Adult: Frons wider than diameter of compound eye; vertex raised. Pronotum as long as wide. Legs short, as long as thorax at most; foreleg with femur nearly twice as long as coxa; maximum width of femur wider than that of tibia; tibial spurs slightly curved, not reaching tarsomere 5. Wings distally rounded. Forewing mediocubital area with some indistinct brown markings. Banksian lines absent. Male pilula axillaris absent. Male gonocoxites 9 as a pair of broad plates in posterior view, apex strongly sclerotized and scaly; gonocoxites 11 arched. Female ventral segment 8 forming an antrum, gonocoxites 8 as a pair of digitiform structures; gonapophyses 8 various, absent in some species, nearly wide rectangular if present; digging setae on gonocoxites 9 long and blunt. — Larva: Body laterally covered with many blunt pale setae. Head wider than long. Dolichasters on anterior margin of clypeo-labrum thick, distally swollen and truncate. Ocular tubercles prominent. Distance between the first and second teeth on mandible longer than that between the second and third teeth. Pair of large spiracles present on laterodorsal abdominal segment 1. Odontoid process on abdominal segment 8 present. Abdominal sternum 9 ventrally with some tapered setae of uneven size; rastrum with four distal pair of tapered digging setae.

The known larvae of Layahima are not fossorial and do not camouflage under debris, living exposed under rocks or on tree trunks.

China; India; Nepal; Thailand; Vietnam (Fig. 18).

According to Zheng et al. (2023), Layahima lacks female gonapophyses 8. However, further examination demonstrated that the absence of gonapophyses 8 only occurs in the L. elegans species group, while this character is present in most other species of the genus. Besides, unlike the related genus Paralayahima Zheng, Badano & Liu, 2023, the gonapophyses 8 of Layahima is broad and subrectangular, whereas the gonapophyses 8 of Paralayahima is thin and ribbon-like.

Adult: Body pale yellowish brown with some black and dark brown markings, parts with numerous black dots at base of setae. Both vertex and postgena each with a pair of distinct black spots (Fig. 3B, C). Frons and clypeus with six black spots (Fig. 3B). Pronotum medially with a black spot and laterally with four pairs of black spots. Forewing rhegma present as two indistinct brownish markings (Figs 3C, 5A–C). Male sternum 9 wide, posterior part narrowed and short (Fig. 3E); gonocoxites 9 as a pair of wide, oblate trapezoid plates, twice as wide as long; gonocoxites 11 are triangularly arched (Figs 3H–L, 5D). Female pregenital plate shaped as an indistinct tiny sclerite (Fig. 5E). Gonocoxites 8 short and wide, digitiform (Fig. 3F, G). — 1st instar larva: Head generally brown and dark brown (Fig. 7A, B). Antenna short and robust. Thoracic setiferous processes tubercular; anterior mesothoracic setiferous processes larger than others (Fig. 7C). Abdomen greenish brown, with many dark markings; dorsal abdominal segment 1 with a pair of large transversal dark markings (Figs 6, 7D).

Figure 2. 

Habitus of Layahima spp. from China, adults: A L. chiangi Banks, 1941, male, Cawarong (Xizang); B L. chiangi, male, holotype (© D. Grimaldi from AMNH); C L. qilin sp. nov., male, holotype, Weixi (Yunnan); D L. haohani sp. nov., female, holotype, Puer (Yunnan); E L. pixiu sp. nov., female, holotype, Hutiaoxia (Yunnan); F L. zhitengi sp. nov., female, holotype, Fugong (Yunnan).

Figure 3. 

Layahima chiangi Banks, 1941, adult: A Habitus, female holotype; B head, frontal view; C head and thorax, dorsal view. D, E male terminalia: D Lateral view, E ventral view. F, G Female terminalia: F Lateral view, G ventral view. H–L Male genitalia: H Ventral view, I anteroventral view, J caudal view, K lateral view, L dorsal view.

Figure 4. 

Layahima chiangi Banks, 1941, living photos and its habitat: A Habitat, Cawarong (Xizang), B living adult. C Newly hatched first instar larva. D Well-fed first instar larva (photos by Y. Zheng).

Figure 5. 

Comparison of different individuals of Layahima chiangi Banks, 1941 and L. qilin sp. nov. A–E L. chiangi, F–J. L. qilin sp. nov. A–C, F–H Head-thorax, dorsal view; D, I male genitalia, ventral view; E, J female pregenital plate, ventral view. Location (A, B, D, E) Cawarong (Xizang). C Holotype, type locality “Loutsechiang”, probably near to Cawarong (© D. Grimaldi from AMNH). F Holotype, type locality “Weixi, Zhonglu, Zhazidi” (Yunnan). G, I Batang (Sichuan). H Pantiange (Yunnan). J Baimaxueshan (Yunnan).

Figure 6. 

Habitus of Layahima chiangi Banks, 1941, first instar larva: A Dorsal view, B ventral view, C lateral view.

Figure 7. 

Layahima chiangi Banks, 1941, first instar larva: A Head, dorsal view; B head, ventral view; C thorax, dorsal view, on black background; D abdominal segments 8–9, ventral view.

