Research Article |
Corresponding author: Zi-Wei Yin ( pselaphinae@gmail.com ) Academic editor: Adam Brunke
© 2021 Wen-Xuan Zhang, Fang-Shuo Hu, Zi-Wei Yin.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Zhang W-X, Hu F-S, Yin Z-W (2021) Six new species of Horniella Raffray from the Oriental region (Coleoptera, Staphylinidae, Pselaphinae). ZooKeys 1042: 1-22. https://doi.org/10.3897/zookeys.1042.66576
|
The Oriental pselaphine genus Horniella Raffray, 1905 currently contains 29 species. In this paper, six new species are described: H. nantouensis Zhang, Hu & Yin, sp. nov. and H. taiwanensis Zhang, Hu & Yin, sp. nov. from Taiwan, China; H. bifurca Zhang & Yin, sp. nov. and H. haucki Zhang & Yin, sp. nov. from Thailand; H. khasiensis Zhang & Yin, sp. nov. from northern India; and H. sabahensis Zhang & Yin, sp. nov. from eastern Malaysia. In addition, H. aculeata Yin & Li, 2015, originally described from Yunnan, China, is newly recorded from Thailand.
Ant-loving beetles, Asia, new record, new taxa, species list, taxonomy
The Oriental pselaphine genus Horniella Raffray, 1905 (Tyrini: Somatipionina) currently includes 29 species distributed in China (12 spp.), Thailand (9 spp.), Malaysia (4 spp.), Nepal and India (1 sp.), Sri Lanka (1 sp.), the Philippines (1 sp.), and Indonesia (1 sp.) (
Based on an examination of additional material deposited in the Muséum d’Histoire Naturelle, Geneva, Switzerland, and the National Museum of Natural Science, Taichung City, Taiwan, China, we describe here six new species from China (2), Thailand (2), India (1), and Malaysia (1). Thus, the total species number of Horniella raises from 29 to 35. Furthermore, new collecting data of Horniella aculeata Yin & Li from Thailand are provided.
The type material of the new species described in this paper is deposited in the Muséum d’Histoire Naturelle, Geneva (
Dissected parts were preserved in Euparal on plastic slides that were placed on the same pin with the specimen. The habitus images of the beetles were taken using a Canon 5D Mark III camera in conjunction with a Canon MP-E 65 mm f/2.8 1–5× macro lens, and a Canon MT-24EX Macro Twin Lite flash was used as the light source. Images of the morphological details were produced using a Canon G9 camera mounted to an Olympus CX31 microscope under reflected or transmitted light. Zerene Stacker v. 1.04 was used for image stacking. All images were modified and grouped into plates using Adobe Photoshop CC 2020.
The abdominal tergites and sternites are numbered following
The collecting data of the material are quoted verbatim. The Chinese translation of each locality is included in parentheses at first appearance in the text. A slash is used to separate different labels. Each type specimen bears the following label: ‘HOLOTYPE [red] (or PARATYPE [yellow]), ♂ (or ♀), Horniella + specific name sp. n., det. Zhang & Yin, 2021,
The following abbreviations are applied: AL = length of the dorsally exposed part of the abdomen (posterior to elytra) along the midline; AW = maximum width of the abdomen; EL = length of the elytra along the suture; EW = maximum width of the elytra; HL = length of the head from the anterior clypeal margin to the anterior margin of the occipital constriction; HW = width of the head across eyes; PL = length of the pronotum along the midline; PW = maximum width of the pronotum. Length of the body is a sum of HL + PL + EL + AL.
Horniella aculeata Yin & Li, 2015: 110.
2 ♂♂, labeled ‘Thailand: Nan prov. Doi Phuka Nat. Park, 28.IV-12.V.2002, Průdek & Obořil lgt.’ (
Horniella aculeata is readily recognizable by the presence of a large spine on the mesal margin of the protibiae (
China: Yunnan; Thailand: Nan, Mae Hong Son. New country record for Thailand.
Male. Head approximately as long as wide, with distinct anterolateral genal projection, anterior margin of projection roundly emarginate; with long, apically forked ocular canthus; scape angularly expanded at anterolateral margin, antennomeres 9–11 moderately enlarged. Pronotum rounded at anterolateral margins. Protrochanter and profemur each with long ventral spine; protibia with small triangular apical spur; mesotrochanter with short but distinct ventral spine. Tergite 1 (IV) with median carina extending posteriorly for approximately 1/4 of tergal length, discal carinae long and thick. Aedeagus with asymmetric median lobe, right half of median lobe greatly protruding apicad, left half strongly curved and forked at apex; endophallus composed of two elongate, twisted sclerites.
