Research Article |
Corresponding author: Wen-Bin Liu ( skyliuwenbin@163.com ) Academic editor: Fabio Laurindo da Silva
© 2024 Wen-Bin Liu, Cheng-Yan Wang, Ya-Ning Tang, Ying Wang, Wen-Xuan Pei, Chun-Cai Yan.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Liu W-B, Wang C-Y, Tang Y-N, Wang Y, Pei W-X, Yan C-C (2024) Six new species of Cryptochironomus Kieffer (Diptera, Chironomidae) from the Nearctic region. ZooKeys 1200: 275-302. https://doi.org/10.3897/zookeys.1200.119225
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Six new species of Cryptochironomus Kieffer, 1918, C. absum Liu, sp. nov., C. beardi Liu, sp. nov., C. dentatus Liu, sp. nov., C. ferringtoni Liu, sp. nov., C. parallelus Liu, sp. nov. and C. taylorensis Liu, sp. nov., are described and illustrated based on adult males. The specimens were collected from various water systems in the United States and preserved by Dr. Leonard Charles Ferrington Jr. An updated key to adult males of all known Cryptochironomus species in the Nearctic region is also provided.
Adult male, diagnostic characters, hypopygium, key, Nematocera, taxonomy, US
The genus Cryptochironomus was erected by Kieffer in 1918, with Chironomus (Cryptochironomus) chlorolobus Kieffer, 1918 as type species. The adult males of this genus are distinguished by having a finger-shaped inferior volsella which lacks microtrichia, and is often completely covered by the small superior volsella (
The systematic review of Cryptochironomus and a key to all known males in the Nearctic region were supported by
The morphology and terminology are based on
Holotype. one male, USA, Chamberlain South, Dakota State, Lake Francis Case Elm Creek, 43°56'57"N, 99°31'40"W, 3.IX.1971, light trap, leg: Patrick l. Huson.
AR 2.58; frontal tubercles absent; posterior margin of tergite IX arc-shaped; anal point slightly constricted at base, wider at apex; superior volsella oval-shaped, covered with microtrichia at 1/3 distance from apex; inferior volsella completely covered by superior volsella; gonostylus straight, parallel-sided, tapering to the apex.
Male (n = 1).
Total length 4.47 mm. Wing length 2.02 mm. Total length/wing length 2.22. Wing length/length of profemur 2.16.
Coloration. Thorax yellowish brown. Femora of front legs yellowish brown, tibiae dark brown, tarsi lost; femora and tibiae of mid and hind legs light yellowish brown; mid and hind legs with tarsi I yellowish brown except for dark yellowish brown at both ends, tarsi II–V dark yellowish brown. Abdomen yellowish brown, hypopygium dark brown.
Head
(Fig.
Thorax
(Fig.
Wing
(Figs
Legs. Front tibia with three subapical setae, 143 μm, the remaining lost. Mid legs with two tibial spurs, 42 μm long, the other lost, tibial combs 34 μm and 56 μm wide. Hind legs with two tibial spurs, 23 μm and 42 μm long, tibial combs 44 μm and 87 μm wide. Tarsus I of mid leg with three sensilla chaetica; tarsus I of hind leg with three sensilla chaetica. Lengths (in μm) and proportions of legs as in Table
Lengths (in μm) and proportions of legs of Cryptochironomus absum Liu, sp. nov., adult male (n = 1).
fe | ti | ta1 | ta2 | ta3 | |
P1 | 934 | 721 | - | - | - |
P2 | 876 | 738 | 516 | 230 | 155 |
P3 | 964 | 973 | 730 | 367 | 301 |
ta4 | ta5 | LR | BV | SV | |
P1 | - | - | - | - | - |
P2 | 102 | 89 | 0.70 | 3.70 | 3.13 |
P3 | 168 | 110 | 0.75 | 2.82 | 2.65 |
Hypopygium
(Figs
From the Latin, absum, absent, referring to the character of frontal tubercles absent, adjective in the nominative singular.
Cryptochironomus absum Liu, sp. nov. is similar to C. conus Mason, 1985 in having anal tergite bands, anal point, and superior volsella with similar shapes. However, it can be distinguished from C. conus by the following combination of characters: wing length of 2.02 mm, absence of frontal tubercles, inferior volsella with a finger-shaped appearance, and gonostylus straight in this new species. In contrast, C. conus has a wing length of 5.1–5.3 mm, distinct frontal tubercles, an inferior volsella with a tuberculate appearance and a protrusion at the base, and a curved gonostylus.
