A species checklist of the subgenus Culicoides (Avaritia) in China, with a description of a new species (Diptera, Ceratopogonidae)

Qiong Qiong Chang; Xiao Hong Jiang; Guo Ping Liu; Xiao Fei Li; Xiao Hui Hou

doi:10.3897/zookeys.706.13535

Abstract

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.

Keywords

Biting midges, Ceratopogonidae , checklist, Culicoides, subgenus Avaritia

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.

Taxonomy

Checklist of the subgenus Avaritia in China in alphabetical order

Subgenus Avaritia Fox, 1955

Avaritia Fox, 1955: 218.

Type species. Ceratopogon obsoletus Meigen, 1818.

Culicoides (Avaritia) abchazicus Dzhafarov, 1964

Culicoides abchazicus Dzhafarov, 1964: 263; Liu et al. 2011: 385. Type Locality: Georgia.

Distribution. China (Liaoning); Georgia.

Culicoides (Avaritia) actoni Smith, 1929

Culicoides actoni Smith, 1929: 255; McDonald and Lu 1972: 411; Lee 1988: 29; Yu et al. 2005: 907. Type locality: India.

Culicoides okumensis Arnaud, 1956: 119. Type locality: Japan.

Culicoides imperceptus 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

Culicoides albifascia 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

Culicoides bawanglingensis Yu, Wang & Chen, in Wang et al., 2012: 283. Type Locality: China: Hainan, Bawangling.

Distribution. China (Hainan).

Culicoides (Avaritia) brevipalpis Delfinado, 1961

Culicoides brevipalpis 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

Culicoides brevitarsis Kieffer, 1917: 187; Yu et al. 2005: 911. Type locality: Australia.

Culicoides robertsi Lee & Reye, 1953: 386. Type locality: Australia.

Culicoides radicitus Delfinado, 1961: 657. Type locality: Philippines.

Culicoides superfulvus Das Gupta, 1962b: 253. Type locality: India.

Distribution. China (Anhui, Taiwan, Hainan); Australia, Philippines, India, Laos, Malaysia.

Culicoides (Avaritia) bubalus Delfinado, 1961

Culicoides bubalus Delfinado, 1961: 658; Yu et al. 2005: 913. Type locality: The Philippines.

Distribution. China (Taiwan), Philippines.

Culicoides (Avaritia) chiopterus (Meigen), 1830

Ceratopogon chiopterus Meigen, 1830: 263. Type locality: Europe.

Ceratopogon amoenus Winnertz, 1852: 35. Type locality: Germany.

Monohelea similis Goetghebuer, 1927: 203. Type locality: Belgium.

Culicoides dobyi 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

Culicoides clavipalpis Mukerji, 1931: 1052; McDonald and Lu 1972: 403; Yu et al. 2005: 916. Type locality: India.

Culicoides candidus 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

Culicoides comparis Liu & Yu, in Yu et al., 2005: 917. Type locality: China: Tibet, Nielamu.

Distribution. China (Tibet).

Culicoides (Avaritia) conaensis Liu & Yu, 1990

Culicoides conaensis Liu & Yu, 1990: 19; Yu et al. 2005: 919. Type locality: China: Tibet, Cuona.

Distribution. China (Tibet).

Culicoides (Avaritia) dentiformis McDonald & Lu, 1972

Culicoides dentiformis McDonald & Lu, 1972: 403; Yu et al. 2005: 921. Type locality: China: Taiwan.

Distribution. China (Taiwan).

Culicoides (Avaritia) elongatus Chu & Liu, 1978

Culicoides elongatus Chu & Liu, 1978: 83; Yu et al. 2005: 922. Type locality: China: Yunnan, Mengla.

Distribution. China (Fujian, Yunnan).

Culicoides (Avaritia) fenggangensis Liu & Hou, sp. n.

Culicoides fenggangensis Liu & Hou, sp. n., this paper. Type locality: China: Guizhou, Fenggang.

Distribution. China (Guizhou).

