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ZOOTAXA ISSN 1175-5334 (online edition)

A review of stalk-eyed fruit flies (Diptera: Tephritidae: Trypetinae) XIAOLIN CHEN1, YANZHOU ZHANG1, JIE LI2 & CHAODONG ZHU1,3 1

Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, China. E-mail: [email protected]; [email protected] Institute of Plant Protection, Shanxi Academy of Agricultural Sciences, Taiyuan 030031, China. E-mail: [email protected] 3 Corresponding author. E-mail: [email protected] 2

Abstract Species of stalk-eyed fruit flies of the genera Pelmatops Enderlein and Pseudopelmatops Shiraki are reviewed and discussed; a key to the two genera and eight species is given. Pelmatops tangliangi Chen sp.nov., Pseudopelmatops indiaensis Chen sp.nov. and Pseudopelmatops yunnanensis Chen sp.nov. from the Oriental Region are described and illustrated. Key words: Trypetinae, Pelmatops, Pseudopelmatops, stalk-eyed, fruit flies

Introduction The stalk-eyed fruit flies comprising Pelmatops Enderlein and Pseudopelmatops Shiraki were proposed as a subtribe of Adramini, namely Pelmatopina, by Wang (1996). And they are arranged under Adramini by Norrbom et al (1999). This group is restricted to the Oriental tropics and subtropics, and most of the species are known to occur in southern China. However, Pelmatops ichneumoneus (Westwood) extends westwards to Nepal and India, Pelmatops tangliangi Chen sp. nov. also occurs in Vietnam and India, and Pseudopelmatops indiaensis Chen sp. nov. is so far known only from India. Pelmatopina are rare in collections, even in China where most species occur. The biology of the species is likewise unknown. One individual of Pelmatops ichneumoneus (Westwood) was observed flying fast on the slope beside thickset shrubs in Sichuan, China by the first author. Except for Tephritidae, stalked eyes is also found in other Diptera acalyptrate families, such as Diopsidae, Richardiidae, Ulidiidae, Platystomatidae and Drosophilidae Several unusual morphological characters of this group have led to its separation from other tephritids and strongly support its monophyly. These characters include: eyes borne at the ends of stalks; abdomen slender, elongate and oval; syntergite 1+2 almost parallel-sided, about equal in length to tergites 3–5 combined; the male epandrium large and broad, with the lateral surstylus comparatively small and short; female terminalia with cylindrical oviscape; 3 elongate-oval or oblong spermathecae present; aculeus nearly rectangular in shape, apically tapered to a point and with a few large, broad serrations, or blunt at apex with a small, round, medial process (Wang, 1996). The systematic position of this group and the relationships between the genera and among their species has been studied, but not completely resolved (Chen & Wang, 2006). Morphological terminology generally follows White et al. (1999) and McAlpine (1981). The abbreviations for specimen depositories used in this paper are as follows: BAU BPBM IZCAS

Beijing Agricultural University, Beijing, China Bernice P. Bishop Museum, Honolulu, Hawaii, USA Institute of Zoology, Chinese Academy of Sciences, Beijing, China

Accepted by D. Bickel: 28 Sep. 2010; published: 22 Oct. 2010

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NTU OMNH USNM

National Taiwan University, Department of Plant Pathology & Entomology, Taipei, Taiwan (Entomological Museum of the Government Research Institute, Taihoku, Formosa) Osaka Museum of Natural History, Nagai Park, Higashi-sumiyoshi-ku, Osaka, Japan National Museum of Natural History, Smithsonian Institution, Washington, DC, USA

Taxonomic treatment Key to known species of Pelmatopina: 1. -

2. 3. 4. 5. 6. 7. -

Orbital and genal setae absent; eye stalk in male at least as long as or longer than abdomen (Figs. 2, 4); palpus spatulate, sparsely covered with fine, yellow setulae................................................................... .2 (Pelmatops Enderlein) Orbital and genal setae present; eye stalk in male shorter than abdomen (Figs. 6, 8, 9); palpus narrow, parallel-sided and densely covered with strong, black setulae (except for Ps. indiaensis Chen sp. nov.)............................................ ..........................................................................................................................................4 (Pseudopelmatops Shiraki) Face with 2 blackish brown transverse bands; lateral vertical and postpronotal setae present; pterostigma about 3/4 as long as cell c (Fig. 26) .................................................................................................... Pe. fukienensis Zia & Chen Face entirely yellow; lateral vertical and postpronotal setae absent; pterosigma about as long as cell c (Figs. 27, 28) ...................................................................................................................................................................................... 3 Ratio between eye stalk length and thorax length 1.2–2.3; Ratio between eye stalk length and wing length 0.4– 0.8 . ...................................................................................................................................... Pe. ichneumoneus (Westwood) Ratio between eye stalk length and thorax length 3.0–4.0; Ratio between eye stalk length and wing length 1.1–1.3. . .......................................................................................................................................... Pe. tangliangi Chen sp. nov. Wing with narrow brownish-black crossband extending from crossvein R-M to middle of cu1 and connected to costal band anteriorly (Figs. 29, 31)................................................................................................................................... 5 Wing without brownish-black crossband on crossvein R-M (Figs. 30, 32, 33) ........................................................... 6 Wing with broad brownish-black costal band and its posterior margin partly in cells br and r4+5 (Fig.31).................... ...........................................................................................................................................Ps. indiaensis Chen sp. nov. Wing with narrow brownish-black costal band and its posterior margin not in cells br and r4+5 (Fig. 29)..................... ................................................................................................................................... .Ps. angustifasciatus Zia & Chen Wing with brownish-black costal band narrowed in cell r2+3 (Fig. 33)......................... .Ps. yunnanensis Chen sp. nov. Wing with brownish-black costal band not narrowed in cell r2+3 (Figs. 30, 32) ........................................................... 7 Frontal and postsutural supra-alar setae present; femora entirely black.................................. Ps. nigricostalis Shiraki Frontal and postsutural supra-alar setae absent; femora broadly yellowish at apices ..... Ps. continentalis Zia & Chen

Pelmatops Enderlein, 1912 Pelmatops Enderlein, 1912: 355. Type-species: Achias ichneumoneus Westwood, 1850, by original designation.