Holotype. one male, USA, New Mexico State, Rio Grande Otowi Bridge between Santa Fe and Los Alamos, 35°87'48"N, 99°31'40"W, 16.VII.1976, sweep net, leg: Melvin. Beard. Paratype. one male, North America, San Juan River at Farmington, 18.VII.1976, malaise trap, M. Beard.
Thorax pale yellow, with yellow-brown spots; anal point widest at base, constricted slightly at 1/3 distance from base, apically rounded; anal tergite bands V-shaped, fused in the middle; superior volsella oval-shaped, stretching upward at base, swelling at apex; inferior volsella columnar, ~ 2× as long as wide, with slender extension at base, bearing two long setae at apex, free microtrichia; gonostylus protruded at base, slightly curved in the middle, tapered to the apex.
Male (n = 2, unless stated).
Total length 5.03–5.10, 5.07 mm. Wing length 2.43–2.65, 2.54 mm. Total length/wing length 1.89–2.10, 2.00. Wing length/length of profemur 2.13 (1).
Coloration. Thorax pale yellow, with yellowish brown spots. Femora of front legs yellowish brown except for dark yellow-brown at ends, tibiae and tarsomeres dark brown; femora of mid and hind legs yellowish brown, tarsi I yellowish brown with dark brown in distal; tarsi II–V dark yellowish brown. Abdomen pale yellow, hypopygium yellowish brown.
Head
(Figs
Thorax
(Fig.
Wing
(Figs
Legs. Front tibia with three subapical setae, 144 (1) μm, 151 (1) μm and 162 (1) μm. Combs of mid tibia 38–42, 40 μm wide with 22–27, 25 μm long spur, and 52–58, 55 μm wide with 30–40, 35 μm long spur; combs of hind tibia 47–48, 48 μm wide with 20–25, 23 μm long spur, 81–88, 85 μm wide with 38–42, 40 μm long spur. Tarsus I of mid leg with three sensilla chaetica. Tarsus I of hind leg with four sensilla chaetica. Lengths (in μm) and proportions of legs as in Table
Lengths (in μm) and proportions of legs of Cryptochironomus beardi Liu, sp. nov. adult males (n = 2).
fe | ti | ta1 | ta2 | ta3 | |
P1 | 1139 (1) | 857 (1) | 1442 (1) | 648 (1) | 528 (1) |
P2 | 998–1121, 1060 | 913–950, 932 | 579 (1) | 266 (1) | 204 (1) |
P3 | 1156–1226, 1191 | 1185–1305, 1245 | 832 (1) | 397 (1) | 299 (1) |
ta4 | ta5 | LR | BV | SV | |
P1 | 498 (1) | 170 (1) | 1.68 (1) | 1.86 (1) | 1.38 (1) |
P2 | 121 (1) | 107 (1) | 0.63 (1) | 3.57 (1) | 3.30 (1) |
P3 | 185 (1) | 112 (1) | 0.70 (1) | 3.20 (1) | 2.81 (1) |
Hypopygium
(Figs
Name after M. Beard, for the collector of the materials; noun in nominative case.
Cryptochironomus beardi Liu, sp. nov. resembles C. maculus Yan & Wang, 2016 in having gonostylus and inferior volsella with similar shapes. However, it can be distinguished from C. maculus by the following combination of characters: mid and hind legs with tarsi I yellowish brown base, dark brown distally, anal point apically rounded, superior volsella stretching upward, gonocoxite concave with gonostylus obviously in the former; whereas in C. maculus, the mid and hind legs have the tarsi I yellowish green with dark brown mark on basal portion, anal point apically pointed, superior volsella not stretching, gonocoxite fused with gonostylus completely.
Holotype. One male, USA, New Mexico State, Guadalupe County, Pecos River, Puerto de Luna below Diversion Dam, 33°04'06"N, 104°26'79"W, 20.VIII.1991, light trap, leg: Lensky & Doles. Paratypes. 8 males, USA, Guadalupe Country, Pecos River, Puerto de Luna below Diversion Dam, 20.VIII.1991, light trap, leg: Lensky & Doles.
