taxonID	type	description	language	source
03E76925FFEF6E0EFF6CFB2756F3F0BF.taxon	diagnosis	Diagnosis. The only group of species of Nearctic Culicoides with the following combination of pupal characters: Dorsal apotome without a dorsomedial tubercle, with dark banding on the apex of the pedicel and the base of the respiratory organ (Fig. 1); respiratory organ elongate and slender, with a dark apex; midlength portion bearing scales (Fig. 2 A – B), and 2 – 5 subbasal pores arranged along its length (Fig. 2 A – B); shagreen present on the anterior and posterior margins of the abdominal segments with varying amounts of shagreen laterally (Figs 6 A – B & 7 A – B); and terminal processes extending posterolaterally and with a dark apex (Fig. 8 A – B).	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFEF6E0EFF6CFB2756F3F0BF.taxon	description	Description. Male. Total length = 2.05 – 3.70 mm. Light to medium brown coloration throughout (Fig. 1). Head: Dorsal apotome (DA) with or without spicules, without dorsomedial tubercle, dorsal margin rounded, lateral margins with distinct points (Figs 3 A – D & 4 A – C), DA length (DAL) = 0.26 – 0.37 mm; DA width (DAW) = 0.17 – 0.22 mm; DAW / DAL = 0.56 – 0.75. DA – 1 – H medium long, thick seta on well – developed tubercle, DA – 2 – H dorsal to tubercle base. Dorsolateral cephalic sclerite with one long, one short seta. Palpus extending posteriorly to posterolateral margin of labium, CL – 1 – H about twice length of CL – 2 – H. Thorax: Cephalothorax length = 0.96 – 1.60 mm. Pedicel of respiratory organ short with dark banding at apex (Fig. 1). Respiratory organ (RO) elongate, slender, apex dark brown, mid-length portion with scales, without annulations, with short membranous base, medium brown coloration on posterior margin, with pores closely abutting in single row at apex, 2 – 5 subbasal pores (Fig. 2 A – B), RO length = 0.27 – 0.45 mm; RO width = 0.03 – 0.06 mm; ROW / ROL = 0.08 – 0.15, tracheal tube with reticulations. Anterolateral sensilla (one long, one short seta) on well – developed tubercle. Mesonotum with small bumps anterior to D – 5 – T, smooth posteriorly, D – 1 – T, D – 2 – T short, stout, each on rounded tubercle, D – 3 – T posterior to small rounded tubercle bearing long, slender D – 4 – T, D – 5 – T short or long, on small rounded tubercle, D – 1 – T, D – 2 – T, D – 5 – T in longitudinal row (Fig. 5 A – B). Wing with apical tubercle; halter apex slightly separated from hind leg. Metathorax completely separated medially, with long, thin M – 1 – T near anterior margin, M – 2 – T, M – 3 – T more posterior. Abdomen: Tergites 2 – 7 each with darker pigmentation as medial group of three patches, with anterolateral pair, sternites 3 – 7 each with two medial patches, with anterolateral pair (Figs 1, 6 A – B, & 7 A – B). Distribution of shagreen of abdominal segments variable, though never excessive. Tergite 1 with long D – 3 – I, short D – 2 – I, D – 7 – I anterior, campaniform D – 4 – I, short D – 8 – I, long, thin D – 9 – I on short tubercle posteriorly, L – 2 – I, L – 3 – I short separated medially by long, thin L – 1 – I. Tergite 2 without elongate tubercles, minute L – 2 – II, L – 4 – II separated medially by long, thin L – 3 – II on anterolateral margin. Chaetotaxy of tergite 2, 3, 5 – 7 similar to that of tergite 4. Tergite 4 with short D – 2 – IV on short tubercle, thin D – 3 – IV on elongate, pointed tubercle, D – 5 – IV, D – 4 – IV, D – 7 – IV, D – 8 – IV, D – 9 – IV in transverse row, arranged medially to laterally, minute D – 5 – IV on slightly formed tubercle, D – 4 – IV on small tubercle, D – 7 – IV on small rounded tubercle, D – 8 – IV, D – 9 – IV each long on pointed tubercle, D – 8 – IV seta thicker than D – 9 – IV (Fig. 7 A – B); lateral tubercles with elongate, slender apices, L – 1 – IV thick seta on bifurcate tubercle, L – 2 – IV, L – 4 – IV thick setae, each on elongate bifurcate tubercle, separated by thin L – 3 – IV on elongate bifurcate tubercle (Figs 6 A – B & 7 A – B); sternite 4 with minute V – 5 – IV on small rounded tubercle, V – 6 – IV thin on pointed tubercle, small V – 7 – IV on pointed tubercle, ventral setae in transverse row, shagreen along anterior margins (Fig. 6 A – B). Segment 8 chaetotaxy with seven sensilla: D – 8 – VIII, D – 9 – VIII, L – 2 – VIII, L – 3 – VIII, L – 4 – VIII, V – 6 – VIII, V – 7 – VIII. Segment 9 with genital lobes extending to posterior margin, terminal process projecting posterodorsolaterally to laterally, tapered to pointed, dark apex (Fig. 8 A – B). Female. Similar to male