Diagnosis. Head of both sexes strongly prolonged laterally, eyes borne at the ends of stalks, the length of which usually is as long as or longer than the abdomen in males or much shorter than the abdomen in females (Fig. 2, 4); antenna shorter than face; arista plumose; chaetotaxy greatly reduced: head with one or two pairs of vertical setae; thorax with only one pair of apical scutellar setae, one pair of scapular and two notopleural setae, postsutural supra-alar, postalar, intra-alar, and anepisternal setae presenr; postpronotal seta present or absent; anatergite covered with fine and long erect microtrichia; wing predominantly hyaline tinged with illdefined pale yellowish-brown markings; cell sc yellow-brown; vein R4+5 sparsely setose up to slightly beyond level of crossvein R-M, the latter situated well beyond middle of cell dm; legs slender and long, femora lacking setae or spines ventrally; abdomen elongate, syntergite 1+2 nearly parallel-sided. Metathoracic postcoxal bridge broadly sclerotised. Female terminalia: oviscape tapering and somewhat cylindrical; aculeus more or less rectangular in ventral view, apically tapered to relatively sharp tip and with 4 large, broad serrations on lateral margin. Male terminalia: epandrium large and broad, nearly oval in posterior view; lateral surstylus small and short, apex rounded in lateral view; medial surstylus with 1 black prensiseta. Pelmatops is very similar to Pseudopelmatops Shiraki in general appearance, especially in having the head being strongly prolonged laterally, eyes borne at the ends of stalks; chaetotaxy of head and thorax

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reduced; femora lacking setae or spines ventrally; and abdomen long and slender, syntergite 1+2 almost parallel-sided, about equal in length to tergites 3-5. It differs by the absence of orbital and genal setae, the eye stalks comparatively long in males, aculeus apically tapered to relatively sharp tip and with 4 large, broad serrations on each side in female. Three species are known and restricted to the Oriental Region.

Pelmatops fukienensis Zia & Chen (Figs. 1, 10, 18, 26, 34, 43, 48) Pelmatops fukienensis Zia & Chen, 1954: 307. Type-locality: Shaowu [Shao-Woo], Fujian [Fukien], China. Holotype male in IZCAS. —Hardy, 1977: 76. —Wang, 1996: 111.

Diagnosis. Head with lateral vertical and postpronotal setae present; eye stalk about as long as abdomen in male, shorter than thorax in female; thorax, legs and abdomen entirely yellowish-brown except for anatergite blackish; wing predominantly hyaline, tinged with pale yellowish-brown; cell sc reddish-brown, about 3/4 as long as cell c (Fig. 26). Aculeus more or less pentagonal and parallel-sided in shape, apically tapered to a point and with 4 large, broad serrations on each side (Fig. 34); three elongate oval, black spermathecae present (Fig. 43). Description. Body length 13.5–16.0 mm; wing length 12.5–14.0 mm. The body usually predominantly orange-yellow and sub-shining (Fig. 1). Head (Figs. 10, 18): Orange-yellow with 5 narrow brown bands: 1 extending across anterior edge of frons and anterodorsal portion of eye stalk; 1 on posterior side of stalk extending from base to midlength; 1 on anteroventral side of stalk; 1 across ventral margin of face and parafacial; and 1 on dorsal margin of face (Fig. 10). Palpus spatulate, sparsely covered with fine, yellow setulae. Chaetotaxy reduced: head with medial vertical, lateral vertical and postocellar setae present; ocellar, orbital, frontal and genal setae absent. Antenna distinctly shorter than face, with 1st flagellomere rounded dorsoapically, about twice as long as wide; arista plumose, longest ray shorter than width of 1st flagellomere. Thorax entirely orange-yellow except for anatergite blackish and scutum with two narrow obscure grayish-white microtrichose longitudinal stripes; 1 postpronotal, 2 notopleural, 1 anepisternal, 1 postsutural supra-alar, 1 postalar, 1 intra-alar and 1 apical scutellar setae. Wing (Fig. 26). Predominantly hyaline tinged with ill-defined pale yellowish-brown markings; pterostigma yellowish-brown, narrow and long, about 0.75–0.92 times as long as cell c. Vein R4+5 setose from base to crossvein R-M. Legs entirely orange-yellow, slender and long, mid tibia with black apical spine. Abdomen predominantly orange-yellow, elongate, syntergite 1+2 slightly longer than the combined length of tergites 3 and 4. Female terminalia: Oviscape 0.57 times as wide as long and 0.58 times as long as tergites 4–6 combined. Aculeus 2.43 times as long as wide, 0.60 times as long as oviscape, apically tapered to relatively sharp tip and with 4 large, broad serrations on lateral margin (Fig. 34). Eversible membrane with 2 pairs taenia; taenia about as long as denticles part; denticles proximally with 2 columns in both dorsally and ventrally side, and denticles larger in medial and gradually decreasing in size marginally (Fig. 48). Three elongate oval, black spermathecae (Fig.43). Male terminalia. Not dissected. Distribution. China (Fujian, Sichuan, Taiwan). Taiwan is a new province record for this species. Type data. Holotype ♂(IZCAS), CHINA: Fujian: Shaowu, 20 April 1943, C. C. Maa (examined, head missing); paratype: 1 ♀(IZCAS), same data as holotype (examined). Other specimens examined. CHINA: Fujian: Chongan, 750 m, 24 June 1960, Y. R. Cheng, 1 ♀(IZCAS). Sichuan: Mt. Emei, 21 July 1957, K. R. Huang; 26 May 1957, L. Y. Zheng, 2 ♀ (IZCAS). Taiwan: Alishan, Chiayi Hsien, 2270m, 8–9 April 1965, C.M. Yoshimoto,1 ♀(BPBM).