Frontal tubercles conical; tergite IX saddle-shaped at the posterior margin; anal point parallel-sided with rounded apex; superior volsella crescent-like; inferior volsella finger-shaped, dentate at base; gonostylus curved slightly at 1/3 distance from base, swelling at distal 1/3.
Male (n = 9, unless stated).
Total length 4.58–5.30, 4.91 mm. Wing length 2.05–2.29, 2.16 mm. Total length/wing length 2.14–2.31, 2.26. Wing length/length of profemur 2.04–2.61, 2.23.
Coloration. Thorax yellowish brown. Femora of front legs yellowish brown, tibiae and tarsi dark brown; femora and tibiae of mid and hind legs yellowish brown, tarsi I yellowish brown, tarsi II–V yellowish brown to dark brown gradually. Abdomen yellowish brown, hypopygium dark brown.
Head
(Figs
Thorax
(Fig.
Wing (Figs
Legs. Front tibia with three subapical setae, 87–111, 98 (6) μm, 110–123, 115 (6) μm, 117–133, 124 μm. Combs of mid tibia 37–47, 43 μm wide with 23–34, 27 μm long spur, and 34–54, 46 μm wide with 32–45, 39 μm long spur; combs of hind tibia 30–49, 40 μm wide with 27–40, 31 μm long spur, 80–90, 83 μm wide with 34–48, 42 μm long spur. Tarsus I of mid leg with seven sensilla chaetica; tarsus I of hind leg with five sensilla chaetica. Lengths (in μm) and proportions of legs as in Table
Lengths (in μm) and proportions of legs of Cryptochironomus dentatus Liu, sp. nov., adult males (n = 9, unless stated).
fe | ti | ta1 | ta2 | ta3 | |
P1 | 877–1002, 963 | 739–797, 770 | 1223–1361, 1270 (4) | 596–638, 621 (4) | 484–524, 505 (4) |
P2 | 831–978, 925 (6) | 788–872, 826 | 461–520, 490 | 234–264, 248 | 175–205, 190 |
P3 | 987–1080, 1038 (6) | 1071–1182, 1116 | 712–795, 748 (6) | 370–413, 392 (6) | 306–343, 318 (6) |
ta4 | ta5 | LR | BV | SV | |
P1 | 362–412, 394 (4) | 174–191, 185 (4) | 1.61–1.71, 1.65 (4) | 1.72–1.81, 1.76 (4) | 1.23–1.46, 1.37 (4) |
P2 | 115–134, 124 | 100–111, 106 | 0.56–0.62, 0.59 | 3.06–3.53, 3.34 (6) | 3.41–3.83, 3.59 (6) |
P3 | 168–188, 179 (6) | 122–126, 124 (6) | 0.66–0.68, 0.67 (6) | 2.82–2.88, 2.86 (6) | 2.82–2.93, 2.88 (6) |
Hypopygium
(Figs
From the Latin, dentatus, dentate, tooth-like, referring to the shape of the base of inferior volsella, adjective in the nominative singular.
Cryptochironomus dentatus Liu, sp. nov. bears resemblance to C. fulvus Johannsen, 1905 due to its similar frontal tubercles, anal point, and superior volsella. However, C. dentatus Liu, sp. nov. can be distinguished from C. fulvus by the following combination of characters: absence of spots on the thorax, tergite IX with a shoulder-shaped posterior margin, inferior volsella with a finger-shape and dentate base; whereas C. fulvus, has dark brown spots on the thorax, tergite IX with a conical posterior margin, and inferior volsella with a tuberculate and non-dentate base.
Holotype. One male, USA, South Dakota State, Springfield, Lewis and Clark Lake, Boat Basin, 42°87'33"N, 97°49'02"W, 13–17.VII.1964, leg: Pat Hadson. Paratypes. 3 males, Ohio, Cincinnati, Federal Water Quality Association, lab on the window, 26.X.1970, leg: W. T. Mason. 2 males, Springfield South Dakota, Boat Basin, Springfield, Lewist Clank Lake, 13–17.VII.1964, leg: Pat Hadson.