Size. Head width: 2.27–3.22 mm; forewing length: 28.11–32.24 mm; hindwing length: 28.08–32.20 mm. — Head. Vertex and postgena with a pair of dark brown spots. Scape mostly dark brown; pedicel pale yellowish brown with a dark brown circular marking; antenna mostly yellowish brown, non-swollen part of flagellomeres each basally dark brown, swollen part of flagellum distally dark brown. A dark brown marking present around scapes, anteriorly concaved. Frons and clypeus with six black spots. Labrum pale yellow. Mandibles pale yellowish brown with distal half dark reddish brown. Maxillary palpus yellowish brown. Labial palpus generally yellowish brown, distal palpomere fusiform (Figs 3B, C, 5A–C). — Thorax. Pronotum medially with a black spot and laterally with four pairs of black spots. Mesoprescutum mostly black, anteriorly with a pair of pale yellowish-brown spots; mesonotum pale yellowish brown, with three pair of black stripes and a pair of black spots, pale yellowish-brown part with some black dots; mesoscutellum anteriorly black, mostly pale yellowish brown, variation present in different individuals. Metanotum pale yellowish brown and black; metascutellum dark brown, laterally with a pair of pale yellowish-brown spots. Pleurae pale yellowish-brown and black (Figs 3B, C, 5A–C). — Legs. Short, as long as thorax at most, with many black and pale setae. All tibial spurs slightly curved; tarsomeres each generally pale yellow with distal part dark brown; tarsomere 5 pale yellow, with distal part dark brown, nearly as long as entire length of tarsomeres 2–4; pretarsal claws curved, dark reddish brown. Foreleg: coxa pale yellow with a black spot; femur dorsally brown to dark brown, ventrally pale yellowish brown; tibia pale yellow with a dark anterior marking, distal part dark brown; tibial spurs reaching base of tarsomere 3. Midleg: Coxa pale yellowish brown, black basally; femur mostly yellowish brown, distally dark brown; tibia pale yellow, with some black dots, distal part dark brown; tibial spurs reaching basal of tarsomere 3. Hind leg: Coxa mostly pale yellowish brown, black basally; femur yellowish brown, with distal part dark brown; tibia pale yellow, distally dark brown; tibial spurs reaching tip of tarsomere 1; tarsomere 1 longer than that of fore- and midleg (Fig. 3A). — Wings. Mostly hyaline, distally rounded. Banksian lines absent. Pterostigma pale. Forewing slightly longer than hindwing. Forewing costal space with an indistinct brown spot proximal to pterostigma; subcostal area hyaline; radial area with some indistinct brown spots; mediocubital area with a few indistinct brown spots; rhegma as two indistinct brown markings; cubital area with a basal indistinct dark brown marking; costal crossveins generally simple on basal 1/3 part, while distal 2/3 part each connect with a short oblique veins; RP originates proximally to MP fork; four to five presectoral crossveins present, with from two to three biareolate cells; RP with eight to nine branches. Hindwing rhegma indistinct; presectoral area with only one crossvein; RP originates anteriorly to MP fork (Figs 2A, B, 3). — Abdomen. Pale yellowish brown with some dark brown markings. Tergum 1 generally pale yellowish brown; tergum 2 with a dark brown marking; terga 3–8 each dark brown on posterior margin. — Male genitalia. Sternum 9 basal half wide, posterior part narrow, narrowed part as long as wide (Fig. 3E); gonocoxites 9 shaped as a pair of widely short trapezoid plates, twice as wide as long; gonocoxites 11 triangularly arched; gonostyli 11 slightly prominent (Figs 3H–L, 5D). Ectoproct slightly protruded (Fig. 3D). — Female genitalia. Pregenital plate weakly sclerotized. Gonocoxites 8 rounded, short digitiform; gonapophyses 8 rounded. Gonocoxites 9 with many tapered stout setae, slightly curved. Ectoproct rounded on distal margin (Fig. 3F, G).

Size. Body length (excluding mandible): 2.13–3.22 mm; head length: 0.65–0.78 mm; head width: 0.69–0.82 mm; mandible length: 0.75–0.89 mm. — Head. Nearly rectangular in dorsal view, wider than long, generally brown and dark brown. Dolichasters thick, distally swollen and truncate, dark and pale, present on clypeo-labrum, dorsal head, and basal mandible. Dorsal head dark brown posteromedially and laterally. Ventral head mostly brown, lateral margin dark brown. Antenna short and robust, dark brown. Ocular tubercles well prominent. Mandible dark brown, basally with 3–4 interdental mandibular setae; first tooth shorter than second tooth; second tooth nearly as long as third tooth; two short interdental mandibular setae between first and second teeth; no seta between second and third teeth. Labial palpi dark brown (Figs 6A, B, 7A, B). — Thorax. Thoracic setiferous processes tubercular, with many pale setae and few dark setae. Pronotum dark brown and yellowish brown, with some dark and pale dolichasters. Mesothorax with spiracles present on short stout brown sclerotized tubercle; anterior process longer than posterior process. Metathorax medially with many dark dots; metathoracic setiferous processes equal in length (Figs 6A, 7C). — Legs. Pale brown, with many slender and short tapered setae (Fig. 6B, C). — Abdomen. Greenish brown, with many dark markings; dorsal abdominal segment 1 with a pair of large transversal dark markings. Abdomen dorsally covered with dark setae, laterally covered with pale setae. Sternum 9 ventrally with seven to eight tapered setae of uneven size; rastrum distally with four pair of tapered digging setae (Figs 6, 7D).

Holotype ♂, “Loutsechiang, Yunnan, China. Accession No. 24141”, the Nujiang dry hot river valley around the Cawarong region near Yunnan (AMNH).

Adult: 18♂10♀, CHINA: Xizang, Nyingchi, Zayu County [察隅县], Cawarong Township [察瓦龙乡], Ridong [日东], 1930 m, 27–29.VI.2023, Yuchen Zheng (CAU). 4♂4♀, same information as above (IZCAS). 1♀, same locality as above, 1930 m, 11.VIII.2021, Xingyue Liu (IZCAS). 2♂1♀, same locality as above, 1900 m, 30.VII.2020, Yejie Lin (CAU). 1♂1♀, same locality as holotype (AMNH). 12♂, “Thibet” = [CHINA: Xizang], Coll. Lacroix (MNHN). — Larva: 5 1st larvae, hatched from the eggs, four preserved in 95% ethyl alcohol, one is still rearing, same locality as above, 27–29.VI.2023, Yuchen Zheng (IZCAS).

China (Xizang) (Fig. 18).

The larvae were obtained from eggs laid by field-collected females, so their natural habitat remains unknown. In the artificial settings, the larvae showed a preference for the bare environment rather than burying under sand like most antlions. The larvae fed on termites and springtails around them or crawling over their bodies, catching them with mandibles.