Culicoides (Avaritia) filicinus Gornostaeva & Gachegova, 1972

Culicoides filicinus Gornostaeva & Gachegova, 1972: 522; Yu et al. 2005: 924. Type locality: Russia.

Distribution. China (Tibet, Heilongjiang); Russia.

Culicoides (Avaritia) gaponus Yu, 1982

Culicoides gaponus Yu, 1982: 202; Yu et al. 2005: 925. Type locality: China: Hainan, Diaoluoshan.

Distribution. China (Hainan).

Culicoides (Avaritia) holcus Lee, 1980

Culicoides holcus Lee, 1980: 85; Yu et al. 2005: 927. Type locality: China: Yunnan.

Distribution. China (Yunnan).

Culicoides (Avaritia) hui Wirth & Hubert, 1961

Culicoides hui Wirth & Hubert, 1961: 16; Yu et al. 2005: 928. Type locality: China: Taiwan.

Distribution. China (Taiwan, Guangdong, Hainan, Yunnan, Hunan); Indonesia, Laos, Malaysia.

Culicoides (Avaritia) imicola Kieffer, 1913

Culicoides imicola Kieffer, 1913: 11; Yu et al. 2005: 930. Type locality: Kenya.

Culicoides pallidipennis Carter, Ingram & Macfie, 1920: 265. Type locality: Ghana.

Culicoides iraqensis Khalaf, 1957: 343. Type locality: Iraq.

Culicoides minutus Sen & Das Gupta, 1959: 622. Type locality: India.

Culicoides pseudoturgidus Das Gupta, 1962a: 537. Type locality: India.

Distribution. China (Hainan); Wide spread in Africa, the Middle and Far East, India, Laos, Sri Lanka, Vietnam.

Culicoides (Avaritia) incertus Yu & Zhang, 1988

Culicoides incertus Yu & Zhang, in Yu, 1988: 136; Yu et al. 2005: 931. Type locality: China: Tibet.

Distribution. China (Tibet).

Culicoides (Avaritia) innoxius Sen & Das Gupta, 1959

Culicoides innoxius Sen & Das Gupta, 1959: 626; Yu et al. 2005: 933. Type locality: India.

Distribution. China (Hainan); India, Cambodia, Indonesia, Laos, Malaysia, Sri Lanka, Thailand.

Culicoides (Avaritia) insignipennis Macfie, 1937

Culicoides insignipennis Macfie, 1937b: 469; Lee 1988: 65; Yu et al. 2005: 935. Type locality: Malaysia.

Distribution. China (Fujian, Taiwan, Yunnan, Hunan); Malaysia, Brunei, Indonesia, Laos, Philippines, Singapore, Thailand.

Culicoides (Avaritia) iphthimus Zhou & Lee, 1984

Culicoides iphthimus Zhou & Lee, 1984: 295; Yu et al. 2005: 936. Type locality: China: Chongqing.

Distribution. China (Chongqing).

Culicoides (Avaritia) jacobsoni Macfie, 1934

Culicoides jacobsoni Macfie, 1934: 215; Yu 1982: 48. Yu et al. 2005: 938. Type locality: Indonesia.

Culicoides buckleyi Macfie, 1937a: 117. Type locality: Malaysia.

Culicoides kitaokai Tokunaga, 1955: 6. Type locality: Japan.

Culicoides unisetiferus Tokunaga, 1959: 236. Type locality: Papua New Guinea.

Distribution. China (Fujian, Taiwan, Guangdong, Guangxi, Hainan, Yunnan, Tibet, Hunan); Indonesia, Malaysia, Japan, New Guinea, Philippines, Thailand.

Culicoides (Avaritia) kepongensis Lee, 1988

Culicoides kepongensis Lee, 1988: 69; Wirth and Hubert 1989: 346; Yu et al. 2005: 940. Type locality: China; Malaysia.

Distribution. China (Fujian, Taiwan); Malaysia, Laos, Thailand.

Culicoides (Avaritia) kinabaluensis Wirth & Hubert, 1989

Culicoides kinabaluensis Wirth & Hubert, 1989: 211; Yu et al. 2005: 941. Type locality: Indonesia.

Distribution. China (Hainan).