Tergite IX with slightly cone-like posterior margin; anal point narrow at basal 1/3, expanded at 1/3 of the apex, apically rounded; superior volsella crescent-like, with hook-like extension at the base, apically rounded; inferior volsella triangular widest at base, apex with a small protrusion; the junction of gonostylus and gonocoxite concaved obviously; gonostylus widest at basal 1/3, tapered to the apex.
Male (n = 6, unless stated).
Total length 3.29–3.91, 3.58 mm. Wing length 1.47–1.69, 1.61 mm. Total length/wing length 2.12–2.37, 2.23. Wing length/length of profemur 1.81–2.24, 2.02.
Coloration. Thorax yellowish brown. Femora of front legs yellowish brown, tibiae and tarsi lost; mid and hind legs yellowish brown. Abdomen pale yellow, hypopygium yellowish brown.
Head
(Fig.
Thorax
(Fig.
Wing
(Figs
Legs. Front tibia with three subapical setae, 113–132, 123 (3) μm, 124–127, 126 (3) μm, 128–144, 135 (3) μm. Mid legs with two spurs, 16–23, 20 μm and 20–34, 27 μm long, tibial comb 20–39, 30 μm and 27–40, 32 μm wide. Spurs of hind tibia 19–30, 26 μm and 30–45, 36 μm long, tibial comb 30–43, 36 μm and 50–83, 67 μm wide. Tarsus I of mid leg with 4–5, 4 sensilla chaetica, tarsus I of metapedes leg with 3–4, 4 sensilla chaetica. Lengths (in μm) and proportions of legs as in Table
Lengths (in μm) and proportions of legs of Cryptochironomus ferringtoni Liu, sp. nov., adult males (n = 6, unless stated).
fe | ti | ta1 | ta2 | ta3 | |
P1 | 724–938, 800 | 483–559, 531 | 960 (1) | - | - |
P2 | 633–730, 691 | 554–662, 613 | 389–468, 430 | 159–186, 172 | 112–134, 125 |
P3 | 610–797, 711 | 710–850, 795 | 566–638, 603 (4) | 245–296, 269 (4) | 204–238, 220 (4) |
ta4 | ta5 | LR | BV | SV | |
P1 | - | - | 1.81 | - | - |
P2 | 73–89, 80 | 62–74, 67 | 0.69–0.71, 0.70 | 3.73–3.99, 3.91 | 2.93–3.08, 3.04 |
P3 | 122–142, 131 (4) | 80–91, 85 (4) | 0.73–0.80, 0.76 (4) | 2.87–3.12, 2.99 (4) | 2.33–2.59, 2.50 (4) |
Hypopygium
(Figs
Name after Dr. Leonard Charles Ferrington Jr., for his outstanding contribution to the knowledge of Chironomidae taxonomy; noun in nominative case.
Cryptochironomus ferringtoni Liu, sp. nov. bears resemblance to C. rostratus Kiffer, 1921 due to its similar shapes of the posterior margin of tergite IX and superior volsella. However, it can be distinguished from C. rostratus by the absence of frontal tubercles, contraction of the anal point at the basal 1/3 and expansion at the apical 1/3, and a triangular inferior volsella in C. ferringtoni Liu, sp. nov.; whereas C. rostratus frontal tubercles, a distally tapering or parallel-sided anal point, and a tuberculate inferior.
Holotype. one male, USA, New Mexico State, Catron Country, San Francisco River at Glenwood, 33°12'52"N, 109°28'01"W, 7.III.1976, reared, leg: Sta. H & M. Beard. Paratypes. 2 males, USA, New Mexico State, Catron Country, San Francisco River, 17.IX.1974, leg: M. Beard. 1 male, USA, New Mexico State, Catron Country, San Francisco River at Glenwood, 7.III.1976, reared, leg: Sta. H & M. Beard.
Thorax brown, with dark brown spots. Femora and tibiae of mid and hind legs dark brown at proximal and distal 1/5. The posterior margin of tergite IX cone-like; anal point slender, parallel-sided, apically rounded; superior volsella parallel-sided at base, slightly curved in the middle, widest at 1/3 distance from apex and apically swollen; inferior volsella thumb-like, with slight extension at base; gonostylus curved at 1/4 distance from base, parallel-sided towards apex, rounded apically.
Male (n = 4, unless stated).