This species is endemic to the Nujiang dry hot river valley near the Cawarong Region. According to Zheng et al. (2023), the absence of an anteromedial marking on pronotum of the holotype of L. chiangi is individual variability. However, a dozen specimens from Yunnan and western Sichuan preserved in CAU show some differences from the holotype, including the presence of the anteromedial markings, but the differences in all other characters are subtle. The type locality of L. chiangi is ambiguous (“Loutsechiang, Yunnan”, now called “Nujiang”, a watershed that throughs Xizang, Yunnan, Myanmar and Thailand), so it was not possible to locate the original collecting site, and the holotype and we can only refer to type photos for comparison.

Further collecting efforts in 2023 in the Nujiang dry hot river valley of Cawarong, Xizang, allowed to find further specimens of L. chiangi, which clarified the taxonomic status of this species through genital comparisons and molecular analyses. The morphology of these specimens is consistent with the holotype of L. chiangi, confirming that the specimens from Yunnan and Sichuan identified as this species in Zheng et al. (2023), belong to an undescribed taxon, i.e., L. qilin sp. nov.

L. chiangi is characterized by dark brown scape, pronotum lacking anteromedial marking, abdominal terga with reduced dark markings, male sternum 9 short and posteriorly narrowed, male gonocoxites 9 twice as wide as long, male gonocoxites 11 triangularly arched, female pregenital plate weakly sclerotized, and shorter female gonocoxites 8. Layahima qilin sp. nov. differs in the pronotum with anteromedial marking, abdominal terga with wide dark markings, male sternum 9 elongate and posteriorly narrowed, male gonocoxites 9 as long as wide, female pregenital plate shaped as an anterior sclerotized prominent papilla and a posterior ridge-like prominence, and female gonocoxites 8 longer than that of L. chiangi.

We also shed new light on the type locality of L. chiangi with better precision. This species inhabits the unique habitat of the Nujiang dry hot river valley in Cawarong. This area represents an isolated “mountainous island”, surrounded by snow-capped mountains and monsoon rainforests, characterized by many endemic species, such as the spoon-winged lacewing Sinonemoptera tibetana Zheng & Liu, 2024 (Zheng and Liu 2024). Therefore, it is likely that the type locality of L. chiangi is the town of Cawarong or its environs, rather than Yunnan. All the examined specimens from the river valleys of belonged to L. qilin sp. nov. (Jinshajiang river valley and Lancangjiang river valley) and L. pixiu sp. nov. (Jinshajiang river valley). Finally, it is worth noting that the type locality of L. chiangi is labeled as “Yunnan” instead of “Xizang” or “Tibet”, because Cawarong Township is close to Gongshan County in western Yunnan Province, and both places are near the Nujiang River. Moreover, Cawarong was visited by European and American missionaries in the mid-19^th early 20^th centuries (Liu 2019), and they often collected insect specimens referring to the churches on the territory as geographic information, resulting in strongly biased locality data.

Adult: Body pale yellow with dark markings. Vertex with three transversal black markings. Pronotum with three longitudinal black stripes, the middle one discontinuous, posterior margin mostly black (Fig. 8C). Forewing radial area distally with two indistinct dark brown markings, rhegma and basal cubital markings distinct. Hindwing radial area distally with an indistinct transversal dark brown stripe, marking of rhegma reaching wing margin (Figs 2D, 8A). Female pregenital plate shaped as a rounded sclerotized plate with a vertical prominence (Fig. 8D, E).

Figure 8. 

Layahima haohani sp. nov., adult: A Habitus, female, holotype; B head, frontal view; C head and thorax, dorsal view. D, E Female terminalia: D Lateral view, E ventral view.

Size. Head width: 2.12 mm; forewing length: 31.42 mm; hindwing length: 31.22 mm. — Head. Vertex pale yellow, with three transversal black markings. Scape mostly yellowish brown internally with a dark marking; pedicel mostly black; non-swollen part of flagellum brown, each flagellomere basally dark brown, swollen part of flagellum mostly black. A dark brown marking present around scape, anteriorly protruded. Frons and clypeus pale yellow. Labrum brown. Mandibles pale yellowish brown with distal half dark reddish brown. Maxillary palpus mostly yellowish brown, distal margin with a dark brown marking. Labial palpus generally yellowish brown (Fig. 8B, C). — Thorax. Pronotum with three longitudinal black stripes, the middle one discontinuous, posterior margin mostly black. Mesoprescutum mostly black, medially with a pale yellowish stripe, laterally with a pair of pale yellowish spots; mesonotum, medially mostly pale yellow, laterally mostly black with two pair of pale yellowish spots, lateral margin pale yellow; mesoscutellum black with a pair of pale yellowish markings. Metanotum black and pale yellow; metascutellum black with a pair of pale yellowish spots. Pleurae pale yellow and black (Fig. 8C). — Legs. Short, as long as thorax at most, with many dark setae. All tibial spurs slightly curved; tarsomeres 1–2 pale yellow; distally brown, tarsomeres 3–4 black; tarsomere 5 pale yellow, distally black, nearly as long as entire length of tarsomeres 2–4; pretarsal claws curved, dark reddish brown. Foreleg: coxa pale yellow with a black spot; femur mostly pale yellow; tibia pale yellow, basally and distally black; tibial spurs reaching tip of tarsomere 3. Midleg: Coxa pale yellow, black basally; femur basally pale yellow, distally dark brown; tibia pale yellow, basally and distally black; tibial spurs reaching tip of tarsomere 3. Hind leg: Coxa mostly pale yellow; femur pale yellow basally, other part black; tibia pale yellow, medially and distally black; tibial spurs reaching base of tarsomere 2; tarsomere 1 longer than that of fore- and midleg (Fig. 8A). — Wings. Mostly hyaline, with some dark brown markings. Banksian lines absent. Pterostigma pale. Forewing slightly longer than hindwing, posterior margin rounded; costal space distally with a few indistinct brown dots; subcostal area with a few dark brown spots; radial area distally with two indistinct dark brown markings; rhegma dark brown; mediocubital area with four indistinct dark brown spots; cubital area basally with a dark brown marking; costal crossveins generally simple; RP originates proximally to MP fork; four presectoral crossveins present, with one biareolate cell; RP with five to six branches. Hindwing radial area distally with an indistinct transversal dark brown stripe, marking of rhegma reaching wing margin; presectoral area with only one crossvein; RP originating anteriorly to MP fork (Figs 2D, 8A). — Abdomen. Dark brown and pale yellow. Tergum 1 pale yellow medially with a subtriangular spot. Terga 2–8 mostly dark brown, each anterior and posterior margin pale yellow. — Male genitalia. Unknown. — Female genitalia. pregenital plate shaped as a rounded sclerotized plate with a vertical prominence. Gonocoxites 8 digitiform, slightly posterior curved; gonapophyses weakly sclerotized. Gonocoxites 9 with many tapered stout setae, slightly curved. Ectoproct slightly truncated (Fig. 8D, E).

