Article. New species of the genus Lebinthus (Orthoptera, Grylloidea, Eneopterinae, Lebinthini) from Indonesia and the Solomon Islands

Zootaxa 2386: 25 48 (2010) www.mapress.com/zootaxa/ Copyright 2010 Magnolia Press Article ISSN 11755326 (print edition) ZOOTAXA ISSN 11755334 (online edition) New species of the genus Lebinthus (Orthoptera, Grylloidea, Eneopterinae, Lebinthini) from Indonesia and the Solomon Islands TONY ROBILLARD Muséum national d'histoire naturelle, Département Systématique et Evolution, UMR 7205 CNRS OSEB, CP 50 (Entomologie), 75231 Paris Cedex 05, France. Email: robillar@mnhn.fr Abstract Lebinthus crickets from Indonesia and the Solomon Islands are analysed. Six new species are described: L. ambonensis n. sp., L. buruensis n. sp., L. cyclopus n. sp., L. greensladei n. sp., L. kolombara n. sp. and L. villemantae n. sp., and one is redescribed (L. truncatipennis Chopard, 1929). Descriptions focus on male and female genitalia and forewing venation. Data are provided for L. villemantae n. sp. fine scaled habitat, behaviour and song repertoire. Key words: Orthoptera, Eneopterinae, Lebinthini, Lebinthus, new species, Indonesia, Solomon Islands Introduction The cricket subfamily Eneopterinae Saussure, 1874 (Grylloidea) is particularly diversified in the Pacific region and in SouthEast Asia (DesutterGrandcolas & Robillard 2006, Otte 2007), where they are represented by two tribes, the Nisitrini Robillard, 2004, and mostly by the Lebinthini Robillard, 2004 (Robillard & DesutterGrandcolas 2004a, 2006, 2008). The Lebinthini are responsible for an important part of the acoustic diversity of Eneopterinae, with originalities laying mainly in the spectral properties of their songs and the use of high frequencies for communication (Robillard & DesutterGrandcolas 2004b; Robillard et al. 2007; Robillard 2009). In this study six new species of Lebinthus are described from different islands of Indonesia and the Solomon Islands and one species is redescribed. Descriptions focus on male and female genitalia, and forewing venation. Although other species of the genus are known for their dense populations (Robillard 2009), most of the present descriptions are based on few specimens only from several museum collections, without information about species habitat, life history and behaviour. However, based on recent fieldwork in South Sulawesi, more data could be obtained for L. villemantae n. sp. about the species finescale habitat, behaviour, and song repertoire. Material and methods Material. Fieldwork was made in several localities in Sulawesi Selatan, Indonesia. Specimens were collected by sight only, by night and day, in order to observe their habitat and type of activity. Newly collected specimens are deposited in the collections of the Museum Zoologicum Bogoriense, Bogor (MZB) and of the Muséum national d Histoire naturelle, Paris (MNHN). For museum specimens, political districts and islands are mentioned for every specimen within lists of material, and square brackets are used for additional information not mentioned on specimen labels. Male tegminal venation. Male tegminal veins and cells (Fig. 1) are named after DesutterGrandcolas (2003) for Ensifera, and Robillard & DesutterGrandcolas (2004a) for the subfamily Eneopterinae more specifically. Accepted by D. Rentz: 24 Dec. 2009; published: 2 Mar. 2010 25