Total length 4.24–5.05, 4.59 mm. Wing length 2.00–2.37, 2.12 mm. Total length/wing length 2.12–2.27, 2.17. Wing length/length of profemur 1.95–2.31, 2.20.
Coloration. Thorax brown, with dark brown spots. Femora and tibiae of front legs dark brown except for yellowish brown at distal 3/4, tarsi I dark brown except for yellowish brown at basal, tarsi II–V dark brown; femora and tibiae of mid and hind legs both dark brown at both proximal and distal 1/5, yellowish brown in the middle, tarsi I yellowish brown except for dark yellowish brown at apex, tarsi II–V dark yellowish brown. Abdomen yellowish brown, hypopygium dark yellowish brown.
Head
(Figs
Thorax
(Fig.
Wing (Figs
Legs. Front tibia with three subapical setae, 114–121, 117.5 (2) μm, 129 (1) μm, 138 (1) μm. Combs of midtibia 41–57, 49 μm wide with 21–29, 24 μm long spur, and 42–61, 54 μm wide with 32–39, 35 μm long spur; combs of hind tibia 28–48, 36 μm wide with 28–36, 34 μm long spur, 80–92, 84 μm wide with 40–50, 44 μm long spur. Tarsus I of mid leg with three sensilla chaetica; tarsus I of hind leg with three sensilla chaetica. Lengths (in μm) and proportions of legs as in Table
Lengths (in μm) and proportions of legs of Cryptochironomus parallelus Liu, sp. nov., adult males (n = 4, unless stated).
fe | ti | ta1 | ta2 | ta3 | |
P1 | 871–1112, 945 | 675–770, 710 | 1091–1125, 1111 (3) | 491–553, 520 (3) | 383–431, 413(3) |
P2 | 728–990, 831 | 753–872, 793 | 440–540, 477 | 224–306, 255 | 160–218, 191 |
P3 | 845–1083, 960 | 922–1124, 990 | 704–710, 707 (3) | 338–382, 353 (3) | 270–296, 281 (3) |
ta4 | ta5 | LR | BV | SV | |
P1 | 300–334, 316 (3) | 153–167, 160 (3) | 1.60–1.62, 1.61 (3) | 1.85–1.99, 1.91 (3) | 1.40–1.45, 1.42 (3) |
P2 | 106–152, 124 | 89–120, 104 | 0.58–0.62, 0.60 | 3.01–3.42, 3.13 | 3.29–3.50, 3.40 |
P3 | 158–180, 168 (3) | 107–109, 108 (3) | 0.72–0.77, 0.75 (3) | 2.81–2.83, 2.83 (3) | 2.52–2.84, 2.64 (3) |
Hypopygium
(Figs
From the Latin parallelus, parallel-sided, referring to the character of anal point, adjective in the nominative singular.
Cryptochironomus parallelus Liu, sp. nov. resembles Cryptochironomus digitatus Malloch, 1915 and Cryptochironomus blarina Townes, 1945 by having similar shapes of the posterior margin of tergite IX, inferior volsella and gonostylus. However, it can be distinguished from C. digitatus by the following combination of characters: small frontal tubercles, AR 1.99–2.37, Wing length 2.00–2.37, 2.12 mm, anal point 54–65, 59 μm long and is slender with parallel sides, apically rounded in C. parallelus Liu, sp. nov.; whereas large frontal tubercles, AR 3.14–3.23, superior volsella with a hook-shaped base, and anal point constricted at the 1/3 distance from the base in C. digitatus; whereas large frontal tubercles, AR 3.7, Wing length 3.5 mm, anal point 230 μm long and is gradually tapered and pointed apically in C. blarina.
Holotype. one male, USA, New Mexico State, Colfax Country, Canadian River at Taylor Springs, 36°32'74"N, 104°49'22"W, 15.VII.1974, sweep net, leg: Sta. C. Paratypes. 2 males, New Mexico State, Colfax Country, Canadian River at Taylor Springs, 15.VII.1974, sweep net, leg: Sta. C.
Frontal tubercles absent; squama with 19 setae; anal point parallel-sided, slightly widening at apex; superior volsella crescent-like, with slender extension at the base, curved in the middle and apically swollen; inferior volsella triangular, widest at base, completely covered by superior volsella; gonostylus gradually tapered and pointed apically.
Male (n = 3, unless stated).
