Review of the genus Superciliaris Meng, Qin & Wang, 2020 (Hemiptera, Fulgoroidea, Issidae): from island endemic to wide Oriental distribution

. Three new species of the genus Superciliaris Meng, Qin & Wang, 2020 are described from southern Vietnam, Malaysia (northern Borneo), and Indonesia (northern Sulawesi). The genus is recorded for the ﬁ rst time from mainland Asia and Sunda Archipelago. A key to species of Superciliaris is given. The relationships of the species are discussed.


Introduction
The genus Superciliaris Meng, Qin & Wang, 2020(in Zhang et al. 2020 belongs to the planthopper family Issidae Spinola, 1839, which is widely distributed and species-rich in the Oriental region (Gnezdilov 2013;Bourgoin 2022). Originally, Superciliaris was included in the monotypic issid subfamily Superciliarinae Meng, Qin & Wang, 2020(in Zhang et al. 2020, subsequently placed in synonymy under the tribe Hemisphaeriini Melichar, 1906 of the subfamily Issinae Spinola, 1839 sensu Gnezdilov et al. (2020) (Gnezdilov 2022). This genus was erected for two species, Superciliaris reticulatus Meng, Qin & Wang, 2020 (type species) and S. diaoluoshanis Meng, Qin & Wang, 2020, both described from Hainan Island (China) (Zhang et al. 2020). During my study of the material on Issidae from Vietnam and the Sunda Archipelago deposited in the Zoological Institute of the Russian Academy of Sciences in Saint Petersburg (Russia) and in the Natural History Museum in London (UK), three new species of Superciliaris were discovered and they are described below. These new fi ndings signifi cantly expand the distribution of the genus in the Oriental Region from Hainan Island to Vietnam, Borneo and Sulawesi.

Material and methods
The terminology of the head and body follows Anufriev & Emeljanov (1988) and that of the male genitalia follows Gnezdilov et al. (2014). The taxonomy of the family Issidae follows Gnezdilov et al. (2020). Photographs were taken using a Canon EOS 5D Mark IV camera with a lens Canon-MP-E-65mm f/2,8 1-5 × Macro and a fl ash Canon Macro Twin-Lite MT-26EX-RT. Images were produced using Helicon Focus ver. 7.6.4 and Adobe Photoshop ver. СС 2019 software. The genital segments were macerated in 10% KOH and fi gured in glycerine jelly (Brunel Micro Ltd, UK) using a Leica MZ9.5 stereo microscope with a camera lucida.
The type specimens of the species described are deposited in the Zoological Institute of the Russian Academy of Sciences, Saint Petersburg, Russia (ZIN) and in the Natural History Museum, London, UK (BMNH).
Label information for the specimens from BMNH is quoted, with '/' indicating a new line and '//' indicating the next label.
Style with nearly straight hind margin (Fig. 7E). Male anal tube distinctly enlarged from its base to its apex (in dorsal view) (

Diagnosis
Generally brown yellowish. Metope and coryphe joined at acute angle. Metope with distinct median and sublateral carinae. Phallobase convex ventrally, without horn-shaped processes apically. Aedeagus with short nearly straight ventral hooks (⅓ of aedeagus length) arise nearly medially on aedeagus.

