Paraleucophenga Hendel, 1914: 114

Zhao, Feng, Gao, Jian-Jun & Chen, Hong-Wei, 2009, Taxonomy and molecular phylogeny of the Asian Paraleucophenga Hendel (Diptera, Drosophilidae), Zoological Journal of the Linnean Society 155 (3), pp. 615-629 : 618-621

Paraleucophenga Hendel, 1914: 114 .

Type species: Helomyza invicta Walker, 1857: 130 .

Diagnosis: Modified from Bächli (1971) and Okada (1988). Frons narrow, less than one third of head width; arista with long dorsal and fine ventral branches; scutellum usually with small setae along margin; katepisternal setae 3; abdominal tergites with silvery white pollinosity; cercus elongated below and slightly constricted submedially ( Figs 16, 18 , 20, 22 ); paramere with tooth comb ( Figs 17, 19 , 21, 23 ); aedeagal apodeme sub-basally narrow, arched, distally broadened ( Figs 17, 19 , 21, 23 ).

Description: Body slender, usually longer than 4.0 mm.

Head: Eye dark red. Ocellar triangle: black, with one pair of small setae above ocellar setae. Frons dark brown, with one row of interfrontal setulae along fronto-orbital plate. Pedicel brown. First flagellomere grey yellow. Face grey yellow, facial carina not prominent. Clypeus grey yellow. Gena: narrow and linear, yellowish. Postgena yellow. Vibrissa prominent; other orals small. Palpus slender, basally grey yellow, distally black, with a few small setae along ventral margin. Occiput glossy, brownish black.

Thorax: yellow, with brown patches and silvery pollinosity. Postpronotal lobe pale yellow, with one long and a few short setae. Acrostichal setulae in approximately ten irregular rows. Prescutellar setae one pair. Anepisternum glabrous. Scutellum usually concolorous with thorax, with one obscure patch. Basal scutellar setae divergent; apicals cruciate.

Wing: brownish. Basal medial–cubital crossvein absent; C 1 setae 2, indistinctly differentiated. R 2+3:

slightly curved to costa at tip. R 4+5 and M 1 nearly parallel. Halters grey white.

Legs: yellow. Fore femur with two or three irregular rows of long setae on posterior surface. Preapical and apical dorsal setae present on all tibiae. Fore- and mid first tarsomeres each as long as other tarsomeres combined; hind first tarsomere as long as the three succeeding tarsomeres together.

Abdomen: Males tergites 1–5 with dense silvery white pollinosity, and with brown to dark brown coloration. Females yellow, with distinct pattern ( Lin & Wheeler, 1972; Fig. 5B ). Sternites usually grey yellow.

Male terminalia: Epandrium slightly constricted mid-dorsally, broadened below, nearly entirely pubescent, except for ventral margin, with dense setae ( Figs 16, 18 , 20, 22 ). Cercus separated from epandrium, entirely pubescent, and setigerous ( Figs 16, 18 , 20, 22 ). Surstylus quadrate, with a few setae on outer surface, and small prensisetae on apical margin ( Figs 16, 18 , 20, 22 ). Membrane between epandrium and cercus pubescent ( Figs 16, 18 , 20, 22 ). Hypandrium narrow, arched, sub-basally with one pair of somewhat triangular plates ( Figs 17, 19 , 21, 23 ). Gonopods fused to each other, forming posteromedian plate, anteriorly forming contiguous to aedeagus (basiphallus) ( Figs 17, 19 , 21, 23 ). Parameres sub-basally and apically with a few sensilla ( Figs 17, 19 , 21, 23 ). Aedeagus: short and thick, membranous tube with pubescence distally ( Figs 17, 19 , 21, 23 ).