Total length 4.65–5.49, 5.01 mm. Wing length 2.31–2.59, 2.42 mm. Total length/wing length 1.98–2.12, 2.07. Wing length/length of profemur 2.00–2.79, 2.38.
Thorax light yellow-brown, with yellow-brown spots. Femora of front legs yellow-brown except for dark yellow-brown at both ends, tibia dark brown, tarsomeres lost; femora and tibiae of mid and hind legs yellow-brown, tarsus I–V yellow-brown to dark brown gradually. Abdomen yellow-brown, hypopygium dark brown.
Head
(Fig.
Thorax
(Fig.
Wing
(Figs
Legs. Fore leg bearing three subapical setae, 142 (1) μm, 144 (1) μm, 146 (1) μm long. Combs of mid tibia 40–60, 52 μm wide with 24–34, 29 μm long spur, and 45–69, 60 μm wide with 43–56, 49 μm long spur; combs of hind tibia 51–56, 53 μm wide with 34–39, 37 μm long spur, 97–105, 100 μm wide with 51–56, 53 μm long spur. Tarsus I of mid leg with 3–5, 4 sensilla chaetica; tarsus I of hind leg with 3–5, 4 sensilla chaetica. Lengths (in μm) and proportions of legs as in Table
Lengths (in μm) and proportions of legs of Cryptochironomus taylorensis Liu, sp. nov., adult males (n = 3, unless stated).
fe | ti | ta1 | ta2 | ta3 | |
P1 | 830–1173, 1034 | 841–857, 848 | 1283 (1) | 669 (1) | 505 (1) |
P2 | 815–1062, 950 | 899–952, 927 | 539–583, 556 | 242–267, 251 | 157–183, 167 |
P3 | 994–1142, 1050 | 1127–1285, 1197 | 807–817, 812 (2) | 381–383, 382 (2) | 302–310, 306 (2) |
ta4 | ta5 | LR | BV | SV | |
P1 | 418 (1) | 203 (1) | 1.53 (1) | 1.65 (1) | 1.30 (1) |
P2 | 103–115, 109 | 88–108, 98 | 0.59–0.61, 0.60 | 3.79–4.03, 3.89 | 3.18–3.49, 3.37 |
P3 | 184–201, 193 (2) | 114–129, 122 (2) | 0.69–0.72, 0.70 (2) | 2.95–2.97, 2.96 (2) | 2.65–2.66, 2.66 (2) |
Hypopygium
(Figs
Name after the type locality, the Canadian River at Taylor Springs; noun in nominative case.
Cryptochironomus taylorensis Liu, sp. nov. bears resemblance to C. curryi Mason, 1986 due to its similar shapes of the anal point, superior volsella, and anal tergite bands. However, it can be distinguished from C. curryi by the following combination of characters: R with 15–18 setae, inferior volsella triangular, and gonostylus pointed apically in this new species; whereas R with 40–46 setae, inferior volsella finger-shaped, and gonostylus rounded apically in C. curryi.
1 | Anal point broad and flat | 2 |
– | Anal point narrow | 3 |
2 | Mesal apical margin of gonostylus emarginate | C. scimitarus Townes, 1945 |
– | Mesal apical margin of gonostylus straight or nearly straight | C. sorex Townes, 1945 |
3 | Mesal apical margin of gonostylus emarginate | C. argus Roback, 1957 |
– | Mesal apical margin of gonostylus not emarginate | 4 |
4 | AR ~ 3.7, Wing length 3.1–3.4 mm | 5 |
– | AR ~ 2.63, Wing length 2.43–2.65 mm | 6 |
5 | Anal point gradually tapered and pointed apically | C. blarina Townes, 1945 |
– | Anal point widest at base | C. beardi Liu, sp. nov. |
6 | Gonostylus widened towards apex, frontal tubercles present | 7 |
– | Gonostylus not distinctly widened towards apex, frontal tubercles mostly absent | 9 |
7 | AR 4.35–5.12, LR1 1.12–1.21, gonostylus strongly widened towards apex | C. stylifera Johannsen, 1908 |
– | AR 1.99–4.03, LR1 1.23–1.62, gonostylus slightly widened towards apex | 8 |
8 | Frontal tubercles large, AR 3.14–3.23, the posterior margin of tergite IX shoulder-shaped | C. digitatus Malloch, 1915 |
– | Frontal tubercles small, AR 1.99–2.37, the posterior margin of tergite IX cone-like | C. parallelus Liu, sp. nov. |