Female. Similar to male in external morphology, profemur each with two ventral spines near base, protibia lacking preapical spur, mesotrochanter lacking ventral spine; genital complex as in Fig.
Male. Body reddish-brown, length 3.35 mm. Head (Fig.
Diagnostic characters of Horniella bifurca sp. nov. A left half of head, in dorsal view B head, in lateral view C protrochanter and profemur D protibia E mesotrochanter and mesofemur F mesotibia G–I aedeagus, in dorsal (G), lateral (H), and ventral (I) view. Scale bars: 0.3 mm (A, C–F); 0.2 mm (B, G–I).
Pronotum as long as wide, PL and PW 0.67 mm, widest at apical 1/3; anterolateral margins rounded; disc moderately convex, finely punctate, with distinct median antebasal and lateral antebasal foveae connected by complete transverse sulcus.
Elytra much wider than long, EL 0.91 mm, EW 1.36 mm; each elytron with two large, setose basal foveae; discal striae extending from outer basal foveae to apical 3/4 of elytral length.
Legs elongate; protrochanter (Fig.
Abdomen slightly broader than long, broadest at lateral margins of tergite 1 (IV), AL 1.08 mm, AW 1.22 mm; tergite 1 (IV) largest, as long as tergites 2 and 3 (V and VI) combined, with short median carina extending to near basal 1/4 of tergal length, discal carinae long and thick, with broad basal impression, tergite 2 (V) lacking carina, tergites 2–4 (V–VII) each with small basolateral foveae. Sternite 2 (IV) with broad basal sulcus, lacking mediobasal foveae, basolateral foveae developed as large cuticular pockets, with two pairs of antebasal protuberances, sternites 3–5 (V–VII) each with basolateral foveae, and one median and two lateral nodules, sternite 7 (IX) with well-sclerotized apical half and membranous basal half.
Aedeagus (Fig.
Female. General morphology similar to male, each eye composed of approximately 30 facets; profemur with two long ventral spines near base, protibia lacking preapical spur, mesotrochanter lacking ventral spine. Measurements (as for male): BL 3.08 mm, HL 0.68 mm, HW 0.61 mm, PL 0.65 mm, PW 0.63 mm, EL 0.79 mm, EW 1.17 mm, AL 0.96 mm, AW 1.2 mm. Genital complex (Fig.
This species is placed as a member of the H. centralis group. It can be readily separated from the other members of the group by the long, apically-forked ocular canthi, as well as by the unique shape of the aedeagus.
Thailand: Chiang Mai.
The new specific epithet bifurca (-us, -um) is a Latin adjective means ‘two-pronged’, referring to the apically-forked ocular canthus of the new species.
Holotype: Thailand: ♂, ‘THAI, N, Mae Hong Son prov., SE of Soppong, 1500 m, 19°27'N, 98°20'E, 23–27.v.1999, D. Hauck leg.’ (
Male. Head longer than wide, with distinct anterolateral genal projection, anterior margin of projection roundly emarginate; with long ocular canthus; scape angularly expanded at basolateral margin, antennomeres 9–11 moderately enlarged. Pronotum rounded at anterolateral margins. Protrochanter and profemur each with long ventral spine; protibia strongly curved near apex, with long apical projection; mesotrochanter with large sharp ventral spine, mesofemur distinctly arched. Tergite 1 (IV) with median carina extending posteriorly for approximately 3/4 of tergal length, discal carinae short and thin. Aedeagus with asymmetric median lobe, left half of median lobe greatly protruding in dorso-ventral view; endophallus composed of three long sclerites.
Male. Body reddish-brown, length 3.49 mm. Head (Fig.
Diagnostic characters of Horniella haucki sp. nov. A left half of head, in dorsal view B head, in lateral view C protrochanter and profemur D protibia E mesotrochanter and mesofemur F mesotibia G–I aedeagus, in dorsal (G), lateral (H), and ventral (I) view. Scale bars: 0.3 mm (A, C–F); 0.2 mm (B, G–I).