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FIGURES 1–9. Pelmatopina species, habitus, dorsal: 1.Pelmatops fukienensis Zia & Chen (female); 2. Pe. ichneumoneus (Westwood) (male); 3. Pe. ichneumoneus (Westwood) (female); 4. Pe. tangliangi Chen sp. nov. (male); 5. Pseudopelmatops angustifasciatus Zia & Chen (female); 6. Ps. continentalis Zia & Chen (male); 7. Ps. indiaensis Chen sp. nov. (female) 8. Ps. nigrocostalis Shiraki (after Shiraki, 1933) (male); 9. Ps. yunnanensis Chen sp. nov. (male).

Pelmatops ichneumoneus (Westwood) (Figs. 2, 3, 11, 12, 19, 20, 27, 35, 37, 40, 45, 49) Achias ichneumoneus Westwood, 1850: 235. Type-locality: India Orientalis. Syntypes (male & female) in BMNH. Pelmatops ichneumoneus: Enderlein, 1912: 355; Hendel, 1927: 33. —Zia, 1937: 113. —Zia & Chen, 1954: 306. — Hardy, 1977: 76. —Kapoor, 1993: 90 —Wang, 1996: 111.

Diagnosis. This species strongly resembles P. fukienensis in general appearance, but differs by lacking the lateral vertical and postpronotal setae; having an anteroventral brown band on the eye stalk and brown

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crossbands on the face; also by the thorax entirely yellowish-brown; wing predominantly hyaline; and pterostigma relatively long, about equal in length to cell c (Fig. 27). Description. Body length 14.0–16.0 mm; wing length 113.0–14.5 mm. The body usually predominantly orange-yellow and sub-shining (Figs. 2, 3). Head (Figs. 11, 12, 19, 20): Orange-yellow except frons and ocellar triangle yellow-brown to dark brown, with 2 brown bands: 1 extending across anterior edge of frons and anterodorsal portion of eye stalk; 1 on posterior side of eye stalk (two bands separated in female, but connected in male at anterior 3/4 of eye stalk). The ratio between eye stalk length to thorax length 1.23–1.61 in female, 2.00–2.26 in male. Palpus spatulate, sparsely covered with fine, yellow setulae. Chaetotaxy reduced: head with medial vertical and postocellar setae present; lateral vertical, ocellar, orbital, frontal and genal setae absent. Antenna distinctly shorter than face, 1st flagellomere rounded dorsoapically, about twice as long as wide; arista plumose, longest ray slightly shorter than width of 1st flagellomere. Thorax almost orange-yellow except apical part of halter brown to dark brown, 2 notopleural, 1 anepisternal, 1 postsutural supra-alar, 1 postalar, 1 intra-alar and 1 apical scutellar setae. Wing (Fig. 27). Predominantly hyaline, tinged with ill-defined pale yellowish-brown markings; pterostigma yellow-brown, narrow and long, about 1.00–1.03 times as long as cell c. Vein R4+5 setose from base to beyond crossvein R-M. Legs entirely orange-yellow, slender and long, mid tibia with black apical spine. Abdomen predominantly orange-yellow, elongate, syntergite 1+2 longer than tergites 3 and 4 combined. Female terminalia: Oviscape orange-yellow, tapering and somewhat cylindrical; width to length ratio 0.52; oviscape length to tergites 4–6 ratio 0.58. Aculeus more or less rectangular in shape, 2.43 times as long as wide, 0.52 times as long as oviscape, apically tapered to relatively sharp tip and with 4 large, broad serrations on each side (Fig. 35). Eversible membrane with 2 pairs taenia; taenia slightly shorter than denticles part; denticles much larger on basal half of membrane than on distal half, larger basal ones nearly divided into 2 columns, smaller denticles on distal half not divided; similar dorsally and ventrally (Fig. 49). Male terminalia: Epandrium large and broad, nearly oval in posterior view (Fig. 37); lateral surstylus small and short, apex rounded in lateral view (Figs. 40); medial surstylus with 1 black prensiseta. Glans very stout, without distinct acrophallus, but with slender and membranous subapical lobe (Fig. 45). Distribution. China (Sichuan, Yunnan, Xizang, Hainan), India, Nepal. Hainan is a new province record for this species. Type data. India Orientalis. Syntypes (male & female) in BMNH (not examined). Other specimens examined. CHINA: Hainan: Jianfengling, 1400 m, D. Zhang, 6 May 2007, 1♀ (IZCAS). Sichuan: Wan Xian, 1200 m, 15 August 1993, X. L. Chen, 1♀ (IZCAS); near Cheng tu, D.C. Graham, 1♀ (USNM). Yunnan: Xishuangbanna, 650 m, 15 May 1958, S. P. Hong, 1♂ (IZCAS). Xizang: Medog, 900–1200 m, 11 April 1982, Y. H. Han, 1♀ (IZCAS). Yunnan: Honghepingbian, Dawei Mountain 1330 m, 24 May 2009, X. S. Yang, 1♀ (BAU). INDIA: Ranikhet, June 1949, I. M.Newell, 1♂ (BPBM); Ranikhet, June 1950,F. A. Bianchi, 1♂ (BPBM); near Chaubattia, 22 June 1949, F. A. Bianchi, 1♀ (BPBM).