Included species: Paraleucophenga semiplumata ( Duda, 1939) ; Paraleucophenga argentosa ( Okada, 1956) ; Paraleucophenga emeiensis Sidorenko, 1998 ; Paraleucophenga invicta ( Walker, 1857) ; Paraleucophenga javana Okada, 1988 ; Paraleucophenga neojavanaii Singh & Negi, 1992 ; Paraleucophenga shimai Okada, 1988 ; Paraleucophenga brevipenis sp. nov.; Paraleucophenga hirtipenis sp. nov.; Paraleucophenga longiseta sp. nov., and Paraleucophenga tanydactylia sp. nov.

KEY TO ALL PARALEUCOPHENGA SPECIES EXAMINED, BASED ON MALE MORPHOLOGICAL DATA

1. Frons narrow, less than one third of head width; arista with long dorsal and fine ventral branches; scutellum usually with small setae along margin; katepisternal setae 3; abdominal tergites with silvery white pollinosity; cercus elongated below and slightly constricted submedially; paramere with tooth comb; aedeagal apodeme sub-basally narrow, arched, distally broadened; (genus Paraleucophenga ). ................................................... 2

2. Scutellum without setae along margin; midleg femur medially with one row of about five strong setae on anterior surface; thorax and abdominal tergites with silvery white pollinosity............................................... P. shimai Scutellum with small setae along margin; midleg femur lacking strong setae on anterior surface; thorax and abdominal tergites with silvery pollinosity..............................................................................................3

3. Fifth sternite without long setae. ......................................................................................................... 4 Fifth sternite with long setae. ............................................................................................................. 6

4. Paramere submedially with longitudinal teeth row.................................................................... P. argentosa Paramere distally with transverse teeth row...........................................................................................5

5. Brown posterior band of third tergite as broad as half of the tergite width; fifth tergite nearly entirely dark brown, with one yellow patch laterally. ............................................................................................. P. emeiensis Brown posterior band of third tergite as broad as a fifth of the tergite width; posterior band of fifth tergite as broad as two thirds of the tergite width, laterally dark.......................................................................... P. invicta

6. First tergite silvery white; fifth sternite with long finger-like processes, lacking spinules along posterior margin.......................................................................................................................................................7 First tergite dark brown; fifth sternite with a few spinules along posterior margin, lacking long finger-like processes.......................................................................................................................................................8

7. Brown posterior band of second tergite submedially as broad as half of the tergite width; fifth sternite with six long, finger-like processes................................................................................................................. P. javana Brown posterior band of second tergite as broad as a third of the tergite width; fifth sternite with 12 long, finger-like processes.......................................................................................................... P. tanydactylia sp. nov.

8. Fifth sternite with 22–23 long and strong setae; gonopods subapically with one sclerotized right process; aedeagal apodeme distally with one sclerotized right process...................................................... P. brevipenis sp. nov. Fifth sternite with 16–18 long and strong setae; gonopods subapically with one sclerotized left process...............9

9. Aedeagal apodeme distally with one sclerotized left process. .......................................... P. hirtipenis sp. nov. Aedeagal apodeme distally with one sclerotized right process. ......................................... P. longiseta sp. nov. KEY TO SEVEN PARALEUCOPHENGA SPECIES BASED ON THE ND2 GENE SEQUENCES