Pronotum longer than wide, PL 0.74 mm, PW 0.69 mm, widest at apical 1/3; anterolateral margins rounded; disc moderately convex, finely punctate, with distinct median antebasal and lateral antebasal foveae connected by complete transverse antebasal sulcus.
Elytra much wider than long, EL 0.95 mm, EW 1.35 mm; each elytron with two large, setose basal foveae; discal striae extending from outer basal foveae to apical 2/3 of elytral length.
Legs elongate; protrochanter (Fig.
Abdomen slightly broader than long, broadest at lateral margins of tergite 1 (IV), AL 1.05 mm, AW 1.32 mm; tergite 1 (IV) largest, as long as tergites 2 and 3 (V and VI) combined, with median carina extending to near basal 3/5 of tergal length, discal carinae short and thin, tergite 2 (V) lacking carina, tergites 2–4 (V–VII) each with small basolateral foveae. Sternite 2 (IV) with broad basal sulcus, lacking mediobasal foveae, basolateral foveae developed as large cuticular pockets, with two pairs of antebasal nodules, sternites 3–5 (V–VII) with basolateral foveae, one median and two lateral nodules, sternite 7 (IX) with well-sclerotized apical half, and membranous basal half.
Aedeagus (Fig.
Female. Unknown.
This new species can be readily separated from all members of the H. centralis group primarily by the characteristic shape of the aedeagus, especially the form of the apical portion of the median lobe, and the configuration of the endophallus.
Thailand: Mae Hong Son.
The new species is named after David Hauck (České Budějovice, Czech Republic), collector of the holotype.
Holotype: India: ♂, ‘INDIA, Meghalaya State (7+9), E Khasi Hills, 11km SW Cherrapunjee, Laitkynsew, 25.iv.2008, 25°12'48"N, 91°39'48"E, 735 m, Fikáček, Podskalská, Šípek lgt. / secondary tropical rainforest with young trees + bamboo, below village, thin layer of leaf litter (sifting).’ (
Male. Head wider than long, with distinct anterolateral genal projection, anterior margin of projection roundly emarginate; with markedly long ocular canthus; scape angularly expanded at middle of lateral margin, antennomeres 9–11 enlarged. Pronotum rounded at anterolateral margins. Protrochanter and profemur each with long ventral spine; mesotrochanter with short, small ventral tubercle. Tergite 1 (IV) with median carina extending posteriorly for approximately 1/4 of tergal length, lacking discal carinae, tergite VIII with large medioapical process. Aedeagus with slightly asymmetric median lobe, apex broadly truncate in dorso-ventral view; endophallus composed of three sclerites.
Male. Body reddish-brown, length 2.84 mm. Head (Fig.
Pronotum as long as wide, PL and PW 0.64 mm, widest anterior to middle; lateral margins rounded; disc moderately convex, finely punctate, with distinct median antebasal and lateral antebasal foveae connected by complete transverse antebasal sulcus.
Elytra much wider than long, EL 0.75 mm, EW 1.2 mm; each elytron with two large, setose basal foveae; discal striae extending from outer basal foveae to near posterior margin of elytra.
Legs elongate; protrochanter (Fig.
Abdomen broader than long, broadest at lateral margins of tergite 1 (IV), AL 0.89 mm, AW 1.21 mm; tergite 1 (IV) largest, as long as tergites 2 and 3 (V and VI) combined, with short median carina extending to near basal 1/4 of tergal length, lacking discal carinae, tergite 2 (V) lacking carina, tergites 2–4 (V–VII) each with small basolateral foveae, tergite 5 (VIII) with large medioapical process. Sternite 2 (IV) with broad basal sulcus, lacking mediobasal foveae, basolateral foveae developed as large cuticular pockets, with two pairs of antebasal nodules, sternites 3–5 (V–VII) with basolateral foveae, one median and two lateral nodules, sternite 7 (IX) nearly oval, with well-sclerotized apical half and less sclerotized basal half.
Aedeagus (Fig.
Female. Unknown.
This species is placed as a member of the H. burckhardti group, and is most similar to H. hongkongensis Yin & Li in having similar spination of the legs and a general aedeagal form. They can be clearly separated by the more distinctly expanded basolateral margin of the scape, tergite VIII with a large medioapical process, and the different structure of the aedeagal endophallus.
India: Meghalaya.
The new species is named after its type locality, the East Khasi Hills.
