Research Article Journal of Orthoptera Research 2022, 31(1): 9-40 Rugabinthus, a new genus of Lebinthina (Orthoptera, Gryllidae, Eneopterinae) from New Guinea MING KAI TAN!, TONY ROBILLARD! 1 Institut de Systématique, Evolution, Biodiversité (ISYEB), Muséum national d'Histoire naturelle, CNRS, SU, EPHE, UA, 57 rue Cuvier, CP 50, 75231 Paris Cedex 05, France. Corresponding author: Ming Kai Tan (orthoptera.mingkai@gmail.com) Academic editor: Klaus-Gerhard Heller | Received 1 September 2021 | Accepted 30 September 2021 | Published 21 February 2022 http://zoobank.org/3906D 111-1849-4F9B-87FD-F70673B1BGOE Citation: Tan MK, Robillard T (2022) Rugabinthus, a new genus of Lebinthina (Orthoptera, Gryllidae, Eneopterinae) from New Guinea. Journal of Orthoptera Research 31(1): 9-40. https://doi.org/10.3897/jor.31.73800 Abstract Brachypterous crickets from the monophyletic group of Lebinthina were traditionally grouped under the genus Lebinthus. However, the mor- phology and calling song are highly diversified, prompting the erection of numerous genera to reclassify the species. Based on the strong char- acteristic fold carrying the diagonal vein of the male forewing, a new ge- nus of cricket from the subtribe Lebinthina is described: Rugabinthus gen. nov. This brachypterous genus is endemic to the island of New Guinea and nearby islands. We redescribe the type species Rugabinthus leopoldi (Chopard, 1931) comb. nov. and describe 12 new species, R. manokwari sp. nov., R. kencana sp. nov., R. maoke sp. nov., R. nabire sp. nov., R. alba- tros sp. nov., R. karimui sp. nov., R. yayukae sp. nov., R. biakis sp. nov., R. mamberamo sp. nov., R. tariku sp. nov., R. faowi sp. nov., and R. baduri sp. nov. We also transferred R. newguineae (Bhowmik, 1981) comb. nov. and provide a key to all known species of Rugabinthus gen. nov. Keywords Grylloidea, Lebinthini, new species, Papua, Southeast Asia, taxonomy Introduction The tribe Lebinthini Robillard, 2004 is comprised of a high- ly speciose clade of crickets with diverse morphologies, ranging from the tiny brachypterous Pixibinthus Robillard & Anso, 2016 to the large and fully winged species of the genus Cardiodactylus Saussure, 1878. Male lebinthines produce high-frequency calls (10-28 kHz), and their call structure can also be highly diverse, ranging from a few chirps to a combination of a series of chirps followed by a trill (e.g., Robillard and Desutter-Grandcolas 2004a, 2004b, 2011, Tan et al. 2021). Lebinthines have been demonstrated to be important species for the study of insect communication systems. A new communication system using vibrational responses (including vibrotaxis) to high-frequency male calls and population-specific microevolution and plastic- ity in acoustic properties was recently discovered (ter Hofstede et al. 2015, Benavides-Lopez et al. 2020, Tan and Robillard 2021a, b). Among the Lebinthini, the subtribe Lebinthina is distributed in the hyper-diverse yet poorly studied Southeast Asia and the western Pacific. This clade includes many taxa that are still being discovered. In particular, two genera of Lebinthina were described from Southeast Asia, Fadinthus Robillard & Tan, 2021, and Falcer- minthus Robillard & Tan, 2021 (see Tan et al. 2021), which include numerous new species (Baroga-Barbecho et al. 2016, 2020, Tan et al. 2019). Continued efforts to sample the region where Lebinthina is distributed and discover new species with diverse morpholo- gies have led to recent insights about how species are related and should be classified. Traditionally, Lebinthus included most brachypterous species of Lebinthini. However, as new informa- tion about these species became available, their classification has been more accurately revised (Tan et al. 2021) and the subtribe Lebinthina Robillard & Tan, 2021 has been defined. Multiple new genera have also been erected, including Centuriarus Robil- lard, 2011 (Robillard 2011), Gnominthus Robillard & Vicente, 2015 (Vicente et al. 2015), Microbinthus Robillard & Dong, 2016, and Macrobinthus Robillard & Dong, 2016 (Robillard et al. 2016) from New Guinea; Pixibinthus Robillard & Anso, 2016 from New Cal- edonia (Anso et al. 2016); Falcerminthus Robillard & Tan, 2021 and Fadinthus Robillard & Tan, 2021 from Southeast Asia (Tan et al. 2021); and Julverninthus Robillard & Su, 2018 from Australia (Robillard and Su 2018). These studies revealed that most Lebin- thus species known before 2010 corresponded to distinct genera, and that the island of New Guinea possesses the highest diversity of genera and species of the Lebinthina clade (Tan et al. 2021, Tan and Robillard 2021c). Pursuing the taxonomic revision of the species described from New Guinea, we found that Lebinthus leopoldi Chopard, 1931 (Fig. 1) differs from the Lebinthina genera that are cur- rently recognized. It shows unique morphological features in the male forewing (FW), including a strong fold carrying the diagonal vein (Fig. 2). In addition, study of undetermined ma- terial from several natural history museums, mostly collected in the western half of New Guinea, revealed numerous new spe- cies that share the characteristics of L. leopoldi (Fig. 3). Hence, JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) 10 we propose a new genus name for these species: Rugabinthus gen. nov. We redescribe the type species Rugabinthus leopoldi (Chopard, 1931) comb. nov. and describe 12 new species, as well as providing a key to all known species of Rugabinthus gen. nov. We also transfer R. newguineae (Bhowmik, 1981) comb. nov. to this new genus. Materials and methods Materials.—Specimens in MNHN were studied in addition to ma- terials loaned from BPBM, MZB, NHMUK, RBINS, RMNH, and ZIN (see abbreviations below). Morphology.—Male tegminal veins and cells follow the terminol- ogy of Robillard and Desutter-Grandcolas (2004a). Male and female genitalia were dissected in softened specimens by cut- ting the membranes between the paraprocts and the subgenital plate, or between the ovipositor and the subgenital plate, re- spectively; they were observed after cleaning with cold KOH and then kept in glycerine. Male genitalia are named according to Desutter (1987), modified in Desutter-Grandcolas (2003) and Robillard and Desutter-Grandcolas (2004a). For abbreviations, see below. Close-up images of habitus and morphological features were obtained using a Canon EOS 6D digital SLR camera with a macro photo lens MP-E 65 mm f/2.8 USM (1-5x). Imaging stacking was done using Helicon Remote version 9.3.1. W and Helicon Focus 6.8.0. Photographs of male and female genita- lia were done with a binocular microscope Leica MZ16 with an AMScope Microscope Eyepiece Camera (MU1000, 10 MP Ap- tina Colour, CMO50) attached via an AmScope FMAO50 fixed microscope adaptor and the software ToupView. Image editing was accomplished using CombineZP version 1.0 and Adobe Photoshop CC2014. To highlight the structural components of genitalia, a water solution containing a drop of JBL Punktol was used. To fix orientations and stabilization of geni talia for pho- tography, a clear and viscous hand sanitizer was used following Su (2016). Abbreviations. — Depositories: BPBM Bernice Pauahi Bishop Museum, Hawaii, U.S.A.; MNHN Muséum national d'Histoire naturelle, Paris, France; MZB Museum Zoologicum Bogoriense, Bogor, Java, Indonesia; NHMUK Natural History Museum, London, U.K.; RBINS Royal Belgian Institute of Natural Sciences, Brussels, Belgium; RMNH Nationaal Natuurhistorisch Museum (formerly Rijksmuseum van Natuurlijke Historie), Lei- den, The Netherlands; ZIN Zoological Institute, Russian Academy of Sci- ences, St. Petersburg, Russia. General morphology: I, U1, il front, median, hind, respectively (femora, legs, tibiae); F femora; M.K. TAN AND T. ROBILLARD FW forewing; Talll-1 basal segment of hind leg tarsomere; T tibiae. Tegminal venation: 1A-4A first to fourth anal veins; CuA anterior cubital vein; CuA1, CuA2,... first, second, ... bifurcations of CuA; CuP posterior cubital vein; M median vein; Sc subcostal vein; R radial vein; c1-3 first to third cells of C alignment; d1 cell (mirror) first cell(s) of D alignment; d2 second cell of D alignment; el first cell of E alignment; ha harp area. Measurements: FIIIL length of hind femora; FITW width of hind femora; FWL forewing length; FWW forewing width (at the level of maximal width); OL ovipositor length; PronL pronotum length; PronW pronotum width; THIL length of hind tibiae. Results Taxonomy Family Gryllidae Laicharting, 1781 Subfamily Eneopterinae Saussure, 1874 Tribe Lebinthini Robillard, 2004 Subtribe Lebinthina Robillard & Tan, 2021 Genus Rugabinthus Robillard & Tan, gen. nov. http://zoobank.org/5239CAF1-F14C-4332-8153-2EC75E03CFOD Type species.—Lebinthus leopoldi Chopard, 1931 Etymology.—Genus named after the Latin word “Ruga” for wrinkle or fold, referring to the characteristic fold on the male FW carrying the diagonal vein. Distribution.—Island of New Guinea: Indonesia (West Papua) and Papua New Guinea (Fig. 3). Diagnosis.—Among the Lebinthina genera, Rugabinthus species are average to large sized and stocky with a dark brown coloration. General shape close to that of Macrobinthus, also from New Guin- ea, from which it differs by male FWs with a narrow triangular harp (shield shaped in Macrobinthus) occupying half of FW width, with a characteristic and strong diagonal fold carrying the diago- nal vein and cell cl, and separating FWs in two distinct areas; file vein area also characterized by a strong bean-shaped sclerotiza- tion; harp with a strong transverse oblique vein, bi- or poly-furcat- ed anteriorly; venation posterior to diagonal fold usually faint and JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) M.K. TAN AND T. ROBILLARD reticulated, longitudinal veins only strong at apex. Eyes prominent and large as in Macrobinthus compared to Agnotecous and Centuri- arus; face as high as wide (higher than wide in Macrobinthus), close to that of Lebinthus. Microptery in both sexes, FWs not reaching mid-length of abdomen. Mirror not differentiated (slightly differ- entiated in Macrobinthus). CuA almost straight (clearly curved in- wards in Macrobinthus). Male genitalia with pseudepiphallus usu- ally elongate, its posterior apex highly variable in size and shape, rami short; pseudepiphallic parameres made of two main lobes variable in shape and orientation; endophallic sclerite very long, trifid posteriorly. Female: FWs shorter than in males, very slight- ly overlapping, generally rounded posteriorly. Ovipositor rather long, its apex slightly denticulate on dorsal edge. Female copula- tory papilla usually rounded, with a C-shaped basal sclerite; apex rounded, generally folded ventrally. Description.