Etymology
The species is named after the collector, Dr Alexandr E. Anichkin (Yoshkar-Ola, Russia), soil biologist, who spent many years studying invertebrates of Vietnam. STRUCTURE. Metope with two large depressions besides of midline above postclypeus and with strong median carina running from upper margin to middle and with weaker sublateral carinae joined to metopial upper margin, but not fused to median carina, and running slightly beyond eyes (Figs 1C, 4B). Postclypeus not large, weakly swollen, smooth, without carinae. Metope and coryphe joined at acute angle in shape of short process (in lateral view) (Fig. 1B). Coryphe wide, transverse, shagreened, with three large bulges (triangular medial one and two rounded lateral ones); anterior margin strongly convex; posterior margin widely concave; lateral margins and posterior margin keel-shaped (Fig. 1A). Pronotum short, 0.5 × as long as coryphe at midline, without carinae but with pair of small bulges besides of midline; anterior margin widely convex; posterior margin straight. Mesonotum 3 × as long as pronotum, with strong median carina and with pair of large bulges laterally. Each forewing with large bulge in claval basement (Fig. 1B). Fore and middle femora not fl attened. Hind legs missed. COLORATION (Fig. 1). General coloration brown yellowish to dark brown. Eyes dark brown. Forewings with yellowish areas and whitish spots in cells except in basal third. Abdominal sternites, pygofer and styles dark brown. Anal tube light brown. MALE TERMINALIA (Fig. 5). Pygofer vertically elongate, with convex hind margins (Fig. 5A). Anal tube wide and short, enlarged and truncate apically, with nearly straight posterior margin (in dorsal view) (Fig. 5G). Anal column long surpassing posterior margin of anal tube. Phallobase tubular-shaped, elongate, curved (in lateral view), apical part infl ated in water after boiling (Fig. 5C). Ventral phallobase lobe long and wide, slightly narrowing apically (Fig. 5D). Ventral aedegal hooks nearly straight, directed basally, slightly enlarged subapically, with pointed apices (Fig. 5B, D). Connective relatively large, with long handle. Style vertically elongate, with straight hind margin and weak comb behind neck (in lateral view) (Fig. 5E). Capitulum of style on distinct neck, with wide lateral tooth. Capitulum narrowing apically (in dorsal view) (Fig. 5F).

Female
Unknown.

Remark
The species was collected in a soil trap in mixed forest with dominating of Pinus dalatensis Ferré (Pinaceae) on a soil surface with a well-developed moss cover (Dr Alexandr E. Anichkin pers. com.). Diagnosis Generally dark brown. Metope and coryphe joined at nearly right angle (in lateral view). Metope without median carina but with weak sublateral carinae. Phallobase with median groove ventrally; each dorsolateral phallobase lobe with short horn-shaped process apically. Aedeagus with pair of long ventral hooks ( 4 /5 of aedeagus length) arising subapically. Style with deeply concave hind margin. Male anal tube equally wide (in dorsal view).

Etymology
The species is named after the type locality. STRUCTURE. Metope slightly depressed below upper margin medially, without median carina but with weak sublateral carinae running from upper margin to slightly beyond eyes and not reaching metopoclypeal suture (Figs 2C, 4D). Postclypeus fl attened dorso-ventrally, with distinct median carina. Metope and coryphe joined at nearly right angle (in lateral view). Coryphe twice as wide as long at midline, with pair of weak lateral bulges and with three folds medially; anterior margin convex, with weak median notch; posterior margin weakly concave ( Fig. 2A). Pronotum 0.5 × as long as coryphe, depressed medially. Mesonotum more than 3 × as long as pronotum, with fi ne median carina and pair of lateral bulges. Each forewing clavus with bulge basally. Femora and tibiae not fl attened. Hind tibia with two lateral spines above middle and with fi ve apical spines. First metatarsomere slightly longer than second one, but distinctly wider, with long setae ventrally, with two latero-apical and fi ve intermediate spines. Second metatarsomere with two latero-apical spines only. Arolium of pretarsus nearly reaching claw apices (in dorsal view). COLORATION (Fig. 2). Eyes light, except dark brown to black in middle. Metope, coryphe, pro-and mesonotum dark brown, with dense light yellow dots. Clypeus, genae, scapus, rostrum, fore and middle legs dark brown. Pedicel dark brown, with light yellow sensory organs. Paranotal lobes of pronotum light brown yellowish, with dark brown margins. Forewings dark brown to black basally, each with light yellow spot at claval apex. Hind legs brown. Apices of leg spines black. Abdominal segments brown to dark brown, except light yellow hind margins of sternites IV-VII. Anal tube, pygofer and styles dark brown.

Female
Unknown.

Remark
Currently Dumoga-Bone National Park is called Bogani Nani Wartabone National Park located in northern Sulawesi.

Diagnosis
Generally dark brown. Metope and coryphe joined at nearly right angle (in lateral view). Metope with very weak and short median carina and weak sublateral carinae. Phallobase with median groove ventrally; each dorso-lateral phallobase lobe with short horn-shaped process apically. Aedeagus with pair of long ventral hooks ( 4 /5 of aedeagus length) arise subapically. Style with nearly straight hind margin. Male anal tube distinctly enlarged from base to apex (in dorsal view).