Pelmatops tangliangi Chen sp. nov. (Figs. 4, 13, 24, 28, 38, 41, 46) Diagnosis. This species generally resembles P. ichneumoneus, but the eye stalk in the male is much longer. The ratio between eye stalk length and thorax length is 3.0–4.0 (but 1.2–2.3 in P. ichneumoneus), and the ratio between eye stalk length and wing length is 1.1–1.3 (but 0.4– 0.8 in P. ichneumoneus). Description. Male: body length 15.8 mm, wing length 14.0 mm. The body predominantly orange-yellow and sub-shining (Fig. 4). Head (Figs. 13, 24): Orange-yellow except frons and ocellar triangle yellow-brown to dark brown, with 2 brown bands: 1 extending across anterior edge of frons and anterodorsal portion of eye stalk; 1 on posterior side of eye stalk (bands connected at anterior 3/4 of eye stalk). Head longer than high, strongly prolonged

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laterally, ratio between eye stalk length and thorax length 3.0–4.0, and ratio between eye stalk length and wing length 1.1–1.3. Palpus spatulate, sparsely covered with fine, yellow setulae. Chaetotaxy reduced: head with medial vertical and postocellar setae present; lateral vertical, ocellar, orbital, frontal and genal setae absent. Antenna distinctly shorter than face, with 1st flagellomere rounded dorsoapically, about twice as long as wide; arista plumose, longest ray slightly shorter than width of 1st flagellomere. Thorax almost orange-yellow except for apical part of halter brown to dark brown. 2 notopleural, 1 anepisternal, 1 postsutural supra-alar, 1 postalar, 1 intra-alar and 1 apical scutellar setae. Wing (Fig. 28). Predominantly hyaline tinged with ill-defined pale yellowish-brown markings; pterostigma yellow-brown, narrow and long, about 1.00–1.09 times as long as cell c. Vein R4+5 setose from base to beyond crossvein R-M. Legs entirely orange-yellow, slender and long, mid tibia with black apical spine. Abdomen predominantly orange-yellow, elongate, syntergite 1+2 longer than tergites 3 and 4 combined. Male terminalia: epandrium large and broad, nearly oval in posterior view (Fig. 38); lateral surstylus small and short, apex rounded in lateral view (Fig. 41); medial surstylus with 1 black prensiseta. Glans very stout, without distinct acrophallus, but with slender and membranous subapical lobe (Fig. 46). Female: Unknown. Distribution. China (Yunnan), India, Vietnam. Type data. Holotype ♂ (IZCAS), CHINA: Yunnan: Xishuangbanna 1750m, November 2008, J. Hu & L. Tang. Paratypes: 1 ♂ (OMNH). VIETNAM: Vinh Phu Prov.: Tam Dao, 500–1000m, 2 June1997, R. Matsumoto. 1♂ (BPBM), INDIA: Ranikhet, June 1949, I. M. Newell. Etymology. This species is named after L. Tang, one of the collectors of the Holotype. Remarks. This species is generally similar to Pe. ichneumoneus, but it differs from it and all other congeners by the longer eye stalk. Males of both species were collected at two localities in China and India, and no specimens with eye stalks of intermediate length have been collected.

Pseudopelmatops Shiraki Pseudopelmatops Shiraki, 1933: 49. Type-species: P. nigricostalis Shiraki, 1933, by original designation.

Diagnosis. This is another distinctive genus of Tephritidae characterized by the head of both sexes prolonged laterally, eyes borne at the ends of stalks, the eye stalk in the male shorter than the abdomen (Fig.6, 8,9); antenna almost equal to face or longer than face; arista plumose; chaetotaxy reduced: head with orbital, medial vertical and genal setae present, frontal setae present or absent; thorax with 1 notopleural, 1 postalar and 1 apical scutellar setae present; anatergite covered with fine erect setulae; wing predominantly hyaline; cell sc yellow-brown; vein R4+5 bare or setose; legs slender and long, femora lacking setae or spines ventrally; abdomen elongate, syntergite 1+2 nearly parallel-sided. Metathoracic postcoxal bridge broadly sclerotised. Female terminalia: oviscape yellow-brown, tapering and somewhat cylindrical, aculeus more or less rectangular in ventral view, with a small and round apex, apicolaterally not serrated. Male terminalia: epandrium large and broad, rounded in posterior view; lateral surstylus small and short, apex rounded in lateral view; medial surstylus with 1 black prensiseta. Pseudopelmatops is closely related to Pelmatops Enderlein in the head being strongly prolonged laterally, eyes borne at the ends of stalks; chaetotaxy of head and thorax reduced; femora lacking setae or spines ventrally; and abdomen long and slender, syntergite 1+2 almost parallel-sided, about equal in length to tergites 3–5. It can be recognized by the presence of orbital setae; the eye stalk in the male distinctly shorter than the abdomen; palpus narrow, parallel-sided and densely covered with strong, black setulae (except for Ps. indiaensis Chen sp. nov.). Five species are known and restricted to the Oriental Region.

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FIGURES 10–17. Pelmatopina species, head, front: 10.Pelmatops fukienensis Zia & Chen (female); 11. Pe. ichneumoneus (Westwood) (female); 12. Pe. ichneumoneus (Westwood) (male); 13.Pe. tangliangi Chen sp. nov. (male); 14. Pseudopelmatops angustifasciatus Zia & Chen (female); 15. Ps. continentalis Zia & Chen (male); 16. Ps. indiaensis Chen sp. nov. (female); 17. Ps. yunnanensis Chen sp. nov. (male).

Pseudopelmatops angustifasciatus Zia & Chen (Figs. 5, 14, 22, 29, 36, 44, 50) Pseudopelmatops angustifasciatus Zia & Chen, 1954: 310. Type-locality: Tianmushan, Zhejiang [Chekiang]; Shaowu [Shao-Woo], Fujian [Fukien], China. Lectotype female and three paralectotypes designated by Wang, 1996 in IZCAS. —Hardy, 1977: 76. —Wang, 1996: 112.