Culicoides (Avaritia) lansangensis Howarth, 1985

Culicoides lansangensis Howarth, 1985: 58; Liu et al. 1996: 38; Yu et al. 2005: 942. Type locality: Laos.

Distribution. China (Guangdong, Hainan, Hunan).

Culicoides (Avaritia) lengi Yu & Liu, 1990

Culicoides lengi Yu & Liu, 1990: 10; Yu et al. 2005: 944. Type locality: China: Guangdong, Shixing.

Distribution. China (Guangdong).

Culicoides (Avaritia) liui Wirth & Hubert, 1961

Culicoides liui Wirth & Hubert, 1961: 20; Yu et al. 2005: 946. Type locality: China: Taiwan.

Distribution. China (Taiwan, Yunnan); Indonesia, Laos, Malaysia, Philippines, Thailand.

Culicoides (Avaritia) longirostris Qu & Wang, 1994

Culicoides longirostris Qu & Wang, 1994: 486; Yu et al. 2005: 947. Type locality: China: Tibet.

Distribution. China (Tibet).

Culicoides (Avaritia) malayae Macfie, 1937

Culicoides malayae Macfie, 1937b: 471; Yu 1982: 190; Yu et al. 2005: 949. Type locality: Malaysia.

Distribution. China (Fujian, Taiwan, Guangdong, Guangxi, Hainan, Yunnan); Indonesia, Malaysia, Philippines, Thailand.

Culicoides (Avaritia) mamaensis Lee, 1979

Culicoides mamaensis Lee, 1979: 101; Yu et al. 2005: 951. Type locality: China: Tibet, Cuona.

Distribution. China (Sichuan, Tibet).

Culicoides (Avaritia) motoensis Lee, 1978

Culicoides motoensis Lee, 1978: 75; Yu et al. 2005: 952. Type locality: China: Tibet, Motuo.

Distribution. China (Tibet).

Culicoides (Avaritia) nielamensis Liu & Deng, 2000

Culicoides nielamensis Liu & Deng, 2000: 246; Yu et al. 2005: 954. Type locality: China: Tibet, Nielamu.

Distribution. China (Tibet).

Culicoides (Avaritia) nigritus Fei & Lee, 1984

Culicoides nigritus Fei & Lee, 1984: 345; Yu et al. 2005: 955. Type locality: China: Inner Mongolia, Linxi.

Distribution. China (Inner Mongolia).

Culicoides (Avaritia) nujiangensis Liu, 1990

Culicoides nujiangensis Liu, 1990: 59; Yu et al. 2005: 956. Type locality: China: Yunnan, Nujiang.

Distribution. China (Yunnan).

Culicoides (Avaritia) obsoletus (Meigen), 1818

Ceratopogon obsoletus Meigen, 1818: 76. Type locality: Europe.

Ceratopogon varius Winnertz, 1852: 35. Type locality: Germany.

Ceratopogon yezoensis Matsumura, 1911: 60. Type locality: Russia.

Culicoides obscuripes Santos Abreu, 1918: 297. Type locality: Spain.

Culicoides lacteinervis Kieffer, 1919a: 47. Type locality: Slovak Republic, Ukraine.

Culicoides rivicola Kieffer, 1921: 56. Type locality: Germany.

Culicoides clavatus Kieffer, 1921: 56. Type locality: Germany.

Culicoides heterocerus Kieffer, 1921: 57. Type locality: Germany.

Culicoides pegobius Kieffer, 1922: 235. Type locality: Germany.

Culicoides kabyliensis Kieffer, 1922: 505. Type locality: Algeria.

Culicoides concitus Kieffer, 1922: 71. Type locality: Germany.

Culicoides intermedius Okada, 1941: 22. Type locality: Japan.

Culicoides sintrensis Cambournac, 1956: 591. Type locality: Portugal.

Culicoides seimi Shevchenko, 1967: 173. Type locality: Ukraine.