—Size variable, medium to large for the subtribe. Dor- sum of head rounded, prolonged by trapezoidal fastigium almost as long as wide (Fig. 4). Head dorsum with six wide dark brown longitudinal bands more or less distinct (Fig. 4). Eyes large and prominent. Head triangular in facial view, almost as wide as high (Fig. 5). Face coloration usually red brown to dark brown or black, with various color patterns; most species with four pale brown or yellow dots forming a square on face (two between scapes, and two above epistomal suture) (Fig. 5). Maxillary palpi brown or dark brown (sometimes black in dark species) with lighter brown rings. Ocelli pale, forming a rather wide triangle; median ocellus slightly oval, lateral ocelli small and rounded. Scapes small. An- tennae usually dark brown with yellow brown rings. Lateral part of head dark brown with a yellow spot below eye and a yellow band behind eye (Fig. 6). Pronotum dorsal disk trapezoidal, wider than long, its posterior margin straight, usually dark brown with lat- eral edges yellow (Fig. 4). Lateral lobes of pronotum longer than high, most often uniformly red brown or black dorsally, its ven- tral margin mostly dark brown, sometimes with a yellow pattern (Fig. 6). Legs: TI with two tympana; inner tympanum covered by a flat sclerotized expansion, its membrane visible along a small longitudinal slit only; outer tympanum ellipsoidal, its membrane transversally plicate in dorsal half. TI with two inner and two outer apical spurs. TII with two inner and two outer spurs. FI muscu- lar. TIII serrulate on their whole length, slightly furrowed dorso- longitudinally and with four pairs of subapical spurs and three pairs of apical spurs; inner spurs long and curved, outer spurs shorter and straight. Legs I and II light brown to yellow brown, femora with brown spots and longitudinal patterns, tibiae with brown rings. TallI-1 with 4-5 spines on dorsal outer edge, with- out spines on dorsal inner edge. TIIIs and tarsomeres brown with spines and spurs with dark apices. FWs short in both sexes, not reaching abdomen mid-length; hind wings absent. Cerci well de- veloped, nearly as long as abdomen. Abdomen: Tergites brown to dark brown, without longitudinal bands. Male. Metanotal glands absent. Dorsal field of FWs not dis- tinctly longer than lateral field. FWs longer than wide, longer than pronotum dorsal disk. FW venation (Figs 2, 7): 1A vein (file) trans- verse part straight, variably curved at basal end, bisinuate anterior- ly to angle, forming a notch in most species. Genus characterized by a large bean-shaped sclerotization posterior to transverse part of file vein (1A), including base of chords. Diagonal vein straight or faintly sinusoidal, strong basally, fainter posteriorly. Harp tri- angular, longer than wide, occupying approximately half of dorsal field surface; with a strong characteristic fold along posterior face, 1] carrying diagonal vein and cell cl and delimiting dorsal field of FW in two distinct parts; harp with a strong transverse oblique vein, bi- or poly-furcated anteriorly; area posterior to diagonal fold with weak reticulated venation, cell alignments almost in- distinct except at apex and main longitudinal veins weak except at apex. CuA anterior part strong throughout, straight, slightly curved inward near apex. Mirror (d1) usually not differentiated. Apical field short, including one or two cell alignments. Lateral field with 5-6 strong longitudinal veins; Sc vein without bifur- cation. Subgenital plate elongated, clog-shaped, slightly pointed; inner side of subgenital plate with lateral swellings. Epiproct and subgenital plate brown. Male genitalia: (Figs 8, 11, 15, 20, 24) Pseudepiphallus trian- gular, variable in length but usually elongate, its basal margin straight or variably indented in the middle, posterior apex usually somewhat truncated, sometimes slightly acute, rarely with paired lophi. Rami very short, usually shorter than half of pseudepiphal- lus length, parallel to slightly diverging anteriorly. Pseudepiphallic parameres average sized and sclerotized, variable in shape, with posterior apex usually enlarged, their basis strong. Ectophallic ap- odemes parallel and long, usually reaching beyond anterior mar- gin of pseudepiphallic sclerite. Ectophallic arc well sclerotized, transverse. Ectophallic fold with two ventral sclerites of variable shapes, sometimes fused together. Endophallic sclerite Y-shaped, comprising a long anterior region and a short median expansion and lateral arms posteriorly. Female. FWs very short, shorter than in male, reaching or slight- ly surpassing posterior margin of first tergite, close together and usu- ally slightly overlapping at their bases. Dorsal field usually oblique posteriorly (more so in some species than others); dorsal field and lateral field with more or less distinct longitudinal veins (Fig. 9). Dorsal field with cells brown and mottled and veins brown; lateral field with cells brown mottled with gray, and veins brown. Female genitalia: Ovipositor most often slightly longer than FI, slightly denticulate on dorsal edge, its apex acute. Female copulatory papilla rounded, almost entirely membranous, some- times with a basal sclerotized ring; apex rounded, generally folded ventrally (Fig. 10). Calling song.—Unknown. Natural history.—The rare information obtained from photographs from iNaturalist suggest that these crickets are found in the leaf litter and in the foliage of low-lying vegetation in forest (Fig. 3). Included species.—(14 in total; ordered by similarity) . leopoldi (Chopard, 1931) comb. nov. mamberamo sp. nov. yayukae sp. nov. faowi sp. nov. . kencana sp. nov. manokwari sp. nov. maoke sp. nov. biakis sp. nov. nabire sp. nov. tariku sp. nov. . albatros sp. nov. . baduri sp. nov. . karimui sp. nov. . newguineae (Bhowmik, 1981) comb. nov. Dm DDD DAD DAD DDD JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) M.K. TAN AND T. ROBILLARD ee Manon. (Nouvelle Guinée) : eI ey pee Prince Leopold | NN HA-ENSIF 4893 Fig. 1. R. leopoldi (Chopard, 1931) comb. nov. male (A, B) and female (C, D) habitus in dorsal (A, C) and lateral (B, D) views. Paratype labels (E). Scale bar: 10 mm. Rugabinthus leopoldi (Chopard, 1931) comb. nov. (Figs 1, 3, 4A, 5A, 6A, 7A, 8A, 9A, 10A, 11A, 11B) Lebinthus leopoldi Chopard, 1931: 7; 1968: 354; Cigliano et al. 2021 (Orthoptera species file online). Material examined.—Holotype: INDONESIA ¢ 4; West Papua, Ma- noi [Sorong Manoi]; 2 March 1929; Prince Leopold leg.; RBINS. Allotype: INDONESIA ¢ 9; same information as holotype; RBINS. Paratypes: INDONESIA ¢ 3, 49; same information as holotype; RBINS ¢ 14; West Papua, Manoi [Sorong Manoi]; 2 March 1929; Prince Leopold leg.; MNHN-EO-ENSIF1441 ¢ 19; same informa- tion as holotype; molecular sample L173; MNHN-EO-ENSIF1443 e 19; same information as holotype; MNHN-EO-ENSIF1442. Type locality.—_INDONESIA: West Papua: Manoi Diagnosis.—This species differs from all congeners by male genita- lia with pseudepiphallus more rectangular, very elongate, its pos- terior part curved posteriorly, its apex truncated with small paired apical lophi; lophi triangular with obtuse apex. Redescription.—Average size among congeners (Fig. 1). Dorsum of head with broad red brown bands barely separated (Fig. 4A). Fastigium red brown (Fig. 4A). Scapes red brown with some dark patches. Fastigium verticis brown with two vertical yellow brown stripes diverging ventrad, frons brown with two yellow brown anterior notch of 1A CuA di Fig. 2. Schematic venation of male FW of Rugabinthus. The gray area represents the diagonal fold carrying the diagonal vein; the yellow area represents the bean-shaped strong sclerotization along 1A vein. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) M.K. TAN AND T. ROBILLARD 13 @ R. leopoldi © R. mamberamo +e R. yayukae R. faowi R. kencana R. manokwan * R. maoke WR. biakis 7 AR. nabire - R. fariku WR. albatros ©) R. baduri ) R. karimui R. newguineae A iNat: 72638321 - | @ iNat: 73044465 Fig. 3. A. Map of New Guinea island showing the distribution of Rugabinthus species; B. Female Rugabinthus species in their natural habitats in Acemo, South Manokwari (https://www.inaturalist.org/observations/73044465) and C. Malagufuk (https://www.inatural- ist.org/observations/72638321). Photo credit: Benoit Segerer. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) 14 M.K. TAN AND T. ROBILLARD mam Fig. 4. Head and pronotum in dorsal views: A. R. leopoldi (Chopard, 1931) comb. nov.; B. R. mamberamo sp. nov.; C. R. yayukae sp. nov.; D. R. faowi sp. nov.; E. R. kencana sp. nov.; F. R. manokwari sp. nov.; G. R. maoke sp. nov.; H. R. biakis sp. nov.; I. R. nabire sp. nov.; J. R. tariku sp. nov.; K. R. albatros sp. nov.; L. R. baduri sp. nov.; M. R. karimui sp. nov.; N. R. newguineae (Bhowmik, 1981) comb. nov. Scale bar: 5 mm. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) M.K. TAN AND T. ROBILLARD 15 Fig. 5. Face: A. R. leopoldi (Chopard, 1931) comb. nov.; B. R. mamberamo sp. nov.; C. R. yayukae sp. nov.; D. R. faowi sp. nov.; E. R. ken- cana sp. nov.; F. R. manokwari sp. nov.; G. R. maoke sp. nov.; H. R. biakis sp. nov.; I. R. nabire sp. nov.; J. R. tariku sp. nov.; K. R. albatros sp. nov.; L. R. baduri sp. nov.; M. R. karimui sp. nov.; N. R. newguineae (Bhowmik, 1981) comb. nov. Scale bar: 2 mm. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) 16 M.K. TAN AND T. ROBILLARD Fig. 6. Head and pronotum in lateral views: A. R. leopoldi (Chopard, 1931) comb. nov.; B. R. mamberamo sp. nov.; C. R. yayukae sp. nov.; D. R. faowi sp. nov.; E. R. kencana sp. nov.; FE. R. manokwari sp. nov.; G. R. maoke sp. nov.; H. R. biakis sp. nov.; I. R. nabire sp. nov.; J. R. tariku sp. nov.; K. R. albatros sp. nov.; L. R. baduri sp. nov.; M. R. karimui sp. nov.; N. R. newguineae (Bhowmik, 1981) comb. nov. Scale bar: 5 mm. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) M.K. TAN AND T. ROBILLARD 17 Fig. 7. Male FW in dorsal views: A. R. leopoldi (Chopard, 1931) comb. nov.; B. R. mamberamo sp. nov.; C. R. yayukae sp. nov.; D. R. faowi sp. nov.; E. R. kencana sp. nov.; F. R. manokwari sp. nov.; G. R. maoke sp. nov.; H. R. biakis sp. nov.; I. R. nabire sp. nov.; J. R. tariku sp. nov.; K. R. albatros sp. nov.; L. R. karimui sp. nov. Scale bar: 2 mm. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) 18 M.K. TAN AND T. ROBILLARD leo mam fao ken mao bia nab . tar | ; | alb Fig. 8. Male genitalia in dorsal view; posterior apex of pseudepiphallus: A. R. leopoldi (Chopard, 1931) comb. nov.; B. R. mamberamo sp. nov.; C. R. yayukae sp. nov.; D. R. faowi sp. nov.; E. R. kencana sp. nov.; FE. R. manokwari sp. nov.; G. R. maoke sp. nov.; H. R. biakis sp. nov.; I. R. nabire sp. nov.; J. R. tariku sp. nov.; K. R. albatros sp. nov.; L. R. karimui sp. nov. Scale bar: 0.5 mm. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) M.K. TAN AND T. ROBILLARD 19 Fig. 9. Female FW in dorsal views: A. R. leopoldi (Chopard, 1931) comb. nov.; B. R. mamberamo sp. nov.; C. R. faowi sp. nov.; D. R. ken- cana sp. nov.; E. R. manokwari sp. nov.; F. R. nabire sp. nov.; G. R. tariku sp. nov.; H. R. albatros sp. nov.; I. R. baduri sp. nov.; J. R. karimui sp. nov.; K. R. newguineae (Bhowmik, 1981) comb. nov. Scale bar: 2 mm. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) 20 M.K. TAN AND T. ROBILLARD Bo mam E man F nab G tar alb Fig. 10. Female copulatory papilla in ventral (all except I, K) and lateral (I, K) views. A. R. leopoldi (Chopard, 1931) comb. nov.; B. R. mamberamo sp. nov.; C. R. faowi sp. nov.; D. R. kencana sp. nov.; E. R. manokwari sp. nov.; F. R. nabire sp. nov.; G. R. tariku sp. nov.; H, I. R. albatros sp. nov.,; J. R. baduri sp. nov.; K, L. R. karimui sp. nov.; M. R. newguineae (Bhowmik, 1981) comb. nov. Scale bar: 0.5 mm. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) M.K. TAN AND T. ROBILLARD 21 Fig. 11. Male genitalia ventral (A, C, E) and lateral (B, D, F) views: A, B. R. leopoldi (Chopard, 1931) comb. nov.; C, D. R. mamberamo sp. nov.; E, F. R. yayukae sp. nov. Scale bar: 1 mm. