Research Article |
Corresponding author: Yue Fu ( fuyue2007915@yahoo.com ) Corresponding author: Yun-Li Xiao ( xiaoyunli0817@126.com ) Academic editor: Fabio Laurindo da Silva
© 2022 Yue Fu, Xiang-Liang Fang, Xin-Hua Wang, Mi Shen, Yun-Li Xiao.
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:
Fu Y, Fang X-L, Wang X-H, Shen M, Xiao Y-L (2022) Corynoneura Winnertz species from Hunan Province, Oriental China, delineated with morphological and 16S rDNA data (Diptera, Chironomidae). ZooKeys 1082: 87-102. https://doi.org/10.3897/zookeys.1082.73019
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The genus Corynoneura Winnertz, 1846 from Hunan Province in Oriental China is reviewed. Four new species, C. enormis Fu sp. nov., C. gibbera Fu sp. nov., C. incuria Fu sp. nov., and C. longshanensis Fu sp. nov. are described and illustrated based on adult males. Sequence data from the 16S rDNA gene were used to infer relationships between these species and complement morphological delineation. Sequences from the mitochondrial large ribosomal subunit (16S rDNA) from these species are uploaded to the National Center for Biotechnology Information (NCBI). Relationships were inferred using the Neighbor-Joining method based on 16S rDNA.
Mitochondrial gene, morphology, non-biting midge, taxonomy
Corynoneura was erected by
Prior to this study, there were 107 valid species in the world, including 47 species from the Palearctic Region, 19 species from the Nearctic Region, 25 species from the Neotropical Region, 27 species from the Oriental Region, four species from the Afrotropical Region, and five species from the Australasian Region (
Previously, only one species, namely Corynoneura prominens Fu, Sæther & Wang, 2009 was recorded in Hunan Province. In this study, four new species are described and illustrated based on the new material from Hunan Province: Corynoneura enormis sp. nov., Corynoneura gibbera sp. nov., Corynoneura incuria sp. nov. and Corynoneura longshanensis sp. nov.. The female of C. incuria sp. nov., associated with the male by 16S rDNA, is described and illustrated.
Adults were mainly collected in the habitats of small streams and lakes next to mountain forests. Adults were collected by light traps near the water body or swept from marginal vegetation beside natal aquatic sites. The specimens were preserved in 85% ethanol, and stored in the dark at 4 °C before molecular analyses. Total genomic DNA of specimens was extracted from the thorax and legs using Qiagen DNA Blood & Tissue Kit. The standard protocol of the Qiagen DNeasy Blood & Tissue Kit was used, except that the final elution volume was 100 𝜇L due to the small specimen size. PCR amplification of the mitochondrial 16S ribosomal RNA gene was carried out with the primers and temperature regimes given in
Measurements are given as ranges followed by the mean, when three or more specimens were measured. The specimens examined in this study are deposited at the College of Biology and Agricultural Resources, Huanggang Normal University (HNU), Huanggang, China.
AR antennal ratio = length of ultimate flagellomere/combined lengths of flagellomeres one to penultimate;
VR venarum ratio;
Cu cubitus;
P1, P2, P3 fore, middle, and hind legs, respectively;
fe femur;
ti tibia;
ta tarsomere;
LR leg ratio (ratio of metatarsus to tibia in front leg);
BV Bein ratio (length of (femur + tibia + ta1) /length of (ta2 + ta3 + ta4 + ta5));
SV Schenkel-Schiene ratio (length of (femur + tibia) / length of ta);
BR bristle ratio (ratio of longest seta on ta1 to minimum width of ta1 measured one third from apex);
HR hypopygium ratio = gonocoxite length / gonostylus length;
HV hypopygium value =body length / gonostylus length × 10.
Measurements and ratios of hind tibia follow
a Maximum width;
b Length of ventral elongation;
c1 Length of strong broad part, measured from apex;
c2 Total length of broadening;
d Width of tibia basally to the apical broadening.
Corynoneura Winnertz, 1846: 12.
Holotype , male (HNU: 17091206HJL), China: Hunan Province, Loudi City, Xinhua County, Xihe Town, Cushi Village, 27°51'45"N, 111°31'51"E, 315 m a. s. l., 29.Ⅶ.2016, sweep net, leg. Jingli Huang.