Distribution. China (Shanxi, Inner Mongolia, Liaoning, Jilin, Heilongjiang, Fujian, Shandong, Sichuan, Chongqing, Yunnan, Xinjiang, Gansu, Tibet); wide distribution in Palaearctic Region, Britain, Germany, Russia, Canary Islands, Algeria, Japan, Portugal, Slovakia, Ukraine.

Culicoides (Avaritia) orestes Wirth & Hubert, 1989

Culicoides orestes Wirth & Hubert, 1989: 222; Yu et al. 2005: 959. Type locality: Malaysia.

Distribution. China (Hainan); Malaysia.

Culicoides (Avaritia) orientalis Macfie, 1932

Culicoides orientalis Macfie, 1932: 490; Lee 1978: 83; Yu et al. 2005: 961. Type locality: Malaysia.

Culicoides nayabazari 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

Culicoides palauensis 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

Culicoides pastus 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

Culicoides pelius 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

Culicoides peregrinus Kieffer, 1910: 191; Yu et al. 2005: 967. Type locality: India.

Culicoides judicandus Bezzi, 1916: 8. Type locality: Philippines.

Culicoides esmoneti Salm, 1917b: 136. Type locality: Indonesia.

Culicoides philippinensis Kieffer, 1921: 564. Type locality: Philippines.

Culicoides assamensis Smith & Swaminath, 1932: 183. Type locality: India.

Culicoides quadratus Tokunaga, 1951: 108. Type locality: Indonesia.

Distribution. China (Hebei, Inner Mongolia, Liaoning, Jiangsu, Fujian, Taiwan, Jiangxi, Henan, Guangdong, Guangxi, Hainan); India, Philippines, Indonesia.

Culicoides (Avaritia) qionghaiensis Yu & Liu, 1990

Culicoides qionghaiensis Yu & Liu, 1990: 4; Yu et al. 2005: 969. Type locality: China: Sichuan, Xichang.

Distribution. China (Sichuan).

Culicoides (Avaritia) ruiliensis Lee, 1980

Culicoides ruiliensis Lee, 1980: 86; Yu et al. 2005: 971. Type locality: China: Yunnan.

Distribution. China (Yunnan).

Culicoides (Avaritia) scoticus Downes & Kettle, 1952

Culicoides scoticus Downes & Kettle, 1952: 65; Yu et al. 2005: 972. Type locality: Great Britain.

Distribution. China (Tibet); Great Britain, France.

Culicoides (Avaritia) sinanoensis Tokunaga, 1937

Culicoides sinanoensis Tokunaga, 1937: 331; Yu et al. 2005: 973. Type locality: Japan.

Culicoides obsoletiformis Amosova, 1957: 233. Type locality: Russia.

Distribution. China (Liaoning, Jilin, Heilongjiang, Yunnan, Shaanxi); Japan, Russia.

Culicoides (Avaritia) suiyangensis Hou, Han, Lv & Jiang, 2014

Culicoides suiyangensis Hou, Han, Lv & Jiang, 2014: 98. Type Locality: China: Guizhou, Suiyang.

Distribution. China (Guizhou).

Culicoides (Avaritia) sumatrae Macfie, 1934

Culicoides sumatrae Macfie, 1934: 190; Yu et al. 2005: 975. Type locality: Malaysia.

Culicoides amamiensis Tokunaga, 1937: 325. Type locality: Japan.

Culicoides kagiensis Tokunaga, 1937: 327. Type locality: Taiwan.

Culicoides ohmorii Takahashi, 1958: 113. Type locality: Japan.

Culicoides assimilis Delfinado, 1961: 660. Type locality: Philippines.

Distribution. China (Fujian, Taiwan, Guangdong, Guangxi, Hainan, Yunnan, Tibet); Malaysia, Japan, Philippines.

Culicoides (Avaritia) suzukii Kitaoka, 1973

Culicoides suzukii Kitaoka, 1973: 212; Lee 1988: 107; Yu et al. 2005: 977. Type locality: Japan.

Distribution. China (Taiwan, Yunnan); Japan.

Culicoides (Avaritia) tainanus Kieffer, 1916

Culicoides tainanus Kieffer, 1916: 114; Yu et al. 2005: 979. Type locality: China: Taiwan.

