2urn:lsid:arphahub.com:pub:45048D35-BB1D-5CE8-9668-537E44BD4C7Eurn:lsid:zoobank.org:pub:91BD42D4-90F1-4B45-9350-EEF175B1727AZooKeysZK1313-29891313-2970Pensoft Publishers10.3897/zookeys.706.1353513535ChecklistArthropodaCeratopogonidaeCulicomorphaDipteraHexapodaInsectaInvertebrataNematoceraCatalogues and ChecklistsTaxonomyCenozoicAsiaA species checklist of the subgenus Culicoides (Avaritia) in China, with a description of a new species (Diptera, Ceratopogonidae)ChangQiong Qiong1JiangXiao Hong1LiuGuo Ping2LiXiao Fei1HouXiao Huihxh19801122@163.comhttps://orcid.org/0000-0002-7713-28081Zunyi Medical University, Zunyi 563000, Guizhou Province, ChinaZunyi Medical UniversityZunyiChinaCenter for Disease Control and Prevention of Shenyang PLA Command, Shenyang 110034, Liaoning Province, ChinaCenter for Disease Control and Prevention of Shenyang PLA CommandShenyangChina
201704102017706117135FFB72F5C-FFE1-DA0D-FF93-FFD8FF8BFFCE42231A0D-0FF4-4E5D-B83D-C5309E4DE9C911498580405201721082017Qiong Qiong Chang, Xiao Hong Jiang, Guo Ping Liu, Xiao Fei Li, Xiao Hui HouThis 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.http://zoobank.org/42231A0D-0FF4-4E5D-B83D-C5309E4DE9C9
A checklist of the subgenus Culicoides (Avaritia Fox) (Diptera: Ceratopogonidae: Culicoides) in China, currently including 57 species, is provided. Their full citations, more detailed locations of the type locality, and distribution of each species by province, and/or state of each species are also provided. Culicoides (Avaritia) fenggangensis Liu & Hou, sp. n. is described and illustrated, based on both male and female specimens from China. The new species is compared with its similar congeners, C. (A.) comparis Liu & Yu, 2005 and C. (A.) dentiformis McDonald & Lu, 1972.
Biting midgesCeratopogonidaechecklist, Culicoidessubgenus AvaritiaNational Natural Science Foundation of China501100001809http://doi.org/10.13039/501100001809Citation
Chang QQ, Jiang XH, Liu GP, Li XF, Hou XH (2017) A species checklist of the subgenus Culicoides (Avaritia) in China, with a description of a new species (Diptera, Ceratopogonidae). ZooKeys 706: 117–135. https://doi.org/10.3897/zookeys.706.13535
Introduction
Biting midges of the genus Culicoides Latreille (Diptera, Ceratopogonidae) are found everywhere in the world and females are the smallest of insect vectors (Mellor et al. 2000). Some species of Culicoides spread disease in humans and livestock as vectors of arboviruses, such as bluetongue virus (BTV), Schmallenberg virus (SBV), and others (Borkent 2005), which leads to direct economic costs for agriculture (Gibbens 2012; Velthuis et al. 2010). In China, Bluetongue has been recorded from several provinces in south, such as Yunnan, Hubei, Anhui, Sichuan, Shanxi and so on (Zhang et al. 2015), and the vector Culicoides of BTV mainly distributed in the south region of north latitude 40° (Zhang et al. 2014). Therefore, members of this genus have great international significance and have attracted more scientific attention in recent years. The genus Culicoides, currently with a total number of species of 1415 in the world, 1368 extant species and 47 fossil species (Borkent, 2016), has 348 species in China. Many of the important vectors are in the subgenus Culicoides (Avaritia Fox). The purpose of this paper is to provide a checklist of this subgenus in China, and describe and illustrate Culicoides (A.) fenggangensis sp. n.