other than sexual differences on segment 9 and the following: total length = 1.81 – 3.94 mm, DA length = 0.23 – 0.34 mm, DA width = 0.18 – 0.24 mm, DAW / DAL = 0.64 – 0.85, Cephalothorax length = 0.96 – 1.55 mm, Ro Length = 0.21 – 0.43 Mm, Ro Width = 0.03 – 0.06 Mm, Row / rol = 0.08 – 0.16. Taxonomic discussion. Comparison with the pupae of other species of C. (Monoculicoides) outside of the Nearctic Region was difficult due to limited material or because they remain undescribed. Of these species, we examined pupae of C. helveticus Callot, C. longicollis Glukhova, C. puncticollis (Becker), and C. nubeculosus (Meigen), and these were found to be consistent with our diagnosis for C. (Monoculicoides). Additionally we observed clear morphological differences between these species and the seven Nearctic species treated herein. Though this subgenus includes 23 species, only 13 have pupal descriptions, and we examined the pupae of ten of those species. The species of C. (Monoculicoides) with previously described pupae not examined here were those of C. cornutus de Meillon (Afrotropical), C. homotomus Kieffer (Oriental), and C. parroti Kieffer (Palearctic). Although descriptions of these species include fewer pupal characters than the current study, we found no contradictory information to either the diagnosis or description here. Of the 12 other subgenera of Culicoides that occur in North America, we have tested our subgeneric diagnosis against all of them. We re-examined Nearctic pupal exuviae from nine subgenera and compared literature descriptions of two subgenera (C. (Avaritia) and C. (Drymodesmyia )). We also tested our diagnosis against previously described pupae of an array of species groups and those considered miscellaneous (Borkent 2016) as listed by Borkent (2014: Table 2), for some based only on descriptions of species outside the Nearctic Region. This is the first diagnosis of the pupa of any subgenus within Culicoides other than that of C. (Selfia) by Atchley (1970) but which included errors (see below).	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFEA6E0EFF6CFCB5565AF5C4.taxon	description	(Fig. 4 B)	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFEA6E0EFF6CFCB5565AF5C4.taxon	diagnosis	Diagnosis. The only species of C. (Monoculicoides) lacking spicules on the dorsal apotome (Fig. 4 B).	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFEA6E0EFF6CFCB5565AF5C4.taxon	description	Description. Male. Unknown. Female. Total length = 2.77 mm (n = 1). Light brown coloration throughout. Head: Dorsal apotome (DA) without spicules (Fig. 4 B), DA length = 0.23 mm (n = 1), ventral margin of the DA not extending past the DA – 1 – H apex; DA width = 0.20 mm (n = 1); DAW / DAL = 0.87 (n = 1). CL – 1 – H long, thin, CL – 2 – H not visible, O – 4 – H not visible. Thorax: 1.05 mm (n = 1). Respiratory organ (RO), elongate, slender, with 3 – 4 subbasal pores, RO length = 0.30 mm (n = 1); RO width = 0.04 mm (n = 1); ROW / ROL = 0.13 (n = 1), D – 1 – T, D – 2 – T stout, each on elongate, rounded tubercle ,, D – 5 – T short, on small rounded tubercle Abdomen: Tergite 1 with long D – 3 – I and D – 2 – I, D – 7 – I anterior, campaniform D – 4 – I, thin D – 8 – I, and long, thin D – 9 – I posteriorly, L – 2 – I, L – 3 – I short separated medially by long, thin L – 1 – I on lateral margin. Shagreen along the anterior margins of segments 2 – 9, without lateral shagreen. Chaetotaxy of tergite 2 difficult to discern. Chaetotaxy of segment 3 similar to that of segment 4. Tergite 4 with D – 2 – IV on small tubercle, thin D – 3 – IV on elongate, pointed tubercle, D – 5 – IV, D – 4 – IV, D – 7 – IV, D – 8 – IV, D – 9 – IV in transverse row, arranged medially to laterally, minute D – 5 – IV on small tubercle, D – 4 – IV on small rounded tubercle, D – 7 – IV on small rounded tubercle, D – 8 – IV, D – 9 – IV each long on elongate pointed tubercle, D – 8 – IV seta thicker than D – 9 – IV; lateral tubercles elongate with short, blunt apices, L – 1 – IV long, stout seta on small rounded tubercle, L – 2 – IV, L – 4 – IV stout setae, each on elongate bifurcate tubercle, separated by thin L – 3 – IV on elongate pointed tubercle,; sternite 4 with minute V – 5 – IV on small rounded tubercle, V – 6 – IV thin on small tubercle, V – 7 – IV on small tubercle, ventral setae in transverse row. Chaetotaxy of segments 5 – 7 similar to that of segment 4. Segment 8 chaetotaxy with seven sensilla on reduced tubercle, without lateral shagreen, Segment 9 with anterior margin not strongly modified Taxonomic discussion. With