Umbrageocoris kondorosyi Kóbor, 2019, by monotypy.

General habitus ovoid, moderately elongate. Integument shiny, with sparse, silvery pubescence at least on abdominal venter. Head pentagonal; compound eyes large, reniform, slightly stylate; posterior edge of compound eyes touching anterior edge of pronotum. Ocular sulcus complete but shallow, slightly visible. Integument of vertex impunctate, with a thin longitudinal furrow of various length, at least present on clypeus. Antenniferous tubercles minute, not visible in dorsal view. Antennomere I shortest, graniform; antennomere II longest, cylindrical; antennomeres III and IV subequal in length; antennomere III cylindrical, antennomere IV fusiform. Clypeus with margins subparallel and apex rounded, surpassing the mandibular plates. Bucculae pointed, surpassing mandibular plates; ventral margins converging, forming a Y- or cup-shaped labial trough which continues in a suture of variable length towards base of head (Figs 1, 2). Labiomeres I–II conspicuously stouter than labiomeres III–IV; labiomere I not reaching anterior margin of propleurite, labiomere II shorter than labiomere III, apex of labiomere IV reaching or slightly surpassing metacoxae. Thorax. Pronotum trapeziform with anterior edges and margin broadly convex, lateral margin slightly constricted. Integument shiny, deeply punctate; punctation variably dense. Pronotal callosities and humeral angles at most moderately bulging. Scutellum elongate triangular, basal width less than length; variably punctate except at trifurcate carina. Trifurcate carina of scutellum complete or apically reduced. Apex of scutellum sharply pointed. Submacropterous, brachypterous and coleopterous wing morphs known. Submacropterous morph: hemelytron with margins of clavus converging apically, claval commissure reduced, indistinct; corium with lines of punctures along claval margin, Cu, and costal margin; punctation variably present between M-R and costal margin in the apical half of corium. M-R of corium branching in apical third. Exocorium mostly narrow, sometimes slightly flared in apical half. Membrane at most slightly surpassing apex of abdomen. Metathoracic wing with hamus partly reduced, level of reduction variable (Figs 3, 4); intervannals present, basally fused. Thoracic pleurites and sternites with dense punctation except prosternal collar, supracoxal lobes and peritreme of metathoracic scent efferent apparatus. Prosternite with a narrow, but conspicuously bulging collar. Peritreme bulbous with dorsal flange protruding, sometimes indented; vestibular scar present, reaching metasternal venter; evaporatorium reduced to extreme surroundings of peritreme (Figs 5–10). Femora of prothoracic legs more incrassate than those of meso- and metathoracic legs. Legs covered with sparse, decumbent pubescence; fore femora with simple trichobothria in a single line, fore tibia with strong, dense setosity ventrally. Tarsomeres I and III of pro- and mesothoracic legs subequal in length, tarsomere II shortest. Tarsomere III of metathoracic leg conspicuously elongate, longer than combined lengths of tarsomeres I and II. Abdomen. Abdominal tergites III–V rugose medially; sutures of tergites 4/5 and 5/6 strongly curved medially; suture 4/5 elongate nearly reaching suture 5/6 (Fig. 15). Abdominal sternites with rugose spots subdorsally. Abdominal trichobothria on sterintes III–IV with simple bothrium bearing a single sensilla, situated submedially, very close to each other in triangular arrangement (Fig. 11); trichobothria on sternites V–VII are subdorsal, either simple bothrium or trichome with microtrichia bearing a single sensilla, arrangement as in Fig. 3B. Genitalia. Posterior opening of male pygophore with pointed, short lateral processes, situated dorsolaterally; paramere with trunk moderately stout and blade slender, evenly curved (Fig. 13); gonoporal process of aedeagus with 10 coils. Female ovipositor short, bisecting only abdominal sternite VII (Fig. 14).