From Latin, enormis, immense, huge, vast, referring to lateral sternapodeme with a large attachment point.
The male imago is characterized by having an antenna with eight flagellomeres, AR 0.51; anterior margin of cibarial pump distinctly concave; hind tibia with hooked spur; superior volsella small rounded and undeveloped; inferior volsella narrow, with dented edge, along the inner margin of gonocoxite; phallapodeme apically curved, placed in lateral position of sternapodeme; sternapodeme curved into a U-shape, and lateral sternapodeme with large caudal attachment point.
Adult males (N = 1). Total length 0.92 mm. Wing length 0.53 mm. Total length/wing length ratio 1.74. Wing length/profemur length ratio 2.69.
Coloration. Head dark brown; thorax dark brown. Legs yellowish. Abdomen brown.
Head.
Antenna with eight flagellomeres, AR 0.51, ultimate flagellomere 115 µm long, with many short apical sensilla chaetica (Fig.
Thorax. Five dorsocentral setae. Scutellum with two setae. One or two prealar setae.
Wing
(Fig.
Legs.
Fore trochanter with dorsal keel. Most of fore- and mid-legs lost. Spurs of hind tibia 25 µm and 10 µm long. Width of hind tibia at apex (a) 29 µm, width of hind tibia 1/3 from apex (d) 18 µm, elongation length (b) 34 µm, length of maximum thickening (c1) 48 µm, total length of thickening (c2) 72 µm; hind tibial ratios: a/d 1.61; b/d 1.89; c1/d 2.61; c2/d 4.00. Hind tibia expanded with comb of 17 setae and S-shaped spur (Fig.
Lengths (in µm) and proportions of leg segments of male Corynoneura enormis sp. nov. (N = 1).
fe | ti | ta 1 | ta 2 | ta 3 | ta 4 | ta 5 | LR | BV | SV | BR | |
---|---|---|---|---|---|---|---|---|---|---|---|
P1 | 197 | 240 | 130 | 71 | 42 | 20 | 28 | 0.54 | 3.52 | 3.36 | 1.80 |
P2 | 216 | 245 | 145 | 70 | 35 | 16 | 27 | 0.59 | 4.09 | 3.17 | 2.00 |
P3 | 216 | 211 | 113 | 67 | 29 | 19 | 29 | 0.54 | 3.75 | 3.78 | 1.80 |
Hypopygium
(Fig.
This species is similar to Corynoneura ascensa Fu & Sæther, 2012 and Corynoneura sesquipedalis Fu & Fang, 2018 by having a large attachment point on the lateral sternapodeme. The new species can be separated from C. ascensa by having antenna with 8 flagellomeres, narrow and undeveloped inferior volsella; and differs from C. sesquipedalis by having a narrow inferior volsella, transverse sternapodeme present and with an oral projection (broad inferior volsella, transverse sternapodeme V-shaped,without transverse part in C. sesquipedalis). The sequence of 16S rDNA from this species is highly similar to Corynoneura tumula Fu & Fang, 2018, but there are distinct morphological differences between them: in C. enormis the antenna has 8 flagellomeres, AR 0.51; inferior volsella narrow, lateral sternapodeme with large caudal attachment point, while C. tumula has an antenna with 9 flagellomeres, AR 0.46; inferior volsella relatively broad, lateral sternapodeme with small caudal attachment point.
Holotype male (HNU: 17090801HJL), China: Hunan Province, Huaihua City, Hecheng County, Wushui River, Xiyi Bridge, 27°33'29"N, 109°57'41"E, 259 m a. s. l., 23.VII.2016, light trap, leg. Haixia Shi. 6 males (HNU: 17090902HJL, 17090904HJL, 17091004HJL, 17091003HJL, 17090804HJL, 17090803HJL), 21–23.VII. 2016, as holotype.
From Latin, gibbera, protuberant, referring to the prominent inferior volsella.
The male imago is characterized by having an antenna with nine or ten flagellomeres, AR 0.43–0.57, 0.52; superior volsella triangular; inferior volsella prominent, like a small rectangle, and placed caudally of gonocoxite; transverse sternapodeme inverted U-shaped; phallapodeme scalpel-like, in caudal position of sternapodeme.
