Epacanthaclisis batangana Yang, 1992

Zheng, Yuchen, Badano, Davide, Aspöck, Ulrike, Aspöck, Horst, Hayashi, Fumio & Liu, Xingyue, 2025, Revision of the antlion genus Epacanthaclisis Okamoto, 1910 (Neuroptera: Myrmeleontidae: Dendroleontinae), Zootaxa 5657 (1), pp. 1-100 : 85-89

Epacanthaclisis batangana Yang, 1992

( Figs. 50–51 )

Epacanthaclisis batangana Yang, 1992: 447 (type locality: China, Sichuan, Garze, Batang county; holotype in CAU). Ao et al., 2010: 49 ( Epacanthaclisis , declaration the holotype lost). Wang et al., 2018: 48 ( Epacanthaclisis , declaration the holotype lost). Yang et al., 2018: 62 ( Epacanthaclisis ).

Diagnosis. Frons generally dark brown, pale yellowish-brown posteriorly. Pronotum dark brown, anteriorly with three pale yellowish-brown spots, posteriorly with a pair of large pale yellowish-brown markings, laterally with a pair of pale yellowish stripes. Mesoprescutum dark brown, laterally with two pair of pale yellowish-brown spots; mesoscutellum anteriorly dark brown, medioposteriorly pale yellowish-brown, posterior margin black. Fore femur generally dark brown.

Re-description of holotype. Size. Head width: 3.22 mm; forewing length: 45.21 mm; hindwing length: 44.46 mm.

Head. Vertex generally dark brown ( Fig. 51C ). Scape and pedicel each yellowish-brown, internally with a dark brown spot; basal flagellomeres each basally with a dark circular marking, distally pink, distally darken; swollen flagellum dark brown. Frons generally dark brown, pale yellowish-brown posteriorly ( Fig. 51B ). Clypeus pale yellowish-brown, covered with dark and pale setae. Labrum yellowish-brown. Maxillary palpus generally yellowish-brown. Labial palpus generally yellowish-brown, distal segment fusiform and generally dark brown, distal part yellowish-brown. Mandibles yellowish-brown with distal half dark brown.

Thorax. Dark brown with pale yellowish-brown markings. Pronotum dark brown, anteriorly with three pale yellowish-brown spots, posteriorly with a pair of pale yellowish-brown markings, laterally with a pair of pale yellowish stripes; laterally covered with pale hairs. Mesoprescutum dark brown, laterally with two pair of pale yellowish-brown spots; mesonotum damaged; mesoscutellum anteriorly dark brown, medioposteriorly pale yellowish-brown, posterior margin black. Metanotum generally dark brown ( Fig. 51C ). Pleurae generally dark brown, with pale yellowish-brown markings.

Legs. Covered with many pale and dark setae. All coxae pale yellowish-brown each with an indistinct dark spot; tibial spurs brown, barely curved; tarsomere 5 as long as entire length of tarsomeres 1–4; pretarsal claws slightly curved, protruded basally. Foreleg: femur generally dark brown; tibia generally dark brown and black, basally and medially with pale yellow markings; tibial spur reaching tip of tarsomere 3; tarsi generally dark brown. Midleg: Similar to foreleg. Hind leg: femur pale yellow, distally dark brown; tibia pale yellow, mediobasally with a dark brown spot, distally dark brown; tibial spur reaching base of tarsomere 3 ( Fig. 51A ).

Wings. Mostly hyaline, with some markings. Forewing longer than hindwing. Pterostigma pale. Forewing costal space with a distinct dark brown spot proximad pterostigma; rhegma as a slightly curved slender stripe; mediocubital area with indistinct scattered spots; cubital area basally with an oblique indistinct dark brown marking; marginal cubital area with scattered dots; costal, distal half of prefork and basal half of posterior crossveins mostly connected by oblique veinlets; four to five presectoral crossveins present, distal presectoral cells sometimes biloculate; RP with 10–11 branches; RP origin basal to MP fork, nearly align at CuA fork; posterodistal margin slightly concave. Hindwing mostly hyaline; rhegma as a dark brown dot; tip of the MP and Cu fused part with a dark brown dot; two presectoral crossveins present ( Figs. 50 , 51A ).