Umbrageocoris displays a remarkable similarity to the Australian genus Stylogeocoris in general facies [studied species: S. biroi Montandon, 1913 (lectotype, HNHM) and S. elongatus (Distant, 1901) (lectotype and paralectotype, BMNH; non-types, PCPK). However, Umbrageocoris can be readily distinguished from the latter genus based on the combination of the following characters: integument of vertex smooth, dorsum of head with a single longitudinal furrow of various extent medially (it might be finely punctate and dorsum of head with arcuate grooves anteriad to ocelli in Stylogeocoris); labial through Y- or cup-shaped (V-shaped in Stylogeocoris); labiomere III longer than II (labiomeres II and III subequal in length in Stylogeocoris); margins of clavus converging gradually towards apex, claval commissure absent (margins of clavus subparallel, claval commissure short but distinct in Stylogeocoris); hamus of hemelytron partly reduced (complete in Stylogeocoris); peritreme rounded, evaporatorium reduced to surroundings of peritreme (peritreme auricular, evaporatorium covering most of the metapleurite in Stylogeocoris); sutures of abdominal tergites 4/5 elongate almost reaching margin of 5/6, apex of suture 5/6 obtuse (sutures of abdominal tergites 4/5 and 5/6 not elongate, apex rounded as in Fig. 15).

Umbrageocoris can be distinguished from representatives of genus Geocoris distributed in the region [studied species: G. ochropterus (Fieber, 1844) (lectotype, BMNH); G. willeyi Kirkaldy, 1905 (syntype, BMNH)] by the combination of following characters: head pentagonal, posterior edge of compound eyes touching anterior edges of pronotum (head lunulate, posterior edge of compound eyes encompassing anterior edges of pronotum in Geocoris); pronotum trapeziform with lateral margins slightly constricted medially (semicircular, lateral margins mostly not constricted in Geocoris); trifurcate carina of scutellum distinct, at most apically reduced (slightly distinct to completely reduced in Geocoris); hamus of metathoracic wing at most partly reduced, intervannals present, fused basally (hamus completely reduced and intervannals absent in Geocoris); apex of suture of abdominal tergites 5/6 obtuse (rounded in Geocoris sp., as in Fig. 18).

Umbrageocoris differs from representatives of Germalus [studied species: G. coloratus Distant, 1918 (syntype, BMNH); G. fuscovittatus Malipatil, 2013 (non-type, BMNH); G. victoriae Bergroth, 1895 (lectotype, BMNH)] by the following characters: ocular sulcus complete, but slightly visible (always well-defined in Germalus); labial through Y- or cup-shaped (U- or V-shaped in Germalus); labiomere III longer than II (labiomeres II and III subequal in length in Germalus); margins of clavus converging gradually towards apex, claval commissure absent (margins of clavus subparallel, claval commissure well-developed, distinct in Germalus); hamus of hemelytron partly reduced (complete in Germalus); peritreme rounded, evaporatorium reduced to surroundings of peritreme (peritreme auricular with dorsal supportive process, evaporatorium covering metapleurite and posterior half of mesopleurite in Germalus); sutures of abdominal tergites 4/5 and 5/6 elongated, suture 4/5 almost reaching margin of 5/6 (sutures of abdominal tergites 4/5 and 5/6 very short, strongly obtuse, as in Fig. 17).

Representatives of the genus are distributed in continental Indomalaya and in the Indo-Australian Archipelago, from Laos to Northeast Australia (Northern Territories, Queensland, New South Wales).

U. kondorosyi can be distinguished from other species of the genus by the entirely blackish pronotum decorated with small ochraceous spots at humeral angles; irregular brownish spots covering most of the corium; confluent pronotal callosities; and a narrow impunctate band at the posterior edge of pronotum, widened at humeral angles.

Distributed across New Guinea and adjacent islands. The species inhabits mostly rainforests, but records from mangroves and grasslands are available too.