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) 22 spots ventral of stripes; clypeus sometimes darkened, mouthparts yellow brown to red brown (Fig. 5A). Pronotal disk red brown with lighter lateral ovular spots near anterior half and some faint lighter spots along posterior margin (Fig. 4A). Lateral lobes of pronotum dark brown, with yellow spot at anterior ventral corner (Fig. 6A). FIs and FIIs generally yellow brown with brown spots and rings; TIs and TIls brown with two yellow brown rings. FIIIs brown, knees slightly darker. Male. FW reaching apex of third tergite; apex rounded. FW col- oration (Fig. 7A): Dorsal field cells and veins mostly brown; area between M and R yellow brown; basal area with a wide cream- colored spot on external corner. Lateral field dark brown. FW ve- nation typical of genus; 1A slightly bisinuate anterior to angle; oblique vein trifurcated, posterior branch straight and transverse. Male genitalia: (Figs 8A, 11A, 11B) Pseudepiphallus form- ing a very elongate rectangle, its basal margin slightly indented in the middle, lateral margins widened basally, forming wide shoulders carrying base of rami; posterior part of pseudepiphal- lus narrow, distinctly curved dorsally posterior to pseudepiphal- lic parameres, with two short dorsal pre-apical expansions (Fig. 8A); posterior apex truncated, posterior apex with small paired lophi, triangular with obtuse apex. Rami wide, very short, way shorter than half the pseudepiphallus length, diverging anteri- orly, with posterior end protruding externally. Pseudepiphallic parameres strongly bent in basal half (~90°), with posterior apex enlarged, bean shaped. Ectophallic apodemes thin, paral- lel, and long, reaching beyond anterior margin of pseudepiphal- lic sclerite. Endophallic sclerite with anterior region very short and not reaching anterior margin of pseudepiphallic sclerite, with lateral arms elongated, tongue-shaped, and longer than median expansion. Female. FW reaching base of third tergite, with basal area with a cream-colored oblong spot without clearly defined margin, M.K. TAN AND T. ROBILLARD forming two indistinct spots near base and apex; lateral field dark brown (Fig. 9A). Female genitalia: Ovipositor about as long as FIII. Copulatory papilla very small, rounded, apex folded ventrally, short, pointed; dorsal face with a sclerotized area; ventro-anterior end base form- ing an oval ring (Fig. 10A). Measurements.—See Table 1. Rugabinthus mamberamo sp. nov. http://zoobank.org/31AB9356-DC05-4203-AD3B-F7257C4B611E (Figs 3, 4B, 5B, 6B, 7B, 8B, 9B, 10B, 11C, 11D, 12) Material examined.—Holotype: INDONESIA ¢ <4; West Papua, Fawi [Faowi] village in upper part of Tariku River (tributary of Mamberamo River), partly low-lying forest and partly forest on hills; 29 January-17 February 2012; A. Gorochov leg.; molecular sample L94; ZIN. Paratype: INDONESIA ¢ 19; same information as holotype; MNHN-EO-ENSIF1758. Type locality.—INDONESIA: West Papua: Faowi Etymology.—This species is named after Mamberamo River; noun in apposition. The name is derived from the main river rather than the tributary Tariku River because it has more elongated and larger male genitalia compared to the sympatric species R. tariku sp. nov. Diagnosis.—This new species differs from all congeners by male genitalia, with pseudepiphallus very slender and very elongate, close to that of R. leopoldi from which it differs by posterior apex forming a long flat spoon slightly curved dorsally near apex, look- ing like a simpler version of R. leopoldi, with thinner pre-apical ex- pansions, apex somewhat truncated, without lophi; pseudepiphal- Fig. 12. R. mamberamo sp. nov. male (A, B) and female (C, D) habitus in dorsal (A, C) and lateral (B, D) views. Scale bar: 10 mm. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) M.K. TAN AND T. ROBILLARD Table 1. Measurements (in mm). 23 PronL PronW FWL FWW FIUIL FTW THIL OL R. leopoldi (Chopard, 1931) Male paratype MNHN1441 3.0 4.9 4.5 3.0 13.7 3.9 10.4 - Female paratype MNHN1443 3.3 4.9 3.8 2.8 14.3 4.5 12.2 14.1 Female paratype MNHN1442 3.2 5.0 4.0 2.6 14,2 4.5 10.9 14.8 R. mamberamo sp. nov. Male holotype 3.0 4.4 4.3 2.8 12.4 3.6 11.0 - Female paratype 3.0 44 3.7 2.6 12.3 3.9 10.7 9.1 R. yayukae sp. nov. Male holotype 2.8 4.8 5.7 3.8 13.9 4.1 10.1 - Males (n = 4) 2.6-3.3 (2.9) 4.7-5.0(4.8) 4.9-6.0 (5.5) 3.8-4.5 (4.0) 13.4-15.6 (14.2) 4.1-4.7 (4.3) 10.1-12.6 (11.7) - R. faowi sp. nov. Male holotype 2.9 3.6 4.6 2.6 11.9 Sf 10.7 - Males (n = 4) 2.9-3.3 (3.0) 3.6-5.2(4.6) 4.6-6.7 (5.8) 2.6-3.5 (3.2) 11.9-15.8(13.9) 3.7-4.5(4.1) 10.7-13.1 (11.9) - Females (n = 4) 3.4-3.6 (3.5) 5.0-5.3(5.1) 4.4-4.6 (4.5) 2.8-2.8(2.6) 16.4-17.0 (16.7) 4.9-5.2 (5.0) 15.0-15.6 (15.4) 19.2-21.0 (20.2) R. kencana sp. nov. Male holotype 3.0 5.0 6.9 5.0 - - - - Males (n = 2) 3.0-3.4 (3.2) 5.0-5.1 (5.13 6.4-6.9 (6.7) 4.3-5.0 (4.7) 14.9 (14.9) 4.6 (4.6) 1329. C1329) - Female paratype 3.2 5.1 4.5 2.8 16.2 4.7 14.1 17.9 R. manokwari sp. nov. Male holotype 2.9 4.4 4.1 372. 11.7 3.8 10.4 - Female paratype 2.9 4.4 3.5 DS 13.4 3.7 12.1 10.9 R. maoke sp. nov. Male holotype 3.4 5.8 6.5 4.7 18.0 512 11.0 - Male paratype 3.6 5.5 6.5 5.0 17.3 4.9 15.8 - R. biakis sp. nov. Male holotype 2.5 3.9 3.9 2.7 10.7 2.7 8.6 - Males (n = 4) 2.4-2.5 (2.5) 3.9-4.0(4.0) 3.7-2.4(3.9) 2.7(2.7) _10.4-11.3 (10.8) _2.7-3.5 (3.1) _7.9-9.1 (8.5) : R. nabire sp. nov. Male holotype cml 4.5 4.4 pal 12.8 4.0 11.1 - Males (n = 3) 2.8-3.1 (3.0) 4.3-4.7(4.6) 4.1-4.4(4.3) 3.1-3.4(3.3) 12.8(12.8) 3.8-4.0(3.9) 11.0-11.1 (11.1) ; Female paratype 3.0 4.6 4.4 2.8 15.2 3.1 11.5 13.6 R. tariku sp. nov. Male holotype 2.7 4.0 4.3 2.8 12.1 3.1 9.8 - Males (n = 4) 2.6-3.0 (2.7) 4.0-4.4 (4.2) 4.2-4.7 (4.4) 2.3-2.8(2.6) 11.8-12.6 (12.2) 3.1-3.8 (3.6) 9.8-10.7 (10.2) : Female (n = 2) 2.8-3.2 (3.0) 4.0-4.7(4.4) 3.5 (3.5) 2.7(2.7) 12.2-13.2(12.7) 3.8 (3.8) 10.8-11.9 (11.4) _9.5-10.8 (10.2) R. albatros sp. nov. Male holotype 2.9 4.7 5.8 3.5 15.7 4.6 13.0 - Males (n = 3) 2.9-3.1 (3.0) 4.5-4.8 (4.7) 5.2-5.8(5.5) 3.0-3.5 (3.3) 14.8-15.7(15.4) 4.6-4.7 (4.6) 12.5-13.7 (13.0) s Female paratype 3.1 4.8 4.7 2.8 14.2 4.6 12.9 19.0 R. baduri sp. nov. Female holotype Si Bee 3.5 2.5 13.0 4.5 10.4 13.4 R. karimui sp. nov. Male holotype 2.0 3.6 3.7 3.0 9.5 3.3 8.0 - Male paratype QL. 3.9 4.2 2.9 8.8 229 8.1 : Female paratype 2.3 3.2 2.4 2.0 7.7 3.0 7.8 8.0 lic parameres stout, weakly curved, apex strongly sclerotized and rounded. From the sympatric species R. tariku sp. nov., the new species differs by larger size and by male and female genitalia. Description. —Average sized among congeners (Fig. 12). Dorsum of head with broad red brown bands narrowly separated (Fig. 4B). Fastigium red brown (Fig. 4B). Scapes dark brown. Fastigium verti- cis and frons dark brown, frons without spots; clypeus and mouth- parts dark brown, labrum yellow brown (Fig. 5B). Pronotal disk dark red brown, lateral parts yellow brown mottled with red brown patterns (Fig. 4B). Lateral lobes of pronotum dark brown, distinct- ly darker than disk (Fig. 6B). FIs and FIIs yellow brown with dense dark spots and patterns; Tls and TlIls dark brown with a cream- colored spot or incomplete ring in middle. FIIs brown, knees dark brown to black. Tergites brown, with posterior margin darker. Male. FWs reaching middle of third abdominal tergite. FW color- ation (Fig. 7B): Dorsal field cells and veins mostly brown; with area between M and R infumate cream-colored; basal area with a me- dium cream-colored spot on external corner. Lateral field brown in dorsal half, gray brown in ventral half. FW venation typical of genus; 1A angle broken by a flat segment; oblique vein posterior branch almost straight. Apical field very small, with only one straight cell alignment posterior to D alignment. Apex of dorsal field rounded. Male genitalia: (Figs 8B, 11C, 11D) Pseudepiphallus very slender and elongate, very sclerotized; slightly concave in lateral view, raised dorsally at base and apex; its basal margin almost straight, with a dorsal sclerotized plate; widened laterally, forming wide shoulders carrying bases of rami; lateral margins substraight; posterior part of pseudepiphallus elongate, forming a flat narrow spoon slightly con- cave, with two thin dorsal pre-apical expansions (Fig. 11C); posterior apex somewhat truncated, without lophi. Rami very short, way short- er than half the pseudepiphallus length. Pseudepiphallic parameres stout, weakly curved, their apex strongly sclerotized and rounded. Endophallic sclerite narrow, Y-shaped, with anterior region short and not reaching anterior margin of pseudepiphallic sclerite, with lateral arms elongated and slender, without median posterior expansion. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) 24 Female. FWs slightly surpassing second tergite. Dorsal field with a cream-colored oval spot at base without clearly defined margin (Fig. 9B). Female genitalia: Ovipositor distinctly shorter than FIII. Copu- latory papilla globular, its basal part with an irregular sclerotiza- tion forming a basal rim, with a large, rounded apex, mostly mem- branous and curved ventrally (Fig. 10B). Measurements.—See Table 1. Rugabinthus yayukae sp. nov. http://zoobank.org/F2E2AF2E-A052-4128-A8F2-2E050707A859 (Fies:324G65G/6C.7C28C. TLE, LIES) Material examined.—Holotype: INDONESIA ¢ <@; Mountain slope above Bernhard Camp, 750 m; 19 March 1939; L. J. Toxopeus leg.; MZB-ORTH10791. Paratypes: INDONESIA ¢ 13; Mountain slope above Bernhard Camp, 100 m; 8? April 1939; L. J. Toxopeus leg.; mo- lecular sample L178; MZB-ORHT9414 e 14; Araucaria Camp 800 m; 3 March 1939; L. J. Toxopeus leg. (MZB-ORHT 9417); molecular sam- ple L212; MNHN-EO-ENSIF1728 ¢ 13); N. Guinea, Bor?, ca. 400 m2; 8 April 1911; Dr P. N. Kampen Ned N.W Guinea Exp. leg.; RMNH. Type locality -INDONESIA: West Papua, Bernhard Camp. Etymology.—The species is dedicated to Prof. Yayuk R. Suhardjono from the Zoological Museum, Cibinong Science Centre in Jakarta- Bogor (Indonesia). M.K. TAN AND T. ROBILLARD Diagnosis.—This new species differs from all congeners by male genitalia, with characteristic shape of pseudepiphallus, its apical part tapering into a subacute apex and pseudepiphallic parameres including a median inner process and a triangular apex. The new species is close to R. faowi sp. nov. by general shape of male genita- lia and shape of endophallic sclerite but differs by apex of pseude- piphallus and pseudepiphallic parameres. Description.—Average to large sized among congeners (Fig. 13). Dorsum of head with broad red brown bands barely separated (Fig. 4C). Fastigium red brown (Fig. 4C). Scapes dark brown. An- tennae yellow brown basally, distally brown with yellow rings. Fastigium verticis red brown with a faint yellow brown T-shaped pattern; frons brown without spots; clypeus and mouthparts brown (Fig. 5C). Pronotal disk red brown with very few irregular yellow brown patterns, with a yellow brown stripe near the late- ro-posterior margin (Fig. 4C). Lateral lobe of pronotum slightly darker than disk (Fig. 6C). FIs and FIIs brown with few spots; TIs and TIls dark brown with one yellow brown ring in middle. FIIs brown, knees dark brown to black. Tergites brown, with posterior margin darker. Male. FWs reaching middle of fourth abdominal tergite. FW coloration (Fig. 7C): Dorsal field cells and veins mostly brown, with a faint spot on external corner. Lateral field red brown, with ventral margin cream-colored. FW venation typical of genus, 1A slightly bisinuate anterior to angle. Harp longer than wide; poste- rior branch of oblique vein almost straight and transverse. Mirror (d1) slightly differentiated. Apical field with three parallel cell align- Fig. 13. R. yayukae sp. nov. male habitus in dorsal (A) and lateral (B) views. Scale bar: 10 mm. