taxonID	type	description	language	source
175587A7FFEEFFD5FF3BFC84FC5A45C2.taxon	diagnosis	Diagnosis. A medium to large species of the C. nubeculosus - stigma complex most closely resembling C. riethi, but differing from that and other Nearctic species in this complex by the following combination of characters: male wing length 1.22 – 1.39 mm, female wing length 1.57 – 1.97 mm; wing membrane and veins dusky brown with indistinct pale spots just beyond apex of costa and over r-m crossvein, and pale streaks in mid portion of cell m 2 and posterior to base of vein CuA 1; femora, tibiae usually without distinct pale bands, fore tibia with poorly developed subapical pale band in some specimens. Male: gonocoxite and gonostylus very short; tergite 9 greatly tapered distally with elongate, conical, divergent apicolateral processes; aedeagus with broad distal portion and rounded apex; parameres fused proximally with bifurcate distal portion. Female: mandible usually with 2 – 7 small teeth on inner margin (a few specimens without teeth); lacinia without teeth; palpal segment 3 very short (palpal ratio 1.46 – 2.17); spermatheca elongate with numerous tiny hyaline punctations and a broad opening. Male. Head: (Fig. 1) Brown. Eyes widely separated by diameter of 4 – 5 ommatidia, with sparse minute inter-ommatidial spicules. Antennal flagellum with flagellomeres 1, 11 – 13 separate, 2 – 10 completely or partially fused; flagellomeres 11 – 13 considerably longer than 2 – 10; all flagellomeres covered with short, fine setae; flagellomeres 1, 5 – 8 with single apical sensilla coeloconica (rarely on 5 or 9; one specimen with 2 sensilla on 8); flagellomere 1 with 2 rows, 2 – 13 with single row of sensilla chaetica, those on 1 – 10 comprising the moderately long, sparse plume; combined total lengths of flagellomeres 0.707 (0.666 – 0.747, n = 14) mm; antennal ratio 0.65 (0.62 – 0.69, n = 14). Palpus similar to female; segment 3 very short, moderately swollen, with well-developed shallow mesoapical pit bearing numerous long capitate sensilla that extend well beyond pit opening; palpal ratio 1.75 (1.46 – 2.00, n = 14). Proboscis short, mandible and lacinia vestigial, without teeth. Thorax: (Fig. 2) Dark brown to golden brown; scutellum paler with 2 marginal, 3 – 4 centrally located large setae. Legs brown, similar to female; claws smaller, gently curved with bent, deeply bifid tips. Wing (Fig. 3) shorter, narrower than female; wing membrane covered with dense, coarse microtrichia; macrotrichia restricted to costa, radius, moderately dense on anterior margin of cell r 3, few or absent on central portions of cells m 1, m 2, sparse along apical margins of m 1, m 2; veins infuscated brown, distal portion of costa, 2 nd radial cell darker; indistinct pale spots just beyond apex of costa and over r-m crossvein, pale streaks in cells m 1, m 2; wing length 1.31 (1.22 – 1.39, n = 18) mm; costal ratio 0.55 (0.52 – 0.57, n = 17). Halter pale brown. Abdomen: Dark brown to medium brown. Genitalia (Fig. 4) dark brown. Tergite 9 abruptly tapering distally to narrow, rounded apex with narrow, deep, V-shaped median notch; apicolateral process elongate, conical, tapered distally with single short apical seta, apices divergent; cercus large, setose, extending to mid-length of apicolateral process. Sternite 9 nearly 3 x broader than long, anterior margin slightly curved, posterior margin with moderately deep V- or U-shaped excavation. Gonocoxite nearly straight, stout, slightly longer than broad with well-developed, heavily sclerotized mesobasal tubercle, apex broad; dorsal root heavily sclerotized, extending to or near base of parameres. Gonostylus as long as gonocoxite; stout proximally, curved, tapering at mid-length to moderately slender apex, tip pointed. Parameres Y-shaped, heavily sclerotized, fused basally; basal arm stout, broad, slightly curved; extreme base tapered, apex slightly pointed; distal portion deeply bifurcate, each half slender, well separated with narrow sharply pointed apex that extends below level of apex of aedeagus. Aedeagus heavily sclerotized on basal arm and mid portion; basal arm stout, recurved nearly 90 ◦, apex rounded; basal arch U-shaped, extending 0.35 – 0.40 of total length; distal portion more lightly sclerotized, heaviest on margins, expanded distally to broadly rounded apex, tips of margins ventrally directed and extend just beyond apex. Female. Head: (Fig. 5) Brown. Eyes moderately separated by diameter of 3 – 4 ommatidia; with sparse fine inter-ommatidial spicules. Antennal flagellum with flagellomeres 9 – 13 slightly to considerably longer than 2 – 8, 13 longest; flagellomeres 1, 6 – 8 (rarely 5 – 8 or 6 – 9) with single apical sensilla coeloconica, rarely with 2 sensilla, sensilla rarely absent on 6 or 7; flagellomeres covered with short, fine setae and single whorl of moderately long sensilla chaetica; total combined