Holotype ♀, CHINA: Yunnan, Puer, Yunxian Township [云仙乡], Mamu Village [骂木村], 980 m, 9.IV.2023 (IZCAS).

China (Yunnan) (Fig. 18).

The new species is dedicated to Mr. Haohan Mao (毛昊涵), who helped us in obtaining the holotype of this rare species, also collecting many myrmeleontid specimens for us.

The new species can be easily distinguished from other Layahima species by the markings on the body and wings, and the shape of female pregenital plate.

Adult: See Zheng et al. (2023). — Larva: Lateral body covered with many pale setae. Head pale yellow, medially with three pair of dark brown markings, laterally with a pair of dark brown stripes fused with the lateral markings in dorsal view. Dolichasters on clypeo-labrum dark and pale. Mandible mostly pale yellow, distally reddish brown. Three short interdental mandibular setae between first and second teeth; one seta between second and third teeth (Fig. 10A, B). Thoracic setiferous processes digitiform; anterior mesothoracic setiferous processes twice as long as others, curved; posterior mesothoracic setiferous processes nearly as long as others on metathorax. Meso- and metanotum, and dorsal abdominal medially pale, each segment with a pair of black markings (Fig. 10C, D). Abdomen with dozens black dots, abdominal setiferous processes tubercular (Fig. 9A, B). Segment 9 rastrum distally with five pair of tapered digging setae (Fig. 10E).

Figure 9. 

Habitus of Layahima lhoba Zheng, Badano & Liu, 2023, third instar larva: A Dorsal view, B ventral view, C lateral view.

Figure 10. 

Layahima lhoba Zheng, Badano & Liu, 2023, third instar larva: A Head, dorsal view; B head, ventral view; C thorax, dorsal view, on black background; D thorax, dorsal view, on pale background; E abdominal segments 8–9, ventral view.

Size. Body length (excluding mandible): 4.32 mm; head length: 2.25 mm; head width: 2.27 mm; mandible length: 2.84 mm. Examined specimen discolored. — Head. Nearly rectangular in dorsal view, as long as wide, pale yellow and dark brown. Clypeo-labrum dark brown, with dark and pale dolichasters. Head pale yellow, medially with three pair of dark brown markings, laterally with a pair of dark brown stripes fused with the lateral markings in dorsal view, covered with pale setae laterally. Ventral head mostly pale yellow. Ocular tubercle prominent, ocular peduncle dark brown. Mandible mostly pale yellow with few irregular brown markings, distally reddish brown, basally with 3–4 interdental mandibular setae; first tooth shorter than second tooth; second tooth nearly as long as third tooth; three short interdental mandibular setae between first and second teeth; one seta between second and third teeth. Labial palp dark brown (Figs 9, 10A, B). — Thorax. Laterally with many pale setae. Thoracic setiferous processes digitiform, with dark brown dots. Pronotum pale yellow, medially with a forked dark brown marking, laterally with a pair of arched dark brown markings, covered with many dark and pale setae. Mesothorax with spiracles present on short, sclerotized tubercle, mostly pale, distally dark brown; anterior mesothoracic setiferous processes twice as long as others, curved. Posterior mesothoracic setiferous processes nearly as long as others on metathorax. Meso- and metanotum medially pale, each with a pair of black markings (Figs 9, 10C, D). — Legs. Pale yellow, with many slender and short tapered setae (Fig. 9B, C). — Abdomen. Pale yellow with many dark brown markings. Dorsally covered with dense black setae, with dozens black dots; medially pale, each segment with a pair of black markings. Abdominal setiferous processes tubercular, distally dark brown. Lateral margin covered with pale setae. Ventrally generally pale yellow. Segment 9 dorsally with two pair of stout tapered setae; ventrally with 17 tapered setae of uneven size, the external row of setae largest, the internal row of setae smallest; rastrum distally with four pair of tapered digging setae; rastrum distally with five pair of tapered digging setae (Figs 9A, B, 10E).

Figure 11. 

Habitus of adults of Layahima spp. associated with larvae: A L. lhoba Zheng, Badano & Liu, 2023; B L. yangi Wan & Wang, 2006.

Holotype ♂, China: Xizang, Nyingchi, Bome County, Yiong Town, Tangmai Township, 2100 m, 6.VIII.2020, Yuchen Zheng (CAU). Other type specimens were examined in Zheng et al. (2023).

Larva. 3rd instar larva preserved in 95% ethyl alcohol, CHINA: Xizang, Nyingchi, Medog County [墨脱县], Ximo Bridge [西莫桥], 726m, 10.VII.2023, Zhiteng Chen (ZCAU).

China (Xizang).

The only known larva of this species was found hiding under a rock by the collector looking for stoneflies (Plecoptera), who turned over a rock not far from the river while collecting stonefly nymphs.

