornell University Press: Ithaca, New York. xv + 941 p.
Taber SW. 2008. A new species of black fly (Diptera: Simuliidae) from Michigan. Southwestern Entomologist 33(3): 223-236.
Merritt RW, Ross DH, Peterson BV. 1978. Larval ecology of some lower michigan black flies (Diptera: Simuliidae) with keys to the immature stages. The Great Lakes Entomologist 11(4):177-208.
Stone A. 1964. Guide to the insects of Connecticut. Part VI. The Diptera or true flies of Connecticut, ninth fascicle. Simuliidae and Thaumaleidae. State Geological and Natural History Survey of Connecticut Bulletin 97. vii + 126 pp.

References - Taxonomy

Adler PH. 1983. Ecosystematics of the Simulium vittatum complex (Diptera: Simuliidae) and associated black flies. Ph.D. thesis. Pennsylvania State University, University Park. 308 pp.
Adler PH. 1990. Two new species of black flies (Diptera: Simuliidae) from North America. Proceedings of the Entomological Society of Washington 92: 431-443.
Adler PH, Currie DC. 1986. Taxonomic resolution of three new species near Simulium vernum Macquart (Diptera: Simuliidae). The Canadian Entomologist 118: 1207-1220.
Adler PH, Kim KC. 1986. The black flies of Pennsylvania (Simuliidae, Diptera): bionomics, taxonomy, and distribution. Pennsylvania State University Agricultural Experiment Station Bulletin 856: 1-88.
Adler PH, Currie DC, Wood DW. 2004. The black flies (Simuliidae) of North America. Cornell University Press: Ithaca, New York. xv + 941 pp.
Anderson JR, Dicke RJ. 1960. Ecology of the immature stages of some Wisconsin black flies (Simuliidae: Diptera). Annals of the Entomological Society of America 53: 386-404.
Anderson JR, DeFoliart GR. 1961. Feeding behavior and host preferences of some black flies (Diptea: Simuliidae) in Wisconsin. Annals of the Entomological Society of America 54: 716-729.
Basnur VR. 1957. Cytotaxonomic studies in black flies and midges. Ph.D. thesis. University of Toronto, Toronto. 49 pp. + 8 pls.
Basnur VR, Rothfels KH. 1959. Triploidy in natural populations of the black fly Cnephia mutata (Malloch). Canadian Journal of Zoology 37: 571-589.
Bedo DG. 1975. Polytene chromosomes of three species of blackflies in the Simulium pictipes group (Diptera: Simuliidae). Canadian Journal of Zoology 53: 1147-1164.
Bennett GF. 1960. On some ornithophilic blood-sucking Diptera in Algonquin Park, Ontario, Canada. Canadian Journal of Zoology 38: 377-389.
Brockhouse CL. 1984. Sibling species and sex chromosomes in Eusimulium vernum (Diptera: Simuliidae). M.S. thesis. University of Toronto, Toronto. 94 pp.
Brockhouse CL. 1985. Sibling species and sex chromosomes in Eusimulium vernum (Diptera: Simuliidae). Canadian Journal of Zoology 63: 2145-2161.
Carlsson G. 1962. Studies on Scandinavian black flies (Fam. Simuliidae Latr.). Opuscula Entomologica 21 (Suppl.): 1-280.
Comstock JH, Comstock AB. 1895. A manual for the study of insects. Comstock Publishing, Ithaca, N.Y. 701 pp.
Coquillett DW. 1898. The buffalo-gnats, or black-flies, of the United States. United States Department of Agriculture Division of Entomology Bulletin (New Series) 10: 66-69.
Coquillett DW. 1902. New Diptera from North America. Proceedings of the United States National Museum 25(1280): 83-126.
Crosskey RW. 1969. A re-classification of the Simuliidae (Diptera) of Africa and its islands. Bulletin of the British Museum (Natural History). Entomology Supplement 14: 1-195.
Crosskey RW. 1990. The natural history of blackflies. John Wiley, Chichester, U.K. 711 pp.
Choate LA. 1984. A cytological description of Eusimulium vernum, E. X, and the E. pugetense complex (Diptera: Simuliidae) in Alaska. M.S. thesis. University of Alaska, Fairbanks. 122 pp.
Craig DA. 1968. The clarification of a discrepancy in descriptions of maxillary musculature in larval Simuliidae. Quaestiones Entomologicae 4: 31-32.
Currie DC. 1997. Black flies (Diptera: Simuliidae) of the Yukon, with reference to the black-fly fauna of northwestern North America, pp. 563-614 in Danks HV, Downes JA (eds.), Insects of the Yukon. Biological Survey of Canada (Terrestrial Arthropods), Ottawa. 1034 pp.
Currie DC, Hunter FF. 2003. A new species of Stegopterna Enderlein and its relationship to the allotripoid species St. mutata (Malloch 1914). (Diptera: Simuliidae). Zootaxa 214: 1-11.
Davies DM. 1949. Description of Simulium euryadminiculum, a new species of blackfly (Simuliidae: Diptera). The Canadian Entomologist 81: 45-49.
Davies DM, Peterson BV, Wood DM. 1962. The black flies (Diptera: Simuliidae) of Ontario. Part I. Adult identification and distribution with descriptions of six new species. Proceedings of the Entomology Society of Ontario (1961) 92: 69-154.
Davis L. 1966. The taxonomy of British black-flies (Diptera: Simuliidae). Transactions of the Royal Entomological Society of London 118: 413-511.
Davis CC. 1971. A study of the hatching process in aquatic invertebrates. XXV. Hatching in the blackfly, Simulium (probably venustum) (Diptera, Simuliidae). Canadian Journal of Zoology 49: 333-336.
DeFoliart GK. 1951. The life histories, identification and control of blackflies (Diptera: Simuliidae) in the Andirondack mountains. Ph.D. thesis. Cornell University, Ithaca, N.Y. 98 pp.
Doby J-M, David F. 1959. Greniera, genre nouveau de Simuliidé (Diptères-Nématocères). Discussion de la postion systèmatique. Comptes Rendus Hebdomadaires des Séances de l'Académie des Sciences 249: 763-765.
Dunbar RW. 1958. The salivary gland chromosomes of two sibling species of black flies included in Eusimulium aureum Fries. Canadian Journal of Zoology 36: 23-44.
Dunbar RW. 1962. Cytotaxonomic studies in Simuliidae. Ph.D. thesis. University of Toronto, Toronto. 101 pp.
Dunbar RW. 1967. The salivary gland chromosomes of six closely related black flies near Eusimulium congareenarum (Diptera: Simuliidae). Canadian Journal of Zoology 45: 377-396.
Dyar HG, Shannon RC. 1927. The North American two-winged flies of the family Simuliidae. Proc. U.S. Natl. Mus. 69 (10) [#2636], 54 pp, 7 pls.
Edwards FW. 1933. Oxford University Expedition to Hudson's Strait, 1931: Diptera Nematocera. With notes on some other species of the genus Diamesa. Annals and Magazine of Natural History (Series 10) 12: 611-620.
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