only one pupal exuviae to examine, we cannot report any variation. For the available specimen, CL – 2 – H was not visible, and it was difficult to discern the shagreen and chaetotaxy of abdominal segment 2. There was a lack of overall shagreen compared to the other members of this subgenus, and it is the only member without any spicules on the dorsal apotome. The tubercles of L – 1 on each segment are rounded whereas in all other species of C. (Monoculicoides) examined; these tubercles are pointed or bifurcate. The tubercles of L – 2 and L – 4 of each segment are similar to most members of the subgenus, contrasting the most with those of C. sonorensis whose lateral tubercle apices are elongate and slender. This species is known only from southeastern Utah.	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFEA6E0EFF6CFCB5565AF5C4.taxon	materials_examined	Material examined: ALLOTYPE female # 3660 (USNM), USA, Utah, Grand Co, near Cisco, margin of alkaline stream, VI. 30.1958, R. H. Jones.	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFE56E01FF6CFF3D5185F53C.taxon	description	(Figs. 3 A, 4 C, 5 B, 6 A, 8 B)	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFE56E01FF6CFF3D5185F53C.taxon	diagnosis	Diagnosis. The only species of C. (Monoculicoides) with no more than 70 % of the dorsal apotome covered in uniform spicules (Fig. 3 A) and bearing three to four clypeal setae (Fig. 4 C).	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFE56E01FF6CFF3D5185F53C.taxon	description	Description. Male. Total length = 2.71 – 3.15 (2.99, 0.196 SD, n = 4) mm. Light brown coloration throughout (Fig. 1). Head: Dorsal apotome (DA) roughly 70 % covered in uniform spicules decreasing in abundance dorsally (Fig. 3 A), DA length = 0.33 – 0.37 (0.35, 0.021 SD, n = 4) mm; DA width = 0.21 – 0.22 (0.216, 0.004 SD, n = 4) mm; DAW / DAL = 0.59 – 0.65 (0.615, 0.027 SD, n = 4), CL – 1 – H about twice length of CL – 2 – H, with CL- 3 - H, with or without campaniform CL- 4 - H (Fig. 4 C); O – 2 – H, O – 4 – H separated medially by O – 1 – H and O – 3 – H. Thorax: Cephalothorax length = 1.30 – 1.37 (1.33, 0.031 SD, n = 4) mm. Respiratory organ (RO) elongate, slender, 3 – 4 subbasal pores (Fig. 2 B), RO length = 0.39 – 0.45 (0.42, 0.035 SD, n = 4) mm; RO width = 0.04 – 0.05 (0.043, 0.005 SD, n = 4) mm; ROW / ROL = 0.08 – 0.12 (0.102, 0.019 SD, n = 4), D – 5 – T short. Abdomen: Tergite 1 with long D – 3 – I, short D – 2 – I, D – 7 – I anterior, campaniform D – 4 – I, short D – 8 – I, and long, thin D – 9 – I on short tubercle posteriorly, L – 2 – I, L – 3 – I short separated medially by long, thin L – 1 – I on lateral margin. Chaetotaxy, shagreen of tergite 2 similar to tergite 4, without elongate tubercles, minute L – 2 – II, L – 4 – II separated medially by long, thin L – 3 – II on anterolateral margin. Chaetotaxy, shagreen of segment 3 similar to that of segment 4. Tergite 4 (Fig. 6 A) with short D – 2 – IV on short tubercle, thin D – 3 – IV on elongate, pointed tubercle, D – 5 – IV, D – 4 – IV, D – 7 – IV, D – 8 – IV, D – 9 – IV in transverse row, arranged medially to laterally, minute D – 5 – IV on slightly formed tubercle, D – 4 – IV on small tubercle, D – 7 – IV on small rounded tubercle, D – 8 – IV, D – 9 – IV each long on pointed tubercle, D – 8 – IV seta thicker than D – 9 – IV; lateral tubercles with elongate, apices, L – 1 – IV thick seta on small bifurcate tubercle, L – 2 – IV, L – 4 – IV thick setae, each on elongate bifurcate tubercle, separated by thin L – 3 – IV on elongate bifurcate tubercle; sternite 4 with minute V – 5 – IV on small rounded tubercle, V – 6 – IV thin on pointed tubercle, small V – 7 – IV on pointed tubercle, ventral setae in transverse row, shagreen along anterior margins, lateral portion with scattered shagreen. Segments 5 and 6 with similar chaetotaxy, shagreen to that of segment 4. Segment 7 with similar chaetotaxy to segment 4, with reduced lateral shagreen. Segment 8 chaetotaxy with seven sensilla, without lateral shagreen. Segment 9 with anterior shagreen, anterior margin not strongly modified (Fig. 8 B). Female. Similar to male other than sexual differences on segment 9 and the following: total length = 2.76 – 3.20 (2.93, 0.234 SD, n = 3) mm, DA – 1 – H apex extending past ventral margin of DA, DA length = 0.29 – 0.32 (0.30, 0.015 SD, n = 3) mm, DA width = 0.23 – 0.25 (0.237, 0.012 SD, n = 3) mm, DAW / DAL = 0.77 – 0.79 (0.78, 0.013 SD, n = 3), Cephalothorax length = 1.30 – 1.37 (1.34, 0.035 SD, n = 3) mm, RO length = 0.42 – 0.43 (0.423, 0.006 SD, n = 