Adult: Body black with some yellowish-brown markings. Vertex mostly black, anterior and posterior margins with a few yellowish-brown markings. Frons yellowish brown, with a crossed black marking (Fig. 12B, C). Pronotum mostly black, medially with four yellowish brown markings, laterally with a pair of curved yellowish-brown stripes, sometimes the middle markings connected with the lateral markings (Fig. 12B). Abdominal terga 3–7 black, each medially with a yellowish-brown marking and posterior margin yellowish brown (Fig. 2E). Male sternum 9 wide, posterior part narrowed and short (Fig. 12E); gonocoxites 9 externally swollen and protruded (Fig. 12H–L). Female pregenital plate shaped as a vertical prominence; gonocoxites 8 short and narrow, posteriorly curved (Fig. 12F, G).

Figure 12. 

Layahima pixiu sp. nov., adult: A Habitus, female holotype; B head, frontal view; C head and thorax, dorsal view. D, E Male terminalia: D Lateral view, E ventral view. F, G Female terminalia: F Lateral view, G ventral view. H–L Male genitalia: H Ventral view, I anteroventral view, J caudal view, K lateral view, L dorsal view.

Size. Head width: 2.68–3.12 mm; forewing length: 29.67–33.43 mm; hindwing length: 28.23–32.14 mm. — Head. Vertex mostly black, anterior and posterior margins with a few yellowish-brown markings. Scape and pedicel mostly dark brown; antenna mostly black, non-swollen part of flagellomeres each distally yellowish brown; swollen part of flagellum with a yellowish-brown marking. A dark brown marking present around the antennal scape, anteriorly slightly concaved. Frons yellowish brown, with a crossed black marking. Clypeus and labrum pale yellowish brown. Mandible pale yellowish brown with distal half dark reddish brown. Maxillary palpus mostly dark brown, only segment 3 brown. Labial palpus generally yellowish brown, distal palpomere fusiform and dark brown (Fig. 12B, C). — Thorax. Pronotum mostly black, medially covered with short black setae, lateral margin with pale setae; medially with four yellowish brown markings, laterally with a pair of curved yellowish-brown stripes, sometimes the middle markings connected with the lateral markings. Mesoprescutum black with a pair of yellowish-brown spots; mesonotum black, medially with an inverted triangular yellowish brown marking, laterally with two pair of yellowish brown stripes and spots; mesoscutellum black, with a pair of yellowish brown spots, posterior margin yellowish brown. Metanotum black and yellowish brown; metascutellum mostly black, posterior margin yellowish brown. Pleurae mostly black (Fig. 12C). — Legs. Short, as long as thorax at most, with many black and pale setae. All tibial spurs slightly curved; tarsomere 1 pale yellow, tarsomere 2 pale yellow with distally black, tarsomeres 3–4 black; tarsomere 5 pale yellow with distally black, nearly as long as entire length of tarsomeres 2–4; pretarsal claws curved, dark reddish brown. Foreleg: coxa pale yellowish brown with a black spot; femur dorsally and ventrally black, laterally yellowish brown; tibia black, basally and medially with two pale yellow markings; tibial spurs reaching base of tarsomere 3. Midleg: Coxa pale yellowish brown with a black spot; femur mostly pale yellowish brown, distally dark brown; tibia black, basally and medially with two pale yellow markings; tibial spurs reaching base of tarsomere 3. Hind leg: Coxa pale yellowish brown with a black spot; femur mostly pale yellowish brown, distally dark brown; tibia pale yellow, mediobasally and distally black; tibial spurs reaching tip of tarsomere 1; tarsomere 1 longer than that of fore- and midleg (Fig. 12A). — Wings. Mostly hyaline, distally rounded. Banksian lines absent. Pterostigma pale. Forewing slightly longer than hindwing. Forewing costal space with a dark brown spot proximal to pterostigma; subcostal area with some indistinct brown spots; radial area with some indistinct brown spots; mediocubital area with a few indistinct brown spots; rhegma as two indistinct brown markings; cubital area with a basal indistinct dark brown marking; costal crossveins generally simple on basal 1/4 part, while distal 3/4 part each connect with a short oblique veins; RP originates proximally to MP fork; four to five presectoral crossveins present, with from two to four biareolate cells; RP with eight to nine branches. Hindwing rhegma as an indistinct brown marking reaching to posterior margin; presectoral area with only one crossvein; RP originates anteriorly to MP fork (Figs 2E, 12A). — Abdomen. Terga mostly black, with some yellowish-brown marking, sterna mostly yellowish brown. Tergum 1 generally pale; tergum 2 mostly dark brown; terga 3–7 black, each medially with a yellowish-brown marking and posterior margin yellowish brown (Fig. 2E). — Male genitalia. Sternum 9 wide, posterior part narrowed and short (Fig. 12E). Gonocoxites 9 shaped as a pair of trapezoid plates, externally swollen and protruded; gonocoxites 11 widely arched; gonostyli 11 rounded protruded in lateral view (Fig. 12H–L). Ectoproct rounded in posterior margin (Fig. 12D). — Female genitalia. Pregenital plate shaped as a vertical prominence. Gonocoxites 8 short and narrow, digitiform, posteriorly curved; gonapophyses 8 indistinct and rounded. Gonocoxites 9 with many tapered stout setae, slightly curved. Ectoproct truncate on distal margin (Fig. 12F, G).

Holotype ♀, CHINA: Yunnan, Diqing, Shangri-La County [香格里拉县], Hutiaoxia Town [虎跳峡镇], Jiangbian Village [江边村], 1830 m, 25.V.2023, Quanyu Ji (IZCAS). Paratypes: 1♀, same locality as above, 24.V.2023, Quanyu Ji (IZCAS); 1♂, CHINA: Sichuan, Garze, Muli County [木里县], Maidilong Township [麦地龙乡], Yalongjiang Town [雅砻江镇], Zhongpuzi Village [中埔子村], 18.V.2024, Hao Xun (CAU); 1♂, CHINA: Yunnan, Kunming, Luquan County [禄劝县], Shangshihuiyao [上石灰窑], 1860 m, 3.VII.2022, Zhixin Yang (IZCAS).