9 | LR1 1.31–1.67, frontal tubercles absent | 10 |
– | LR1 1.45–2.02, when lower than 1.7 frontal tubercles present | 12 |
10 | Wing length 4.99–5.78 mm; LR1 1.48–1.67 | C. eminentia Mason, 1986 |
– | Wing length 2.30–4.75 mm; LR1 1.31–1.55 | 11 |
11 | Wing length 3.01–4.75 mm, AR 2.64–3.40, LR1 1.34–1.55 | C. ramus Mason, 1986 |
– | Wing length 2.31–2.59 mm, AR 2.68–2.90, LR1 1.53 | C. taylorensis Liu, sp. nov. |
12 | Wing length 1.47–1.8 mm, gonostylus only ~ 2× as long as wide | 13 |
– | Wing length 1.9–5.6 mm, gonostylus at least 2.5× as long as wide | 14 |
13 | Wing length 1.8 mm, AR 2.75, frontal tubercles present | C. parafulvus Beck & Beck, 1964 |
– | Wing length 1.47–1.69 mm, AR 2.29–2.49, frontal tubercles absent | C. ferringtoni Liu, sp. nov. |
14 | Gonostylus ~ 2.8–3.2× as long as wide, anal point slightly spatulate apically, frontal tubercles absent | C. ponderosus (Sublette, 1964) |
– | Gonostylus ~ 2.7× as long as wide, anal point tapering parallel-sided or slightly spatulate, frontal tubercles present or absent | 15 |
15 | Frontal tubercles absent, wing length 2.5–5.6 mm | 16 |
– | Frontal tubercles present, wing length 5.1–5.3 mm | 17 |
16 | Gonostylus rounded apically, wing length 2.5–5.6 mm | C. curryi Mason, 1986 |
– | Gonostylus tapering to the apex, wing length 2.0 mm | C. absum Liu, sp. nov. |
17 | Wing length 5.1–5.3 mm, AR 2.5–2.6 | C. conus Mason, 1986 |
– | Wing length 1.8–3.2 mm; AR 2.5–3.4 | 18 |
18 | LR1 1.56–1.73, anal point slightly spatulate apically | C. imitans Sæther, 2009 |
– | LR1 1.60–2.02, anal point parallel-sided | 19 |
19 | Thorax with dark brown spots, inferior volsella not dentate at base | C. fulvus Johannsen, 1905 |
– | Thorax without spots, inferior volsella dentate at base | C. dentatus Liu, sp. nov. |
We want to thank Dr. Leonard Charles Ferrington Jr. (Department of Entomology, University of Minnesota) for checking the specimens and providing much input at various levels of this work. We are grateful to Dr. Alexander Egan (University of Minnesota, USA), who corrected some grammatical errors on a previous version of this manuscript. Financial support from the National Natural Science Foundation of China (32370489, 32170473, 32100402), Natural Science Foundation of Tianjin Science and Technology Correspondent (23KPHDRC00240, 22KPXMRC00070, 20JCQNJC00420) and Yinshanbeilu Grassland Eco-hydrology National Observation and Research Station, China Institute of Water Resources and Hydropower Research, Beijing 100038, China, Grant NO. YSS202308. Are acknowledged with thanks.
The authors have declared that no competing interests exist.
No ethical statement was reported.
Financial support from the National Natural Science Foundation of China (32370489, 32170473, 32100402), Natural Science Foundation of Tianjin Science and Technology Correspondent (23KPHDRC00240, 22KPXMRC00070, 20JCQNJC00420) and Yinshanbeilu Grassland Eco-hydrology National Observation and Research Station, China Institute of Water Resources and Hydropower Research, Beijing 100038, China, Grant NO. YSS202308.
Wenbin Liu: Conceptualization, Funding acquisition, Writing – original draft. Chengyan Wang: Formal analysis, Data curation. Yaning Tang: Software. Ying Wang: Data curation, Visualization, Wenxuan Pei Data curation. Chuncai Yan: Writing – review and editing.
Wen-Bin Liu https://orcid.org/0000-0001-6916-0692
All of the data that support the findings of this study are available in the main text.