Holotype: China: ♂, ‘TAIWAN: Nantou County, Huisun Forest Reserve [惠荪林场], track to Xiaochushan Mt., 24.0745N, 121.0366E; 1150 m, 4.v.2019; Damaška, Fikáček, Hu & Liu lgt., 2019-TW14 / primary forest on the slope with sparse understory; sifting of small accumulations of leaves / Huisun Leaf Litter Beetles Project, Additional specimen: HS1-034 / HORNIELLA sp., P. Hlaváč det., 2019’ (
Male. Head longer than wide, with weakly indicated anterolateral genal projection, anterior margin of projection oblique; with short ocular canthus; lateral margin of scape straight, antennomeres 9–11 slightly enlarged. Pronotum rounded at anterolateral margins. Ventral margin of profemur with one short and acute, and one tiny spine at base; protibia with one small preapical denticle. Tergite 1 (IV) with median carina extending posteriorly for approximately 1/3 of tergal length, lacking discal carinae. Aedeagus with asymmetric median lobe, apical part of median lobe narrowed and protruding apicad, apex nearly rounded in dorsal view. Female. Similar to male in external morphology, profemur with two ventral spines near base; genital complex as in Fig.
Male. Body reddish-brown, length 3.68 mm. Head (Fig.
Diagnostic characters of Horniella nantouensis sp. nov. A left half of head, in dorsal view B head, in lateral view C protrochanter and profemur D protibia E mesotrochanter and mesofemur F mesotibia G–I aedeagus, in dorsal (G), lateral (H), and ventral (I) view. Scale bars: 0.2 mm (A, B, G–I); 0.3 mm in (C–F).
Pronotum longer than wide, PL 0.77 mm, PW 0.71 mm; widest at apical 1/3; anterolateral margins rounded; disc moderately convex, finely punctate, with distinct median antebasal and lateral antebasal foveae connected by complete transverse antebasal sulcus.
Elytra much wider than long, EL 0.85 mm, EW 1.33 mm; each elytron with two large, setose basal foveae; discal striae extending from outer basal foveae to middle of elytral length.
Legs elongate; protrochanter (Fig.
Abdomen slightly longer than broad, broadest at lateral margins of tergite 1 (IV), AL 1.32 mm, AW 1.28 mm; tergite 1 (IV) slightly longer than tergites 2 (V), with median carina extending to near basal 1/3 of tergal length, lacking discal carinae, tergite 2 (V) lacking carina, tergites 2–4 (V–VII) each with small basolateral foveae. Sternite 2 (IV) with broad basal sulcus, lacking mediobasal foveae, basolateral foveae developed as large cuticular pockets, with two pairs of antebasal nodules, sternites 3–5 (V–VII) with basolateral foveae, one median and two lateral nodules.
Aedeagus (Fig.
Female. General morphology similar to male, each eye composed of approximately 40 facets; profemur each with two distinct ventral spines near base, protibia lacking spur. Measurements (as for male): BL 3.68–3.72 mm, HL 0.74 mm, HW 0.63–0.65 mm, PL 0.72–0.73 mm, PW 0.74–0.75 mm, EL 0.87–0.9 mm, EW 1.33 mm, AL 1.35 mm, AW 1.32–1.33 mm. Genital complex (Fig.
This species is placed as a member of the H. hirtella group. The new species is similar to H. simplaria Yin & Li by the male having similar anterolateral genal projections, and presence of two ventral spines of profemur. They can be otherwise clearly separated by the larger body size (3.68 mm vs 3.23 mm), lack of a mesal hook-like spine of the protibia (present in H. simplaria), and the different shape and structure of the aedeagus of the new species.
China: Taiwan.
The new specific is named after its type locality, Nantou County.
Holotype: East Malaysia: ♂, ‘Borneo: Sabah, Batu Punggul Resort, primary forest, 24.vi.–1.vii.96, Kodada lgt. / vegetation debris and forest floor litter accumulated around large trees near river.’ (
Male. Head longer than wide, anterolateral genal projections weakly developed, anterior margin of projection oblique; scape lacking expansion at lateral margin, antennomeres 9–11 moderately enlarged, forming distinct club. Pronotum rounded at lateral margins. Profemur with two tiny ventral spines near base; metatibia with preapical triangular denticle. Tergite 1 (IV) with median carina extending posteriorly for approximately 3/4 of tergal length, lacking discal carinae, tergite 2 (V) with short median carina. Aedeagus with slightly asymmetric median lobe, apical portion of median lobe narrowed, apex truncate in dorso-ventral view; endophallus lacking sclerite, composed of elongate membranous structure with many small denticles.