Adult male (N = 7). Total length 0.82–1.10, 0.95 mm. Wing length 0.45–0.66, 0.57 mm. Total length/wing length ratio 1.67–1.82, 1.70. Wing length/profemur length ratio 2.32–3.04, 2.79.
Coloration. Head brown, with dark brown eyes; thorax dark brown; legs yellowish; tergites I-V yellowish, VI-IX brownish.
Head.
Antenna with nine or ten flagellomeres, AR 0.43–0.57, 0.52, ultimate flagellomere 98–144, 124 µm long, slightly expanded apically, with many short apical sensilla chaetica (Fig.
Thorax
(Fig.
Wing
(Fig.
Legs.
Fore trochanter with dorsal keel. Spurs of fore tibia 17–24, 19 µm long and 7–12, 10 µm long, spurs of mid tibia 7–10, 8 and 10–12, 11 µm long, and spurs of hind tibia 22–29, 24 µm long and 12–14, 13 µm long. Width of fore tibia at apex 17–22, 19 µm, of mid tibia 12–19, 15 µm, of hind tibia (a) 17–24, 22 µm. Width of hind tibia 1/3 from apex (d) 17–22, 20 µm, elongation length (b) 31–43, 36 µm, length of maximum thickening (c1) 60–72, 66 µm, total length of thickening (c2) 84–120, 95 µm; hind tibial ratios: a/d 1.41–2.00, 1.60; b/d 1.41–2.39, 2.10; c1/d 2.73–4.00, 3.75; c2/d 3.83–6.67, 5.62. Hind tibia expanded, with comb of 15–19, 16 setae, with S-shaped spur (Fig.
Lengths (in µm) and proportions of legs segments of male Corynoneura gibbera sp. nov. (N = 7).
fe | ti | ta 1 | ta 2 | ta 3 | ta 4 | |
---|---|---|---|---|---|---|
P1 | 194–225, 208 | 225–255,245 | 120–146,131 | 70–82, 73 | 38–46,42 | 19–22, 20 |
P2 | 265–323,297 | 245–284, 265 | 144–176, 162 | 65–79, 72 | 34–36, 35 | 14–19, 17 |
P3 | 225–265, 243 | 235–284, 255 | 120–146, 132 | 70–77, 75 | 29–31, 30 | 17–19, 18 |
ta5 | LR | BV | SV | BR | ||
P1 | 26–31, 29 | 0.51–0.56,0.53 | 3.35–3.75, 3.55 | 3.32–3.68, 3.48 | 1.30–2.20, 1.70 | |
P2 | 26–31, 29 | 0.59–0.62, 0.61 | 4.61–4.88, 4.74 | 3.40–4.16, 3.64 | 1.50–2.20, 1.80 | |
P3 | 26–31, 29 | 0.51–0.54, 0.52 | 3.91–4.46, 4.15 | 3.56–3.92, 3.76 | 2.00–2.20, 2.10 |
Hypopygium
(Fig.
This species is closely related to Corynoneura macula Fu & Sæther, 2012 by having similarly shaped inferior volsella and an inverted U-shaped sternapodeme. The new species can be separated from the latter by having AR 0.43–0.57, 0.52, gonostylus relatively long and slender, apically curved, while C. macula has a yellowish antenna with a dark brown apical spot, AR 0.27–0.37, and the gonostylus is relatively short and strongly curved. The new species is also similar to Corynoneura aurora Makarchenko & Makarchenko, 2010 by having similar inferior volsella, the same shaped sternapodeme and phallapodeme, but differs from the latter by the antenna having 12 flagellomeres, and the gonostylus being slightly convex on the outer edge in C. aurora.
Holotype male (HNU: 17090903HJL), China: Hunan Province, Huaihua City, Hecheng County, Wushui River, Xiyi Bridge, 29°33'29"N, 109°57'41"E, 259 m a. s. l., 23.VII.2016, light trap, leg. Jingli Huang. Paratype: 1 female (HNU:17091205HJL), CHINA: Hunan Province, Loudi City, Xinhua County, Xihe Town, Cushi Village, 27°51'45"N, 111°31'51"E, 315 m a. s. l., 29.Ⅶ.2016, sweep net, leg. Jingli Huang.
From Latin, incuria, neglect, referring to the inferior volsella being absent and fused with the inner margin of gonocoxite.