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) M.K. TAN AND T. ROBILLARD ments (E-F-G) posterior to mirror. Apex of dorsal field obliquely rounded. Lateral field with projections of Sc slightly sinuous. Male genitalia: (Figs 8C, 11E, 11F) Pseudepiphallus elongate, not convex dorsally, its basal margin straight, slightly widened lat- erally at base of rami and raised dorsally; lateral margin at basal half parallel and wider, tapering in middle into apical half; api- cal half faintly tapering then straightening at apex; posterior apex truncated and faintly indented in the middle. Rami very short, much shorter than half the pseudepiphallus length. Pseudepiphal- lic parameres C-shaped, with inner lobe in middle with apical half triangular, tapering into a subacute apex; inner lobe straight, perpendicular to paramere. Ectophallic apodemes parallel and long, usually reaching beyond anterior margin of pseudepiphallic sclerite. Endophallic sclerite with anterior region elongate, almost reaching anterior margin of pseudepiphallic sclerite; posterior part with short triangular lateral arms, without median expansion. Female. Unknown. Measurements.—See Table 1. Rugabinthus faowi sp. nov. http://zoobank.org/40095825-464D-47BA-8171-19AE6950E3D9 (Figs 3, 4D, 5D, 6D, 7D, 8D, 9C, 10C, 14, 15A, 15B) Material examined.—Holotype: INDONESIA ¢ 3; West Papua, Fawi [Faowi] village in upper part of Tariku River (tributary of Mamberamo River), partly low-lying forest and partly forest on hills; 29 January-17 February 2012; A. Gorochov leg.; molecular sample L92; ZIN. Paratypes: INDONESIA ¢ 19; same information C 25 as holotype; ZIN ¢ 44, 62; same information as holotype; ZIN 14, 12; same information as holotype; MZB e 14’, 19; same infor- mation as holotype; MNHN-EO-ENSIF11144-ENSIF11145. Type locality. INDONESIA: West Papua: Faowi. Etymology.—This species is named after the type locality Faowi; noun in apposition. Even though three species of Rugabinthus are found in Faowi, this species was the most abundantly collected. Diagnosis.—This new species is close to R. manokwari sp. nov., R. kencana sp. nov., and R. maoke sp. nov. in terms of male FW ve- nation with 1A not indented, and by male genitalia with pseude- piphallus triangular. This new species differs from R. manokwari sp. nov. by larger size, darker coloration, apex of pseudepiphallus not indented, and slight difference in shape of pseudepiphallic parameres. This new species differs from R. kencana sp. nov. and R. maoke sp. nov. by lateral margins of pseudepiphallus more in- dented in the middle and shape of pseudepiphallic parameres. Description.—Large sized among congeners (Fig. 14). Dorsum of head with broad red brown bands narrowly separated (Fig. 4D). Fastigium red brown (Fig. 4D). Scapes red brown. Fastigium ver- ticis and frons brown, frons with very faint light spots; clypeus and mouthparts brown, labrum distinctly cream-colored (Fig. 5D). Pronotal disk red brown with some irregular yellow brown patterns laterally and near the anterior margin, with lighter lat- eral ovular spots near anterior half and with a yellow brown stripe near the latero-posterior margin (Fig. 4D). Lateral lobes not darker D' & : , Be ‘ Ce, Fig. 14. R. faowi sp. nov. male (A, B) and female (C, D) habitus in dorsal (A, C) and lateral (B, D) views. Scale bar: 10 mm. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) 26 M.K. TAN AND T. ROBILLARD Fig. 15. Male genitalia ventral (A, C, E, G) and lateral (B, D, F, H) views: A, B. R. faowi sp. nov.; C, D. R. kencana sp. nov.; E, F. R. ma- nokwari sp. nov.; G, H. R. maoke sp. nov. Scale bar: 1 mm. than disk (Fig. 6D). FIs and FIs yellow brown with dense dark Male. FWs reaching middle of fourth abdominal tergite. FW spots and patterns (less dense in inner surface of FIs); TIs dark coloration (Fig. 7D): Dorsal field cells and veins mostly brown; brown with a broad cream-colored ring in middle, THs with one with area between M and R infumate cream-colored; basal area or two incomplete rings. FIIIs brown, knees dark brown to black. with a cream-colored spot on external corner. Lateral field with Tergites brown, with posterior margin darker. brown veins, cells with dorsal part brown, otherwise cream-color- JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) M.K. TAN AND T. ROBILLARD ed. FW venation typical of genus; 1A forming a right angle, slightly bisinuate anterior to angle; oblique vein posterior branch almost straight, anterior one bifurcated. Apical field with two cell align- ment posterior to D alignment. Apex of dorsal field rounded. Male genitalia: (Figs 8D, 15A, 15B) Pseudepiphallus triangular, slightly convex dorsally in lateral view; its basal margin straight, not indented, slightly widened laterally at base of rami; basal third much wider and lateral margins tapering to a narrower apical third; posterior third forming a thick spoon-like finger, gently ta- pering into a narrow posterior apex, truncated and without lophi. Rami short, shorter than half of pseudepiphallus length, slightly diverging anteriorly. Pseudepiphallic parameres stout, somewhat straight, and not bent or curved, with small internal lobule in middle, apex not swollen and obtuse. Ectophallic fold triangular, membranous apically, with parallel lateral sclerites. Ectophallic apodemes parallel and long, surpassing beyond anterior margin of pseudepiphallic sclerite with lateral arms elongated triangular. Endophallic sclerite with anterior region triangular, barely reach- ing anterior margin of pseudepiphallic sclerite; its posterior apex trident-like, with tongue-shaped lateral arms not surpassing large median posterior expansion. Female. FW slightly surpassing second tergite, with a small cream- colored rounded spot at base and one fainter at apex (Fig. 9C). Female genitalia: Ovipositor distinctly longer than FIII. Cop- ulatory papilla globular, its base with a semi-complete ring-like sclerotization; apex folded ventrally, short and rounded, slightly sclerotized on apical face (Fig. 10C). Measurements.—See Table 1. Rugabinthus kencana sp. nov. http://zoobank.org/DE3097D6-3COD-41 70-9657-807576C2BC19 (Figs,3,.4E; SEXOR;. 7B, SE;.9D; JOD s15Cs 15 D316) Material examined.—Holotype: INDONESIA ¢ <@; West Papua, Timika (Irian Jaya Freeport Concession Timika); lowland forest, 27 Kuala Kencana nr. sewage plant; 4°26.21'S, 136°51.84'W, 100 m; 17-24 March 1997; S. Miller, Peggie, Yaku, Ubaidillah leg.; mo- lecular sample L46; MZB-ORHT97020. Paratypes: INDONE- SIA ¢ 192; West Papua, Timika (Irian Jaya Freeport Concession Timika); lowland rainforest, Kuala Kencana Light Ind. Park 4°26.21'S 136°52.59'W, 100 m, Malaise trap 1 (site 5); (MZB 97024); MNHN-EO-ENSIF3552 @ 14; West Papua, Freeport MT2 plot #5; 12-25 March 1997; molecular sample L11LFRE1; (MZ- BORTH97025-5); MNHN-EO-ENSIF1760. Type locality.—_INDONESIA: West Papua: Kuala Kencana. Etymology.—The species is named after the type locality: Kuala Kencana; noun in apposition. Diagnosis.—This new species differs from all congeners by its large size, the large triangular shape of male pseudepiphalllus, and stout pseudepiphallic parameres, gently curved but not bent in middle, apex swollen and bilobate. The new species is very similar to R. maoke sp. nov. but differs by overall smaller size, male FW venation with two cell alignments in apical field (instead of one in R. maoke sp. nov.), shape of oblique vein, and male genitalia with distinctly smaller pseudepiphallus and endophallic sclerite shorter anteriorly (forming a Y-shape in R. maoke sp. nov.), with lateral arms pointing more posteriorly. The male genitalia are also similar to R. manokwari sp. nov. but differ by the absence of in- dentation at base of the apical third of lateral margin and shape of pseudepiphallic parameres. Description.—Large sized among congeners (Fig. 16). Dorsum of head with broad red brown bands faintly to narrowly separated (Fig. 4E). Fastigium red brown (Fig. 4E). Scapes dark brown, pos- terior part yellow with a dark spot. Fastigium verticis and frons black, and black beneath scapes; clypeus and mouthparts pale brown with darker patterns (Fig. 5E). Pronotal disk red brown with some faint and irregular paler brown patterns (Fig. 4E). Fig. 16. R. kencana sp. nov. male (A, B) and female (C, D) habitus in dorsal (A, C) and lateral (B, D) views. Scale bar: 10 mm. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) 28 Lateral lobes slightly darker than disk, dark brown except a yel- low spot on ventral margin (Fig. 6E). FIs and FIIs brown with a few dark spots near knees, TIs and TIIs dark with pale yellow rings. FIIs brown, knees dark brown. Tergites brown, with poste- rior margin darker. Male. FWs reaching apex of fourth abdominal tergite. FW coloration (Fig. 7E): Dorsal field cells and veins mostly brown; with area between M and R dark brown; basal area with a small cream-colored spot on external corner. Lateral field red brown, with ventral region more yellow brown. FW venation typical of genus; 1A forming a right angle, without notch anterior to angle; oblique vein bifurcated, posterior branch almost straight. Cells of D alignment homogeneous. Apex of dorsal field obliquely round- ed. Apical field with two cell alignments. Male genitalia: (Figs 8E, 15C, 15D) Pseudepiphallus trian- gular, its basal margin straight, not indented; basal third much wider, lateral margin tapering to a narrower apical third; apical third tapering into a narrow posterior apex, not narrowed pre- apically, apex subacute, without lophi. Rami short, slightly shorter than half of pseudepiphallus length, slightly diverging anteriorly. Pseudepiphallic parameres stout, gently curved but not bent in middle, apex swollen and bilobate. Ectophallic apodemes parallel and long, widened apically, surpassing beyond anterior margin of pseudepiphallic sclerite. Endophallic sclerite with anterior region triangular, barely reaching anterior margin of pseudepiphallic M.K. TAN AND T. ROBILLARD sclerite; its posterior apex trident-like, with short lateral arms not surpassing median posterior expansion. Female. FW faintly surpassing apex of third tergite, homogene- ously brown, without a basal spot (Fig. 9D). Female genitalia: Ovipositor slightly longer than FIII. Copula- tory papilla very small, somewhat rectangular, its basal part with sclerotization forming a basal rim, apex almost as wide as base, ended by a narrow stout and truncated lobule (Fig. 10D). Measurements.—See Table 1. Rugabinthus manokwari sp. nov. http://zoobank.org/AB112D2B-497F-4577-A9EE-42160922A12C (Fiss"3, 4h. oF OR e/E8E SE AOE SE, 15k 17) Material examined.—Holotype: INDONESIA e¢ 3; West Papua, Manokwari, Vogelkop, 75 m; 24 July 1957; D. Elmo Hardy leg,; BPBM. Paratype: INDONESIA e 19; West Papua, Manokwari town, primary forest on hills near sea; 4-6 November 2004; A. Gorochov leg.; molecular sample L88; ZIN. Type locality INDONESIA: West Papua: Manokwari Etymology.—The species is named after the type locality Manok- wari; noun in apposition. Fig. 17. R. manokwari sp. nov. male (A, B) and female (C, D) habitus in dorsal (A, C) and lateral (B, D) views. Scale bar: 10 mm. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) M.K. TAN AND T. ROBILLARD Diagnosis.—This new species differs from all congeners by lighter coloration, including pronotal disk yellow brown with sparse red brown patterns, and shape of male genitalia. Among species group, R. manokwari sp. nov. differs by male genitalia with pseudepiphal- lus indented dorsally at base of apical region and tapering to a narrower apex. This species has similar pale coloration as R. biakis sp. nov. but differs by absence of longitudinal median stripe in the pronotal disk and by shape of male genitalia. Description.