Male. Body reddish-brown, length 3.41 mm. Head (Fig.
Diagnostic characters of Horniella sabahensis sp. nov. A left half of head, in dorsal view B head, in lateral view C protrochanter and profemur D protibia E mesotrochanter and mesofemur F mesotibia G metatibia H–J aedeagus, in dorsal (H), lateral (I), and ventral (J) view. Scale bars: 0.2 mm (A, B, H–J); 0.3 mm (C–G).
Pronotum distinctly longer than wide, PL 0.71 mm, PW 0.6 mm; widest at middle; lateral margins rounded; disc moderately convex, finely punctate, with distinct median antebasal and lateral antebasal foveae connected by complete transverse antebasal sulcus.
Elytra much wider than long, EL 0.77 mm, EW 1.2 mm; each elytron with two large, setose basal foveae; discal striae extending from outer basal foveae to apical 2/3 of elytral length.
Legs elongate; protrochanter (Fig.
Abdomen slightly longer than broad, broadest at lateral margins of tergite 1 (IV), AL 1.25 mm, AW 1.16 mm; tergite 1 (IV) largest, slightly shorter than tergites 2 and 3 (V and VI) combined, with median carina extending to near basal 3/4 of tergal length, lacking discal carinae, tergite 2 (V) with median carina extending to near basal 1/4 of tergal length, tergites 2–4 (V–VII) each with small basolateral foveae. Sternite 2 (IV) with broad basal sulcus, lacking mediobasal foveae, basolateral foveae developed as large cuticular pockets, with two pairs of antebasal nodules, sternites 3–5 (V–VII) with basolateral foveae, one median and two lateral nodules, sternite 7 (IX) nearly oval, with well-sclerotized apical half and less sclerotized basal half.
Aedeagus (Fig.
Female. Unknown.
Horniella sabahensis sp. nov. is placed as a member of the H. hirtella group. Males of this species share with H. prolixo Yin & Li from Thailand the weakly developed anterolateral genal projections, lack of an expansion at the lateral margin of the scape, and a moderately expanded preapical portion of the metatibia. They can be best separated by the larger body size (3.41 mm vs 2.95–3.02 mm), tergite V with a short median carina (lacking in H. prolixo), as well as the much narrower apex of the aedeagus of the new species.
East Malaysia: Sabah.
The new species is named after its type locality, Sabah, East Malaysia.
Holotype: China: ♂, ‘TAIWAN: Taoyuan City, Northern Cross-island Highway 35.7 k (北横公路35.7 k), Fusing Township, 15-IV-2018, leg. K. X. Zhan’ (
Male. Head longer than wide, with distinct anterolateral genal projections, anterior margin of projection narrowly emarginate, with long ocular canthus; scape roundly expanded at basolateral margin, antennomeres 9–11 slightly enlarged. Pronotum rounded at anterolateral margins. Protrochanter, profemur and mesotrochanter each with ventral spine; protibia and mesotibia with large apical projection. Tergite 1 (IV) with median carina extending posteriorly for approximately 1/4 of tergal length, lacking discal carinae. Aedeagus with asymmetric median lobe, right half of median lobe greatly protruding apicad, apical margin nearly rounded in dorsal view.
Male. Body reddish-brown, length 4.05–4.15 mm. Head (Fig.
Diagnostic characters of Horniella taiwanensis sp. nov. A left half of head, in dorsal view B head, in lateral view C protrochanter and profemur D protibia E mesotrochanter and mesofemur F mesotibia G–I aedeagus, in dorsal (G), lateral (H), and ventral (I) view. Scale bars: 0.2 mm (A, B, G–I); 0.3 mm (C–F).
Pronotum slightly longer than wide, PL 0.78–0.82 mm, PW 0.76–0.77 mm; widest at apical 1/3; anterolateral margins rounded; disc moderately convex, finely punctate, with distinct median antebasal and lateral antebasal foveae connected by complete transverse antebasal sulcus.
Elytra much wider than long, EL 0.94–1.01 mm; EW 1.51 mm; each elytron with two large, setose basal foveae; discal striae extending from outer basal foveae to apical 2/3 of elytral length.