lengths of flagellomeres 0.640 (0.594 – 0.699, n = 17) mm; antennal ratio 0.85 (0.80 – 0.91, n = 17). Palpus 5 - segmented; segment 3 very short, swollen, with large shallow subapical pit bearing numerous long capitate sensilla that extend well beyond pit opening; palpal ratio 1.79 (1.46 – 2.17, n = 17). Proboscis short, proboscis / head ratio 0.65 (0.60 – 0.70, n = 12); mandible with 2 – 7 small apical teeth on inner margin, rarely without teeth; lacinia vestigial, without teeth. Thorax: (Fig. 6) Mesonotum with scutum medium dark brown to golden brown with 2 narrow to moderately broad submarginal golden stripes; humerus, lateral margins golden; scutellum light brown or golden with 7 – 13 large setae arranged more or less linearly along mid-line; postscutellum medium brown to golden brown; pleural regions light brown to golden; katepisternum medium brown. Legs medium to light brown; femora, tibiae generally without distinct pale bands, fore tibia with poorly developed subapical pale band in some specimens; femorotibial joints dark brown; femora, tibiae with sparse coarse setae, most numerous dorsally; tarsomeres 1 – 3 with 2 apical spines; hind tibial comb with 6 spines, 2 nd longest, slightly longer than 3 rd, spur massive, bifurcate; claws small, curved distally, tips pointed, entire. Wing (Fig. 7) membrane covered with dense, coarse microtrichia; macrotrichia sparse, longest on costa, radius, much shorter macrotrichia on distal portions of vein M 1, present or absent on M 2, CuA 1, CuA 2, also present on distal half of cell m 1, in a few more or less linear rows or absent in cells m 2, cua 1; radial cells well developed but narrow, or reduced to sutures or completely coalesced, distal portion of costa, 2 nd radial cell dark, posterior veins paler; membrane infuscated with indistinct pale spots in cell r 3 just beyond end of costa and over r-m crossvein, a pale streak in mid portion of cell m 1, m 2 and posterior to base of vein CuA 1, poorly defined pale areas may also be present in cell CuA 1 and anal cell; wing length 1.74 (1.57 – 1.97, n = 18) mm; costal ratio 0.58 (0.55 – 0.62, n = 18). Halter stem pale to light brown; knob whitish, but may appear blackish in some slide mounted specimens. Abdomen: (Fig. 8) Brown, light brown or golden in gravid specimens; segments 7 – 10 brown to medium dark brown; tergite 8 with short anterior, heavily sclerotized, crescent-shaped band, sternite 8 with wrinkled posterior margin; segment 9 a continuous band without well-defined pleurae; sternite 10 broadly triangular with 2 pairs of large ventral setae. Spermatheca elongate, ovoid to egg-shaped; heavily sclerotized with numerous tiny hyaline punctations; distal portion may be narrowed or constricted, with broad opening; neck and duct non-sclerotized but readily visible; total length 0.088 (0.077 – 0.102, n = 18) mm.	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFEEFFD5FF3BFC84FC5A45C2.taxon	distribution	Distribution. Southern and central Alberta and North Dakota, however, it may also occur in other northern Great Plains states in the United States and in Saskatchewan and Manitoba, Canada (see Bionomics).	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFEEFFD5FF3BFC84FC5A45C2.taxon	etymology	Etymology. The specific epithet is a patronym in honor of Joseph A. Shemanchuk, mentor of TJL at Lethbridge Research Centre, and in recognition of his pioneering work on Culicoides in Alberta.	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFEEFFD5FF3BFC84FC5A45C2.taxon	materials_examined	Type material. Holotype male, CANADA, Alberta, Fort Macleod, 20. VI. 1955, (J. A. D. 194 / 6 / 14); allotype female, same data, (J. A. D. 194 / 8 / 5); paratypes, 15 males, 18 females, same locality data except J. A. Downes, 10. VI. 1955 (males, J. A. D. 184 / 1 / 351; females, J. A. D. 184 / 1 / 15 - 19, 352) and 20. VI. 1955 (males, J. A. D. 194 / 8 / 4, 17 - 24, 32 - 35, 194 / 11 / 4; females, J. A. D. 194 / 5 / 5, 194 / 6 / 24,37, 194 / 8 / 25 - 31, 194 / 11 / 1 - 3). Holotype, allotype and paratypes deposited in CNCI. Other paratypes: UNITED STATES, North Dakota, Pierce Co., Pleasant Lake, June 1969, W. W. Wirth, alkali lake, 11 males, 9 females with associated pupal exuviae. Other material examined. Additional pinned specimens, probably the same species, but are not designated paratypes, with same locality data as holotype and allotype: 10. VI. 1955 (28 females, J. A. D 184 / 1 / 321 - 343), 20. VI. 1955 (8 males, J. A. D. 194 / 7 / 4,15,17 - 22; 10 males, J. A. D. 194 / 7 / 2, 5 - 13), and 22. VII. 1955 (17 males J. A. D. 220 / 5 / 52,53,55,57,59,62,65,66,86; 220 / 1 / 45 - 48,50,51,53; 221 / 4 / 108; 13 females, J. A. D. 220 / 5 / 54,56,58,60,63,64; 220 / 1 / 42 - 44,49,52,54; 221 / 4 / 72).	