Materials and methods
The specimens were collected with light traps near households in the mountains of Fenggang County of Guizhou Province. For microscopic observation, specimens were preserved in 100% ethanol and then slide-mounted in Canada balsam following the technique described by Yu et al. (2005). Diagnostic features were microphotographed using an Imaging System of Upright Research Microscope adapted to a microscope (Nikon Eclipse Ni-E) and a Digital System of Large depth-of-field 3D Digital Microscope (Keyence VHX-1000C), and Photoshop CS4 was used to obtain the final images. Morphological terms are from the chapter on Ceratopogonidae by Yu et al. (2005). Terms of structures specific to Culicoides follow those described by Santarém et al. (2014) and Han et al. (2017). Measurements of wings, flagellar segments, palpus, and legs are given in millimeters, and measurements of spermathecae are given in microns. Meristic information is presented as ranges of values, followed by mean and sample size. The type specimens are deposited in Insect Collection of Zunyi Medical University, Guizhou Province, China.
TaxonomyChecklist of the subgenus Avaritia in China in alphabetical order
Subgenus Avaritia Fox, 1955
Avaritia Fox, 1955: 218.
Type species. Ceratopogonobsoletus Meigen, 1818.
Culicoides (Avaritia) abchazicus Dzhafarov, 1964
Culicoidesabchazicus Dzhafarov, 1964: 263; Liu et al. 2011: 385. Type Locality: Georgia.
Distribution. China (Liaoning); Georgia.
Culicoides (Avaritia) actoni Smith, 1929
Culicoidesactoni Smith, 1929: 255; McDonald and Lu 1972: 411; Lee 1988: 29; Yu et al. 2005: 907. Type locality: India.
Culicoidesokumensis Arnaud, 1956: 119. Type locality: Japan.
Culicoidesimperceptus Das Gupta, 1962a: 538. Type locality: India.
Distribution. China (Heilongjiang, Jiangsu, Anhui, Fujian, Taiwan, Shandong, Hubei, Guangdong, Guangxi, Hainan, Sichuan, Yunnan, Tibet, Shaanxi, Hunan); India, Indonesia, Malaysia, Philippines, Vietnam, Japan, Thailand.
Culicoides (Avaritia) albifascia Tokunaga, 1937
Culicoidesalbifascia Tokunaga, 1937: 319; Lee 1978: 28; Yu et al. 2005: 908. Type locality: China: Taiwan.
Distribution. China (Heilongjiang, Tibet, Sichuan, Yunnan, Taiwan).
Culicoides (Avaritia) bawanglingensis Yu, Wang & Chen, 2012
Culicoidesbawanglingensis Yu, Wang & Chen, in Wang et al., 2012: 283. Type Locality: China: Hainan, Bawangling.
Distribution. China (Hainan).
Culicoides (Avaritia) brevipalpis Delfinado, 1961
Culicoidesbrevipalpis Delfinado, 1961: 654; Yu et al. 2005: 910. Type locality: Philippines.
Distribution. China (Taiwan, Guangdong, Hainan, Yunnan); Australia, Indonesia, Malaysia, Japan, Sri Lanka, Thailand, Philippines.
Culicoides (Avaritia) brevitarsis Kieffer, 1917
Culicoidesbrevitarsis Kieffer, 1917: 187; Yu et al. 2005: 911. Type locality: Australia.
Culicoidesrobertsi Lee & Reye, 1953: 386. Type locality: Australia.
Culicoidesradicitus Delfinado, 1961: 657. Type locality: Philippines.
Culicoidessuperfulvus Das Gupta, 1962b: 253. Type locality: India.
Distribution. China (Anhui, Taiwan, Hainan); Australia, Philippines, India, Laos, Malaysia.
Culicoides (Avaritia) bubalus Delfinado, 1961
Culicoidesbubalus Delfinado, 1961: 658; Yu et al. 2005: 913. Type locality: The Philippines.
Distribution. China (Taiwan), Philippines.
Culicoides (Avaritia) chiopterus (Meigen), 1830
Ceratopogonchiopterus Meigen, 1830: 263. Type locality: Europe.
Ceratopogonamoenus Winnertz, 1852: 35. Type locality: Germany.
Monoheleasimilis Goetghebuer, 1927: 203. Type locality: Belgium.
Culicoidesdobyi Callot & Kremer, 1969: 610. Type locality: France.
Distribution. China (Hebei, Inner Mongolia, Liaoning, Jilin, Heilongjiang, Tibet); Germany, France, Belgium, Britain, Russia.
Culicoides (Avaritia) clavipalpis Mukerji, 1931
Culicoidesclavipalpis Mukerji, 1931: 1052; McDonald and Lu 1972: 403; Yu et al. 2005: 916. Type locality: India.