In the following key to seven Paraleocophenga species, each of the numbers indicates the position of a diagnostic character in the affiliated sequences, followed by its status (i.e. nucleotide type, given after a colon) and (after a slash) corresponding amino acid type (all shown in standard three-letter abbreviations; in the case of more than two types of amino acid being present, they are bracketed together, and are separated from each other using an additional slash). 1. 251: T/Met; 596: T/Ile; 653: A/Tyr; 671: G/Ser; 689: T/Phe; 827: C/Thr; 839: T/Val........................... P. emeiensis 251: C/(Ser/Thr); 596: C/Thr; 653: T/Phe; 671: A/Asn; 689: A/His; 827: T/Met; 839: G/Ser................................2 2. 266: T/Leu; 278: C/Thr; 284: G/Ser; 440: A/Asn; 608: A/Tyr; 995: T/(Met/Ile).................................................3 266: C/Ser; 278: T/Ile; 284: T/Met; 440: C/Thr; 608: T/Phe; 995:G/Ser..........................................................4 3. 46: A/Thr; 223: G/Val; 452: T/Ile; 637: T/Leu; 955: A/Met; 1019: T/Met. ............................................ P. javana 46: G/Ala; 223: A/Ile; 452: C/Thr; 637: A/Met; 955: G/Val; 1019: C/Thr........................ P. tanydactylia sp. nov. 4. 106: G/Val; 275: G/Ser; 682: A/Thr; 926: A/Asn; 967: T/Phe; 1015: A/Ile. ...................................................... 5 106: A/Ile; 275: A/Asn; 682: T/Ser; 926: T/Phe, 967: A/Met; 1015: T/Phe.......................................................6 5. 452: C/Thr; 799: T/Ser; 809: A/Asn; 883: G/Ala; 905: T/Phe. ....................................... P. brevipenies sp. nov. 452: T/Ile; 799: A/Thr; 809: G/Ser; 883: T/Ser; 905 A/Tyr.............................................. P. hirtipenis sp. nov. 6. 70: G/Ala; 205: A/Thr; 314: G/Ser; 604: G/Val; 932: C/Ser; 935: A/Lys; 941: C/Thr.............. P. longiseta sp. nov. 70: T/Ser; 205: T/Ser; 314: C/Thr; 604: A/Met; 932: T/Leu; 935: G/Ser; 941: A/Asn. ......................... P. argentosa

PARALEUCOPHENGA ARGENTOSA ( OKADA, 1956)

( FIGS 1 , 9 )

Leucophenga (Trichiaspiphenga) argentosa Okada, 1956: 24 (synonymized as P. invicta by Okada, 1988: 620).

Paraleucophenga shanyinensis Chen & Toda, 1994: 71 syn. nov.

Specimens examined: Japan: paratypes, two males, one female, Setagawa , Tokyo, 2.iv.1954, T . Okada ( NSMT) ; two males, Ueinohara , Yamanashi, 12.viii.2002, ex. around human eyes, H.W. Chen ; two males, one female, Tomochi , Kumamoto, 17.viii.2002, ex. around human eyes, H.W. Chen . China: one male, Huajiang, Guilin , Guangxi, 9.ix.2001, H. Watabe .

Distribution: Japan (Honshu and Kyushu) and China (Anhui, Zhejiang, and Guangxi).

Remarks: Okada (1988) checked a specimen of P. invicta ( Walker, 1857) from the type locality (Taiwan), and thought that P. argentosa was a synonym of P. invicta . After examining these specimens, we confirmed that the two forms are actually different species. In fact, Okada (1988) mistook the anteroventral processes of the gonopod for the paramere, and thus concluded that P. argentosa was a synonym of P. invicta . On the other hand, no morphological diagnostic character has been found to discern between P. argentosa and P. shanyinensis , thereby supporting P. shanyinensis as a new synonym.

PARALEUCOPHENGA EMEIENSIS SIDORENKO, 1998

( FIGS 2 , 10 )

Specimens examined: China: holotype male, Emeishan , Sichuan, 550 m a.s.l., M.J. Toda ( SEHU) . China: three males, two females, Wuyishan, Fujian, 700 m a.s.l., 17.viii.2001, on tree trunks, H.W. Chen; two males, Guanshan, Yifeng , Jiangxi, 500 m a.s.l., 8.vi.2004, ex. around human eyes, J.J. Gao; one male, Nonggang , Longzhou , Guangxi, 550 m a.s.l., 5.vi.1993, ex. around human eyes, Y.S. Cui; two males, Emeishan , Leshan , Sichuan, 1700 m a.s.l., 4.vii.2007, ex. around human eyes, H.Z. Cao and J.J. Jiang; one male, Fengtongzhai , Baoxing , Sichuan, 1600 m a.s.l., 18.ix.2005, H.W. Chen; one male, Leigongshan , Kaili , Guizhou (~ 26°24′N, 108°19′E), 800 m a.s.l., 16.vii.2000, ex. around human eyes, J.J. Gao; five males, Menglun, Xishuangbanna, Yunnan, 14.ix.2002, 24–26.xii.2004, 17–20.iv.2007, 600– 900 m a.s.l., ex. around human eyes, H.W. Chen and J.J. Gao .