3) mm, RO width = 0.04 – 0.05 (0.043, 0.006 SD, n = 3) mm, ROW / ROL = 0.09 – 0.11 (0.102, 0.012 SD, n = 3). Taxonomic discussion. This species has 3 - 4 subbasal pores on the respiratory organ and along with C. shemanchuki, are the only members of C. (Monoculicoides) with four subbasal pores on the respiratory organ. We have not observed any specimens of C. sonorensis or C. variipennis, the other members of the C. variipennis complex, with more than three subbasal pores. This could potentially be a useful identifying character within this complex, however, more specimens are needed to confirm this as high variation has been observed in the number of subbasal pores within other species (Lawson 1951). This species and C. sonorensis are the only members of C. (Monoculicoides) examined where some specimens had a 4 th ocular seta, O – 4 – H (Shults et al. 2016). Of the members of the subgenus with spicules on the dorsal apotome, C. occidentalis has the lowest spicule density (Fig. 3 A). Pupae of this species had 1 or 2 extra campaniform clypeal sensilla, CL – 3 – H and CL – 4 – H (Fig. 4 C). One extra CL sensilla is also present in C. riethi but C. occidentalis is the only species observed with a CL – 4 – H.	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFE56E01FF6CFF3D5185F53C.taxon	materials_examined	Material examined: 1 male, 2 females (CNCI), Canada, B. C., Kamloops, alkaline lake, 21. VII. 1994, Grogan & McKinnon. 4 males, 4 females, (1 male, 1 female TAMU, voucher series # 732; 3 males, 3 females CNCI), Canada, B. C., White Lake, 5 km SW Okanagan Falls, 49 ° 18 ’ 27.51 " N, 119 ° 138 ’ 00.23 " W, 4. V. 2014, A. Borkent.	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFE16E05FF6CFF3D5185F45C.taxon	description	(Figs. 1, 3 B, 6 B, 8 A)	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFE16E05FF6CFF3D5185F45C.taxon	diagnosis	Diagnosis. The only species of C. (Monoculicoides) with anterior portion of segment 9 greatly swollen (Fig. 8 A).	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFE16E05FF6CFF3D5185F45C.taxon	description	Description. Male. Total length = 3.30 – 3.70 (3.5, 0.283 SD, n = 2) mm. Medium brown coloration throughout (Fig. 1). Head: Dorsal apotome (DA) roughly 90 % covered in uniform spicules (Fig. 3 B). DA length = 0.32 – 0.33 (0.325, 0.007 SD, n = 2) mm; DA width = 0.21 – 0.22 (0.215, 0.007 SD, n = 2) mm; DAW / DAL = 0.636 – 0.688 (0.662, 0.036 SD, n = 2). DA – 1 – H apex extending past ventral margin of DA. CL – 1 – H about twice length of CL – 2 – H, CL – 3 – H campaniform sensilla medial. Thorax: Cephalothorax length = 1.00 – 1.13 (1.08, 0.057 SD, n = 2) mm. Respiratory organ (RO) elongate, slender, 1 – 2 subbasal pores, RO length = 0.21 – 0.26 (0.23, 0.016 SD, n = 2) mm; RO width = 0.03 – 0.035 (0.030, 0.001 SD, n = 2) mm; ROW / ROL = 0.1 1 – 0.15 (0.13, 0.01 SD, n = 2), D – 5 – T short. Abdomen: Tergite 1 with long D – 3 – I, short D – 2 – I, D – 7 – I anterior, short D – 4 – I, and long, thin D – 9 – I separated medially by campaniform D – 8 – I on short tubercle posteriorly, L – 2 – I, L – 3 – I short separated medially by long, thin L – 1 – I on lateral margin. Chaetotaxy, shagreen of tergite 2 similar to tergite 4, without elongate tubercles, minute L – 2 – II, L – 4 – II separated medially by long, thin L – 3 – II on anterolateral margin. Chaetotaxy, shagreen of segment 3 similar to that of segment 4. Tergite 4 (Fig. 6 C) with short D – 2 – IV on short tubercle, thin D – 3 – IV on elongate, pointed tubercle, D – 5 – IV, D – 4 – IV, D – 7 – IV, D – 8 – IV, D – 9 – IV in transverse row, arranged medially to laterally, minute D – 5 – IV on slightly formed tubercle, D – 4 – IV on small tubercle, D – 7 – IV on small rounded tubercle, D – 8 – IV, D – 9 – IV each long on pointed tubercle, D – 8 – IV seta thicker than D – 9 – IV; lateral tubercles with elongated apices, L – 1 – IV thick seta on small bifurcate tubercle, L – 2 – IV, L – 4 – IV thick setae, each on elongate, bifurcate tubercle, separated by thin L – 3 – IV on elongate, bifurcate tubercle; sternite 4 with minute V – 5 – IV on small rounded tubercle, V – 6 – IV thin on pointed tubercle, small V – 7 – IV on pointed tubercle, ventral setae in transverse row, shagreen along anterior margins, with lateral shagreen. Segments 5 – 7 with similar chaetotaxy, shagreen to that of segment 4. Segment 8 slightly swollen, with seven sensilla, with lateral shagreen. Segment 9 with anterior shagreen, anterior margin greatly swollen. Female. S imilar to male other than sexual differences on segment 9 