Diagnosis. This species is readily differentiated from other Pseudopelmatops by wing largely hyaline with a narrow brownish-black costal band and a narrow brownish-black band extending from crossvein R-M to middle of cu1 and connecting to the costal band anteriorly (Fig. 29). Description. Body length 12.0–15.0 mm; wing length 9.5–11.0 mm. The body usually predominantly dark brown to black and sub-shining (Fig. 5). Head (Figs. 14, 22): dark brown to black except middle of frons with a very narrow yellow longitudinal stripe, the ventral 1/2 of face, parafacial and palpus yellow to yellow-brown. Palpus narrow, parallel-sided and densely covered with strong, black setulae. Chaetotaxy reduced: head with orbital, medial vertical and genal setae present; postocellar, lateral vertical, ocellar, frontal setae absent. Antenna almost equal to face, with 1st flagellomere about 1/4–1/3 as wide as long, and dorsoapically slightly sharp; arista plumose, longest ray slightly shorter or about equal to width of 1st flagellomere. Thorax entirely dark brown to black except scutum with two slightly broad grayish-white microtrichose longitudinal stripes, halter yellow. Chaetotaxy reduced: only 1 notopleural, 1 postalar and 1 apical scutellar setae present. Wing (Fig. 29). Largely hyaline with a narrow brownish-black costal band (posterior margin not in cells br and r4+5) and a narrow brownish-black crossband extending from crossvein R-M to middle of cu1 and connect to the costal band anteriorly; pterostigma yellow-brown, narrow and long, about 0.97 –1.00 times as long as cell c. Vein R4+5 bare or only with 1–2 setulae beyond crossvein R-M.

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Legs. Slender and long. Coxa, trochanter and the basal 2/3 of femora dark brown, other parts yellow brown. Abdomen elongate with tergites 1–2 dark brown and other tergites reddish-brown or yellow-brown; tergites 1–2 nearly parallel-sided and almost equal to the combined length of tergites 3 and 4. Female terminalia: Oviscape yellow-brown, tapering and somewhat cylindrical; width to length ratio 0.35; oviscape length to tergites 4–6 ratio about 0.55. Aculeus parallel-sided and nearly rectangular in general shape, 2.9 times as long as wide, 0.40 times as long as oviscape, with a small and blunt apex, apicolaterally not serrated (Fig. 36). Eversible membrane with 2 pairs taenia; taenia slender, about as long as denticles part; denticles larger on basal half of membrane, gradually decreasing in size distally, larger basal ones nearly divided into 2 columns, smaller denticles on distal half not divided; similar dorsally and ventrally (Fig. 50). Three pale, oblong spermathecae present (Fig. 44). Male. Unknown. Distribution. China (Hubei, Hunan, Zhejiang, Fujian, Sichuan, Guizhou). Guizhou is a new province record for this species. Type data. Lectotype ♀ (IZCAS), CHINA: Zhejiang: Tianmushan, 13 July 1937 (examined). Paralectotypes: 1 ♀ (IZCAS), same locality as lectotype, 8 July 1936 (examined); 1 ♀ (IZCAS), as above, 25 July 1937 (examined); 1 ♀ (IZCAS), Fujian: Shaowu, 20 April 1942 (examined). Other specimens examined. CHINA: Fujian: Shaowu, 900–1170 m, 28 May 1960, Y. R. Zhang, 1♀ (IZCAS). Guizhou: Xishui, 800m, 24–28 September 2000, W. Xiao, 1♀ (IZCAS). Hubei: Badong, 1500m, 14 July 2006, Y. L. Chen,1♀ (IZCAS); Hefeng, 1400 m, 31 July 1989, S. Y. Wang, 1 ♀ (IZCAS). Hunan: Tianpingshan, 18 August 1981, 1♀ (IZCAS). Sichuan: Mt. Emei, May–July 1957, K. R. Huang, 2♀ (IZCAS); Mt. Wawu, 1750m, N29.40.031 E102.56.498, 11 August 2007, F. Liu, 1♀ (IZCAS). Zhejiang: Tianmushan, 26 June 1961, E. Suenson, 1♀ (USNM).

FIGURES 18–25. Pelmatopina species, head, lateral view: 18. Pelmatops fukienensis Zia & Chen (female); 19. Pe. ichneumoneus (Westwood) (female); 20. Pe. ichneumoneus (Westwood) (male); 21. Pseudopelmatops yunnanensis Chen sp. nov. (male); 22. Ps. angustifasciatus Zia & Chen (female); 23. Ps. indiaensis Chen sp. nov. (female); 24. Pe. tangliangi Chen sp. nov. (male); 25. Ps. continentalis Zia & Chen (male).

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FIGURES 26–33. Pelmatopina species, wing: 26.Pelmatops fukienensis Zia & Chen; 27. Pe. ichneumoneus (Westwood); 28. Pe. tangliangi Chen sp. nov. (male); 29. Pseudopelmatops angustifasciatus Zia & Chen; 30. Ps. continentalis Zia & Chen; 31. Ps. indiaensis Chen sp. nov.; 32. Ps. nigrocostalis Shiraki (after Shiraki, 1933); 33. Ps. yunnanensis Chen sp. nov.

Pseudopelmatops continentalis Zia & Chen (Figs. 6, 15, 25, 30, 39, 42, 47) Pseudopelmatops nigricostalis continentalis Zia & Chen, 1954: 310. Type-locality: Tianmushan, Zhejiang; Chongan [not Shao-Woo as originally recorded by Zia & Chen (1954)], Fujian, China. Lectotype male and three paralectotypes designated by Wang, 1996 in IZCAS. —Hardy, 1977: 76. —Wang, 1996: 112.