—Size average for the genus (Fig. 17). Dorsum of head yellow brown with five well-separated red brown longitudinal bands, middle one corresponding to incomplete fusion of two bands (Fig. 4F). Fastigium yellow brown with red brown patterns. Scapes yellow brown with dark patterns. Antennae yellow basally, distally brown with pale yellow rings. Fastigium verticis black with a yellow cross-shaped pattern; frons dark brown to black with two lateral yellow spots, black beneath scapes; clypeus and mouthparts black dorsally, pale yellow ventrally (Fig. 5F). Head lateral side red brown, genae slightly lighter with a triangular pale spot. Pronotal disk yellow brown mottled with dark brown (Fig. 4F). Lateral lobes of pronotum distinctly darker than disk, dark red brown, with a paler stripe near ventral margin (Fig. 6F). FIs and FIIs mostly cream-colored with a few well-defined brown spots, knee areas brown; TIs and TiIs dark brown with a cream- Fig. 18. R. maoke sp. nov. male habitus in dorsal (A) and lateral (B) views. Scale bar: 10 mm. 29 colored ring in middle. FIIs yellow brown, knees brown. Tergites unicolorous brown. Male. FW reaching base of fourth abdominal tergite. FW col- oration (Fig. 7F): Cells and veins of dorsal field mostly brown; area between M and R creamed-colored; basal area with a cream- colored spot on external corner. Lateral field red brown, with ven- tral region more yellow brown. FW venation typical of genus; 1A slightly bisinuate anterior to angle; oblique vein bifurcated, poste- rior branch slightly sinuous. Apex of dorsal field rounded; apical parts of its longitudinal veins very strong. Apical field reduced, including few cells of E alignment only. Male genitalia: (Figs 8F, 15E, 15F). Pseudepiphallus triangular, its basal margin slightly indented in the middle, basal third much wider, lateral margins indented and tapering to a narrower apical third, elongate as characteristic of the species group. Apical third narrowed in dorsal view, slightly concave dorsally, tapering into a Narrow posterior apex; slightly truncated and indented apically. Rami short, slightly shorter than half of pseudepiphallus length, slightly diverging anteriorly. Pseudepiphallic parameres stout, not bent in middle, apical half finger-like with subacute apex. Ec- tophallic apodemes parallel and long, surpassing beyond anterior margin of pseudepiphallic sclerite. Endophallic sclerite with ante- rior region short and barely reaching anterior margin of pseude- piphallic sclerite, with posterior lateral arms tongue-shaped and slightly longer than median expansion. Female. FW reaching middle of second tergite, with an apical indistinct cream-colored spot (Fig. 9E). Female genitalia: Ovipositor shorter than FIII. Copulatory pa- pilla conical, smaller and stout; apex folded ventrally, short, point- ed; dorsal face with a sclerotized area; ventro-anterior end forming an oval to pyriform rim (Fig. 10E). Measurements.—See Table 1. Rugabinthus maoke sp. nov. http://zoobank.org/DA4BAC81-EDD 1-4F86-921B-4AC602F47257 (Figs 3, 4G, 5G, 6G, 7G, 8G, 15G, 15H, 18) Material examined.—Holotype: INDONESIA ¢ 3; West Papua, Star Range; 25 m; 10 September 1959; Neth. New Guinea Exped 1959 leg.; RMNH. Paratype: INDONESIA ¢ 1; West Papua, Star Range; 25 m; 10 September 1959; Neth. New Guinea Exped 1959 leg; molecular sample L179; MNHN-EO-ENSIF1757. Type locality. INDONESIA: West Papua: Star Range Etymology.—This species is named after the Maoke Mountain Range where Star Range was situated within; noun in apposition. Diagnosis.—This new species is very similar to R. kencana but dif- fers by overall larger size, distinctly larger pseudepiphallus, and endophallic sclerite with lateral arms more elongated and point- ing more externally. Description.—Large sized among congeners (Fig. 18). Dorsum of head with broad red brown bands faintly to narrowly separated (Fig. 4G). Fastigium red brown (Fig. 4G). Scapes dark red brown. Fastigium verticis and frons dark red brown, black beneath scapes; clypeus and mouthparts pale brown with darker patterns (Fig. 5G). Pronotal disk red brown with some faint and irregular paler brown patterns (Fig. 4G). Lateral lobes faintly darker than disk, dark brown except for a yellow spot on ventral margin (Fig. 6G). JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) 30 FIs and FIIs yellow brown with a few dark spots near knees, TIs dark with a pale yellow ring in middle, THs mostly dark. FIIs brown, knees dark brown. Tergites brown, with posterior margin darker. Male. FWs reaching base of fourth abdominal tergite. FW col- oration (Fig. 7G): Dorsal field cells and veins mostly brown; with area between M and R dark brown; basal area with a small cream- colored spot on external corner. Lateral field red brown, with ven- tral region more yellow brown. FW venation typical of genus; 1A forming a right angle, without notch anterior to angle; oblique vein bifurcated, posterior part sinuated. Cells of D alignment wid- ened posteriorly. Apex of dorsal field obliquely rounded. Male genitalia: (Figs 8G, 15G, 15H) Pseudepiphallus very simi- lar to R. kencana, triangular with an indented, narrow apical third with subacute apex, but differs by being distinctly larger. Rami, pseudepiphallic parameres and ectophallic apodemes similar. En- dophallic sclerite with anterior region triangular, not reaching an- terior margin of pseudepiphallic sclerite, with long posterior lat- eral arms pointing diagonally and surpassing median expansion. Female. Unknown. Measurements.—See Table 1. Rugabinthus biakis sp. nov. http://zoobank.org/D074F7D3-D6C5-463C-8D3E-4EFD304AD4C9 (Fies 3, 4H. 5H, GH;-7H-8H, 19;20A;°20B) Material examined.—Holotype: INDONESIA ¢ <@; West Papua, Biak Island (not far from N. coast of W New Guinea), secondary forest on hill near vill. Corem; 14-15 November 2004; A. Goro- chov leg.; ZIN. Paratypes: INDONESIA ¢ 14; same information as holotype; molecular sample L86; MZB ¢ 10; Biak Island (not far from N. coast of W New Guinea), secondary forest on hill near Fig. 19. R. biakis sp. nov. male habitus in dorsal (A) and lateral (B) views. Scale bar: 10 mm. M.K. TAN AND T. ROBILLARD vill. Corem; 14-15 November 2004; A. Gorochov leg.; MNHN- EO-ENSIF1745. Other material—INDONESIA e 1 juvenile; West Papua, Biak Is- land, Biak Town, forest on not high hill near airport; 17-20 Janu- ary 2012; leg. A. Gorochov; molecular sample L85; ZIN. Type locality -INDONESIA: West Papua: Biak. Etymology.—The species is named after the type locality: Biak Island. Diagnosis.—This new species differs from all congeners by its small size; dorsum of head yellow brown with five well-separated and well- defined dark brown longitudinal bands; pronotal disk yellow brown with a median dark brown longitudinal band and sparse but well defined dark brown spots of different sizes; lateral lobe contrasting dark in coloration. It also differs from all congeners by male genitalia with pseudepiphallus rectangular, stouter than congeners, apex trun- cated forming a short posterior plate with a small median nodule. Description.—Small sized among congeners (Fig. 19). Dorsum of head yellow brown with five well-separated and well-defined dark brown longitudinal bands, middle band widest resulting from fu- sion of two bands, partly split anteriorly (Fig. 4H). Fastigium yellow brown with dark brown patterns. Scapes brown. Antennae yellow brown basally, distally brown with yellow rings. Fastigium verticis and frons black, frons with two faint yellow brown spots, eyes un- derlined with yellow; clypeus and mouthparts dark brown to black, labrum distinctly cream-colored (Fig. 5H). Maxillary palpi cream- colored with brown patterns. Pronotal disk yellow brown with a median dark brown longitudinal band and sparse but well defined dark brown spots of different size (Fig. 4H). Lateral lobes of pro- notum entirely dark brown, distinctly darker than disk (Fig. 6H). Legs pale yellow brown with well-defined brown spots and rings. FIs and FIIs mostly cream-colored with a few well-defined brown spots, knees brown; TIs and THs dark brown with a cream-colored ring in middle. FHIs cream-colored with numerous oblique dark brown bands, knees dark brown. Tergites unicolorous dark brown. Male. FWs reaching middle of fourth abdominal tergite. FW coloration (Fig. 7H): Dorsal field cells and veins mostly brown; some veins near basal area cream-colored; M vein orange brown; area between M and R infumate cream-colored; basal area with a large cream-colored spot on anterior corner and another one near 1A and 2A. Lateral field dark red brown in dorsal half, gray brown in ventral half. FW venation typical of genus, 1A bisinuate anterior to angle; oblique vein base faded, both its posterior and anterior branches almost straight. Apex of dorsal field obliquely rounded. Male genitalia: (Figs 8H, 20A, 20B) Pseudepiphallus rectan- gular, stouter, and shorter than in congeners, very wide laterally at base of rami, its basal anterior margin almost straight, lateral margin generally parallel, apex truncated, forming a short pos- terior plate, slightly concave dorsally, with a small median nod- ule. Rami short, but longer than half of pseudepiphallus length. Pseudepiphallic parameres strongly bent 90° in the middle, apex strongly sclerotized and forming two stout lobules. Ectophallic apodemes surpassing anterior margin of pseudepiphallic scler- ite. Endophallic sclerite short, barely reaching anterior margin of pseudepiphallus, with thin posterior lateral arms and a very small medio-posterior expansion. Female. Unknown. Measurements.—See Table 1. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) M.K. TAN AND T. ROBILLARD 3] Fig. 20. Male genitalia ventral (A, C, E) and lateral (B, D, F) views: A, B. R. biakis sp. nov.; C, D. R. nabire sp. nov.; E, F. R. tariku sp. nov. Scale bar: 1 mm. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) 32 Rugabinthus nabire sp. nov. http://zoobank.org/EB99A2A4-2D8A-46A9-89B8-247023253B58 (Figs 3, 41, 51, 6L, 71, 81, 9E 10K 20C, 20D, 21) Material examined.—Holotype: INDONESIA ¢ 3; West Papua, Na- bire, S. Geelvink Bay, 5-50 m; 25 August-2 September 1962; J. Sedlacek leg.; BPBM. Paratypes: INDONESIA ¢ 19; West Papua, Nabire, S. Geelvink Bay, 0-30 m; 2-9 September 1962; J. L. Gres- sitt leg; BPBM ¢ 14; West Papua, Nabire, S. Geelvink Bay, 10- 40 m; 13 October 1962; N. Wilson leg.; molecular sample L162; BPBM e¢ 14; West Papua, Nabire, S. Geelvink Bay; 16 September 1962; Malaise trap, in jungle; H. Holtmann leg.; MNHN-EO-EN- SIF11332 ¢ 14; West Papua, Nabire, S. Geelvink Bay, 10-40 m; sweeping; 12 October 1962; N. Wilson leg.; BPBM. Type locality.—_INDONESIA: West Papua: Nabire. Etymology.—The species is named after the type locality, Nabire; noun in apposition. Diagnosis.—This new species differs from all congeners by male subgenital plate rounded apically (pointed in all congeners) and male genitalia with stout pseudepiphallus, slightly convex dor- sally, its posterior part very short, slightly concave dorsally, with apex narrowed forming two small lophi with subacute apices; basal margin of pseudepiphallus strongly indented in the mid- dle; pseudepiphallic parameres stout, posterior half forming an M.K. TAN AND T. ROBILLARD internal lobe, apex truncated. Except for the respective original characters of each species, in particular in the shape of the apical part of the pseudepiphallus, the male genitalia of the new species resemble that of R. karimui sp. nov. and R. albatros sp. nov. Description.