and the following: total length = 3.50 – 3.94 (3.66, 0.241 SD, n = 3) mm, DA length = 0.31 – 0.34 (0.32, 0.017 SD, n = 3) mm, DA width = 0.21 – 0.23 (0.22, 0.01 SD, n = 3) mm, DAW / DAL = 0.65 – 0.74 (0.689, 0.048 SD, n = 3), Cephalothorax length = 1.35 – 1.40 (1.38, 0.025 SD, n = 3) mm, RO length = 0.37 – 0.41 (0.387, 0.021 SD, n = 3) mm, RO width = 0.05 – 0.06 (0.057, 0.006 SD, n = 3) mm, ROW / ROL = 0.13 – 0.16 (0.147, 0.015 SD, n = 3). Taxonomic discussion. Pupae of C. riethi are medium brown in coloration (Fig. 1), similar to that of C. shemanchuki, but have much more lateral shagreen than that species. The density of the spines on the DA (Fig. 3 B), abundance of lateral shagreen on all abdominal segments, and the swollen anterior most region of segment 9 (Fig. 8 A) make this the most distinctive species of C. (Monoculicoides). Like C. occidentalis, this species has an additional clypeal seta not seen in other members of the subgenus, although no specimen has been observed with a CL – 4 – H. The pupa of C. riethi has been redescribed by many authors since its original description by Kieffer (1914) (Rieth 1915, Thienemann 1928, Mayer 1934, Kettle & Lawson 1952, Dzhafarov 1964, Damian-Georgescu & Spătaru 1971, and Glukhova 1989) and we found no contradictions in those papers to our description. This species is Holarctic in distribution. In North America it ranges from Alaska to Manitoba, south to British Columbia, Wyoming and Nebraska (Borkent & Grogan 2009 (as C. gigas), Grogan & Lysyk 2015).	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFE16E05FF6CFF3D5185F45C.taxon	materials_examined	Material examined: 2 males, 3 females, (1 female TAMU, voucher series # 732; 2 males, 2 females CNCI), Canada, B. C., White Lake, 5 km SW Okanagan Falls, 49 ° 18 ’ 27.51 " N, 119 ° 138 ’ 00.23 " W, 4. V. 2014, A. Borkent.	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFE16E07FF6CF92B55C4F1D7.taxon	description	(Figs. 2 A, 5 A)	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFE16E07FF6CF92B55C4F1D7.taxon	diagnosis	Diagnosis. The only species of C. (Monoculicoides) with 5 pores on the subbasal portion of the respiratory organ (Fig. 2 A) and D – 5 – T equal in length to D – 1 – T and D – 2 – T (Fig. 5 A).	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFE16E07FF6CF92B55C4F1D7.taxon	description	Description. Male. Total length = 2.95 mm (n = 1). Medium brown coloration throughout. Head: Dorsal apotome (DA) covered in short spicules. DA length = 0.32 mm (n = 1); DA width = 0.18 mm (n = 1); DAW / DAL = 0.563 (n = 1). Thorax: Cephalothorax length = 1.26 mm (n = 1). Respiratory organ (RO) elongate, slender, 5 subbasal pores (Fig. 2 A). RO length = 0.34 mm (n = 1); RO width = 0.04 mm (n = 1); ROW / ROL = 0.118 (n = 1). D – 1 – T, D – 2 – T, D – 5 – T long, stout, equal in length, each on rounded tubercle in longitudinal row (Fig. 5 A). Abdomen: Tergite 1 with long D – 3 – I, short D – 2 – I, D – 7 – I anterior, short D – 4 – I, and long, thin D – 9 – I separated medially by campaniform D – 8 – I on short tubercle posteriorly, L – 2 – I, L – 3 – I short separated medially by long, thin L – 1 – I on lateral margin. Chaetotaxy, shagreen of tergite 2 similar to tergite 4, without elongate tubercles, small L – 2 – II, L – 4 – II separated medially by thin L – 3 – II on anterolateral margin. Chaetotaxy, shagreen of segment 3 similar to that of segment 4. Tergite 4 with short D – 2 – IV, thin D – 3 – IV on short tubercle, D – 5 – IV, D – 4 – IV, D – 7 – IV, D – 8 – IV, D – 9 – IV in transverse row, arranged medially to laterally, minute D – 5 – IV on slightly formed tubercle, D – 4 – IV on small tubercle, D – 7 – IV on small rounded tubercle, D – 8 – IV, D – 9 – IV each long on rounded tubercle, D – 8 – IV seta thicker than D – 9 – IV; lateral tubercles with elongate, slender apices, L – 1 – IV thick seta on bifurcate tubercle, L – 2 – IV, L – 4 – IV thick setae, each on elongate bifurcate tubercle, separated by thin L – 3 – IV on elongate bifurcate tubercle; sternite 4 with minute V – 5 – IV on small rounded tubercle, V – 6 – IV thin on rounded tubercle, V – 7 – IV on rounded tubercle, ventral setae in transverse row, shagreen along anterior margins, no lateral shagreen. Segments 5 – 7 with similar chaetotaxy, shagreen to that of segment 4, D – 9 tubercles becoming more pointed with subsequent segments. Segment 8 chaetotaxy with seven sensilla, without lateral shagreen. Segment 9 with anterior shagreen, anterior margin not strongly modified. Female. Similar to male other than sexual differences