Ceratopogon maculatus Shiraki, 1913: 294. Type locality: China: Taiwan.

Culicoides kii Tokunaga, 1937: 284. Type locality: Japan.

Culicoides sigaensis Tokunaga, 1937: 322. Type locality: Japan.

Culicoides kyotoensis Tokunaga, 1937: 329. Type locality: Japan.

Culicoides suborientalis Tokunaga, 1951: 106. Type locality: Indonesia.

Distribution. China (Fujian, Hainan, Yunnan, Shaanxi, Shandong); Indonesia, Japan, Laos, Malaysia, Philippines, Thailand, Vietnam.

Culicoides (Avaritia) tibetensis Chu, 1977

Culicoides tibetensis Chu, 1977: 102; Yu et al. 2005: 980. Type locality: China: Tibet.

Distribution. China (Sichuan, Tibet).

Culicoides (Avaritia) trimaculatus McDonald & Lu, 1972

Culicoides trimaculatus McDonald & Lu, 1972: 415; Yu et al. 2005: 982. Type locality: China: Taiwan.

Distribution. China (Taiwan).

Culicoides (Avaritia) wadai Kitaoka, 1980

Culicoides wadai Kitaoka, 1980: 14; Yu et al. 2005: 984. Type locality: China: Taiwan.

Distribution. China (Fujian, Guangxi, Taiwan).

Culicoides (Avaritia) wandashanensis Wang & Liu, 1999

Culicoides wandashanensis Wang & Liu, 1999: 328; Yu et al. 2005: 985. Type locality: China: Heilongjiang, Raohe.

Distribution. China (Heilongjiang).

Culicoides (Avaritia) yamii Lien, Lin & Weng, 1998

Culicoides yamii Lien, Lin & Weng, 1998: 57; Yu et al. 2005: 987. Type locality: China: Taiwan, Lanyu.

Distribution. China (Taiwan).

Culicoides (Avaritia) yuchihensis Lien, Lin & Weng, 1998

Culicoides yuchihensis Lien, Lin & Weng, 1998: 58; Yu et al. 2005: 988. Type locality: China: Taiwan.

Distribution. China (Taiwan).

Culicoides (Avaritia) fenggangensis Liu & Hou, sp. n.

http://zoobank.org/52A53C15-0F18-4040-B83C-B678BD2E9A66

Figs 1, 2, 3

Diagnosis

Male: only species of Culicoides in China with the following combination of features: the 3^rd segment of the palpus is slender, PR 3.11; the apex of 9^th 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 3^rd 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. 3^rd 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).

Figure 1.

Culicoides (A.) fenggangensis sp. n. Left Wing. a Female b Male.

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 1^st, proximal 1/2 of 2^nd radial cell in dark spot; three pale spots near anterior margin: 1^st over base of cell r₁ and r-m crossvein extending from below M₁ to margin of costa, 2^nd over distal of cell r₂ from dorsal portion of M₁ to costa, 3^rd 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: 1^st proximal to CuA, 2^nd between medial and mediocubital forks, 3^rd and 4^th 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, 1^st spine is longest.

Table 1.

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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. (♀).

Leg	TR	F-T
Foreleg	2.12	85 : 88 : 53 : 25 : 16 : 10 : 11
Midleg	2.31	109 : 114 : 60 : 26 : 16 : 10 : 11
Hindleg	1.73	109 : 115 : 57 : 33 : 17 : 11 : 12

Abdomen. Brown. Two subequal-size ovoid spermathecae (Fig. 2i), measuring 65.0–72.5 × 50.0–57.5 (n = 3) μm, 65.0–72.5 × 50.0 (n = 2) μm, slender sclerotized necks with 2.5–7.5 (n = 3) μm; third slender, elongate rudimentary spermatheca, length 20.0–25.0 (n = 3) μm.

Figure 2.

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).

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): 9^th 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.

Figure 3.

Culicoides (A.) fenggangensis sp. n. a Genitalia and aedeagus (parameres absent), ventral view b parameres, ventral view.

Table 2.

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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 3^rd 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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