Culicoidescandidus Sen & Das Gupta, 1959: 620. Type locality: India.
Distribution. China (Jiangsu, Fujian, Shandong, Hainan, Sichuan); India, Indonesia, Laos, Malaysia, Philippines, Thailand.
Culicoides (Avaritia) comparis Liu & Yu, 2005
Culicoidescomparis Liu & Yu, in Yu et al., 2005: 917. Type locality: China: Tibet, Nielamu.
Distribution. China (Tibet).
Culicoides (Avaritia) conaensis Liu & Yu, 1990
Culicoidesconaensis Liu & Yu, 1990: 19; Yu et al. 2005: 919. Type locality: China: Tibet, Cuona.
Culicoidesorestes Wirth & Hubert, 1989: 222; Yu et al. 2005: 959. Type locality: Malaysia.
Distribution. China (Hainan); Malaysia.
Culicoides (Avaritia) orientalis Macfie, 1932
Culicoidesorientalis Macfie, 1932: 490; Lee 1978: 83; Yu et al. 2005: 961. Type locality: Malaysia.
Culicoidesnayabazari Das Gupta, 1963: 35. Type locality: India.
Distribution. China (Fujian, Taiwan, Hainan, Sichuan, Yunnan, Tibet, Hunan); Malaysia, India, Indonesia, Philippines, Thailand, Vietnam, the Solomon Islands.
Culicoides (Avaritia) palauensis Tokunaga, 1959
Culicoidespalauensis Tokunaga, in Tokunaga & Murachi, 1959: 348; Yu 1982: 57; Yu et al. 2005: 963. Type locality: USA.
Distribution. China (Guangdong, Hainan, Yunnan); Oceania.
Culicoides (Avaritia) pastus Kitaoka, 1980
Culicoidespastus Kitaoka, 1980: 11; Lee 1988: 91; Yu et al. 2005: 964. Type locality: Japan.
Distribution. China (Sichuan, Yunnan); Japan.
Culicoides (Avaritia) pelius Liu & Yu, 1990
Culicoidespelius Liu & Yu, 1990: 23; Liu and Deng 2000: 245; Yu et al. 2005: 966. Type locality: China: Tibet, Cuona.
Distribution. China (Tibet).
Culicoides (Avaritia) peregrinus Kieffer, 1910
Culicoidesperegrinus Kieffer, 1910: 191; Yu et al. 2005: 967. Type locality: India.
Culicoidesjudicandus Bezzi, 1916: 8. Type locality: Philippines.
Culicoidesesmoneti Salm, 1917b: 136. Type locality: Indonesia.
Culicoidesphilippinensis Kieffer, 1921: 564. Type locality: Philippines.
Culicoidesassamensis Smith & Swaminath, 1932: 183. Type locality: India.
Culicoidesquadratus Tokunaga, 1951: 108. Type locality: Indonesia.
Distribution. China (Hebei, Inner Mongolia, Liaoning, Jiangsu, Fujian, Taiwan, Jiangxi, Henan, Guangdong, Guangxi, Hainan); India, Philippines, Indonesia.
Male: only species of Culicoides in China with the following combination of features: the 3rd segment of the palpus is slender, PR 3.11; the apex of 9th tergite has lateral processes; parameres with apical portion elongate, bent abruptly; aedeagus nearly triangular, with a long ovoid process at its apex. Female: only species of Culicoides in China with the following combination of features: cell m2 with four sparsely distributed pale spots; the 3rd segment of the palpus is slender, PR 3.20–3.75.
Description.
Female.Head (Fig. 2a). Brown. Eyes (Fig. 2b) contiguous, abutting medially for length of 1.5 ommatidia, with interfacetal hairs. Antennal pedicel brown; Lengths of antennal flagellomeres in proportion of 19: 14: 14: 15: 15: 15: 16: 17: 25: 25: 25: 31: 47; AR 1.22–1.28 (1.24, n = 3); sensilla coeloconica on flagellomeres 1, 9–13. 3rd segment of palpus slender, slightly swollen at apical 1/3, with small, rounded sensory pit (Fig. 2c–d); PR 3.20–3.75 (3.45, n = 3). Mandible with 14–18 (16, n = 4) teeth (Fig. 2e); Maxilla with 17 teeth (n = 3); P/H ratio 1.04–1.17 (1.11, n = 3).