on segment 9 and the following: total length = 3.57 mm (n = 1), DA length = 0.27 mm (n = 1), DA width = 0.21 mm (n = 1), DAW / DAL = 0.77 (n = 1), Cephalothorax length = 1.35 mm (n = 1), RO length = 0.36 (n = 1), RO width = 0.04 mm (n = 1), ROW / ROL = 0.11 (n = 1). Taxonomic discussion. The coloration of the pupa of C. shemanchuki is similar to that of C. riethi (Fig. 1) but it has much less lateral shagreen on its abdominal segments. This species has 4 - 5 subbasal pores on the respiratory organ, and is the only C. (Monoculicoides) we examined to have 5. However, we only examined two specimens and there is a significant possibility that there could be greater variation in the number of subbasal pores (Lawson 1951). Both specimens examined had folded dorsal apotomes, which made it difficult to obtain accurate measurements, descriptions, and photographs of this structure. This species is known only from southern Alberta and north-central North Dakota (Grogan & Lysyk 2015).	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFE16E07FF6CF92B55C4F1D7.taxon	materials_examined	Material examined: 1 male, 1 female (CNCI), USA, North Dakota, Pierce Co, Pleasant Lake, June 1969, W. W. Wirth, alkaline lake.	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFE36E19FF6CFC5D5600F2A7.taxon	description	(Fig. 4 A)	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFE36E19FF6CFC5D5600F2A7.taxon	diagnosis	Diagnosis. The only species of C. (Monoculicoides) with short respiratory organ (± 0.26 mm in length) (Fig. 9) and short uniform spicules on the dorsal apotome (Fig. 4 A).	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFE36E19FF6CFC5D5600F2A7.taxon	description	Description. Male. Total length = 2.05 – 2.60 (2.39, 0.137 SD, n = 17) mm. Light brown coloration throughout. Head: Dorsal apotome (DA) roughly 90 % covered in short, uniform spicules (Fig. 4 A), DA length = 0.28 – 0.32 (0.299, 0.014 SD, n = 17) mm; DA width = 0.17 – 0.19 (0.179, 0.004 SD, n = 17) mm; DAW / DAL = 0.58 – 0.64 (0.601, 0.024 SD, n = 17). Thorax: Cephalothorax length = 1.00 – 1.13 (1.08, 0.057 SD, n = 17) mm. Respiratory organ (RO) slender, 1 – 2 subbasal pores, RO length = 0.21 – 0.26 (0.23, 0.016 SD, n = 17) mm; RO width = 0.03 0 – 0.035 (0.03, 0.001 SD, n = 17) mm; ROW / ROL = 0.11 – 0.15 (0.13, 0.01 SD, n = 17). Abdomen: Tergite 1 with long D – 3 – I, short D – 2 – I, D – 7 – I anterior, short D – 4 – I, and long, thin D – 9 – I separated medially by campaniform D – 8 – I on short tubercle posteriorly, L – 2 – I, L – 3 – I short separated medially by long, thin L – 1 – I on lateral margin. Chaetotaxy, shagreen of tergite 2 similar to tergite 4, without elongate tubercles, minute L – 2 – II, L – 4 – II separated medially by long, thin L – 3 – II on anterolateral margin. Chaetotaxy, shagreen of segment 3 similar to that of segment 4. Tergite 4 with short D – 2 – IV on short tubercle, thin D – 3 – IV on elongate, pointed tubercle, D – 5 – IV, D – 4 – IV, D – 7 – IV, D – 8 – IV, D – 9 – IV in transverse row, arranged medially to laterally, minute D – 5 – IV on slightly formed tubercle, D – 4 – IV on small tubercle, D – 7 – IV on small rounded tubercle, D – 8 – IV, D – 9 – IV each long on pointed tubercle, D – 8 – IV seta thicker than D – 9 – IV; lateral tubercles with short, blunt apices, L – 1 – IV thick seta on small bifurcate tubercle, L – 2 – IV, L – 4 – IV thick setae, each on elongate, bifurcate tubercle, separated by thin L – 3 – IV on elongate, bifurcate tubercle; sternite 4 with minute V – 5 – IV on small rounded tubercle, V – 6 – IV thin on pointed tubercle, small V – 7 – IV on pointed tubercle, ventral setae in transverse row, shagreen along anterior margins, lateral shagreen reduced or absent. Segments 5 – 7 with similar chaetotaxy, shagreen to that of segment 4. Segment 8 chaetotaxy with seven sensilla, without lateral shagreen. Segment 9 with anterior shagreen, anterior margin not strongly modified. Female. Similar to male other than sexual differences on segment 9 and the following: total length = 1.94 – 2.86 (2.04, 0.248 SD, n = 15) mm, DA – 1 – H apex extending past ventral margin of DA, DA length = 0.23 – 0.3 (0.267, 0.018 SD, n = 15) mm, DA width = 0.17 – 0.215 (0.197, 0.016 SD, n = 15) mm, DAW / DAL = 0.65 – 0.79 (0.737, 0.035 SD, n = 15), Cephalothorax length = 1.00 – 1.15 (1.07, 0.059 SD, n = 15) mm, RO length = 0.21 – 0.26 (0.237, 0.015 SD, n = 15) mm, RO width = 0.03 – 0.035 (0.031, 0.003 SD, n = 15) mm, ROW / ROL = 0.11 – 0.15 (0.133, 0.019 SD, n = 15). Taxonomic discussion. This species