The species name “pixiu” is derived from the Chinese mythological creature “Pixiu” (貔貅), which is a legendary animal known for its ability to attract wealth and protect against evil. According to the Chinese legend, the Pixiu only devours wealth (especially gold and silver treasures) and does not excrete, symbolizing wealth accumulation, not letting the loss of wealth. The Pixiu has a striking resemblance to both the mythical antlion, —associated with riches—and the real insect, known for not excreting at the larval stage. The name is treated as a noun in apposition.

China (Sichuan, Yunnan) (Fig. 18).

L. pixiu sp. nov. can be easily distinguished from other Layahima species by its mostly black body with reduced yellowish-brown markings, and the swollen and prominent external male gonocoxites 9. L. pixiu sp. nov. is distributed in some transition regions between the Mt. Hengduanshan Mountain and Ailaoshan Mountain ranges.

Adult: Body pale yellowish brown with some black and dark brown markings, parts with numerous black dots at base of setae. Vertex anteriorly with four median small black spots, posteriorly with a pair of transversal black markings. Frons and clypeus mostly pale yellowish brown with a transversal stripe (Fig. 13B, C). Pronotum anteromedially with a dark brown marking, anteromedially with a pair of dark brown spots, medially with a dark brown spot, and laterally with four pairs of dark brown spots (Figs 5F–H, 13C). Male sternum 9 wide, posterior part narrowed and elongate (Fig. 13D, E); gonocoxites 9 are as long as wide. Female pregenital plate shaped as an anterior sclerotized papilla prominence and a posterior ridge-like prominence.

Figure 13. 

Layahima qilin sp. nov., adult: A Habitus, female holotype; B head, frontal view; C head and thorax, dorsal view. D, E Male terminalia: D Lateral view, E ventral view. F, G Female terminalia: F Lateral view, G ventral view. H–L Male genitalia: H Ventral view; I anteroventral view; J caudal view; K lateral view; L dorsal view.

Size. Head width 3.09–3.31 mm; forewing length 27.45–32.35 mm; hindwing length 25.32–33.78 mm. — Head. Vertex anteriorly with four median small black spots, posteriorly with a pair of transversal black markings. Scape largely pale yellow, internal margin black; pedicel dark brown; flagellum mostly pale brown, each flagellomere brown basally. A dark brown marking present around scapes, anteriorly concaved. Frons and clypeus mostly pale yellowish brown with a transversal stripe. Labrum pale yellow. Maxillary palpus pale yellow. Labial palpus generally pale, distal flagellomere dark brown, fusiform. Mandible pale yellow with distal half dark reddish brown (Figs 5F–H, 13B, C). — Thorax. Pronotum anteromedially with a dark brown marking, anteromedially with a pair of dark brown spots, medially with a dark brown spot, and laterally with four pairs of dark brown spots. Mesoprescutum mostly black, anteriorly with a pair of pale yellowish-brown spots; mesonotum pale yellowish brown, with three pair of black stripes and a pair of black spots, pale yellowish-brown part with some black dots; mesoscutellum anteriorly black, mostly pale yellowish brown, variation present in different individuals. Metanotum pale yellowish brown and black; metascutellum mostly pale yellowish brown. Pleurae pale yellowish-brown and black (Figs 5F–H, 13C). — Legs. Short, as long as thorax at most, with many black and pale setae. All tibial spurs slightly curved; tarsomeres each generally pale yellow with distal part dark brown; tarsomere 5 pale yellow, with distal part dark brown, nearly as long as entire length of tarsomeres 2–4; pretarsal claws curved, dark reddish brown. Foreleg: coxa pale yellow with a black spot; femur dorsally brown to dark brown, ventrally pale yellowish brown; tibia pale yellow with a dark anterior marking, distal part dark brown; tibial spurs reaching base of tarsomere 3. Midleg: Coxa pale yellowish brown, black basally; femur mostly yellowish brown, distally dark brown; tibia pale yellow, with some black dots, distal part dark brown; tibial spurs reaching basal of tarsomere 3. Hind leg: Coxa mostly pale yellowish brown, black basally; femur yellowish brown, with distal part dark brown; tibia pale yellow, distally dark brown; tibial spurs reaching tip of tarsomere 1; tarsomere 1 longer than that of fore- and midleg (Fig. 13A). — Wings. Mostly hyaline, distally rounded. Banksian lines absent. Pterostigma pale. Forewing slightly longer than hindwing. Forewing costal space with an indistinct brown spot proximal to pterostigma; subcostal area hyaline; radial area with some indistinct brown spots; mediocubital area mostly hyaline; rhegma as two indistinct brown markings; cubital area with a basal indistinct dark brown marking; costal crossveins generally simple on basal 1/6 part, while distal 5/6 part each connect with a short oblique veins; RP originates proximally to MP fork; four to five presectoral crossveins present, with from two to three biareolate cells; RP with eight to nine branches. Hindwing rhegma indistinct; presectoral area with 1 crossvein; RP originates anteriorly to MP fork (Fig. 2D, 13A). — Abdomen. Pale yellowish brown with some dark brown markings. Tergum 1 generally pale yellowish brown; tergum 2 mostly dark brown; terga 3–8 each dark brown on posteromedial part (Fig. 2D). — Male genitalia. Sternum 9 wide, posterior part narrowed and elongate (Fig. 13E); gonocoxites 9 shaped as a pair of widely short trapezoid plates, as long as wide; gonocoxites 11 rounded arched; gonostyli 11 slightly prominent (Fig. 13H–L). Ectoproct slightly protruded (Fig. 13D). — Female genitalia. Pregenital plate shaped as an anterior sclerotized papilla prominence and a posterior ridge-like prominence; gonocoxites 8 slightly curved, short digitiform, with an anterior rounded depressed groove; gonapophyses 8 widely triangular. Gonocoxites 9 with many tapered stout setae, slightly curved. Ectoproct rounded on distal margin (Fig. 13F, G).