Legs elongate; protrochanter (Fig.
Abdomen approximately as long as broad, broadest at lateral margins of tergite 1 (IV), AL 1.45–1.49 mm, AW 1.47–1.49 mm; tergite 1 (IV) slightly longer than tergites 2 (V), with short median carina extending to near basal 1/4 of tergal length, lacking discal carinae, tergite 2 (V) lacking carina, tergites 2–4 (V–VII) each with small basolateral foveae. Sternite 2 (IV) with broad basal sulcus, lacking mediobasal foveae, basolateral foveae developed as large cuticular pockets, with two pairs of antebasal nodules, sternites 3–5 (V–VII) with basolateral foveae, one median and two lateral nodules, sternite 7 (IX) with well-sclerotized apical half and less sclerotized basal half.
Aedeagus (Fig.
Female. Unknown.
This species is placed as a member of the H. centralis group and is most similar to H. sichuanica Yin & Li in the shapes of the anterolateral genal projections and spination of the legs. They can be clearly separated by the larger body size (4.05–4.15 mm vs 3.58–3.77 mm), the more distinct apical projections of protibia and mesotibia, and the dilated apex of the aedeagal median lobe of the new species.
China: Taiwan.
The new specific is named after Taiwan.
H. aculeata Yin & Li, 2015: 110. China: Yunnan; Thailand: Nan, Mae Hong Son.
H. asymmetrica Yin & Li, 2014: 42. Thailand: Prachin Buri, Chanthaburi.
H. awana Yin & Li, 2014: 65. West Malaysia: Pahang.
H. bifurca Zhang, & Yin, sp. nov. Thailand: Chiang Mai.
H. burckhardti Yin & Li, 2014: 45. Thailand: Chiang Mai.
H. centralis Yin & Li, 2014: 11. China: Shaanxi.
H. cibodas Yin & Li, 2014: 74. Indonesia: West Java.
H. confragosa Yin & Li, 2014: 14. China: Guangxi, Guizhou.
H. dao Yin & Li, 2014: 17. China: Sichuan.
H. falcis Yin & Li, 2014: 18. China: Guizhou.
H. gigas Yin & Li, 2014: 66. East Malaysia: Sabah.
H. haucki Zhang, & Yin, sp. nov. Thailand: Mae Hong Son.
H. himalayica Yin & Li, 2014: 35. Nepal: Bāgmatī añcal; India: Uttarakhand.
H. hirtella (Raffray, 1901: 30). Sri Lanka: Northern, North Central, Central, Uva.
H. hongkongensis Yin & Li, 2014: 21. China: Hong Kong.
H. intricata Yin & Li, 2014: 47. Thailand: Mae Hong Son, Chiang Mai.
H. jinggangshana Yin & Li, 2015: 113. China: Jiangxi.
H. kaengkrachan Yin & Li, 2014: 50. Thailand: Phetchaburi.
H. khaosabap Yin & Li, 2014: 51. Thailand: Chanthaburi.
H. khasiensis Zhang, & Yin, sp. nov. India: Meghalaya.
H. loebli Yin & Li, 2014: 54. Thailand: Chiang Mai.
H. nakhi Yin & Li, 2014: 25. China: Yunnan.
H. nantouensis Zhang, Hu & Yin, sp. nov. China: Taiwan.
H. philippina Yin & Li, 2014: 63. Philippines: Laguna.
H. phuphaman Yin & Li, 2014: 56. Thailand: Khon Kaen.
H. pilosa Yin & Li, 2014: 69. East Malaysia: Sabah.
H. prolixo Yin & Li, 2014: 60. Thailand: Chiang Mai.
H. sabahensis Zhang, & Yin, sp. nov. East Malaysia: Sabah.
H. schuelkei Yin & Li, 2014: 25. China: Yunnan.
H. schwendingeri Yin & Li, 2014: 60. Thailand: Chiang Mai.
H. sichuanica Yin & Li, 2014: 28. China: Sichuan.
H. simplaria Yin & Li, 2014: 28. China: Guangxi.
H. smetanai Yin & Li, 2014: 72. East Malaysia: Sabah.
H. taiwanensis Zhang, Hu & Yin, sp. nov. China: Taiwan.
H. tianmuensis Yin & Li, 2014: 32. China: Zhejiang.
We are grateful to Giulio Cuccodoro (