is one of the smaller members of C. (Monoculicoides), along with C. sonorensis and C. grandensis, but has a proportionally smaller respiratory organ (Fig. 9) in comparison to its body size. The total range of the ratio of respiratory organ length to cephalothorax length of all other C. (Monoculicoides) examined was 0.27 – 0.31 (n = 54), whereas in C. stigma, it is significantly lower 0.19 – 0.22 (n = 32). This species is Holarctic. However, in North America it is currently known only from central Alberta. The pupa of this species was originally described by Kettle & Lawson (1952) and redescribed in two other studies (Dzhafarov 1964; Spătaru 1971), and our study did not find any contradictions with these studies. Although this species is reported as Holarctic, our description of the pupa is based solely on Palearctic material. It is possible that there are morphological differences between populations from the two regions.	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFE36E19FF6CFC5D5600F2A7.taxon	materials_examined	Material examined: 12 males, 7 females (ZIN), Russia, Kaliningrad Province, Rybachii village, Curonian Spit National Park [at the Baltic Sea coast], 16 – 17.07.1989, Glukhova. 1 female (ZIN), Russia, Leningrad Province, Luga district, forest ditch with water, 27.07.1979, Brodskaya. 1 female (ZIN), Russia, Leningrad Province, environs of Luga town, ditch with moss, 13.08.1979, Brodskaya. 1 female (ZIN), Russia, Leningrad Province, Luga district, floodplain of river Obla, 24.08.197, Brodskaya. 3 males, 3 females (ZIN), Luga district, in ditch, 12 – 14. VIII. 1979. 1 male, 1 female (ZIN), Russia, Republic of Karelia, 1966. 1 male, 1 female (ZIN), Russia, Republic of Karelia, 2 – 4. VII. 54.	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFFD6E18FF6CFEB75570F5ED.taxon	description	(Figs. 3 D, 7 B)	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFFD6E18FF6CFEB75570F5ED.taxon	diagnosis	Diagnosis. The only species of C. (Monoculicoides) with lateral areas of all abdominal segments mostly bare or without any shagreen (Fig. 7 B), and lateral tubercles with blunt apices (Fig. 7 B).	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFFD6E18FF6CFEB75570F5ED.taxon	description	Description. Male. Total length = 2.60 – 4.15 (3.49, 0.621 SD, n = 7) mm. Light brown coloration throughout. Head: Dorsal apotome (DA) roughly 90 % covered in spicules of various sizes decreasing in abundance dorsally (Fig. 3 D), DA length = 0.29 – 0.38 (0.32, 0.031 SD, n = 7) mm; DA width = 0.20 – 0.25 (0.21, 0.018 SD, n = 7) mm; DAW / DAL = 0.58 – 0.75 (0.663, 0.051 SD, n = 7). Thorax: Cephalothorax length = 1.25 – 1.65 (1.51, 0.150 SD, n = 7) mm. Respiratory organ (RO) elongate, slender with 2 – 3 subbasal pores, RO length = 0.36 – 0.43 (0.39, 0.023 SD, n = 7) mm; RO width = 0.04 – 0.06 (0.051, 0.009 SD, n = 7) mm; ROW / ROL = 0.10 – 0.15 (0.13, 0.019 SD, n = 7). Abdomen: Tergite 1 with long D – 3 – I, short D – 2 – I, D – 7 – I anterior, campaniform D – 4 – I, short D – 8 – I, and long, thin D – 9 – I on short tubercle posteriorly, L – 2 – I, L – 3 – I short separated medially by long, thin L – 1 – I on lateral margin. Chaetotaxy, shagreen of tergite 2 similar to tergite 4, without moderate tubercles, minute L – 2 – II, L – 4 – II separated medially by long, thin L – 3 – II on anterolateral margin. Chaetotaxy, shagreen of segment 3 similar to that of segment 4. Tergite 4 (Fig. 7 D) with short D – 2 – IV on short tubercle, thin D – 3 – IV on elongate, pointed tubercle, D – 5 – IV, D – 4 – IV, D – 7 – IV, D – 8 – IV, D – 9 – IV in transverse row, arranged medially to laterally, minute D – 5 – IV on slightly formed tubercle, D – 4 – IV on small tubercle, D – 7 – IV on small rounded tubercle, D – 8 – IV, D – 9 – IV each long on pointed tubercle, D – 8 – IV seta thicker than D – 9 – IV; lateral tubercles short with, blunt apices (Fig. 7 B), L – 1 – IV thick seta on small bifurcate tubercle, L – 2 – IV, L – 4 – IV thick setae, each on elongate, bifurcate tubercle, separated by thin L – 3 – IV on elongate, bifurcate tubercle; sternite 4 with minute V – 5 – IV on small rounded tubercle, V – 6 – IV thin on pointed tubercle, small V – 7 – IV on pointed tubercle, ventral setae in transverse row, shagreen along anterior margins, lateral shagreen reduced or absent. Segments 5 – 7 with similar chaetotaxy, shagreen to that of segment 4. Segment 8 chaetotaxy with seven sensilla, without lateral shagreen. Segment 9 with anterior shagreen, anterior margin not strongly modified. Female. Similar to male