The male imago is characterized by having antenna with eleven flagellomeres, AR 0.31; anterior margin of cibarial pump strongly concave; superior volsella developed and with right-angled corner; inferior volsella almost absent, fused with the inner margin of gonocoxite; transverse sternapodeme curved into U-shaped; phallapodeme scalpel-like, apical slightly curved, placed caudal position of sternapodeme. The female imago is characterized by coxosternapodeme with a single transparent, well-developed lamella.
Ault male (N = 1). Total length 1.08 mm. Wing length 0.63 mm. Total length/wing length ratio 1.70.
Coloration. Head and thorax brown, eyes dark brown. Legs pale yellow. Abdominal tergites I-V yellowish, VI-IX yellow-brown.
Head.
Antenna with eleven flagellomeres, AR 0.31, ultimate flagellomere 96 µm long, slightly expanded apically, with many apical sensilla chaetica (Fig.
Thorax
(Fig.
Wing
(Fig.
Legs.
Fore trochanter with dorsal keel. Spurs of fore tibia 12 µm long and 7 µm long, spurs of mid tibia 7 and 9 µm long, and spurs of hind tibia 31 µm long and 10 µm long. Width of fore tibia at apex 17 µm, of mid tibia 17 µm, of hind tibia (a) 36 µm. Width of hind tibia 1/3 from apex (d) 19 µm, elongation length (b) 38 µm, length of maximum thickening (c1) 60 µm, total length of thickening (c2) 103 µm; hind tibial ratios: a/d 1.89; b/d 2.00; c1/d 3.16; c2/d 5.42. Hind tibia expanded, with comb of 16 setae, with S-shaped spur (Fig.
Lengths (in µm) and proportions of legs segments of male Corynoneura incuria sp. nov. (N = 1).
fe | ti | ta 1 | ta 2 | ta 3 | ta 4 | ta 5 | LR | BV | SV | BR | |
---|---|---|---|---|---|---|---|---|---|---|---|
P1 | 228 | 255 | 134 | 72 | 41 | 22 | 26 | 0.53 | 3.83 | 3.60 | 1.90 |
P2 | 323 | 265 | 158 | 73 | 34 | 16 | 28 | 0.60 | 4.96 | 4.35 | 2.00 |
P3 | 255 | 265 | 134 | 77 | 31 | 17 | 31 | 0.51 | 4.19 | 3.88 | 2.00 |
Hypopygium
(Fig.
Adult female (N = 1). Total length 0.83 mm. Wing length 0.59 mm. Total length/wing length ratio 1.41. Wing length/profemur length ratio 3.61.
Coloration. Head, eyes, and thorax brown. Legs pale yellow. Abdomen yellowish brown.
Head. Tentorium 72 µm long; 7 µm wide. Clypeus with four setae.
Thorax
(Fig.
Wing
(Fig.
Legs.
Fore trochanter with dorsal keel. Spurs of fore tibia 10 µm long, spurs of mid tibia 7 and 12 µm long, and spurs of hind tibia 22 µm and 12 µm long. Width fore tibia at apex of 17 µm, of mid tibia 14 µm, of hind tibia (a) 29 µm. Width of hind tibia 1/3 from apex (d) 22 µm, elongation length (b) 31 µm, length of maximum thickening (c1) 60 µm, total length of thickening (c2) 96 µm; hind tibial ratios: a/d 1.32; b/d 1.41; c1/d 2.72; c2/d 4.36. Hind tibia expanded, with comb of 14 setae, with S-shaped spur (Fig.
Lengths (in µm) and proportions of legs segments of female Corynoneura incuria sp. nov. (N = 1).
fe | ti | ta 1 | ta 2 | ta 3 | ta 4 | ta 5 | LR | BV | SV | BR | |
---|---|---|---|---|---|---|---|---|---|---|---|
P1 | 183 | 214 | 106 | 65 | 36 | 17 | 26 | 0.50 | 3.35 | 3.56 | 2.50 |
P2 | 314 | 230 | 146 | 67 | 31 | 17 | 24 | 0.63 | 4.96 | 3.73 | 1.70 |
P3 | 199 | 216 | 113 | 70 | 27 | 17 | 24 | 0.52 | 3.83 | 3.67 | 1.80 |
Genitalia
(Fig.