—Average sized among congeners (Fig. 21). Dorsum of head with 6 broad dark red brown bands barely separated (Fig. 41). Fastigium dark red brown (Fig. 41). Scapes yellow brown with red brown bands. Fastigium verticis dark brown with a faint M- shaped yellow brown pattern, frons dark brown with two faint yellow brown spots; clypeus and mouthparts yellow brown with dark brown patterns (Fig. 51). Pronotal disk and lateral lobe uni- colorous dark red brown, pronotal disk with two faint lateral spots along posterior margin, lateral lobe not darker than disk (Figs 41, 61). Fls and Fils cream-colored with dark spots, TIs and TIIs dark brown with one median pale ring. FIIIs yellow brown in ventral half to red brown in dorsal half, knees dark brown. Tergites brown. Male. FWs slightly surpassing apex of fourth abdominal ter- gite. FW coloration (Fig. 71): Dorsal field cells and veins mostly brown; area between M and R whitish; basal area with a small cream-colored well-defined spot on external corner. Lateral field dark red brown, with ventral margin cream-colored. FW venation typical of genus; 1A bisinuate anteriorly to angle; oblique vein bifurcated, posterior branch almost straight. Apex of dorsal field rounded. Apical field with only few cells in E alignment. Apex of longitudinal veins in dorsal field strong. Subgenital plate short, its apex rounded. Fig. 21. R. nabire sp. nov. male (A, B) and female (C, D) habitus in dorsal (A, C) and lateral (B, D) views. Scale bar: 10 mm. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) M.K. TAN AND T. ROBILLARD Male genitalia: (Figs 81, 20C, 20D) Pseudepiphallus triangular, stouter, convex dorsally, its basal margin straight but strongly in- dented in the middle and slightly raised dorsally. Lateral margins generally parallel, tapering near apex into a narrow posterior end. Posterior part of pseudepiphallus short, concave dorsally, its apex indented in the middle, forming two small lophi with subacute apices; lophi more developed ventrally, slightly setose. Rami very short, shorter than half of pseudepiphallus length. Pseudepiphal- lic parameres stout, posterior half forming an internal lobe, apex truncated. Ectophallic apodemes surpassing anterior margin of pseudepiphallic sclerite. Endophallic sclerite with anterior region slightly surpassing anterior margin of pseudepiphallic sclerite; posterior apex of endophallic sclerite with long, rounded and di- verging arms, median expansion not distinct. Female. FW reaching apex of second tergite, brown with two clearly defined triangular, cream-colored spots laterally, at base and apex (Fig. 9F). Female genitalia: Ovipositor shorter than FIII. Copulatory pa- pilla with a large rounded apical part strongly plicate; ventro-ante- rior end forming a rounded rim (Fig. 10F). Measurements.—See Table 1. Rugabinthus tariku sp. nov. http://zoobank.org/D6961759-141B-40F6-95ED-524D310B8CE2 (Figs 3, 4J, 5J, 6J, 7J, 8J, 9G, 10G, 20E, 20F, 22) Material examined.—Holotype: INDONESIA e 3; West Papua, Fawi [Faowi] village in upper part of Tariku River (tributary of Mam- beramo River), partly low-lying forest and partly forest on hills; 29 33 January—17 February 2012; A. Gorochov leg.; molecular sample L93; ZIN. Paratypes: INDONESIA ¢ 14, 12; same information as holo- type; ZIN ¢ 14, 19; same information as holotype; MNHN-EO-EN- SIF11142-ENSIF11143 ¢ 14, same information as holotype; MZB. Type locality.—_INDONESIA: West Papua: Faowi. Etymology.—This species is named after the Tariku River; noun in apposition. This species is named after the tributary river rather than the main Mamberamo River, because it has smaller and stout- er male genitalia compared to the sympatric species R. mamberamo. Diagnosis.—This new species differs from all congeners by male FW venation with extremely indented 1A vein, with its transverse part restricted to inner half of FW, and by male genitalia with pseudepiphallus rectangular, its basal margin strongly indented in the middle, slightly widened laterally near base of rami. Pos- terior part of pseudepiphallus short and trilobate, with two short stout lophi linked by a thin sclerotized plate forming a median lobe; pseudepiphallic parameres very stout, strongly bent 90° in the middle. General shape of male genitalia similar to that of R. biakis, from which the new species differs by apex of pseude- piphallus, base of rami, and parameres; the two species also differ in body coloration. From the sympatric species R. mam- beramo, the new species differs by smaller size and by male and female genitalia. Description.—Small to average sized among congeners (Fig. 22). Dorsum of head with broad red brown bands barely separated (visible only near posterior margin) (Fig. 4J). Fastigium dark red Fig. 22. R. tariku sp. nov. male (A, B) and female (C, D) habitus in dorsal (A, C) and lateral (B, D) views. Scale bar: 10 mm. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) 34 brown (Fig. 4J). Scapes red brown. Fastigium verticis and frons dark brown, frons with faint yellow brown spots; clypeus and mouthparts dark brown, labrum yellow brown (Fig. 5J). Pronotal disk dark brown with anterior margin yellow brown and posterior margin with some faint irregular yellow brown spots (Fig. 4J). Lat- eral lobes not darker than disk (Fig. 6J). Legs pale brown with few dark brown spots and patterns. FIIIs brown, knees dark brown to black. Tergites brown, with posterior margin darker. Male. FWs reaching middle of third abdominal tergite. FW col- oration (Fig. 7J): Dorsal field cells and veins mostly brown; with area between M and R infumate cream-colored; basal area with a cream-colored spot on lateral corner. Lateral field dark brown, more gray—brown near ventral margin. FW venation typical of genus, with 1A vein strongly bisinuate, forming a big notch restricting transverse part of 1A to inner half of FWs. Oblique vein posterior branch al- most straight; anterior branch simple. Apex of dorsal field rounded. Male genitalia: (Figs 8J, 20E, 20F) Pseudepiphallus short and rectangular, its basal margin strongly indented in the middle, slightly raised dorsally, widened laterally at base of rami. Lat- eral margins parallel; posterior part of pseudepiphallus short, apex truncated, forming three lobules, including two stout lophi linked by a thin sclerotized plate forming the median lobe. Rami short, shorter than half of pseudepiphallus length. Pseudepiphal- lic parameres very stout, strongly bent 90° in the middle, apex strongly enlarged and sclerotized and forming two stout lobules. Ectophallic fold forming a wide plate apically, with thin paral- lel sclerites. Ectophallic apodemes surpassing anterior margin of pseudepiphallic sclerite, with lateral arms elongated. Endophallic sclerite very elongated anteriorly, forming a wide triangular plate carrying a narrow crest dorsally; its posterior apex with thin lateral arms but without median expansion. M.K. TAN AND T. ROBILLARD Female. FW slightly surpassing second tergite, with a distinct cream-colored rounded spot at base (Fig. 9G). Female genitalia: Ovipositor distinctly shorter than FIII. Copu- latory papilla globular, its basal part with an irregular sclerotiza- tion forming a basal rim with a short basal plate curved dorsally; apex rounded, folded ventrally and slightly sclerotized (Fig. 10G). Measurements.—See Table 1. Rugabinthus albatros sp. nov. http://zoobank.org/520A61FC-E62E-4AD0-99E3-238E844FAFA7 (Figs 3, 4K, 5K, 6K, 7K, 8K, 9H, 10H, 101, 23, 24A, 24B) Material examined.—Holotype: INDONESIA ¢ <@; West Papua, Albatros Bivak V. Mamberamo [Mamberamo River, Albatros Bivouac]; 1926; v. Leeuwen leg.; molecular sample L177; MZB- ORTH1974. Paratypes: INDONESIA ¢ 1; same information as holotype; MZB-ORTH8968 ¢ 13; same information as holotype; (MZB-ORTH8957); molecular sample L211; MNHN-EO-EN- SIF11141 ¢ 19; West Papua, Kasonaweja Village on Mamberamo River near Van Rees range, forest on not-high hills; 25-27 January 2012; A. Gorochov leg.; molecular sample L95; ZIN. Other material.—INDONESIA e 1 juvenile; same information as holotype; ZIN. Type locality —INDONESIA: West Papua: Mamberamo River, Alba- tros Bivouac. Etymology.—The species is named after the type locality: Albatros; noun in apposition. Fig. 23. R. albatros sp. nov. male (A, B) and female (C, D) habitus in dorsal (A, C) and lateral (B, D) views. Scale bar: 10 mm. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) M.K. TAN AND T. ROBILLARD 35 kar Fig. 24. Male genitalia ventral (A, C) and lateral (B, D) views: A, B. R. albatros sp. nov.; C, D. R. karimui sp. nov. Scale bar: 1 mm. Diagnosis.—This new species differs from all congeners by male genitalia with pseudepiphallus forming at the posterior end two long straight lophi with subacute apices and by female copulatory papilla with a very thin elongate apex. Description.—Average to large sized among congeners (Fig. 23). Dorsum of head with broad red brown bands narrowly separated (Fig. 4K). Fastigium red brown (Fig. 4K). Scapes yellow brown with red brown bands. Fastigium verticis brown with faint yel- low vertical stripes, frons light brown without spots in the mid- dle, dark brown ventral of scapes; clypeus and mouthparts red brown (Fig. 5K). Pronotal disk red brown with some irregular yel- low brown patterns, lighter lateral ovular spots near anterior half and with a yellow brown stripe near latero-posterior margin (Fig. 4K). Lateral lobe of pronotum unicolorous dark red brown, not distinctly darker than disk (Fig. 6K). Fis, FIIs, TIs, and TIls dark brown with yellow brown spots and rings. FIIIs brown, knees and posterior third of FIIIs dark brown. Tergites brown with posterior margin darker. Male. FWs reaching middle of fourth abdominal tergite. FW coloration (Fig. 7K): Dorsal field cells and veins mostly brown with area between M and R infumate cream-colored; basal area with a small cream-colored spot on external corner. Lateral field hyalinous brown. FW venation typical of genus; 1A notch ante- riorly to angle strong, making anterior part of harp wider; harp elongate, almost twice as long as wide. Oblique vein bifurcated, posterior branch slightly bisinuate, anterior branch bifurcated near 1A angle. Apical field with one cell alignment posterior to mirror. Apex of dorsal field rounded. Male genitalia: (Figs 8K, 24A, 24B) Pseudepiphallus triangu- lar, stouter, not convex dorsally in lateral view, its basal margin slightly indented in the middle, lateral margin generally parallel, not widened basally at base of rami, gently tapering into apex; apex forming two long straight lophi with subacute apices. Rami very short, about half the pseudepiphallus length. Pseudepiphallic parameres stout. Endophallic sclerite with a main Y shape; anteri- or region short and not reaching anterior margin of pseudepiphal- lic sclerite; posterior apex trilobate, with stick-like lateral arms and with a large median posterior expansion. Female. FWs slightly surpassing second tergite. Dorsal field with a distinct cream-colored rounded spot at base and a tiny api- cal spot (Fig. 9H). Female genitalia: Ovipositor distinctly longer than FHI. Copula- tory papilla cupular basally, circled by a narrow sclerite expanded in apical region as parallel sclerotization; apex elongate and fold- ed ventrally, terminated by a rounded area (Figs 10H, 101). Measurements.—See Table 1. Rugabinthus baduri sp. nov. http://zoobank.org/OFD67CC5-65D2-4831-82BF-81 BE5102408E (Figs 3, 4L, 5L, 6L, 91, 10J, 25) Material examined.