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFEEFFD5FF3BFC84FC5A45C2.taxon	discussion	Discussion. This new species most closely resembles C. riethi, which differs from C. shemanchuki by its darker wing with numerous light and dark maculations, and a longer, more slender palpal segment 3. Males of C. riethi differ from males of C. shemanchuki in being larger (wing length 1.46 – 1.74 mm; vs. 1.22 – 1.39 mm for males of C. shemanchuki), with overall longer genitalia, an aedeagus with bifurcate distal portion, a longer gonostylus that is abruptly curved at midlength with a very slender distal half, and a much broader tergite 9 with more widely spaced apicolateral processes. Females of C. riethi further differ from females of this new species in having a mandible with 9 – 14 larger teeth, a lacinia with 18 – 25 large teeth, and a broader spermatheca. Bionomics. In his article on assembly and mating in biting nematocerous Diptera, Downes (1958 a) provided details of the unusual mating behavior in the related Palearctic species, C. nubeculosus (Meigen) in a section entitled “ Some Examples of Other Patterns of Mating ” that include “ A number of species which depart widely and conspicuously from the normal are now to be considered. It is a remarkable fact that these species, differing so markedly in certain aspects of their behavior, economy and structure, are nevertheless closely related to normal species and frequently fall within the same sub-genus ” [Monoculicoides]. Downes (1958 a) continued with “ This section can best be introduced by a further account of Culicoides nubeculosus, a blood-sucking and anatomically normal species that can exhibit the whole of the typical behavior of swarming and mating in flight. Unlike most others, however, it is also able to mate in an entirely different manner, in a confined space, without flight, and with antennal setae decumbent and thus without an auditory response to the female. When a male, not in flight, encounters and touches a female, there is an immediate response and normal mating takes place. The female may be crawling, or at rest, or in the act of blood-sucking. She is not recognized at a distance, and direct contact, probably by the legs of the male, is necessary. This mating response is not specific; it is elicited not only by females but also by other males and by the females of other species of Culicoides. It may perhaps be supposed that it represents the final stage of the process of mating during the swarming flight, when the male has moved towards the female and the first contact occurs. ” “ This second manner of mating in C. nubeculosus has been studied only in the laboratory, but it may occur in nature also since the species breeds at high densities in restricted environments (farms, sewage plants, etc.) and the adults would have a certain chance of encountering each other on the surface of the larval habitat. In Russia, however, it has been recorded that the males are attracted to the host of the females, cattle, and so encounter them as they are feeding. ” Similar behavior was noted in the primary vector of bluetongue in the Nearctic region, C. sonorensis by Gerry and Mullens (1998), who observed males swarming near cattle, and then landing and mating with females while they fed upon cattle in southern California. In his next paragraph, Downes (1958 a) describes details of “ … a new species of Culicoides, belonging to the nubeculosus group, from southern Alberta. ” This species is C. shemanchuki, new species, about which Downes wrote “ The larval habitat is very localised, being confined to the zone of damp mud that often surrounds alkaline prairie sloughs, and the larvae occur at very high densities. It is not a blood-sucking species – the mouthparts of the female are reduced – and the ovaries develop within two or three days of emergence from the pupa at the expense of internal reserves. These characteristics, by eliminating the need for dispersal and searching, probably determine its ability to maintain the high level of population. ” “ Both sexes fly in the immediate neighbourhood of the larval habitat, although probably to an unusually small extent; male swarms have not been detected. Both sexes also, however, have the habit, not recorded in any other species of Culicoides, of running about rapidly in a random exploratory manner on the surface of the mud; and since the surface is flat and unencumbered by vegetation it provides optimum conditions for accidental meeting. Mating occurs on contact exactly as in the second manner described for C. nubeculosus; it does not depend on the flight of the female or the erection of the antennal setae of the male and contact occurs only by chance; the claspers of the male, moreover, are somewhat enlarged and heavily sclerotized, and presumably unusually efficient. The mating response is elicited by females and males. ” Finally, Downes (1958 a: fig. 5) included a photo of the antenna of a male C. shemanchuki (as C. n. sp.), in which the plume is “ … much