2 m, 2 f (BPBM): „Thailand: Mae Klang / Chiang Mai Prov. / 340 m, 11. VI. 1965 // K. Morimoto / Collector // U. S.-Japan / Coop. Sci. / Program”; 1 f (BPBM): “Thailand / Nakhon Nayok Prov. / Khao Yai Nat. Park / 6. VI. 1965 // K. Morimoto / Collector // U. S.-Japan / Coop. Sci. / Program”; 1 m (BPBM): “Laos: / Vientiane Prov. / Ban Van Eue / 20. IX. 1967 // Native Collector / RONDON - - BISHOP MUS. / COLLECTION”.

U. maai maai is similar to U. boonei sp. nov. in general aspect, but the following remarkable differences enable its reliable separation from the latter species: vertex with blackish marking (lacking in U. boonei); decoration of pronotum distinctly delimited, with brownish spots at humeral angles (indistinctly delimited, somewhat blurred, spots at humeral angles lacking in U. boonei, compare Figs 8A, 10A); punctation of pronotum sparse and coarse, pronotum with impunctate “collar” anteriad to callosities (with fine, dense punctation and without significant impunctate areas in U. boonei).

The species is distributed from Thailand to Borneo and Java and has data from the following ecoregions: Northern Khorat Plateau moist deciduous forests, Southeast Indochina dry evergreen forests, Peninsular Malaysian rainforests, Kinabalu montane alpine meadows, Borneo lowland rainforests, Sundaland heath forests, and Eastern Java-Bali rainforests.

The subspecies differs from the nominotypical U. maai maai in having less extended pronotal markings and larger impunctate areas on pronotum (a single transversal line of punctures anteriad to callosities, 3–4 transversal rows posteriad to them).

The species is known from Ermera, Timor (Timor and Wetar deciduous forest ecoregion).

Holotype, m (BPBM): “Thailand / Chiangmai Prov.: / Chiangmai / (Arboretum), 300m / 12. VI. 1965 // P. D. Ashlock / Collector / BISHOP Mus. // U. S.-Japan / Coop. Sci. / Program”.

Paratypes: 2 m (BPBM): “Laos: Sedone Prov. / Paksong, 18. V. 1965 // P. D. Ashlock / Collector / BISHOP MUS.”; 1 m (BPBM): “Laos: Ban Theoung / 18km NW from Xieng / Khouang, 1035 m / 2–6. VIII. 1960 // Sweeping grasses // R. F. Leech / Collector / BISHOP”; 1 m (BPBM) “Thailand / Chiangmai Prov.: / Doi Suthep / / 12. VI. 1965 // P. D. Ashlock / Collector / BISHOP Mus. // U. S.-Japan / Coop. Sci. / Program”; 1 m (BPBM): “Thailand: Tang Kao / Chiang Mai Prov. / 10. VI. 1965 // P. D. Ashlock / Collector / BISHOP Mus. // U. S.-Japan / Coop. Sci. / Program”; 1 f (BPBM): Thailand / Bangkok / June 3, 1965 // P. D. Ashlock / Collector / BISHOP Mus.”; 1 m (PCPK): “LAOS: Sedone Prov. / Paksong, 18. V. 1965 // P. D. Ashlock / Collector / BISHOP MUS.”.

Colouration: Head ochraceous with dark brownish irregular spot at base of eye stalks posteriorly. Antennomere I ochraceous with apex irregularly brownish; antennomeres II and III entirely dark brownish; antennomere IV dark brownish with apex ochraceous. Labiomeres uniformly ochraceous. Thorax. Pronotum ochraceous with dark brownish punctation, a dark brownish or dark fuscous transversal band across callosities and irregular infuscate spot of various extent posteriad to callosities. Scutellum entirely dark brownish or blackish. Hemelytra semi-hyaline, ochraceous with dark brownish punctation and an irregular infuscate spot at apical half of corium. Thoracic pleurites and sternites dark brownish except ochraceous prosternal collar, supracoxal lobes and peritreme. Abdomen dark brownish, undecorated.