Etymology
The species is named after the type locality. STRUCTURE. Metope slightly depressed below upper margin, with very weak and short median carina and weak sublateral carinae running from upper margin to slightly beyond eyes, but not reaching metopoclypeal suture (Figs 3C, 4F). Metope and coryphe joined at nearly right angle (in lateral view) (Fig. 3B). Coryphe transverse, nearly 2.5 × as wide as long at midline; anterior margin nearly straight; posterior margin concave (Fig. 3A). Coryphe longitudinally striated medially, with pair of weak bulges laterally. Postclypeus with smooth median carina. Pronotum 0.5 × as long as coryphe at midline, without bulges or carinae. Mesonotum 4 × as long as pronotum, with fi ne median carina and pair of lateral bulges. Forewings without bulges. Hind tibia with two lateral spines above middle and with fi ve apical spines. First metatarsomere slightly longer than second one, but distinctly wider, with long setae ventrally and with two latero-apical and fi ve intermediate spines. Second metatarsomere with two latero-apical spines only. COLORATION (Fig. 3). General coloration dark brown, with black punctuation. Eyes light. 3 rd segment of rostrum light brown outside and dark brown behind. Forewings with light yellow spots in cells of apical part of clavus and ventro-apical half of corium. Forewing appendix (hardly visible on left wing on Fig. 3A) light brown yellowish from apex of clavus to dorso-lateral angle. Hind femora and tibiae brown. Apices of leg spines black. MALE TERMINALIA (Fig. 7). Pygofer vertically elongate, with convex hind margins. Anal tube nearly twice as long as wide medially, distinctly enlarged from base to apex (in dorsal view) (Fig. 7C). Anal column long and narrow, not surpassing posterior margin of anal tube (Fig. 7C-D). Phallobase vertically elongate, narrow, curved (in lateral view), with median groove ventrally; each dorso-lateral phallobase lobe truncate apically, with short horn-shaped process (Fig. 7A-B). Ventral phallobase lobe short and wide, widely and deeply concave apically (Fig. 7C). Aedeagus with pair of long ventral hooks ( 4 /5 of aedeagus length) arising subapically and with several notches apically. Style with nearly straight hind margin (in lateral view) (Fig. 7E). Capitulum on distinct neck, with wide lateral tooth, not narrowing apically (in dorsal view) (Fig. 7F).

Female
Unknown.

Discussion
According to the data on the type locality and the method of collecting (soil trap) of Superciliaris anichkini sp. nov., this group of planthoppers probably lives in leaf litter or moss cover. Some of them, S. reticulatus, S. diaoluoshanis, and S. anichkini sp. nov. are even superfi cially similar to members of the bug family Peloridiidae (Heteroptera) in having spread forewings (in dorsal view) which may be explained by living in similar habitats with moss. However, no ecological data on species of Superciliaris is yet available.
As many other Issidae with coleopterous and subbrachypterous forewings and rudimentary or reduced hind wings, species of Superciliaris could disperse between the islands and the mainland of southeastern Asia apparently only via the land bridges formed during volcanic activity or decline and rise in sea level in different periods of the Cenozoic (Hantoro et al. 1995;Hall 1996) or via current human activity with any plant cargo which is postulated for several issid species that became adventive in the New and Old World (Gnezdilov & O'Brien 2006;Gnezdilov 2013;Chan et al. 2013;Gnezdilov & Bartlett 2022).
Within Superciliaris, there are two groups of species (see the key above). One group uniting S. reticulatus, S. diaoluoshanis, and S. anichkini sp. nov. from Hainan Island and southern Vietnam and another group uniting S. celebensis sp. nov. and S. tawaiensis sp. nov. from Borneo and Sulawesi. In the Indochinese group of species, S. anichkini sp. nov. from Vietnam is very distinct in having an elongate phallobase and almost straight ventral aedeagal hooks while S. reticulatus and S. diaoluoshanis from Hainan are closely related and differ from each other mainly by the shape of the male anal tube. Superciliaris celebensis sp. nov. and S. tawaiensis sp. nov. from Sunda are closely related and differ mainly in the shape of the style and the male anal tube while both clearly differ from the Indochinese species in head structure and length and attachment of the ventral aedeagal hooks. Thus, apparently the age difference between the species from mainland Indochina and the species from Hainan, Borneo, and Sulawesi is much greater than it is between the two Hainan species or the two species of Sunda. In particular, the species from the Sunda Archipelago may only have separated from each other after the last glacial period during the rise in sea level between the islands.