Distribution: China (Fujian, Jiangxi, Guangxi, Sichuan, Guizhou, and Yunnan).

PARALEUCOPHENGA INVICTA ( WALKER, 1857)

( FIG. 3 )

Helomyza invicta Walker, 1857: 130 (Borneo?).

Leucophenga (Trichiaspiphenga) invicta: Duda, 1924: 185 (Taiwan) .

Paraleucophenga invicta: Lin & Wheeler, 1972: 248 ( Taiwan; Nepal); Okada, 1988: 620 ( Taiwan; Malaysia, Sarawak).

Paraleucophenga triseta Hendel, 1914: 114 (Taiwan) .

Specimens examined: China: one male, one female, Kuantsuling, Tainan , Taiwan, 6.vi.1965, T. Saigusa

(NSMT). Malaysia: one male , Sarawak, 25–27. vi.1975, H. Shima ( NSMT) .

Distribution: Taiwan; Nepal (Kathmandu); Malaysia ( Sarawak); Indonesia (Java and Sumatra).

PARALEUCOPHENGA JAVANA OKADA, 1988

( FIGS 4 , 11 )

Paraleucophenga javana Okada, 1988: 621 .

Specimens examined: Holotype male and one male paratype: Indonesia, Tugu , Java, 17–30.vii.1971, T. Okada ( NSMT). China: one male, Dinghushan, Zhaoqing, Guangdong, 6.v.2005, M.F. Xu; three males, Nonggang, Longzhou, Guangxi, 4.vi.1993, 23.viii.2004, ex. around human eyes, Y.S. Cui and H.W. Chen; two males, Menglun, Xishuangbanna, Yunnan, 12.viii.2002, M.J. Toda.

Distribution: China (Guangdong, Guangxi, and Yunnan), Indonesia (Java).

Figures

Figures 16–19. 16, 17, Paraleucophenga brevipenis sp. nov. ♂; 18, 19, Paraleucophenga hirtipenis sp. nov. ♂. 16, 18, epandrium (epand), cercus (cerc) and surstylus (sur); 17, 19, hypandrium (hypd), paramere (pm), gonopod (gon), aedeagus (aed), and aedeagal apodeme (aed a). Scale bars: 0.1 mm.

Figures 20–23. 20, 21, Paraleucophenga longiseta sp. nov. ♂; 22, 23, Paraleucophenga tanydactylia sp. nov. ♂. 20, 22, epandrium, cercus, and surstylus; 21, 23, hypandrium, paramere, gonopod, aedeagus, and aedeagal apodeme. Scale bars: 0.1 mm.

Figures 1–8. Abdominal tergite patterns in males. 1, Paraleucophenga argentosa (Okada, 1956); 2, Paraleucophenga emeiensis Sidorenko, 1998; 3, Paraleucophenga invicta (Walker, 1857); 4, Paraleucophenga javana Okada, 1988; 5, Paraleucophenga shimai Okada, 1988; 6, Paraleucophenga brevipenis sp. nov.; 7, Paraleucophenga longiseta sp. nov.; 8, Paraleucophenga anydactylia sp. nov.

Figures 9–15. Abdominal fifth sternite in males. 9, Paraleucophenga argentosa (Okada, 1956); 10, Paraleucophenga emeiensis Sidorenko, 1998; 11, Paraleucophenga javana Okada, 1988; 12, Paraleucophenga brevipenis sp. nov.; 13, Paraleucophenga hirtipenis sp. nov.; 14, Paraleucophenga longiseta sp. nov.; 15, Paraleucophenga tanydactylia sp. nov. Scale bars: 0.1 mm.