Structure: Head. Eyes slightly stylate; ocular sulcus complete, but slightly visible, rather indistinct in apical half. Ocelli situated near ocular sulci, closer to each other than to compound eyes; ratio of ocellar distance to eye ocellus distance: 1: 1.16. Dorsum of head with median longitudinal furrow extending from apex of clypeus to base of head. Relative lengths of antennomeres: 1.00: 2.40: 1.82: 4.36. Relative lengths of labiomeres: 1.00: 0.57: 0.81: 0.98. Thorax. Pronotum trapeziform with lateral margin not constricted; integument densely and deeply punctate at anterior margin and narrow pronotal callosities; pronotum length to basal width: 1.00: 1.35. Scutellum with integument densely punctate except slightly bulging, complete trifurcate carina; scutellum length to width: 1.00: 1.07. Corium submacropterous, membrane slightly surpassing apex of abdomen; corium punctate along margin of clavus, along Cu and between M-R and costa at apical half; exocorium uniformly narrow at entire length; membrane wrinkled. Thoracic sternites and pleurites densely punctate except narrow prosternal collar, supracoxal lobes and peritreme. Peritreme bulbous, situated ventrolaterally; dorsal flange with distinct peritremal surface; evaporatorium restricted to surroundings of peritreme. Abdomen. Abdominal venter with sparse, decumbent pubescence; integument of abdominal sternites III-VII corrugate dorsolaterally.

Measurements (holotype). Body length: 3.11; head length: 0.62; head width: 1.51; lengths of antennomeres I–IV: 0.14–0.33–0.25–0.60; lengths of labiomeres I–IV: 0.47–0.27–0.38–0.46; pronotum length: 0.83; pronotum width: 1.39; scutellum length: 0.72; scutellum width: 0.79.

U. boonei gen. nov. resembles both subspecies of U. maai in colouration and markings. The new species can be distinguished from other representatives of the genus by lacking an impunctate “collar” on the pronotum and its highly extended trifurcate carina on the scutellum.

The species is known from continental Indomalaya (Laos and Thailand) from the following ecoregions: Kayah-Karen montane rainforests, Luang Prabang montane rainforests, Southern Annamites montane rainforests, Chao Praya freshwater swamp forests, Indochina mangroves.

Holotype, f (MNHN): “MUSEUM PARIS / NOUV. GALLES DU SUD / COLL. NOUALHIER 1898”.

Colouration: Head ochraceous, posterior part of eye stalks with irregular brownish marking. Eyes reddish, ocelli hyaline. Antennae missing in the single examined specimen. Labiomeres dark ochraceous. Thorax. Pronotum ochraceous with brownish punctation and an infuscate transversal band across pronotal callosities. Scutellum entirely dark brownish. Hemelytra semi-hyaline, ochraceous with brownish punctation and an irregular infuscate spot on the corium. Thoracic pleurites and sternites dark brownish except prosternal collar, supracoxal lobes and peritreme; propleurite with an ochraceous spot dorsally. Abdomen dark brownish both dorsally and ventrally with a whitish longitudinal band at dorsal part of sternites.

Structure: Head. Median furrow of vertex short, hardly surpassing base of clypeus. Antennomeres missing. Labial trough Y-shaped with suture reduced, reaching approximately midline of head. Relative lengths of labiomeres: 1.00: 0.85: 1.32: 1:38. Thorax. Pronotum with dense and deep punctation except anterior margin, pronotal callosities and posterior margin with humeral angles. Pronotal callosities confluent. Pronotum length to basal width: 1.00: 1.76. Scutellum with trifurcate carina apically reduced. Hemelytra submacropterous. Scutellum length to basal width: 1.00: 1.06. Clavus with an incomplete row of punctures at corial margin. Corium punctate along margin of clavus, Cu, between M-R and costa in apical half. Exocorium slightly widened subapically. Membrane short, not surpassing apex of abdomen.