reduced, and the pedicel, which contains the vibration-sensitive sensilla, is much smaller than usual. Both features suggest that the antenna has become non-functional as an auditory organ, in relation to the novel habits of this species. ” In a subsequent article, Downes (1971) provided photographs of the head and mouthparts of a female C. shemanchuki, that he referred to as “ … an autogenous biting midge (Culicoides sp. nov.) closely related to C. variipennis. ” He also wrote, “ Note the shortened proboscis and other changes in head structure and the feebly sclerotized mandibles, lacinia, and labrum, with armature vestigial or absent. ”	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFEDFFD8FF3BFC04FB2D4242.taxon	description	(Fig. 9 – 21)	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFEDFFD8FF3BFC04FB2D4242.taxon	diagnosis	Diagnosis. A medium to large species of the C. nubeculosus - stigma complex that differs from other Nearctic species in that complex by the following combination of characters: male wing length 1.46 – 1.74 mm, female wing length 1.55 – 1.94 mm; wing veins and membrane dark dusky brown with distinct pattern of large pale spots over r-m crossvein and on much of the membrane; fore tibia with sub-basal pale band. Male: tergite 9 tapering slightly distally, apex very broad with wide shallow V-shaped median notch, and slightly to moderately divergent apicolateral processes; gonocoxite short; gonostylus much longer, proximal half stout, abruptly tapered and bent at mid-length with very slender distal half; parameres fused proximally, distal portion bifurcate, each prong nearly straight to sinuate with sharply pointed apex; aedeagus with distal portion deeply bifurcate, each prong apically bifid. Female: mandible well developed, with 9 – 14 coarse retrorse teeth on inner margin; lacinia with 18 – 25 coarse, retrorse teeth on outer margin; papal segment 3 moderately slender (palpal ratio 2.20 – 2.83); fore, mid tibiae with sub-basal pale band; spermatheca ovoid to elongate pyriform, with numerous tiny hyaline punctations. Male. Head: (Fig. 9, 15) Brown. Eyes widely separated, with fine sparse setae along margins but apparently absent between ommatidia. Antennal flagellum (Fig. 9, 16) with flagellomeres 1, 11 – 13 separate, 2 – 10 fused; flagellomeres 11 – 13 considerably longer than 2 – 10, 11 longest; flagellomeres 1, 6 – 8 with single apical sensilla coeloconica; flagellomeres 1 with 2 rows, 2 – 13 with single row of sensilla chaetica, those on 2 – 10 comprising the long, moderately dense plume; combined total lengths of flagellomeres 0.806 (0.742 – 0.877, n = 5) mm; antennal ratio 0.55 (0.54 – 0.57, n = 5). Palpal segment 3 slender with small, shallow subapical pit bearing a few capitate sensilla; palpal ratio 2.61 (2.27 – 2.88, n = 5). Proboscis moderately short, extending to palpal segment 4, proboscis / head ratio 0.76 (0.67 – 0.86, n = 11); mandible, lacinia vestigial, without teeth. Thorax: Scutum medium dark brown to golden brown without trace of pattern; scutellum lighter brown to golden with 2 submarginal, 3 – 4 central large setae; postscutellum brown. Femora, tibiae light brown to medium brown; fore tibia with narrow sub-basal pale band; mid, hind tibiae with or without poorly developed sub-basal pale band; claws small, proximal 3 / 4 straight, distal 1 / 4 curved, tips bent, apices bifid. Wing (Fig. 10, 17) similar to but narrower than female wing, pattern of dark and light spots less distinct in some specimens; most of membrane infuscated, covered with coarse microtrichia, which are densest in dark spots; macrotrichia restricted to costa, radius and anterior marginal region of cell r 3, occasionally present on veins M 1, M 2, CuA 1 and in cell m 1; radial cells usually well developed, 1 st narrow, up to twice as long as 2 nd, 2 nd quadrate, dark brown; wing length 1.54 (1.43 – 1.74, n = 5) mm; costal ratio 0.55 (0.53 – 0.56, n = 5). Halter brown to golden. Abdomen: Medium dark brown; pleurae light brown. Genitalia (Fig. 11, 18) dark brown. Tergite 9 slightly tapering distally; apex very broad with shallow, wide V-shaped median notch; apicolateral process conical, moderately slender, extending beyond apex of gonocoxite, with single small apical seta, apices slightly to moderately divergent (dependent on compression from cover slip); cercus setose with pair of large ventrolateral setae, arising anterior to posterior margin, when flattened due to mounting, may extend well beyond posterior margin. Sternite 9 3 x broader than long; anterior margin straight, posterior margin with moderately narrow V- or U-shaped excavation as deep as ½ of total length of sternite. Gonocoxite straight, 1.5 x longer than broad, tapering slightly distally, dorsal root moderately massive, extending to or beyond proximal level of basal arm of parameres, slightly to moderately recurved laterally. Gonostylus