—Holotype: INDONESIA ¢ 9; West Papua: Yapen Island, Aiam Range, Mount Baduri, Japen Seroei Camp 1, 1000 ft. a.s.1.; September 1938; B.M. 1938-593; molecular sample L172; L. E. Cheesman leg; NHMUK. Type locality —INDONESIA: West Papua: Yapen Island, Mount Baduri. Etymology.—This species is named after the type locality, Mount Baduri; noun in apposition. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) 36 M.K. TAN AND T. ROBILLARD Fig. 25. R. baduri sp. nov. female habitus in dorsal (A) and lateral (B) views; and female abdominal apex in dorsal view (C). Scale bars: 10 mm (A, B), 1 mm (C). Diagnosis.—This new species differs from all congeners by its stocky shape, frons mostly cream-colored with some dark pat- terns, scapes whitish, and by last abdominal tergite forming a me- dian sinuate expansion above suranal plate. Description.—Average sized among congeners, but stocky (Fig. 25). Dorsum of head with broad red brown bands narrowly separated (Fig. 4L). Fastigium yellow-brown mottled with brown (Fig. 4L). Scapes whitish with some faint brown patterns. Fastigium verti- cis cream-colored with two lateral rectangular dark spots beside scapes; two median oval brown spots expanded ventrad to frons as band. Frons yellow brown with brown bands widening and diverging from each other; slightly brownish beneath scapes. Cl- ypeus yellow-brown and brown in middle, mouthparts brown to dark; maxillary palpi mostly yellow (Fig. 5L). Pronotal disk brown with faint light-colored patterns laterally (Fig. 4L). Lateral lobes distinctly darker than pronotal disk, dark brown except red brown near ventral margin (Fig. 6L). FIs and FIIs yellow brown with large dark spots and patterns, knees dark-brown; TIs and TIls brown with very faint pale bands. FIIs brown, knees dark brown to black. Male. Unknown. Female. Last abdominal tergite forming a median sinuate ex- pansion partly covering suranal plate (Fig. 25C). FW slightly sur- passing second tergite, not clearly overlapping. Dorsal field with a very faint cream-colored rounded spot at base (Fig. 91). Female genitalia. Ovipositor about as long as FIIs. Copulatory papilla globular, its base with two hemi-circular ring-like sclerites; apex short, rounded with tip slightly pointed, folded ventrally, and slightly sclerotized dorsally (Fig. 10J). Measurements.—See Table 1. Remark.—This new species is known only by the female type specimen, which makes it difficult to place it in a particular genus among the Lebinthina. Its stocky shape first suggested it could be- long to Gnominthus, but a molecular phylogenetic study in prepa- ration revealed that this species belongs to the clade correspond- ing to Rugabinthus without ambiguity. Rugabinthus karimui sp. nov. http://zoobank.org/D299D902-F486-4B1B-8E75-D3E52967B681 (Figs 3, 4M, 5M, 6M, 7L, 8L, 9J, 10K, 10L, 24C, 24D, 26) Material examined.—Holotype: PAPUA NEW GUINEA e <4; Ka- rimui, 1000 m; 2-3 June 1961; J. L. Gressitt leg.; BPBM. Paratypes: PAPUA NEW GUINEA e 19; Karimui, South of Goroka, 1000 m; 3 June 1961; G. L. and M. Gressitt leg.; molecular sample L210; BPBM e 16; Karimui; 4 June 1961; J. L. Gressitt leg.; malaise trap; molecular sample L49; (BPBM); MNHN-EO-ENSIF11333. Type locality.—PAPUA NEW GUINEA: Karimui. Etymology.—The species is named after the type locality: Karimui; noun in apposition. Diagnosis.—This new species differs from all congeners by smaller size, short male FWs without apical field, and shape of male and female genitalia. Male subgenital plate more elongate than in con- geners, with a short apical expansion. Male genitalia very differ- ent from all congeners, characterized by elongate apical part of pseudepiphallus forming a long spoon-like finger. Description.—Small sized among congeners (Fig. 26). Dorsum of head yellow brown with 6 narrow red brown irregular longitu- dinal bands (Fig. 4M). Fastigium unicolorous red brown (Fig. 4M). Scapes yellow brown. Antennae basally yellow brown, dis- tally dark brown with some light rings. Fastigium verticis brown with a pale yellow-brown n-shaped pattern; frons brown in mid- dle with 2 small yellow spots, black beneath scapes; clypeus and mouthparts dark brown dorsally, labrum and ventral part of mouthparts yellow (Fig. 5M). Pronotal disk yellow brown with some irregular red-brown patterns most prominent near anterior and posterior margins (Fig. 4M). Lateral lobe of pronotum red brown, distinctly darker than disk, with 2 light spots in antero- ventral corner (Fig. 6M). Legs yellow brown with dark brown spots. FUIs brown, knees dark brown. Tergites brown, with pos- terior margin darker. JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) M.K. TAN AND T. ROBILLARD 37 Fig. 26. R. karimui sp. nov. male (A, B) and female (D, E) habitus in dorsal (A, D) and lateral (B, E) views; and male subgenital plate in ventral view (C). Scale bars: 10 mm (all except C); 1 mm (C). Male. FW very short, reaching apex of third abdominal ter- gite. FW coloration (Fig. 7L): Dorsal field cells and veins mostly brown; some veins near basal area cream-colored; with area between M and R infumate cream-colored; basal FW area with a large cream-colored spot including base of CuA, 1A and 2A. Lateral field brown, with ventral margin cream-colored. FW venation typical of genus; 1A notch anteriorly to angle strong; oblique vein bifurcated, posterior and anterior branches slightly bisinuate and almost parallel. Apex of dorsal field very short and rounded; D alignment limited to 2 cells; apical field absent. Sub- genital plate very elongate, pointed, with a short apical expan- sion (Fig. 26C). Male genitalia: (Figs 8L, 24C, 24D) Pseudepiphallus elongate, slightly convex dorsally, its basal margin slightly convex, pro- longed anteriorly by a sclerotized plate; lateral margin tapering into apical third, most narrow at apical third; posterior of api- cal third elongated spoon shaped, its apex rounded. Rami very short, way shorter than half the pseudepiphallus length. Pseude- piphallic parameres small, posterior half broadly bulbous and stout. Ectophallic apodemes parallel and long, usually reaching anterior margin of pseudepiphallic sclerite. Endophallic sclerite elongate, with anterior region long but barely reaching anterior margin of pseudepiphallic sclerite; with lateral arms of sclerite narrowly triangular. Female. FW reaching middle of second tergite; brown, without basal spot (Fig. 9J). Female genitalia: Ovipositor slightly longer than FII. Copulato- ty papilla conical, smaller and stout; apex folded ventrally, short, pointed; dorsal face with a sclerotized area; ventro-anterior end forming an oval to pyriform rim (Figs 10K, 10L). Measurements.—See Table 1. Rugabinthus newguineae (Bhowmik, 1981) comb. nov. (Figs 3, 4N, 5N, 6N, 9K, 10M, 27) Larandopsis newguineae Bhowmik, 1981[1979]: 39 - Desutter- Grandcolas and Jaiswara 2012: 31, according to Bhowmik’s JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) 38 M.K. TAN AND T. ROBILLARD descriptions and illustrations belonging to Lebinthus [but not Key to species of Rugabinthus moved to Lebinthus| - Cigliano et al. 2021 (Orthoptera Species Files Online). 1 Frons mostly whitish/cream-colored with some dark patterns; scapes Macrobinthus newguineae - Robillard et al. 2016: 178. RICUMT SEN cores FMM eecdseddDataausd serasustelsbeasds emisallsaeeshamelseasle eet aasmshl R. baduri - Frons mostly dark colored; scapes dark colored ..........ceeeeeeeeeeteees 2 Material examined.—Holotype: INDONESIA e 9; Dutch New 2. Distinctly larger, PronL > 4.5 MM uw... R. newguineae Guinea: Cyclops Mts, Sabron ‘Camip 2; 2000 ft; July 1936; identi- = .Smallét-Promh,<"4:0. mimi. ules. nade leiden 3 fied Larandopsis newguineae Bhowmik by H. K. Bhowmik, 1973; L. 3 —Dorsum of head with broad red brown bands narrowly separated. E. Cheesman leg; BMNH-1936-271. Pronotum dorsal disk generally red brown, sometimes with few Li Shrerepatteniig 5: Bets. Ae ree Bie io ee tenet ars ee ees ei a los 4 Other material—INDONESIA ¢ 19; N. New Guinea, “Boven- - — Dorsum of head yellow brown with five well-separated red brown Jemomaissin” ca. 400 m; 4 April 1911; molecular sample L199; longitudinal bands. Pronotum dorsal disk yellow brown with dark Dr P. N. v. Kampen, Ned. Nw. Guinea Exp leg; RMNH ¢ 19; N. ALLE TIIS a Mao alcds tes tech Ted, iat tde ose Cath dsc oN Mayestlo a tlh cect 12 New GuineasLolleDisPsNeveKampem, “Ned: Nw -GuineasExp leg: <4, Bron Pag wis a2... go ieiyulen da sets adenasl abel ajataes ade saatiasiaas ca tou aasteagheet 5 RMNH e 19; N. New Guinea; April-May 1911; Dr P.N.v. Kampen, - From other parts of New Guinea and not in FaoWi .........cccceeeeeeees 7. Ned. Nw. Guinea Exp leg; RMNH. 5 Pronotum dorsal disk with lateral parts entirely yellow brown or cream-colored, appearing like a lateral bands (even with brown spots Type locality -INDONESIA: Cyclops Mts. WALCHii Chis ATI Juss kor teas8 aesstos Wen, Re advend awlaaadrn la cpiaemles sige kespe Renews 6 - Pronotum dorsal disk unicolorous brown or only with sparse yellow Emended_ diagnosis.—Among congeners, R. newguineae comb. Browiispatrerns ic! Grete: Orne Nh pA ntl Galt ven eke Ltts R. tariku nov. is characterized by its large size, FWs not reaching abdomen 6 __ Face black. Labrum yellow brown. Male FWs longer, FWL > 5.5 mm, mid-length but well developed, coloration homogeneously dark reaching middle of third abdominal tergite ........0..0.... R. mamberamo brown, and very long ovipositor. - Face brown. Labrum cream-colored. Male FWs shorter, FWL < 4.5 mm reaching middle of fourth abdominal tergite............ R. faowi Description.—See Robillard et al. (2016). Z, Pseudepiphallus with apex forming two long straight lophi with sub- ACLILE~ALILGOS caret amcugectareene satel mactactewee (aeat tet ila cherte arnt! caged R. albatros Remark.—This species is known only by female specimens until - — Pseudepiphallus with apex truncated or tapering, not producing into now, which makes it difficult to place it in a genus among the Leb- twolone lophirniccee ete kth shed eae enke ieee Ei tE Er ee. 8 inthina because several genera share the same general shape, such 8 Pseudepiphalluswith-apextruncatedat 00 cei ish lice adil 9 as Rugabinthus and Macrobinthus, while being mostly characterized - Pseudepiphallus with apextapered... 2c cincateweatd chs teclo ete 10 by male characters (FW venation and genitalia). A molecular phy- 9 Male FWs shorter, < 5.0 mm. Fastigium verticis brown with two logenetic study in preparation revealed that this species belongs to vertical yellow brown stripes diverging ventrad. Pseudepiphallic the clade corresponding to Rugabinthus, justifying the new combi- parameres strongly bent in basal half (~90°), with posterior apex nation proposed here. enlarged, bean-shaped ..........cceseeeeeee R. leopoldi (Chopard, 1931) - Male FWs longer, > 5.0 mm. Fastigium verticis red brown with a faint yellow brown T-shaped pattern. Pseudepiphallic parameres with in- ner process in middle with apical half triangular................ R. yayukae 10 Male FWs longer, > 6.0 mm. Pseudepiphallus elongated, slenderer... 