Holotype ♂, CHINA: Yunnan, Diqing, Weixi County [维西县], Zhonglu Township [中路乡], Zhazidi [扎子底], 1885 m, 6.VII.2022, Zhihao Qi (IZCAS). — Paratypes: 1♂1♀ CHINA: Sichuan, Garze, Batang County [巴塘县], Suwalong Township [苏哇龙乡], 2410 m, 2.VII.2020, Yuezheng Tu & Di Li (IZCAS); 2♀, CHINA: Xizang, Qamdo, Markam County [芒康县], Yanjing Town [盐井乡], 2600 m, 7.VI.1976, Yinheng Han (CAU); 1♀, CHINA: Xizang, Qamdo, 19.IX.1982, Wang (CAU); 1♀, CHINA: Yunnan, Diqing, Deqen County [德钦县], Mt. Baimaxueshan [白马雪山], 23.VI.2020, Weidong Huang (CAU); 1♂1♀, CHINA: Yunnan, Diqing, Deqen County, Mt. Meilixueshan [梅里雪山], 22.VII.1982, Shuyong Wang (CAU); 1♀, CHINA: Yunnan, Diqing, Deqen County, Xiaruo Township [霞落乡], 16.V.1976, Xiangqun Yuan (CAU); 3♂1♀, CHINA: Yunnan, Diqing, Deqen County, Yanmen Township [雁门乡], Cizhong Village [茨中村], 16–18.VI.2004, Hao Huang (CAU); 1♂ CHINA: Yunnan, Diqing, Weixi County, Pantiange Township [攀天阁乡], 9.VII.2010 (IZCAS); CHINA: Yunnan, Diqing, Weixi County, Yongchun Township [永春乡], Baipa [白帕], 2300 m, 1.VII.2023, Yuchen Zheng (CAU); 1♀, CHINA: Yunnan, Lijiang, Yulong County [玉龙县], Tacheng Township [塔城乡], 10.VII.2004, Hao Huang (CAU); CHINA: Yunnan, “Mekong” (Lancangjiang) valley, 35 km S Deqen, 3000 m, 14.VI.2000, Enrico Gallo (MSNG).

This new species is named after the “Qilin” (麒麟), a mythical creature in Chinese mythology known for its kindness and auspiciousness. The name is treated as a noun in apposition.

China (Sichuan, Xizang, Yunnan) (Fig. 18).

L. qilin sp. nov. is endemic to the middle elevations (1800–3000 m) of Shaluli and Mangkang mountain systems between Lanchangjiang and Jinshajiang River.

Adult: See Zheng et al. (2023). — Larva: Head pale yellow, anteromedially with a pair of connected dark brown markings, and other part with many dark brown dots in dorsal view; lateral margin with a pair of oblique dark brown markings. Dolichasters on clypeo-labrum mostly pale. Mandible pale yellow with reddish brown markings. Basally with six to seven interdental mandibular setae; three short interdental mandibular setae between first and second teeth; one seta between second and third teeth (Fig. 15A, B). Thoracic setiferous processes long digitiform, meso- and metathorax each with anterior setiferous processes posteriorly curved, nearly twice as long as the posterior one. Mesothoracic posterior setiferous processes anteriorly curved (Fig. 15C). Abdomen rounded; setiferous processes short digitiform, processes on further posterior segment shorter; dorsally pale blue and pale yellow with many dark markings; ventrally without marking, green and pale yellow (Fig. 14A, B); Segment 9 dorsally with four digging setae, rastrum distally with four pair of tapered digging setae (Fig. 15D).

Figure 14. 

Habitus of Layahima yangi Wan & Wang, 2006, third instar larva: A Dorsal view, B ventral view, C lateral view.

Figure 15. 

Layahima yangi Wan & Wang, 2006, third instar larva: A Head, dorsal view; B head, ventral view; C thorax, dorsal view, on black background; D abdominal segments 8–9, ventral view.

Size. Body length (excluding mandible): 10.20 mm; head length: 2.36 mm; head width: 2.58 mm; mandible length: 3.24 mm. — Head. Nearly rectangular in dorsal view, wider than long, pale yellow with some dark markings. Clypeo-labrum dark brown, with pale dolichasters. Head pale yellow, anteromedially with a pair of connected dark brown markings, and other part with many dark brown dots in dorsal view; lateral margin with a pair of oblique dark brown markings; ventrally without marking. Antenna slender. Ocular tubercles well prominent, ocular peduncle dark brown. Mandible pale yellow with reddish brown markings; basally with six to seven interdental mandibular setae; first tooth shorter than second tooth; second tooth nearly as long as third tooth; three short interdental mandibular setae between first and second teeth; one seta between second and third teeth. Labial palps pale brown (Figs 14, 15A, B). — Thorax. Laterally with pale setae. Thoracic setiferous processes long digitiform. Pronotum pale yellow, medially with a pair of closed black stripes, laterally with irregular black markings. Mesothorax with spiracles present on short sclerotized tubercle, basally dark brown and pale yellow, distally pale, tip dark brown. Meso- and metanotum medially pale blue and laterally pale yellow, with many black markings; meso- and metathorax each with anterior setiferous processes posteriorly curved, mostly pale yellow, nearly twice as long as the posterior one; posterior setiferous processes mostly dark brown. Mesothoracic posterior setiferous processes anteriorly curved (Figs 14A, 15C). — Legs. Pale yellow, with many slender and short tapered setae (Fig. 14B, C). — Abdomen. Rounded, dorsally pale blue and pale yellow with many dark markings; ventrally green and pale yellow, without marking. A pair of large spiracles present on laterodorsal abdominal segment 1. Setiferous processes short digitiform, processes on further posterior segment shorter (Fig. 14A, B). Segment 9 dorsally with four digging setae; ventrally with seven tapered setae of uneven size, the external row of setae largest, the internal row of setae smallest; rastrum distally with four pair of tapered digging setae (Fig. 15D).