slightly longer than gonocoxite, proximal ½ stout, moderately curved distally, abruptly tapered and bent up to 90 ◦ at mid-length, distal ½ slender, slightly curved, apex roundly pointed. Parameres fused basally, more or less Y-shaped; basal arm moderately to heavily sclerotized, nearly straight, apex folded ventrad; mid portion moderately to lightly sclerotized; distal portion lightly sclerotized, bifurcate, each prong nearly straight to sinuate, apex sharply pointed. Aedeagus Y-shaped; membrane spiculate; basal arm heavily sclerotized, recurved; mid portion heavily sclerotized on margins, ventral surface bare, basal arch curved, extending ¼ of total length of aedeagus; distal portion lightly sclerotized, bifurcate, apex of each half bifid. Female. Head: (Fig. 12, 19) Brown. Eyes bare, separated by diameter of 2 – 3 ommatidia. Antennal flagellomeres brown, 1 – 8 often paler than 9 – 13; 9 – 13 slightly or moderately longer than 2 – 8, 13 longest; flagellomeres 1, 6 – 8 with apical single (rarely multiple) large sensilla coeloconica; flagellomeres 1 – 13 with single sub-basal whorl of moderately long sensilla chaetica, scattered shorter sensilla chaetica and covering of fine pubescence; combined total lengths of flagellomeres 0.655 (0.581 – 0.699, n = 12) mm; antennal ratio 0.83 (0.71 – 0.95, n = 12). Palpus with segment 3 slender to slightly swollen, slightly longer than segments 4 – 5 combined, with single small round shallow subapical pit (rarely with 2 smaller pits) bearing numerous capitate sensilla that extend well beyond pit opening; palpal ratio 2.37 (2.20 – 2.83, n = 13). Proboscis moderately long; proboscis / head ratio 0.76 (0.67 – 0.86, n = 11); mandible well developed, with 9 – 14 coarse retrorse teeth on inner margin; lacinia with 18 – 25 coarse, retrorse teeth on outer margin. Thorax: Golden brown to dark brown; scutum with two yellowish sutures originating from humeri; scutellum pale to light brown with 6 – 8 large setae. Fore, mid tibiae with narrow sub-basal pale bands; femoro-tibial joints darkened; tarsomeres 1 – 3 pale to light brown, 4 – 5 darker brown; hind tibial comb with 5 – 6 spines, 2 nd or 3 rd from spur longest. Wing (Fig. 13, 20) moderately slender; membrane darkly infuscated, covered with fine pale microtrichia in light spots and streaks, darker coarse microtrichia in dark spots and on veins; macrotrichia sparsely distributed in all cells, densest in distal regions of cells r 3, m 1 and on most veins; pale spots basad to dark stigma and continuing onto r-m crossvein, a larger spot posterior to stigma and in middle of distal dark spot in cell r 3, elongate pale streaks in cells m 1, m 2 and cua 1, marginal pale spots in cells m 1, m 2, cua 1 and along margin and basal region of anal cell; wing length 1.78 (1.55 – 1.94, n = 13) mm; costal ratio 0.57 (0.55 – 0.58, n = 13). Halter light brown. Abdomen: Light to medium brown. Spermatheca (Fig. 14, 21) ovoid to elongate pyriform, with numerous tiny hyaline punctations and large opening; spermatheca length 0.074 (0.064 – 0.082, n = 12) mm.	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFEDFFD8FF3BFC04FB2D4242.taxon	distribution	Distribution. Holarctic; in North America from Alaska to Manitoba, south to British Columbia, Wyoming and Nebraska (Borkent and Grogan 2009). Types. Type locality, C. riethi, Germany, “ Im Salswasser von Sassendorf und Salzkotten, Westfalen. ” (Kieffer 1914). Type locality of holotype female, C. gigas, Fort a la Corne, Saskatchewan, 53 ◦ 17 ’ N, 104 ◦ 30 ’ W (near Kinistino, SE of Prince Albert), 17 July 1925, Kenneth M. King, CNCI (pinned). Specimens examined. CANADA, Alberta, Brooks, 2 June 1955, J. A. Downes, 1 female, 1 male; Lethbridge, 20 July 1955, J. A. Downes, at light (JAD 218 / 213 / 18), 1 female. UNITED STATES: MON- TANA, Sheridan Co., Medicine Lake, 9 June 1969, W. W. Wirth, 1 female, 1 male, with pupal exuviae. NEBRASKA, Cherry Co., Pelican Lake, 2 June 1969, W. W. Wirth, light trap, 1 female; same data except Twin Lake, alkali lake, 1 female, 1 male with pupal exuviae. Garden Co., Crescent Lake Refuge nr. Pellin Lake, 28 - VI- 1994, F. R. Holbrook, 1 female, 1 male. NORTH DAKOTA, Burleigh Co., Long Lake, 4 June 1969, W. W. Wirth, 4 females. SOUTH DAKOTA, Campbell Co., 21 - 27 - VIII- 2002, BTSPP, 46056, ABADRL, 2 females, 1 male. WYOMING, Park Co., Roberts Ranch nr. Cody, 1 st place on E side of hwy 120, 21 - VII- (19) 86, 1 female. FRANCE, Ardennes, Viller-Le Tourneur, Light trap 08 PL 15, 3 - 4 October 2011, coll. J. C. Delécolle, 1 male. Moselle, Marsal, Emergence 4494 P 175, 12 May 2011, coll. J. C. Delécolle, 1 male. Vosges, Frenelle-La Grande, Light trap 88 PL 2, 1 - 2 August 2011, coll. J. C. Delécolle, 1 female. POLAND, Dolina Bedkowska, k. Krakowa, 3 - VII- 1983, R. Szadziewski, 1 female. Plw. Helski, n. Zatoka Wladyslawowo, 23 - VI- 1979, R. Szadziewski, light trap, 2 females, 4 males. Silec, Distr. Ketrzyn, 2 - 14 - VIII- 1979, R. Szadziewski, light trap near buildings, 1 female.	