Diagnosis. This species is similar to P. nigricostalis Shiraki in wing markings, but differs by lacking frontal and postsutural supra-alar setae; and femora yellow except the basal 1/2–3/5 dark brown. It also closely resembles P. angustifasciatus in body coloration and chaetotaxy of the head and thorax, but is easily distinguished by having a much broader brownish-black costal band extending proximally to the base of cell r1 (Fig. 30). Description. Body length 13.5–16.0 mm; wing length 10.5–13.0 mm. The body usually predominantly dark brown to black and sub-shining (Fig. 6).

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FIGURES 34–42. Pelmatopina species, aculeus: 34.Pelmatops fukienensis Zia & Chen; 35. Pe. ichneumoneus (Westwood); 36. Pseudopelmatops angustifasciatus Zia & Chen. 37–39. Pelmatopina species, epandrium and surstyli, posterior: 37. Pe. ichneumoneus (Westwood); 38.Pe. tangliangi Chen sp. nov.; 39. Ps. continentalis Zia & Chen. 40–42. Pelmatopina species, epandrium and surstyli, lateral: 40. Pe. ichneumoneus (Westwood); 41.Pe. tangliangi Chen sp. nov.; 42. Ps. continentalis Zia & Chen.

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FIGURES.43–50. Pelmatopina species, spermathecae: 43. Pelmatops fukienensis Zia & Chen (after Wang,1996); 44. Pseudopelmatops angustifasciatus Zia & Chen (after Wang,1996); Pelmatopina species, glans: 45. Pe. ichneumoneus (Westwood), 46. Pe. tangliangi Chen sp. nov.; 47. Ps. continentalis Zia & Chen; Pelmatopina species, eversible membrane: 48. Pe. fukienensis Zia & Chen; 49. Pe. ichneumoneus (Westwood); 50. Ps. angustifasciatus Zia & Chen.

Head (Figs. 15, 25): dark brown to black except the middle of frons with a very narrow yellow longitudinal stripe; ventral 1/2 of face, parafacial and palpus yellow to yellow-brown. Palpus narrow, parallelsided and densely covered with strong, black setulae. Chaetotaxy reduced: head with orbital, medial vertical and genal setae present; postocellar, lateral vertical, ocellar, and frontal setae absent. Antenna almost equal to face, with 1st flagellomere about 1/4–1/3 as wide as long, and 1st flagellomere dorsoapically slightly sharp; arista plumose, longest ray slightly shorter or about equal to width of 1st flagellomere. Thorax entirely dark brown to black except for scutum with two slightly broad grayish-white microtrichose longitudinal stripes, halter yellow. Chaetotaxy reduced: only 1 notopleural, 1 postalar and 1 apical scutellar setae present. A REVIEW OF STALK-EYED FRUIT FLIES

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Wing (Fig. 30). Largely hyaline with a broad brownish-black costal band extending from the base of cell r1 to the wing tip, all the posterior margin in the cells br and r4+5; pterostigma yellow-brown, narrow and long, about 0.97–1.00 times as long as cell c. Vein R4+5 bare. Legs. Slender and long. Coxa, trochanter and the basal 2/3 of femora dark brown, other parts yellow brown. Abdomen elongate with tergites yellow-brown to dark brown; tergites 1–2 combined nearly parallel-sided and almost as long as tergites 3 and 4 combined. Male terminalia: Epandrium large and broad, rounded in posterior view (Fig. 39); lateral surstylus small and short, apex rounded in lateral view (Fig. 42); medial surstylus with 1 black prensiseta. Glans slightly expanded medially, without distinct acrophallus and membranous subapical lobe (Fig. 47). Female. Unknown. Distribution. China (Hunan, Hubei, Zhejiang, Fujian, Sichuan, Yunnan). Hunan and Hubei are new province records for this species. Type data. Lectotype ♂ (IZCAS), CHINA: Zhejiang: Tianmushan, 8. July 1936 (examined); paraletotypes: 1 ♂ , same data with lectotype (examined); 1 ♂, same locality as lectotype, 20 July 1936 (examined); 1 ♂ (IZCAS), CHINA: Fujian: Chongan, 24 July 1944 (examined). Other specimens examined. CHINA: Hunan: Sangzhi, 1200m, 14 August 2001, J. Wen & N.T. Huang, 1 ♂ (IZCAS). Hubei: Shennongjia, 900m, 11 August 2004, Z. Wang, 1♂ (IZCAS). Sichuan: Mt. Emei, 27 July 1957, Y. C. Lu, 1♂ (IZCAS). Yunnan: Kunming, 1200 m, 7 July 1956, K. R. Huang, 1♂ (IZCAS). Remark. The type specimens of Ps. angustifasciatus (females) and Ps. continentalis (males) were collected in the same days and locality. Considering sexually dimorphic of wing pattern in this group, a possibility that they are conspecific has been considered seriously. However, their unpublished DNA sequence data from author’s recent molecular experiment didn’t support this suppose so far.

FIGURES.51–52. The collecting site environment of Pelmatopina species: 51. Pelmatops ichneumoneus (Westwood); 52. Pe. tangliangi Chen sp. nov.