Holotype ♀, CHINA: Guangxi, Chongzuo, Pingxiang City [凭祥市], 16.V.1963, Chikun Yang (CAU). Other type specimens were examined in Zheng et al. (2023).

Larva: 1 3rd instar larva, CHINA: Guangxi, Guilin, Lingchuan County [灵川县], Sanjie Town [三街镇], 18.IV.2024, Haofei Fan (ZCAU).

China (Guangxi); Vietnam (Hoa Binh) (Fig. 18).

The only known larva of this species was found anchored on a vertical limestone surface, with its body attached to green mosses (Fig. 16). This behavior resembles that of some owlfly larvae (e.g., Puer Lefèbvre, 1842) (Badano & Pantaleoni, 2014b), and some antlion larvae (Gatzara Navás, 1915, Nepsalus Navás, 1914, and Jaffuelia Navás, 1918) (Zheng et al. 2022, 2024; Stange et al. 1994, 2003).

Figure 16. 

Living third larva of Layahima yangi Wan & Wang, 2006: A Habitat, limestone in Guilin (Guangxi); B Living 3rd larva on the limestone surface (photos by Haofei Fan).

Body generally dark grey. Posterior margin of vertex and postgena orange. Pronotum with a pair of “F” shaped pale yellowish markings. Mesonotum with a few pale yellowish markings (Fig. 17C). Forewings long, narrow, nearly four times as long as maximum width, with dense indistinct markings (Figs 2F, 17A). Abdomen mostly dark grey, terga 1 and 3–5 each medially with a pale yellowish marking (Fig. 2F). Female pregenital plate shaped as a sclerotized cone; gonocoxites 8 wide, thumb-shaped (Fig. 2D, E).

Figure 17. 

Layahima zhitengi sp. nov., adult: A Habitus, female, holotype; B head, frontal view; C head and thorax, dorsal view. D, E Female terminalia: D Lateral view, E ventral view.

Size. Head width: 3.08 mm; forewing length: 34.23 mm; hindwing length: 34.42 mm. — Head. Mostly dark grey. Vertex dark grey, posterior margin orange. Postgena orange. Antenna mostly black. Frons with a pair of indistinct curved pale yellowish markings. Clypeus anteriorly dark brown, posteriorly pale. Labrum yellowish brown, with some yellowish-brown setae. Maxillary and labial palpus yellowish brown. Mandibles pale yellowish brown with distal half dark reddish brown (Fig. 17B, C). — Thorax. Pronotum mostly dark grey, with a pair of “F” shaped pale yellowish markings, with some dark and pale setae. Mesoprescutum and mesoscutellum mostly dark grey; mesonotum with a few pale yellowish markings. Metanotum and metascutellum mostly dark grey. Pleurae dark grey (Fig. 17C). — Legs. Short, as long as thorax at most, with many black and pale setae. All tibial spurs slightly curved; tarsomere 1 basally pale yellow with distally dark brown, tarsomeres 1–4 generally black, tarsomere 5 pale yellow with distally black, nearly as long as entire length of tarsomeres 2–4; pretarsal claws curved, pale yellowish brown. Foreleg: Coxa yellowish brown; femur mostly dark brown; tibia black, basally and medially with two pale yellow markings; tibial spurs reaching tip of tarsomere 2. Midleg: Coxa pale yellowish brown with a black spot; femur mostly dark brown; tibia black, basally and medially with two pale yellow markings; tibial spurs reaching base of tarsomere 3. Hind leg: Coxa pale yellowish brown with a black spot, femur mostly dark brown; tibia black, basally and medially with two pale yellowish markings; tibial spurs reaching base of tarsomere 2; tarsomere 1 longer than that of fore- and midleg (Fig. 17A). — Wings. Long, narrow. Banksian lines absent. Pterostigma pale. Forewing nearly four times as long as maximum width, with dense indistinct markings; costal space with a dark brown spot proximal to pterostigma; costal crossveins generally simple on nearly basal half, while nearly distal half each connected by a short oblique vein; RP originates proximally to MP fork; four to five presectoral crossveins present, almost with biareolate cells; RP with 10 branches. Hindwing nearly 3.5 times as long as maximum width; rhegma indistinct; presectoral area with only one crossvein; RP originates anteriorly to MP fork (Figs 2F, 17A). — Abdomen. Mostly dark grey, terga 1 and 3–5 each medially with a pale yellowish marking (Fig. 2F). — Male genitalia. Unknown. — Female genitalia. Pregenital plate shaped as a vertical prominence. Gonocoxites 8 wide, thumb-shape; gonapophyses 8 indistinct and rounded. Gonocoxites 9 with many tapered stout setae, slightly curved. Ectoproct rounded on distal margin (Fig. 2D, E).

Holotype ♀, CHINA: Yunnan, Nujiang, Fugong County [福贡县], Haizha [害扎], 1200 m, 2.VII.2022, Zhiteng Chen (IZCAS).

The new species is dedicated to the plecopteran researcher Dr. Zhiteng Chen (陈志腾), who donated us the holotype of this new species and a larva of L. lhoba.

China (Yunnan) (Fig. 18).

Figure 18. 

Distribution map of Layahima spp.

This new species is similar to L. valida (Yang, 1997), which occurs in South and Central China. Both species are dark grey and have long, narrow wings. However, L. zhitengi sp. nov., is characterized by orange posterior vertex, mesonotum with few pale yellowish markings, abdomen with some pale yellowish spots, female pregenital plate shaped as a vertical prominence, and female gonocoxites 8 thumb-shaped; while L. valida differs in the entirely dark grey vertex, mesonotum, and abdomen, female pregenital plate shaped as a pair of strongly sclerotized subtriangular plates, and female gonocoxites 8 narrowed and digitiform.