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFEDFFD8FF3BFC04FB2D4242.taxon	discussion	Discussion. Root and Hoffman (1937) described the legs of the holotype female of C. gigas as being “ uniformly yellowish brown ”; however, our examination of Nearctic males and females revealed pale bands on at least the fore tibia in males, and fore and mid tibiae in females. We compared our Nearctic specimens of C. gigas with specimens of C. riethi from France (Figs. 15 – 21) and Poland and found them to be virtually identical in size, coloration, pale tibial bands, wing pattern, male genitalia and female spermatheca. Therefore, we consider Culicoides gigas Root and Hoffman a junior synonym of Culicoides riethi Kieffer (new synonym). The possibility that C. gigas and C. riethi might be conspecific was mentioned in a letter by Willis Wirth to Antony Downes of 31 October 1993 in which Wirth wrote “ In the dim recesses of my memory I recalled that someone, probably you, mentioned the similarity of Culicoides gigas to riethi, so I dug out all of my slides and gave them a good look. I really can find no substantial differences including pupae. I have about half a dozen good slides of riethi from Europe … and a dozen slides of gigas. …. Do you think gigas and riethi are conspecific? ” Through the courtesy of Art Borkent and Jeff Cumming, we have a handwritten letter that Downes apparently sent to Wirth ca. late 1993 or 1994, prior to Dr. Wirth’s death on 3 September 1994. Although it is 2.5 pages in length, Downes’s letter only deals with their possible collaboration in describing two undescribed species of C. (Monoculicoides) from North America: “ sp. 113 ” that Grogan and Phillips (2008) described as C. grandensis (see below), and, C. shemanchuki, new species, described above. Bionomics. Downes (1958 b) also provided detailed observations of reproduction in several species of Culicoides in southern Alberta (Brooks and Macleod) and northeastern Manitoba (Churchill). He included a photograph of a highly saline slough at Brooks, Alberta, a breeding site of C. riethi (as C. gigas) and another species in the subgenus Monoculicoides, C. sonorensis (as C. variipennis albertensis Wirth and Jones). We include some of Downes’ (1958 b) observations below. “ The evidence obtained by sampling the larvae at intervals during the season suggests that at least two of the species occurring at Churchill have a life cycle of two years, with the first winter spent as a young, and the second as a full-grown, larva. “ The prairie fauna is also extensive, and again includes a number of undescribed species. Larvae of several species are abundant around the margins of sloughs, often of high salt content … At Brooks and Macleod, in southern Alberta, repeating the experience at Churchill, four autogenous species were discovered; and these four produced very high populations. Perhaps in the prairies there is commonly a sufficiency of rich larval habitats together with a scarcity, or only occasional abundance, of vertebrate hosts; certainly, moreover flight of the adults is severely restricted by the prevailing wind conditions. “ In Culicoides gigas the first ovarian cycle is completed without a blood meal, but since (at Brooks) the species also bites man very readily, it seems probable that it then proceeds to a second ovarian cycle which depends on this blood meal, thus resembling Culex molestus Forskål among the mosquitoes. A very distinctive oviposition flight takes place before sunset, during which the insects fly along a narrow zone of the margin of the pool at the height of a few inches only, ultimately alighting on the blackish soil to deposit their eggs … “ At some of the alkaline sloughs in southern Alberta, C. gigas is accompanied by the related C. variipennis albertensis Wirth and Jones and by a new species of the same group. The latter is not only autogenous but has markedly reduced mouthparts. ... its mating behavior is also aberrant, and the species provides an interesting example of correlation of atypical features of structure, physiology, ecology, and behavior, … ” This autogenous species with reduced mouthparts refers to C. shemanchuki, new species. Regarding the larval habitat of C. riethi in Great Britain, Edwards (1939) wrote “ C. riethi, as far as known at present, is confined to coastal marshes. ” and “ The larvae are recorded by continental observers as occurring in salt water, presumably on the muddy margins of tidal creeks and pools. ” In his review of the distribution and biology of British Culicoides, Boorman (1986) noted that “ This species is often found in coastal salt marshes, although it is by no means confined to such habitats. It has been maintained as a laboratory colony, under which conditions it was autogenous. ”	