Culicoides (A.) fenggangensis sp. n. Left Wing. a Female b Male.
https://binary.pensoft.net/fig/159425
Thorax (Fig. 2f). Dark brown. Scutellum without distinct pattern in slide-mounted specimens. Wing (Fig. 1a) with contrasting pattern of pale, dark spots; distal 1/2 of 1st, proximal 1/2 of 2nd radial cell in dark spot; three pale spots near anterior margin: 1st over base of cell r1 and r-m crossvein extending from below M1 to margin of costa, 2nd over distal of cell r2 from dorsal portion of M1 to costa, 3rd morphological variation, from dorsal portion of M1 to just below costa; cell m1 with two separated pale spots: one spot small, ovoid, another large, triangle, far from distal portion of wing; cell m2 with four different shapes, sizes pale spots: 1st proximal to CuA, 2nd between medial and mediocubital forks, 3rd and 4th below M2, latter reaching wing margin; cua1 with a large, ovoid pale spot abutting wing posterior margin; anal cell with two pale spots: proximal pale spot on Cu2 and CuA, distal pale spot near mediocubital fork; wing base with faint pale spot on M; macrotrichia sparsely distributed on distal 1/3 of wing, but not in basal cell; wing length 1.45–1.60 (1.52, n = 3) mm, width 0.68–0.73 (0.70, n = 3) mm; CR 0.55–0.59 (0.57, n = 3). TR and F-T of legs (Fig. 2f) are given as Table 1, metatibial distal bristles (Fig. 2g–h) with 5 or 6 spines, 1st spine is longest.
Tarsal ratios (TR) and measurements of leg segments and tarsomeres from femur to tarsomere 5 (F-T) of all legs of C.fenggangensis sp. n. (♀).
a–jCulicoides (A.) fenggangensis sp. n. a Head, anterior view b Eye contiguous, anterior view c Left palpus, anterior view d Right palpus, anterior view e Mandibular teeth f Thorax and legs, lateral view g Apex of hind tibia and base of first tarsomere, posterior view. h Apex of hind tibia and base of first tarsomere, posterior view. i Spermathecae j Genitalia, ventral view (Female: a–c, e–i; Male: d, j).
https://binary.pensoft.net/fig/159426
Male. Similar to female with usual sexual differences. Sensilla coeloconica on flagellomeres 1, 11–13; AR 0.87 (n = 1); PR 3.11 (n = 1). Wing with pattern of pale spots as in Fig. 1b, wing length 1.58 (n = 1), width 0.55 (n = 1); CR 0.57 (n = 1). TR and F-T of legs are given as Table 2. Genitalia (Fig. 2j): 9th tergite squarish, distal portion flat with short, conical processes at apicolateral. Ninth sternite with broad, deep, semicircle caudomedian excavation. Gonocoxite twice as long as broad, sclerotized; gonostylus tapering distally, distal portion curved. Parameres (Fig. 3b) separate, contiguous in midportions; each with moderately long, slender basal arm, swollen at base, stem long, slightly curved near base; apical portion tapered, elongate, abruptly bent without lateral fringe of spicules. Aedeagus (Fig. 3a) nearly triangle, basal arms short, unciform, tapering toward end, basal arch low, extending to 1/5 of total length, distal process 1/6 total length, long ovoid process at apex.
Culicoides (A.) fenggangensis sp. n. a Genitalia and aedeagus (parameres absent), ventral view b parameres, ventral view.
https://binary.pensoft.net/fig/159427
TR and F-T of all legs of C.fenggangensis sp. n. (♂).
Leg
TR
F-T
Foreleg
2.67
95 : 92 : 56: 21 : 15 : 8 : 9
Midleg
1.94
115 : 112 : 64 : 33 : 16 : 8 : 9
Hindleg
1.49
117 : 117 : 55 : 37 : 19 : 9 : 10
Type material. Holotype female, Chongxin village, Yong’an town, Fenggang county, Zunyi city, Guizhou province, China (28°06'31.49"N, 107°35'57.94"E), 12. IV. 2016, alt. 908m, Qiongqiong Chang col. Paratypes: 2 males and 3 females, same data as holotype.
Distribution.
China (Guizhou Province).
Etymology.