Measurements. Body length: 4.20; head length: 0.53; head width: 1.66; lengths of labiomeres I–IV: 0.36–0.30–0.47–0.49; pronotum length: 0.88; pronotum width: 1.54; scutellum length: 0.88; scutellum width: 0.83.

U. malipatili sp. nov. resembles U. boonei sp. nov. and both subspecies of U. maai in colouration and markings, but its cup-shaped labial trough and the presence of a minute furrow on its clypeus readily distinguishes it from the Indomalayan representatives of the genus.

The single known specimen is from an unknown locality in New South Wales, Australia.

Lectotype, m (BMNH): circular blue syntype label // circular red type label // “elegantulus / Dist. [hw2] // 1903 – 322. // UNSW_ENT 00011372 // BMNH(E) / 1231237 // NHMUK 010591767 // Townsville, Qld / 16.12.02 / F. P. Dodd.”

1 m (SEMC): “NEW Guinea: Papua / Koitake / 2. X. 1958 / J. L. Gressitt / Collector”.

Additions and corrections to the redescription of Malipatil (1994). Head. Median longitudinal furrow extends from middle of clypeus to middle of vertex. Labial trough cup-shaped with suture reaching only middle of venter of head. Thorax. Peritreme bulbous, dorsal flange indented.

U. elegantulus resembles U. woodwardi in general appearance but can be distinguished from the latter species based on the following characters: vertex with extended blackish markings (vertex only marked basally in U. woodwardi); dorsal flange of peritreme indented (Fig. 9.) (not indented in U. woodwardi).

The species was recorded from the northern parts of Australia in the following ecoregions: Arnhem land tropical savanna, Cape York Peninsula tropical savanna, Queensland tropical rainforests, Brigalow tropical savanna (Malipatil 1994). A specimen from Koitake, New Guinea (Southeastern Papuan rain forest ecoregion) was examined.

This species was described as Geocoris elegantulus (Distant 1904) and cited as such by subsequent authors (Malipatil 1994; Cassis and Gross 2002). A careful examination of both type and non-type specimens available concluded that the species displays synapomorphies used to define Umbrageocoris, including the presence of a median groove on vertex, a Y- or cup-shaped labial trough, and the arrangement of sutures at abdominal tergites 4/5 and 5/6, thus the following new combination is here proposed: Umbrageocoris elegantulus (Distant, 1904), comb. nov.

2 f (BPBM): „New Guinea: NE. / Moife 2100 m. / 7–14. X. 1959 // Swept in / clearing // T. C. Maa / Collector / BISHOP”; 1 f BPBM: „NE. / Moife 2100 m. / 15km NW of Okapu / 7–14. X. 1959 // T. C. Maa / Collector / BISHOP”.

Additions and corrections to the description of Malipatil (1994). Head. Median longitudinal furrow of vertex minute, hardly surpassing base of clypeus. Labial trough cup-shaped with suture short, hardly reaching middle of venter of head. Thorax. Peritreme bulbous, dorsal flange not indented.

U. woodwardi resembles U. elengantulus in general aspect, but it can be distinguished from the latter species based on the combination of the following characters: vertex with dark marking only basally (with more extensive dark markings in U. elegantulus); dorsal flange of peritreme simple (indented in U. elegantulus).

Most of the published records of the species are from Western Australia, where the species inhabits Brigalow tropical savanna and Eastern Australian temperate forest habitats (Malipatil 1994). In the material deposited in the BPBM three specimens from the Central Range montane rain forests of New Guinea are present.

Based on the specimens available as well as the original description and illustrations provided by Malipatil (1994) the species possesses a combination of characters diagnostic of Umbrageocoris, including the presence of a median longitudinal furrow on vertex, a cup- or Y-shaped labial trough, and the arrangement of sutures of abdominal tergites 4/5 and 5/6. The following new combination is accordingly proposed: Umbrageocoris woodwardi (Malipatil, 1994), comb. nov.