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Pseudopelmatops indiaensis Chen sp. nov. (Figs. 7, 16, 23, 31) Diagnosis. This species is readily differentiated from other Pseudopelmatops by wing with broad brownishblack costal band and a narrow brownish-black crossband extending from crossvein R-M to middle of cell cu1 and connect to the costal band anteriorly, and palpus covered with fine, yellow setulae. Description. Wing length 10.5–13.0 mm (abdomen missing). The whole body usually predominantly dark brown to black and sub-shining (Fig. 7). Head (Figs. 16, 23): dark brown to black except the middle of frons with a very narrow yellow longitudinal stripe; the ventral 1/2 of face, parafacial and palpus yellow to yellow-brown. Palpus spatulate, sparsely covered with fine, yellow setulae. Chaetotaxy reduced: head with orbital, medial vertical and genal setae present; postocellar, lateral vertical, ocellar, and frontal setae absent. Antenna almost equal to face, with 1st flagellomere about 1/4–1/3 as wide as long, dorsoapically rounded; arista plumose, longest ray slightly shorter or about equal to width of 1st flagellomere. Thorax entirely dark brown to black except scutum with two slightly broadened grayish-white microtrichose longitudinal stripes, halter yellow. Chaetotaxy reduced: only 1 notopleural, 1 postalar and 1 apical scutellar setae present. Wing (Fig. 31). Largely hyaline with broad brownish-black costal band extending from the base of cell r1 to the wing tip (posterior margin of costal band partly in cells br and r4+5), and a narrow brownish-black crossband extending from crossvein R-M to middle of cu 1 and connect to the costal band anteriorly; pterostigma yellow-brown, narrow and long, about 0.90 times as long as cell c. Vein R4+5 setose from base to crossvein R-M. Legs. Coxa, trochanter and basal 3/5 of femora dark brown, other parts yellow; slender and long. Abdomen missing ( a label “♀”pinned under the specimen. The first author has observed the abdomen kept on a slide separately. However, the slide missed lately because of transportation ). Male. Unknown. Distribution. India (Ranikhet). Type data. Holotype ♀ (BPBM), India: Ranikhet, 22 June 1949, I. M. Newell. Etymology. This species is named after the type locality. Remark. The wing pattern of this species is distinctive. It has a broad brownish-black costal band that is similar to that of Ps. continentalis Zia & Chen, and a narrow brownish-black crossband extending from crossvein R-M to middle of cu1 and connect to the costal band anteriorly that similar to Ps. angustifasciatus Zia & Chen. The palpus of this species is sparsely covered with fine, yellow setulae which more like Pelmatops species than Pseudopelmatops.

Pseudopelmatops nigricostalis Shiraki, 1933 (Figs. 8, 32) Pseudopelmatops nigricostalis Shiraki, 1933: 51. Type-locality: Arisan, Taiwan. Syntypes (2 males) in NTU. —Zia & Chen, 1954: 309. —Hardy, 1977: 77. —Wang, 1996: 113.

Diagnosis. This species differs from other congeners by the presence of 1 pair of frontal setae and 1 postalar seta in combination with entirely black femora. Redescription. Body length 12mm; wing length 11.5mm. The body usually predominantly black and subshining (Fig. 8). Head: Black except the middle of frons with a very narrow reddish longitudinal stripe, face with a narrow ill-defined grayish-yellow transverse band just dorsal to ventral margin, yellowish lunula and palpus, and reddish-yellow proboscis. Palpus narrow, parallel-sided and densely covered with strong, black setulae. Chaetotaxy reduced: head with orbital, frontal, medial vertical and genal setae present; postocellar, lateral

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vertical, and ocellar absent. Antenna distinctly longer than face, 1st flagellomere 1/4–1/3 as wide as long, dorsoapically rounded; arista plumose, longest ray slightly shorter or about equal to width of 1st flagellomere. Thorax entirely dark brown to black except scutum with two slightly broad grayish-white microtrichose longitudinal stripes, halter yellow. Chaetotaxy reduced: only 1 notopleural, 1 postalar and 1 apical scutellar setae present. Wing (Fig. 32). Largely hyaline with broad brownish-black costal band extending from the base of cell r1 to the wing tip (posterior margin of costal band partly in cells br and r4+5); pterostigma yellow-brown, narrow and long, about as long as cell c. Halter yellow. Legs. Slender and long,with the tibiae and tarsi reddish-brown, femora entirely black,. Abdomen brown and elongate; tergites 1–2 nearly parallel-sided Female. Unknown. Distribution. China (Taiwan). Type data. Syntypes 2♂ (Entomological Museum of the Government Research Institute, Taihoku, Formosa), CHINA: Taiwan: Arisan, Vll, X. (not examined). Remark. The redescription of this species is based on the original description (Shiraki, 1933). We were unable to borrow the syntypes.

Pseudopelmatops yunnanensis Chen sp. nov. (Figs. 9, 17, 21, 33) Diagnosis. This species is readily differentiated from other Pseudopelmatops by its wing pattern, which includes a brownish-black costal band that is broad from the base of cell r1 to the midlength of the marginal part of that cell, then narrow to the wing tip, a brownish-black band over crossvein R-M and connected with the costal band, and a small obscure ill-defined mark on the subapical section of vein R4+5. Description. Body length 13.5–16.0 mm; wing length 10.5–13.0 mm. The body usually predominantly dark brown to black and sub-shining. Head (Figs. 17, 21): dark brown to black except the middle of frons with a very narrow yellow longitudinal stripe, the ventral 1/2 of face, parafacial and palpus yellow to yellow-brown. Palpus narrow, parallel-sided and densely covered with strong, black setulae. Chaetotaxy reduced: head with orbital, medial vertical and genal setae present; postocellar, lateral vertical, ocellar, and frontal setae absent. Antenna almost equal to face, with 1st flagellomere about 1/4–1/3 as wide as long, and 1st flagellomere dorsoapically slightly pointed; arista plumose, longest ray slightly shorter or about equal to width of 1st flagellomere. Thorax entirely dark brown to black except scutum with two slightly broad grayish-white microtrichose longitudinal stripes, halter yellow. Chaetotaxy reduced: only 1 notopleural, 1 postalar and 1 apical scutellar setae present. Wing (Fig. 33). Largely hyaline with brownish-black costal band extending from cell c to the wing tip, the band broad to slightly beyond crossvein R-M, then strongly narrowed, cell r2+3 mostly hyaline except for apical margin, but slightly gradually expanded to apex of vein R 4+5 , a short brownish-black crossband covering crossvein R-M and connected with costal band, and a small obscure ill-defined mark on the subapical section of vein R4+5. Pterostigma yellow-brown, narrow and long, 0.98 times as long as cell c. Vein R4+5 bare. Legs. Slender and long. Coxa, trochanter and the basal 2/3 of femora dark brown, other parts yellow brown. Abdomen elongate with tergites 1–2 dark brown and other tergites yellow-brown; tergites 1–2 nearly parallel-sided and almost equal to the combined length of tergites 3 and 4. Male terminalia not dissected. Female. Unknown. Distribution. China (Yunnan).