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFE0FFD9FF3BFB84FBFD44E2.taxon	diagnosis	Diagnosis. A small species of the C. nubeculosus - stigma complex most closely resembling C. shemanchuki, new species, but differing from that and other Nearctic species in this complex by the following combination of characters: male wing length 1.09 mm, female wing length 1.33 – 1.39 mm; wing with indistinct, poorly developed dark markings; femora and tibiae pale, non-banded. Male with very short genitalia; gonocoxite and gonostylus very short, gonostylus tapering at mid-length to moderately slen- der pointed tip; parameres Y-shaped, fused proximally, distal portion slender, bifurcate; distal half of aedeagus tapered, triangular with undivided pointed tip. Female proboscis very short (proboscis / head ratio 0.6); mandible and lacinia vestigial, without teeth; and a very small short, ovoid spermatheca (length 0.054 mm) with minute hyaline punctations and a broad opening.	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFE0FFD9FF3BFB84FBFD44E2.taxon	distribution	Distribution. Known only from southeastern Utah (Grand Co.), but probably also inhabits other sections of the Upper Colorado Plateau in northeastern Arizona, western Colorado, northwestern New Mexico and extreme southwestern Wyoming.	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFE0FFD9FF3BFB84FBFD44E2.taxon	materials_examined	Type. Type locality, Utah, Grand Co., near Cisco.	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFE0FFD9FF3BFB84FBFD44E2.taxon	discussion	Discussion. The original description, illustrations and photographs are sufficient for the identification of this species; therefore, we only provide a diagnosis of adults and included C. grandensis in our key of Nearctic species in the C. nubeculosus - stigma complex. We recommend that interested workers consult Grogan and Phillips (2008) for details of morphological characters and numerical data of this species. In their discussion section of C. grandensis, Grogan and Phillips (2008) included quotes from letters between Willis Wirth and Antony Downes “ … about collaborating on describing two undescribed species of C. (Monoculicoides) from southern Alberta. ” One of these undescribed species was “ … n. sp. near stigma, … ” which Grogan and Phillips assumed to be C. grandensis, and which Wirth and Downes (in litt.) also referred to as “ n. sp. 113 ”, as this number was on the slide labels of the holotype male and allotype female of that species. It is now clear that this was not the case because Art Borkent eventually found Downes’ specimens of “ n. sp. near stigma ” in the CNCI shortly after Grogan and Phillips (2008) published their description and illustrations of C. grandensis, but none of them were labeled n. sp. 113. However, all of these specimens from Alberta, Canada are in fact C. stigma, which we discuss below. Bionomics. As noted by Grogan and Phillips (2008), Jones (1961) reared the holotype male and allotype female from pupae he collected on 30 May 1958 on the non-vegetated margin of a small stream with white salt deposits near Cisco, Grand County, Utah (in USNM). The only other known specimen of this apparently rare species is a female paratype collected with a CDC light trap by Phillips on 18 September 2001, 4 km southwest of Moab, Grand County, Utah (in CNCI).	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFE1FFDEFF3BFCE4FD8445C2.taxon	description	(Fig. 22 – 36)	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFE1FFDEFF3BFCE4FD8445C2.taxon	diagnosis	Diagnosis. A medium to large species of the C. nubeculosus - stigma complex that differs from all other Nearctic species in this complex species by the following combination of characters: male wing length 1.26 mm, female wing length 1.41 mm (1.28 – 2.01 mm in European specimens); wing (Fig. 24 – 25, 28 – 29, 32, 35) membrane and most veins hyaline with only a single dark spot (stigma) over 2 nd radial cell. Male genitalia (Fig. 26, 33) with tergite 9 tapering slightly to broad apex, with a narrow, deep Vshaped median notch, slender apicolateral processes and a pair of submedian bluntly rounded caudal lobes that are nearly as long as the apicolateral processes; gonocoxite elongate, gonostylus long, tapered evenly to midpoint, distal half slender, curved; ventral membrane spiculate; distal portion of aedeagus bifurcate, each prong apically bifid. Female proboscis (Fig. 27, 34) moderately short (proboscis / head ratio 0.67 – 0.71); palpal segment 3 (Fig. 27, 34) moderately slender (palpal ratio 2.67 – 3.00), with a small shallow subapical pit; spermatheca (Fig. 30, 36) small (length 0.061 mm), spherical with slender appendix-shaped extension opposite the spermathecal opening, without minute hyaline punctations, but with lightly sclerotized ring.	