This species is named in tribute to Fenggang county, where the specimens were collected.
Taxonomic discussion
Culicoides (Avaritia) fenggangensis sp. n. is very similar to C. (A.) comparis Liu & Yu, 2005 and C. (A.) dentiformis McDonald & Lu, 1972 based on the interfacetal hairs on the eyes and sensilla coeloconica on the flagellomeres. C.fenggangensis sp. n. can be distinguished from these two congeners by the number and distribution of pale spots on the wing (cell m2 with two and three pale spots respectively in C.dentiformis and C.comparis) (Yu et al. 2005), elongate and cylindrical third palpus segment (third palpus segment is swollen in C.comparis and C.dentiformis). Females of C.fenggangensis sp. n. have a different wing size (wing length 1.33 mm and width 0.63 mm in C.comparis and wing length 1.78 mm and width 0.76 mm in C.dentiformis), distribution of the heavy macrotrichia on the wing (respectively distal 4/5 and 1/4 of the wing in C.comparis and C.dentiformis), they have a more slender 3rd palpus segment compared to the most species of subgenus Avaritia (only 12 species PR > 3.2), PR 3.2–3.75 (respectively PR 2.27 and 2.5 in C.comparis and C.dentiformis), different size of spermathecae (measuring 45.0 × 40.0 μm in C.comparis and 60.0 × 45.0 μm in C.dentiformis). Because male of C.comparis is unknown, the new species will only compare with C.dentiformis. Males of C.fenggangensis sp. n. have a different shape and structure of genitalia, with two lateral processes on the distal portion of ninth tergite (without lateral process in C.dentiformis), parameres apical portion tapered and abruptly bent (linear in C.dentiformis), long ovoid process at apex of aedeagus (diamond-shape process in C.dentiformis). Therefore, the distinctive features to separate C.fenggangensis sp. n. from others are cell m1 and m2 with 2 and 4 pale spots respectively.
The male of C.fenggangensis keys to C.dentiformis in Yu et al. (2005) where it may be distinguished by the presence of conical apicolateral processes on tergite 9 which are absent in C.dentiformis. The female of C.fenggangensis keys to the couplet with C.dentiformis and C.comparis in Yu et al. (2005) where it may be distinguished by the presence of 4 pale spots in cell m2 which are absent in C.dentiformis and C.comparis.
The biogeographical territory of China spans the Palaearctic and Oriental Regions, which results in a rich diversity of biting midge species. The species of the subgenus Culicoides (Avaritia) are distributed in most provinces of China, except Qinghai and Ningxia. There are 15 species distributed in the Palaearctic Region, accounting for 26.3% of the total (C.abchazicus, C.actoni, C.albifascia, C.chiopterus, C.comparis, C.incertus, C.longirostris, C.nielamensis, C.nigritus, C.obsoletus, C.peregrinus, C.scoticus, C.sinanoensis, C.tainanus, C.wandashanensis). There are 49 species present in the Oriental Region, accounting for 86.0% of the total (C.actoni, C.albifascia, C.bawanglingensis, C.brevipalpis, C.brevitarsis, C.bubalus, C.clavipalpis, C.conaensis, C.dentiformis, C.elongates, C.filicinus, C.gaponus, C.holcus, C.hui, C.longirostris, C.imicola, C.innoxius, C.insignipennis, C.iphthimus, C.jacobsoni, C.kepongensis, C.kinabaluensis, C.lansangensis, C.lengi, C.liui, C.malayae, C.mamaensis, C.motoensis, C.nujiangensis, C.obsoletus, C.orestes, C.orientalis, C.palauensis, C.pastus, C.pelius, C.peregrinus, C.qionghaiensis, C.ruiliensis, C.sinanoensis, C.suiyangensis, C.sumatrae, C.suzukii, C.tainanus, C.tibetensis, C.trimaculatus, C.wadai, C.yamii, C.yuchihensis, C.fenggangensis). Finally, there are seven species present in both Regions, accounting for 12.3% of the total. This geographical distribution of biting midges in China is consistent with the distribution of other animals (Yan and Yu 1998).
Acknowledgements
This research was financially supported by the grant from the National Natural Science Foundation of China (No. 81360257), the Program for Innovative Research Team in Zunyi City (No. ZSKH-201540) and the visiting scholar of western light Project.
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