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Type data. Holotype ♂ (IZCAS), China: Yunnan, 195?. Etymology. This species is named after the type locality.

Discussion In order to better understand the phylogenetic position and relationships of the Pelmatopina, a more detailed phylogenetic study might be needed in the future. In addition, no host information is known for this group, but some implications might be deduced from the environments of the collecting sites. One specimen of Pelmatops ichneumoneus was swept in 2009 from the tree shown in the middle of Fig. 51. This tree probably belongs to the family Lauraceae. The site where a specimen of P. tangliangi was collected in 2008 is shown in Fig. 52. Both sites are in Yunnan, China. Pelmatops and Pseudopelmatops are mainly distributed in China and India, so their host plants might be restricted to these two areas. However, the host information of them is still unknown, so how they evolve with host plant or if the origin of eye stalk having relationship with their host might need further study in future when more data available.

Acknowledgements We sincerely thank Allen Norrbom (Systematic Entomology Lab., USDA,Washington, DC,USA), Amnon Freidberg (Department of Zoology, The George S. Wise Faculty of Life Sciences, Tel Aviv University, Israel) for critically review an early version of this paper. We also thank the following scientists, curators and institutions who kindly loaned or provided specimens for study: Ding Yang and Xinli Wang (BAU), Neal Evenhuis, Shepherd Myers and Keith Arakaki (BPBM), Bernhard Merz (MHNG), Rikio Matsumoto (OMNH), Allen Norrbom (USNM). We thank Rong Li (Kunming Institute of Botany, Chinese Academy of Sciences, China) for possible plant identification, and Martin Hauser (Plant Pest Diagnostics, California Department of Food & Agriculture, USA) for locating the valuable specimen of Pelmatops tangliangi Chen sp. nov., Allen Norrbom helped to take some images of terminalia, Guoqing Mai and Liang Tang helped to take that of habitus, Lucrecia Rodriguez helped to prepare wing imagines, Liang Tang and Xiushuai Yang took the imagines of collecting sites, Xuelin Chen prepared some plates for publication. Chuanren Li, Feng Yuan, Hui Dong, Liang Tang, Meicai Wei, Tingting Zhang, Dong Zhang, Xiushuai Yang and Xingjing Wang helped to find specimens or provide valuable information. This project was supported by the National Natural Science Foundation of China (30770267), K. C. Wong Education Foundation, Hong Kong, National Science Fund for Fostering Talents in Basic Research (special subjects in animal taxonomy, NSFC-J0630964/J0109), Shanxi Science and Technology Fund from government (2007031040-1) and Academy of Agriculture Sciences (YGG0930) to screen and mass-rear natural enemies.

References Chen, X.L. & Wang, X.J. (2006) A cladistic analysis of the subtribe Pelmatopina (Diptera: Tephritidae: Trypetinae: Adramini) and its phylogeographic implications. Instrumenta Biodiversitatis, VII, 157–165. Enderlein,G. (1912) Zur Kenntnis orientalischer Ortalinen und Loxoneurinen. Zoologische Jahrbucher Systematik, 33, 347–362. Hardy, D.E. (1977) Family Tephritidae (Trypetidae, Trupaneidae). In: Delfinado, M. D. & D. E. Hardy, (eds). A Catalog of the Diptera of the Oriental Region, 3, 76. Hendel, F. (1927) 49. Trypetidae. In: Lindner, E. (eds). Die Fliegen der palaearktischen Region, 5, 129–221. Kapoor, V.C. (1993) Indian fruit flies (Insecta: Diptera: Tephritidae). Oxford & IBH Publishing Co. Pvt. Ltd., New Delhi, 228 pp: 90. McAlpine, F.J. (1981) Morphology and terminology. Adults 2. In: In: McAlpine JF, Peterson BV, Shewell GE, Teskey HJ, Vockeroth JR, Wood DM, editors. (eds), Manual of the Nearctic Diptera. Volume 1. Agriculture Canada Monograph, 27, 9–63.

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TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Norrbom, A.L., Carroll, L.E., Thompson, F.C., White, I.M. & Freidberg, A. (1999) [dated 1998] Systematic Database of Names. Pp. 65–252. In Thompson, F.C. (ed.), Fruit Fly Expert Identification System and Systematic Information Database. Myia 9, vii + 524 pp. Shiraki, T. (1933) A systematic study of Trypetidae in the Japanese Empire. Memoirs of the Faculty of Science and Agriculture Taihoku Imperial University, 8, 1–509. Wang, X.J. (1996) The fruit flies (Diptera: Tephritidae) of the East Asian Region. Acta Zootaxonomica Sinica, 21(Supplement), 1–338. White, I.M., Headrick, D.H., Norrbom, A.L. & Carroll, L.E. (1999) Glossary, p. 881–924. In, ‘Fruit flies (Tephritidae): phylogeny and evolution of behavior’. (Eds M. Aluja & A. L. Norrbom.) (CRC Press, Boca Raton.) [16] + 944 p. Zia, Y. (1937) Study on the Trypetidae or fruit-flies of China. Sinensia, 8(2), 113. Zia, Y. & Chen, S.H. (1954) Notes on Chinese trypetid flies I. Acta Entomologica Sinica, 4(3), 299–314 (In Chinese with English summary).

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