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFE1FFDEFF3BFCE4FD8445C2.taxon	distribution	Distribution. Holarctic; in North America currently known only from central Alberta, Canada.	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFE1FFDEFF3BFCE4FD8445C2.taxon	materials_examined	Type. Type locality, Europe, exact location unknown, but probably Germany. Specimens examined. CANADA, Beaver River, Alta. (Alberta), 6. VI. 1971, (J.) Shemanchuk, (J. A. D. 857 / 12 / 1, 4 - 5) 3 males, and (J. A. D. 857 / 12 / 2, 4 - 8) 6 females. FRANCE, Aube, Chauffour-Les Bailly, La Loge Bailly, Light-trap 10 PL 1, 6 - 7 October, 2009, coll. J. C. Delécolle, 1 male. Haut-Rhin, Orbey, Les Champs Simon, 68 PL 1, 20 - 26 April 2011, coll. J. C. Delécolle, 1 female. Marne, Mosains, Lighttrap 51 OL 1, 17 - 18 August 2009, coll. J. C. Delécolle, 1 male. Oise, Caisnes, Light-trap 60 PL 3, 11 - 12 April 2011, coll. J. C. Delécolle, 1 female. POLAND, Barciany, 11 - VII- 1981, light trap, R. Szadziewski, 1 male. Brzyno, 22 - V- 1982, staw, mui do hodowli, R. Szadziewski, 1 male. Dolina Bedkowska, k. Krakowa, 3 - VII- 1983, R. Szadziewski, 1 female. Sosnowska, Karkonosze, 4 - VIII- 1982, net, R. Szadziewski, 1 female; same data except 5 - VIII- 1982, light trap, 3 males.	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFE1FFDEFF3BFCE4FD8445C2.taxon	discussion	Discussion. We compared our Nearctic specimens of C. stigma from Alberta, Canada (Figs. 22 – 30) with Palearctic specimens of C. stigma from France (Figs. 31 – 36) and Poland and found them to be conspecific. Our North American specimens from Canada are virtually identical with these Palearctic specimens in size, coloration, wing pattern, male genitalia, and, especially the unique female spherical spermatheca with a slender appendix-shaped extension opposite the spermathecal opening. Bionomics. Edwards (1939) noted that “ In the south of England C. stigma is a decidedly uncommon insect; the very small number of specimens in the British Museum are from the counties of Herts, Middlesex, Devon and Notts. It may be commoner in the north as Mr. H. Britten has taken numerous examples in and around Manchester. Dates of capture are mostly in May and June, with one record in September. Some specimens were taken biting at sunset, the bite being painful. Some of the specimens from Manchester were reared from larvae found among floating green algae at the edge of a small pond. ” Boorman (1986) noted that C. stigma is “ Usually found in association with livestock, breeding in mud contaminated with manure. ”	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFE6FFDFFF3BF967FF784301.taxon	description	–– Wing membrane and veins infuscated with several faint to very dark maculations; spermatheca ovoidal to elongate ovoid, with numerous minute hyaline punctations, without slender appendix-shaped extension opposite the spermathecal opening ............................................ 3	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFE6FFDFFF3BF967FF784301.taxon	description	Larger species, wing length 1.55 – 1.97 mm; wing with several or numerous dark maculations, posterior veins darkly infuscated; palpal segment 3 with subapical pit bearing numerous captitate sensilla; mandible with several well-developed teeth (rarely absent in some specimens of C. shemanchuki, new species); spermatheca more elongate ovoid, larger, length 0.064 – 0.102 mm .............................................................................................................................................. 4	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFE6FFDFFF3BF967FF784301.taxon	description	Palpal segment 3 stout, palpal ratio 1.70 (1.46 – 2.17, n = 17); mandible with 2 – 7 small, poorly developed teeth (rarely without teeth); lacinia vestigial, without teeth; wing veins and membrane dusky brown with a few pale maculations; spermatheca longer, length 0.077 – 0.102 mm .......................................................................................... C. shemanchuki, new species	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFE6FFDFFF3BF967FF784301.taxon	description	Gonostylus short, gradually tapering at distal 2 / 3 to apex; aedeagus with distal portion broad or slender, not bifurcate ............................................................................................................ 7	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
175587A7FFE6FFDFFF3BF967FF784301.taxon	description	Wing veins and membrane dark brown with several light spots; tergite 9 caudal margin with wide, shallow, V-shaped median notch .......................................................... C. riethi Kieffer	en	William L. Grogan, Jr., Lysyk, Timothy J. (2015): A revision of the biting midges in the Culicoides (Monoculicoides) nubeculosus-stigma complex in North America with the description of a new species (Diptera: Ceratopogonidae). Insecta Mundi 2015 (441): 1-24, DOI: 10.5281/zenodo.5182560