Abdomen. Female abdomen dark brown. Female genitalia. Pregenital plate slender tapered. Anterior branches of gonocoxites 8 weakly developed, tuberculate, covered with setae; posterior branches of gonocoxites 8 short digitiform, externally concave, covered with many long setae; gonapophyses 8 as transversal bands, internally curved and sharp. Gonocoxites 9 covered with short stout setae. Ectoproct appears narrowed and semicircular in lateral view ( Figs. 51D–E ).

Type material. Holotype ♀, CHINA: Sichuan, Garze, Batang County [OE塘县], 2650 m, 13.VIII.1982, Xuezhong Zhang ( CAU).

Additional material examined. Adult: 1♀, “Batang. (Tibet). Im Tal des Yangtze (ca. 2800 m), 5.7.1936. H. Höne ” = CHINA: Sichuan, Garze, Batang County ( ZFMK) .

Distribution. China (Sichuan).

Remarks. Ao et al. (2010) and Wang et al. (2018) mentioned that the holotype of E. batangana was lost. However, based on our comparison of original description and photo ( Yang 1992) with all Epacanthaclisis specimens from CAU, we found that the holotype of E. batangana was not lost. Instead, this holotype was misidentified as the only specimen of E. moiwana (a Japanese endemic species not distributed in China) in CAU. Additionally, the collection label of E. batangana was somehow switched with that of an E. banksi specimen. In Ao et al. (2010) and Wang et al. (2018), the collection label of E. batangana was switched to “Jiuzhaigou County, Nanping Town”, matching the collection information of some E. banksi specimens. Meanwhile, a collection label of one E. banksi specimen matched the collection information in the original description of E. batangana . Furthermore, E. banksi is endemic to the Qionglai and Minshan mountain ranges and is not distributed in Batang, which is in the Shaluli mountain range. Therefore, we are confident that the only Chinese specimen identified as E. moiwana by Ao et al. (2010) is the holotype of E. batangana . Notably, Yang (1992) also incorrectly identified the female holotype as a male.

To date, male specimens of this species have yet to be recorded. The female holotype has short, external, concave digitiform posterior branches of gonocoxites 8, and its female gonocoxites 9 have short stout setae, which are like those of females in the E. moiwana group and the E. tuyuezhengi group. Until male specimens and fresh specimens are available, we cannot determine to which species this species belongs.

Key to species of Epacanthaclisis based on adult characters

1. Pronotum anteriorly with three pale yellowish-brown spots, posteriorly with a pair of large pale yellowish-brown markings, laterally with a pair of pale yellowish stripes............................................ E. batangana Yang, 1992

- Pronotum with other patterns............................................................................ 2

2. Hindwing 0.85 times as long as forewing. Male wings narrowed, blade-like. Male abdominal terga 4–5 barely swollen, without any special tuft setae or bristles. Male gonocoxites 9 as a pair of strongly sclerotized plates, internally with short tapered setae. Male ectoproct laterally expanded into digital projection, at least twice as wide as long....... 3 (the E. tuyuezhengi group)

- Hindwing at least 0.9 times as long as forewing. At least one of male abdominal terga 4–5 swollen, mostly with special tuft setae or bristles. Male gonocoxites 9 modified, divided into thick or strongly sclerotized external gonocoxites 9 and slightly sclerotized internal gonocoxites 9. Male ectoproct slightly or barely expanded..................................... 4

3. Pronotum laterally with three pairs of yellowish-brown spots. Male ectoproct twice as wide as long.......................................................................................... E. tuyuezhengi Zheng & Liu sp. nov.

- Pronotum laterally with a pair of discontinuous yellowish-brown stripes and a pair of continuous yellowish-brown stripes. Male ectoproct 2.5 times as wide as long............................................. E. zhihaoi Zheng & Liu sp. nov.