11 - Male FWs shorter, < 5.0 mm. Pseudepiphallus triangular, stouter htc teal tl et ate Mee SN NER SNARE BND ht SN Oe OR at Ac R. nabire 11. Smaller in size, male FUIL < 17.0 mm. Male genitalia distinctly smaller, endophallic apodeme with lateral lamellas pointing posteri- orly Distributions. Kuala Kem eands, scenes dvwwessdhescvmeyseas R. kencana - Larger in size, male FIIIL > 17.0 mm. Male genitalia distinctly larg- er, endophallic apodeme with lateral lamellae pointing diagonally more externally. Distribution: Star Range ............c eee R. maoke 12 Fastigium verticis and frons black. Labrum cream-colored .... R. biakis - Fastigium verticis black but with some yellow patterns. Labrum dark- alga 0116) C06 bene OAR Oe mS UAE ILD SENS CRE OC LOR AUR <9 JR 13 13. Smaller in size, FIL < 10 mm. Fastigium verticis with a pale yellow- biowaianshiapedspatterin,.ce. an. tel oe yuk poh cecal Re stn datea etna R. karimui - Larger in size, FIIL > 11 mm. Fastigium verticis with a (+) cross- shapedsyellow: browit Patter te. timvsaineu darwoiaessreressiuaiucrs R. manokwari Discussion The new genus Rugabinthus presented in this study includes 14 species in total, from different parts of the New Guinea Is- land (mostly in the western part) and from closely related islands Fig. 27. R. newguineae (Bhowmik, 1981) comb. nov. female habi- (Japen I., Biak I.). Except for two new combinations, most spe- tus in dorsal (A) and lateral (B) views. Scale bar: 10 mm. Modified cies belonging to the genus are new and are currently known from Robillard et al. (2016) only from a single locality. Given the low numbers of specimens JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) M.K. TAN AND T. ROBILLARD for most of these species, and given that Lebinthini are usually abundant when specifically collected, it would appear that the material studied here originated primarily from anecdotal side collections. Thus, this suggests that a large diversity of crickets from New Guinea remains undocumented; we postulate that the restricted distributions may be due to largely incomplete sam- pling on the island and that the true distribution of each species is still unknown. Because close-by localities host very different species, with one locality (Faowi) hosting up to three sympatric species, we hypothesize that the diversity of the genus is underestimated. Evidence also suggests that Rugabinthus may show a high level of endemism across New Guinea, similar to or higher than the endemism documented in New Caledonia for Agnoteccous, another genus of Lebinthina (Desutter-Grandcolas and Robillard 2005, Nattier et al. 2012). Adding Rugabinthus to the already high number of Lebinthini reported from New Guinea, in tandem with their morphological and acoustic diversity (Robillard et al. 2016, Tan et al. 2021, Tan and Robillard 2021c), suggests that this large island may have played a major role in the diversification of these crickets, as found at the scale of the genus Cardiodactylus (Dong et al. 2018). Furthermore, there is little biological data existing for spe- cies of Rugabinthus. The calls and natural history for this genus are completely unknown, since most of the materials examined here were based on old museum collections. This highlights the importance of continued survey of the vast areas of New Guinea, some parts of which remain unexplored. Emphasis should also be placed on collecting natural history data, including the recording of calls and behaviors, given the importance of Lebinthina species in the study of communication systems. Acknowledgements We thank Andrej Gorochov (ZIN) for providing samples from the collections of Saint Petersburg; Rob de Vries, Jéro6me Constant, and Carole Paleco (RBINS) for their help during the study of the collections of the Royal Belgian Institute of Natural Sciences, Brus- sels, funded by the SYNTHESYS European program (BE-TAF-6640); Caroline Pepermans and Luc Willemse (RMNH) for their help with the Leiden collections; Judith Marshall and George Beccaloni (NHMUK) for their help during the study of the Natural History Museum of London collections (funded by the SYNTHESYS Eu- ropean program GB-TAF-531); Oscar Effendi and Erni Ernawati (MZB, Indonesia) for their help during the study of Eneopterine crickets in Cibinong, Indonesia (MZB); and others for helping us in the collection of crickets and in other aspects. We also thank Si- mon Poulain for making stacked images of some of the species and Benoit Segerer for allowing the use of his in-situ photographs. This study was supported by the Biodiversity Exploration and Discovery National Geographic Grant [NGS-73188R-20]. The work of MKT was supported by the Fyssen Foundation Postdoctoral Fellowship. References Anso J, Barrabe L, Desutter-Grandcolas L, Jourdan H, Grandcolas P, Dong J, Robillard T (2016) Old Lineage on an old Island: Pixibinthus, a new cricket genus endemic to New Caledonia shed light on gryllid diversi- fication in a hotspot of biodiversity. PLoS ONE 11: e0150920. https:// doi.org/10.1371/journal.pone.0150920 Baroga JB, Yap SA, Robillard T (2016) Two new species of Eneopterinae crickets (Orthoptera: Gryllidae) from Luzon, Philippines. Zootaxa 4139: 93-105. https://doi.org/10.11646/zootaxa.4139.1.5 39 Baroga-Barbecho JB, Tan MK, Yap SA, Robillard T (2020) Taxonomic study of Lebinthus Stal, 1877 (Orthoptera: Gryllidae: Eneopterinae) with description of six new species in the Philippines. Zootaxa 4816: 401- 438. https://doi.org/10.11646/zootaxa.4816.4.1 Benavides-Lopez JL, Ter Hofstede H, Robillard T (2020) Novel system of communication in crickets originated at the same time as bat echolocation and includes male-male multimodal communication. The Science of Nature 107: 1-6. https://doi.org/10.1007/s00114-020- 1666-1 Bhowmik HK (1981[1979]) Studies on some Australo-Oriental Gryllidae (Orthoptera) in the collection of British Museum (Natural History), London. Journal of the Zoological Society of India 32: 35-49. Chopard L (1931) Résultats scientifiques du voyage aux Indes orientales neerlandaises de LL. AA. RR. le prince et la princesse Leopold de Bel- gique. Gryllidae et Gryllacridae. Mémoires du Musée royal d'histoire naturelle de Belgique 4: 1-22. Chopard L (1968) Family Gryllidae: Subfamilies Mogoplistinae, Myreco- philinae, Scleropterinae, Cachoplistinae, Pteroplistinae, Pentacen- trinae, Phalangopsinae, Trigonidiinae, Eneopterinae; Family Oecan- thidae, Gryllotalpidae. In: Beier M (Ed.) Orthopterorum Catalogus. Volume 12. Uitgeverij Dr. W. Junk N. V's, Gravenhage, 215-500. Cigliano MM, Braun H, Eades DC, Otte D (2021) Orthoptera Species File. Version 5.0/5.0 (01/01/2021). http://Orthoptera.SpeciesFile.org Desutter L (1987) Structure et évolution du complexe phallique des Gryl- lidea (Orthoptera) et classification des genres néotropicaux de Gryl- loidea. 1 partie. Annales De La Société Entomologique De France (N.S.) 23: 213-239. Desutter-Grandcolas L (2003) Phylogeny and the evolution of acous- tic communication in extant Ensifera (Insecta, Orthoptera). Zoologica Scripta 32: 525-561. https://doi.org/10.1046/j.1463- 6409.2003.00142.x Desutter-Grandcolas L, Jaiswara R (2012) Phalangopsidae crickets from the Indian Region (Orthoptera, Grylloidea), with the descriptions of new taxa, diagnoses for genera, and a key to Indian genera. Zootaxa 3444: 1-39. https://doi.org/10.11646/zootaxa.3444.1.1 Desutter-Grandcolas L, Robillard T (2005) Phylogenetic systematics and evolution of Agnotecous in New Caledonia (Orthoptera: Grylloidea, Eneopteridae). Systematic Entomology 31: 65-92. https://doi. org/10.1111/j.1365-3113.2005.00299.x Dong J, Kergoat GJ, Vicente N, Rahmadi C, Xu S, Robillard T (2018) Bio- geographic patterns and diversification dynamics of the genus Cardio- dactylus Saussure (Orthoptera, Grylloidea, Eneopterinae) in Southeast Asia. Molecular Phylogenetics and Evolution 129: 1-14. https://doi. org/10.1016/j.ympev.2018.06.001 Nattier R, Grandcolas P, Elias M, Desutter-Grandcolas L, Jourdan H, Cou- loux A, Robillard T (2012) Secondary sympatry caused by range ex- pansion informs on the dynamics of microendemism in a biodiver- sity hotspot. PLoS ONE 7: e48047. https://doi.org/10.1371/journal. pone.0048047 Robillard T (2011) Centuriarus n. gen., a new genus of Eneopterinae crick- ets from Papua (Insecta, Orthoptera, Grylloidea). Zoosystema 33: 49-60. https://doi.org/10.5252/z2011n1a2 Robillard T, Desutter-Grandcolas L (2004a) Phylogeny and the modalities of acoustic diversification in extant Eneopterinae (Insecta, Orthop- tera, Grylloidea, Eneopteridae). Cladistics 20: 271-293. https://doi. org/10.1111/j.1096-0031.2004.00025.x Robillard T, Desutter-Grandcolas L (2004b) High-frequency calling in Ene- opterinae crickets (Orthoptera, Grylloidea, Eneopteridae): an adap- tive radiation revealed by phylogenetic analysis. Biological Journal of the Linnean Society 8: 577-584. https://doi.org/10.1111/j.1095- 8312.2004.00417.x Robillard T, Desutter-Grandcolas L (2011) Evolution of calling songs as multicomponent signals in crickets (Orthoptera: Gryl- loidea: Eneopterinae). Behavior 148: 627-672. https://doi. org/10.1163/000579511X572044 Robillard T, Su YN (2018) New lineages of Lebinthini from Australia (Or- thoptera: Gryllidae: Eneopterinae). Zootaxa 4392: 241-266. https:// doi.org/10.11646/zootaxa.4392.2.2 JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1) 40 Robillard T, Dong J, Legendre F, Agauvoa S (2016) The brachypterous Leb- inthini crickets from Papua New Guinea, with description of two new genera and four new species (Orthoptera: Gryllidae: Eneopterinae). In: Robillard T, Legendre FE, Villemant C, Leponce M (Eds) Insects of Mount Wilhelm, Papua New Guinea. 109. Mémoires du Muséum na- tional d'Histoire naturelle, Paris, 149-202. Saussure H De (1878) Mélanges orthoptérologiques. Vle fascicule. Gryl- lides (2° partie). Mémoires de la Société de Physique et d'Histoire na- turelle de Genéve 25: 369-702. Su YN (2016) A simple and quick method of displaying liquid-preserved morphological structures for microphotography. Zootaxa 4208: 592- 593. https://doi.org/10.11646/zootaxa.4208.6.6 Tan MK, Japir R, Chung AYC, Robillard T (2019) Crickets of the subfamily Eneopterinae (Orthoptera: Grylloidea) from Sandakan, Sabah: one new species and calling songs of a sympatric species. Zootaxa 4619: 347-363. https://doi.org/10.11646/zootaxa.4619.2.9 Tan MK, Malem J, Legendre F, Dong J, Baroga-Barbecho JB, Yap SA, Wahab RA, Japir R, Chung AYC, Robillard T (2021) Phylogeny, systematics and evolution of calling songs of the Lebinthini crickets (Orthoptera, Gryl- loidea, Eneopterinae), with description of two new genera. Systematic Entomology 46: 1060-1087. https://doi.org/10.1111/syen.12510 M.K. TAN AND T. ROBILLARD Tan MK, Robillard T (2021a) Population divergence in the acoustic proper- ties of crickets during the COVID-19 pandemic. Ecology (The Scien- tific Naturalist) 102: e03323. https://doi.org/10.1002/ecy.3323 Tan MK, Robillard T (2021b) Highly diversified circadian rhythms in the calling activity of eneopterine crickets (Orthoptera: Grylloidea: Gryl- lidae) from Southeast Asia. Bioacoustics. https://doi.org/10.1080/09 524622.2021.1973562 Tan MK, Robillard T (2021c) New taxa of the subtribe Lebinthina (Or- thoptera: Gryllidae: Eneopterinae: Lebinthini) from New Guinea and nearby islands. Zootaxa 5082(6): 583-599. https://doi.org/10.11646/ zootaxa.5082.6.5 ter Hofstede HM, Schéneich S, Robillard T, Hedwig B (2015) Evolution of a communication system by sensory exploitation of startle be- havior. Current Biology 25: 3245-3252. https://doi.org/10.1016/j. cub.2015.10.064 Vicente NM, Olivero P, Lafond A, Dong J, Robillard T (2015) Gnominthus gen. nov., a new genus of crickets endemic to Papua New Guinea with novel acoustic and behavioural diversity (Insecta, Orthop- tera, Gryllidae, Eneopterinae). Zoologischer Anzeiger—A. Journal of Comparative Zoology 258: 82-91. https://doi.org/10.1016/j. jcz.2015.06.005 JOURNAL OF ORTHOPTERA RESEARCH 2022, 31(1)