other than sexual differences on segment 9 and the following: total length = 2.96 – 3.75 (3.33, 0.404 SD, n = 4) mm, DA – 1 – H apex extending past ventral margin of DA, DA length = 0.27 – 0.28 (0.279, 0.006 SD, n = 4) mm, DA width = 0.22 – 0.24 (0.233, 0.009 SD, n = 4) mm, DAW / DAL = 0.80 – 0.85 (0.834, 0.029 SD, n = 4), Cephalothorax length = 1.32 – 1.55 (1.413, 0.122 SD, n = 4) mm, RO length = 0.36 – 0.42 (0.388, 0.025 SD, n = 4) mm, RO width = 0.04 – 0.05 (0.043, 0.005 SD, n = 4) mm, ROW / ROL = 0.10 – 0.11 (0.109, 0.007 SD, n = 4). Taxonomic discussion. Shults et al. (2016) reported distinguishing features between C. sonorensis and C. variipennis based on the shape of the apices of the lateral tubercles on the abdominal segments, and further illustration of these characters can be found in that study. Those of C. sonorensis are long and slender (Fig. 7 A), whereas in C. variipennis, they are short and blunt (Fig. 7 B). In comparison to the other Nearctic members of C. (Monoculicoides), these two species seem to represent the extremes of the variation, with the bluntness / elongation of the lateral tubercle apices of the other species falling between these two species. Here we also report the overall reduced shagreen on all abdominal segments in C. variipennis and in particular the lack of lateral shagreen on abdominal segments 4 – 7 as a diagnostic character state. This species is known from British Columbia, Alberta, Washington, and Montana to Nova Scotia, and the United States primarily east of the Mississippi river, Texas and Louisiana (Wirth & Jones 1957; Holbrook et al. 2000; Borkent & Grogan 2009; Grogan & Lysyk 2015; Shults 2015; Jewiss-Gaines et al. 2016). Moderate to large numbers of pupae have been reared and collected from fresh water polluted with manure (Wirth & Jones 1957, Jamnback 1965). Additionally, specimens have been collected from saltmarshes (Blanton & Wirth 1979). The pupa of C. variipennis was briefly described by several authors (Malloch 1915, Thomsen 1937, Fox 1942, Jones 1955, Jamnback 1965, Blanton & Wirth 1979, Weber 2001) but it is difficult to determine whether these descriptions agree with ours as we are uncertain which species of the C. variipennis complex were actually studied by these authors. The specimens we examined from Vernon, B. C. account for the large amount of size variation seen in fig. 10, as they are much smaller (roughly the size of C. occidentalis) than the other specimens of C. variipennis examined. The pupae from Vernon key to C. variipennis but have slightly longer lateral tubercles. It is unclear if these measurements and features fall within the normal geographic variation of this species, represent intermediate character states, or even infer a new cryptic species.	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
03E76925FFFD6E18FF6CFEB75570F5ED.taxon	materials_examined	Material examined: 3 males, 2 female (CNCI), Canada, Ontario, Alfred, 30 – VI – 1985, Dillion & Oliver. 2 males, 2 females (CNCI), Canada, British Columbia, Vernon, Coldstream Ranch, 19. VII. 1994, Grogan & McKinnon. 2 males (NYSM), USA, Missouri, Petersburg, Boone’s Lick salt spring, 11. III. 1955, P. J. Spangler. Morphometric results (Figs. 9, 10, 11, Tables 1 & 2) Morphometric analyses of the Nearctic species of C. (Monoculicoides) show general trends among the size of each species (Figs. 9 & 10) as well as sexual dimorphism within the species examined (Fig. 11). Culicoides stigma has a statistically significant smaller respiratory organ in comparison to the length of the cephalothorax (RO length / cephalothorax length) than all other species of C. (Monoculicoides) (ANOVA with Tukey’s, p <0.001). There was no significant difference in this ratio between any other species. The dorsal apotome of male pupae tend to be longer and narrower than those of female pupae. This is most easily observable in the DA ratio (DAW / DAL) (Fig. 11). Culicoides riethi is the only species examined to show no statistically significant difference in the DA ratio between males and females (Fig. 11) (Student’s t, p = 0.57). All other species examined showed a sexual dimorphism (Student’s t, p <0.05). There is however a sexual dimorphism in the length of the cephalothorax in C. riethi (Student’s t, p <0.05) that does not exist in any other species examined (Fig. 10).	en	Shults, Phillip, Borkent, Art (2018): Pupae of the Nearctic species of Culicoides Latreille subgenus Monoculicoides Khalaf (Diptera: Ceratopogonidae). Zootaxa 4504 (4): 451-472, DOI: 10.11646/zootaxa.4504.4.1