4. Forewing slightly longer than hindwing. Male abdominal tergum 4 with a pair of short stout tufts of bristles, the tip of the pair bristles mixed up in a ball of brown wool. Male gonocoxites 9 plate-like; gonocoxites 11 thin, arched............................................................................................... 5 (the E. continentalis group)

- Forewing subequal to or slightly longer than hindwing. Male abdominal tergum 4 without pair of short stout tufts of bristles. Male gonocoxites 9 as a pair of slender curved structures in ventral view; gonocoxites 11 extremely thick, as long as wide... .................................................................................................. 13

5. Anterior Banksian line of forewing present................................................................. 6

- Anterior Banksian line of forewing absent................................................................. 10

6. Male abdominal tergum 4 with less dark markings........................................................... 7

- Male abdominal tergum 4 medially and posteriorly dark....................................................... 9

7. Male abdominal terga 5–6 with many short dark brown stripes. Male external gonocoxites external gonocoxites 9 thrice as wide as the internal part; gonostyli 11 flattened in lateral view.................................................................................................. E. liuyingqiae Zheng, Badano, U. Aspöck, H. Aspöck & Liu sp. nov.

- Male abdominal terga 5–6 mostly dark brown. Male external gonocoxites external gonocoxites 9 twice as wide as the internal part; gonostyli 11 not flattened in lateral view............................................................... 8

8. Pronotum with a pair of lateral pale yellowish-brown stripes reaching the anterolateral margin of pronotum; thicker part of male external gonocoxites 9 twice as wide as the internal part; internal gonocoxites 9 posteriorly not concave............................................................... E. darman Zheng, Badano, U. Aspöck, H. Aspöck & Liu sp. nov.

- Pronontum with a pair lateral pale yellowish-brown stripes not reaching the anterolateral margin of pronotum; thicker part of male external gonocoxites 9 thrice as wide as the internal part; internal gonocoxites 9 posteriorly concave................................................................................... E. victor Badano, Zheng & Liu sp. nov.

9. Pronotum anterolaterally with a pair of curved pale yellowish-brown markings, posterolaterally with a pair of pale yellowish-brown spots. Male abdominal tergum 5 pale, medially with a longitudinal dark brown marking. Male external gonocoxites 9 not protruded, and internal gonocoxites 9 barely concave in anteroventral view................................................................................... E. zhiweii Zheng, Badano, U. Aspöck, H. Aspöck & Liu sp. nov.

- Pronotum medially with five yellowish spots, laterally with a pair of yellowish stripes. Male abdominal tergum 5 mostly dark brown. Male external gonocoxites 9 slightly protruded, and internal gonocoxites 9 concave in anteroventral view.......................................................... E. zuqii Zheng, Badano, U. Aspöck, H. Aspöck & Liu sp. nov.

10. Vertex posteriorly with a triangular black marking. Female gonocoxites 8 only one pair....... E. alaica Krivokhatsky, 1998

- Vertex marking not triangular. Female gonocoxites 8 two pairs................................................ 11

11. Pronotum pale yellowish-brown, medially with a pair of adjacent longitudinal dark brown stripes; laterally with a pair of short dark lines; lateral margin black. Forewing rhegma as an oblique slender pale brownish stripe.............................................................................................. E. continentalis Esben-Petersen, 1935

- Pronotum pale yellowish-brown, with scattered dark brown markings. Forewing generally hyaline.................... 12

12. Pronotum anteriorly with a pair of curved transversal dark brown bands, medially with a pair of longitudinal short dark brown stripes, posterolaterally with a pair of indistinct short longitudinal dark brown stripes, and lateral margin with a pair of longitudinal dark brown stripes. Pterostigma pale................................... E. hamata Krivokhatsky, 1998

- Pronotum anteriorly with a pair of black spots, posteriorly with a pair of closed black markings, laterally with two pairs of longitudinal black stripes. Pterostigma yellowish-brown.......................... E. kuldurguch Krivokhatsky, 1998

13. Male abdominal segments 4–5 swollen; tergum 4 medially concave, and concave part with dense, relatively long setae. Male external gonocoxites 9 strongly sclerotized with a pair of a pair of membrane tubercles exteriorly.................................................................................................... 14 (the E. maculosa group)

- Only male abdominal segment 5 swollen. Male external gonocoxites 9 strongly sclerotized but without membrane tubercle................................................................................ 19 (the E. moiwana group)

14. Antenna and legs mostly black. Wings with many dark markings............................................... 15

- Antenna dark and pink, legs pale yellow and dark brown. Wings scattered with a few small markings.................. 18

15. Wings with many mostly distinct oblique thin markings...................................................... 16

- Wings with many mostly indistinct rounded markings....................................................... 17

16. Pronotum generally pale yellow, anteriorly with a pair of dark brown spots, laterally with two pairs of short dark brown stripes. Forewing rhegma and basal marking on cubital area fragmentary and smaller.................. E. maculata ( Yang, 1986)

- Pronotum generally dark brown, laterally with a pair of oblique pale yellowish stripes. Forewing rhegma and basal marking on cubital area continuous and larger...................................... E. wuchaoi Zheng, Badano & Liu sp. nov.