This species is closely related to Corynoneura tokarapequea Sasa & Suzuki, 1995 by having antenna with eleven flagellomeres, the same shaped sternapodeme and phallapodeme, and a similar gonostylus. The new species can be separated from the latter by having AR 0.31, the inferior volsella almost absent and fused with the inner margin of the gonocoxite, while C. tokarapequea has AR 0.62–0.70, the inferior volsella obvious and near rectangular. The new species is also similar to Corynoneura floridaensis Fu & Sæther, 2012 by the antenna with eleven flagellomeres, AR 0.36, same shaped sternapodeme and phallapodeme, but differs from the latter by having a thick transverse sternapodeme, and the gonostylus is strongly curved in C. floridaensis.
Holotype male (HNU: 17091204HJL), China: Hunan Province, Loudi City, Lianyuan County, Longshan National Forest Park, 27°31'20"N, 111°45'23"E, 674 m a. s. l., 26.Ⅶ.2016, sweep net, leg. Jingli Huang.
Named after the type locality.
The male imago is characterized by having an antenna with seven flagellomeres, AR 0.55; superior volsella undeveloped, and inferior volsella with right-angular corner, fused with the inner margin of gonocoxite; sternapodeme inverted U-shaped; phallapodeme scalpel-like, apical curved, in caudal position of sternapodeme.
Adult male (N = 1). Total length 0.92 mm. Wing length 0.45 mm. Total length/wing length 2.04.
Coloration. Head and thorax brown, legs and abdomen yellowish.
Head.
Antenna with seven flagellomeres, AR 0.55, ultimate flagellomere 89 µm long, ultimate flagellomere distinctly expanded apically, with about 10 apical sensilla chaetica (Fig.
Thorax. Five dorsocentral setae. Scutellum with two setae.
Wing
(Fig.
Legs.
Fore legs lost. Spurs of mid tibia 5 µm and 8 µm long, and spurs of hind tibia 19 µm and 10 µm long. Width of mid tibia at apex 14 µm, of hind tibia (a) 29 µm. Width of hind tibia 1/3 from apex (d) 14 µm, elongation length (b) 36 µm, length of maximum thickening (c1) 60 µm, total length of thickening (c2) 79 µm; hind tibial ratios: a/d 2.07; b/d 2.57, 2.10; c1/d 4.29; c2/d 5.64. Apex of hind tibia obvious expanded, with comb of 16 setae, with S-shaped spur (Fig.
Lengths (in µm) and proportions of legs segments of male Corynoneura longshanensis sp. nov. (N = 1).
fe | ti | ta 1 | ta 2 | ta 3 | ta 4 | ta 5 | LR | BV | SV | BR | |
---|---|---|---|---|---|---|---|---|---|---|---|
P1 | 160 | 180 | 106 | 48 | 26 | 16 | 24 | 0.59 | 3.91 | 3.21 | 1.90 |
P2 | 209 | 182 | 110 | 48 | 24 | 14 | 22 | 0.60 | 4.64 | 3.55 | 1.70 |
P3 | 175 | 199 | 91 | 50 | 19 | 12 | 24 | 0.46 | 3.86 | 4.11 | 1.80 |
Hypopygium
(Fig.
This new species is similar to Corynoneura hortonensis Fu & Sæther, 2012 by having the same shaped sternapodeme and phallapodeme. The new species can be separated from the latter by the broad and thick transverse sternapodeme, and the median part of the gonostylus expanded with a rugged inner edge.
The primary structure of the mitochondrial large ribosomal subunit (16S rDNA) gene is conservative, while the secondary structure shows spiral differences, which are more suitable for systematic studies of species and genera (
Neighbor-joining Kimura 2 parameter tree based on 16S rDNA of five Corynoneura species and Thienemanniella convexa Fu in
The four new species referred in this study share the same morphological features: a transverse sternapodeme inverted U-shape, and the attachment point for the phallapodeme is placed in a caudal position of the sternapodeme. According to
The project was supported by the National Natural Science Foundation of China (NSFC) (Grant No. 32070483, 31101624, 31460572), Natural Science Foundation of Hubei Province (Grant No. 2020CFB757), Scientific Research Starting Foundation for Ph. D. of Huanggang Normal University (Grant No.2020010), Huanggang Normal University Team Project (4022019006).