17. Pronotum generally dark brown, medially and posterior margin indistinct yellow or orange. Wings long ovoid; forewing costal space with two rows of spots on basal half, distal half with one row of spots; marking on basal cubital area large and distinct. Male tergum 4 without dark spot. Male lateral ectoproct slightly expanded transversally, lateral margin with a longitudinal dark brown stripe..................................... E. jiazhii Zheng, U. Aspöck, H. Aspöck, Hayashi & Liu sp. nov.

- Pronotum mostly yellow or orange with laterally dark brown. Forewing costal space with a row of spots on basal half; marking on basal cubital area as an indistinct dark brown spot. Sunk part of male tergum 4 with a dark brown marking. Male lateral ectoproct barely expanded, without marking............................................ E. maculosa ( Yang, 1986)

18. Pronotum generally yellowish-brown, laterally with a pair of dark brown spots, posterolateral margin with a pair of subtriangular dark brown marking. Sunk part of male tergum 4 with a dark brown marking and covered with dense long setae. Male external gonocoxites 9 oblique rectangular in ventral view......................................... E. minana ( Yang, 1999)

- Pronotum generally dark brown, laterally with a pair of pale yellowish dots and a pair of curved pale yellowish stripes. Sunk part of male tergum 4 without marking, covered with dense long black setae. Male external gonocoxites 9 slightly curved in ventral view, with a pair of curved projection in caudal view...................... E. xiaohongae Zheng & Liu sp. nov.

19. Male abdominal tergum 5 without anterior pair of tufts of bristles....................... E. moiwana ( Okamoto, 1905)

- Male abdominal tergum 5 with anterior pair of tufts of bristles................................................. 20

20. Pale yellowish-brown part on pronotum wide. Male abdomen mostly pale yellow. Male internal gonocoxites 9 transversal in ventral view................................................................. E. banksi Krivokhatsky, 1998

- Pale yellowish-brown part on pronotum narrow. Male abdomen dark brown and pale yellow. Male internal gonocoxites 9 oblique............................................................................................ 21

21. Vertex anteriorly with three black spots fused with a medial transversal black band, posteriorly with three transversal spots, the middle one fused with the medial band. Wings infuscate with markings. Male abdominal tergum 3 pale yellowish-brown anteriorly and dark brown posteriorly................................... E. longwai Zheng, Hayashi & Liu sp. nov.

- Vertex anteriorly with three black spots, medially with an indistinct black marking, posteriorly with three transversal spots. Wings hyaline with markings. Male abdominal tergum 3 generally pale yellowish-brown, with two pair of brown markings.......................................................................... E. ningshana Wan & Wang, 2010

Larva identification by molecular analysis

Larvae that were not identified through the rearing to the adult stage, were assigned to a species based on the results of COI barcodes ( Fig. 52 ; Table 2). This approach eliminated the uncertainty due to sympatric species. The COI interspecific genetic divergence among the four sequenced Epacanthaclisis varies from 0.101 to 0.041. The intraspecific genetic divergence within E. zuqii sp. nov. specimens are always 0.000. The intraspecific genetic divergence within E. moiwana based on Matsumoto et al. (2016) from 0 to 0.012. The intraspecific genetic divergence within E. banksi varies from 0.000 to 0.002. The intraspecific genetic divergence within E. ningshana from 0.000 to 0.007.

Figures

FIGURE 50. Habitus of Epacanthaclisis batangana Yang, 1992, holotype female, Batang (Sichuan).

FIGURE 51. Epacanthaclisis batangana Yang, 1992, adult, holotype female, Batang (Sichuan). A. Habitus; B. head, frontal view; C. head and thorax, dorsal view; female terminalia: D. lateral view; E. ventral view.

FIGURE 52. Molecular phylogeny of selected species of Myrmeleontidae based on COI barcoding sequence data. Tree topologies are identical among the Neighbour-joining analyses.