Male and female genitalia. Male and female genitalia have been dissected in softened specimens by cutting the membranes between the paraprocts and the subgenital plate, or between the ovipositor and the subgenital plate respectively; they have been observed after cleaning with cold KOH using a binocular microscope Leica MZ16 at magnifications up to 160, and then kept in glycerine in vials pinned under specimens. Male genitalia are named according to Desutter (1987), modified in DesutterGrandcolas (2003) and Robillard & DesutterGrandcolas (2004a). Dotted parts in figures correspond to membranous areas. Abbreviations: see below. Acoustic data. The basic cricket song terminology follows Ragge & Reynolds (1998). One song unit is called a syllable and corresponds to one openingclosure cycle of the male forewings; a repeated group of syllables is called an echeme and corresponds to the calling unit of a species; a series of echemes constitutes an echemesequence. The species Lebinthus villemantae n. sp. has been studied in the field and subsequently in the laboratory. The recordings were made with a modified Condenser Microphone Capsule CM16 (Avisoft Bioacoustics, Berlin), with a flat frequency response from 3 to 150 khz (R. Specht pers. comm.), connected to a TASCAM HDP2 digital recorder (96 khz sampling frequency, 16 bit). Acoustic analyses were performed using the computer software AvisoftSASLab Pro version 4.40 (Specht 2008). Song features were measured using the automatic commands under AvisoftSASLab Pro. Mean values and standard deviation per individual and per species were calculated after adjustment for a common recording temperature of 20 C. All recording files have been deposited in the Sound Library of the Muséum national d Histoire naturelle, Paris. Abbreviations. Descriptions. General morphology: FW, forewing; Tarsomere III1: basal segment of hind leg tarsomere. Male genitalia: ec arc, ectophallic arc; ec ap, ectophallic apodeme; ec f, ectophallic fold; en ap, endophallic apodeme; en s, endophallic sclerite; ps l, pseudepiphallic lophi; ps p, pseudepiphallic paramere; ps r, pseudephiphallic dorsal ridges; r, rami. Tegminal venation: 1A4A, first to fourth anal veins; CuA, anterior cubitus; CuA1, CuA2,... first, second,... bifurcations of CuA; CuP, posterior cubitus; MA, MP, anterior, posterior media veins; R, radial vein; c13, first to third cells of C alignment; d1 cell (mirror), first cell(s) of D alignment; d2, second cell of D alignment; ha, harp area; ra, rounded area in harp. Institutions. BMNH: Natural History Museum, London, UK (formerly British Museum of Natural History). MNHN: Muséum national d'histoire naturelle, Paris, France. MZB: Museum Zoologicum Bogoriense, Bogor, Indonesia. RMNH: Nationaal Natuurhistorisch Museum Leiden, The Netherlands (formerly Rijksmuseum van Natuurlijke Historie),. ZRC: Raffles Museum of Biodiversity Research Entomology, Singapore. Systematic part Genus Lebinthus Stål, 1877 Type species: Lebinthus bitaeniatus Stål, 1877 Diagnosis. Among Lebinthini genera, Lebinthus is more closely related to Agnotecous Saussure, 1878, to which it resembles by microptery and FW venation. It is characterised by its rather smaller size, microptery in both sexes (FW short and hind wings absent), and male FW venation with mirror almost not differentiated from apical field, dorsal field as long or longer than lateral field (it is shorter in Agnotecous), median fold short, triangular and located on dorsum. List of Lebinthus species addressed in this study L. ambonensis n. sp. L. buruensis n. sp. L. cyclopus n. sp. 26 Zootaxa 2386 2010 Magnolia Press ROBILLARD

L. greensladei n. sp. L. kolombara n. sp. L. truncatipennis Chopard, 1929 L. villemantae n. sp. Lebinthus ambonensis n. sp. (Figs. 1A, 2, 3A, 4A, 5A C) Type material. Holotype male: Indonesia: Province Maluku: Ambon [Is.], Gunung Seliman, 3IX1995 (R. de Jong) (RMNH). Type locality. Indonesia, Ambon Island. Etymology. Species named after the type locality of Ambon Island, Indonesia. Distribution. Indonesia, Ambon Island. Diagnosis. Species of small size, characterised by dark colouration, male FWs black with yellow intermedian area, no complete harp vein, harp without a distinctive rounded area; male genitalia with a narrow mediodorsal crest on pseudepiphallic sclerite. Description. Size small. Colouration contrasted mostly dark brown and orange brown. Head dorsum with 6 brown longitudinal bands (Fig. 2). Eyes well protruding. Fastigium wider than long, setose, darker than vertex and slightly carenated posteriorly to yellow median ocellus. Scapes yellow with dark brown patterns, antennae yellowish brown. Cheeks black, except a yellow area posterior to eyes and 3 yellow spots on ventral margin of eyes (Fig. 3A). Face dark brown, with a median Tshaped yellow line, 2 yellow spots between scapes and 2 above epistomal suture; dorsal part of front head with two black spots. Labrum yellowish, its base dark brown; clypeus yellowish; mandibles yellowish with dark brown bases. Palpi yellow. Pronotum: Dorsal disk slightly trapezoidal, straight posteriorly; orange brown mottled with yellow and dark brown; lateral margins yellow; posterior margin dark brown. Lateral lobes black except two yellow spots on ventral margin (Fig. 3A). Legs: median coxae yellow, with dark brown spots; fore and median legs orange brown and yellow. Hind femora almost homogeneous orange brown, knees dark brown, 5 6 dark spots on each ventral edge; hind tibiae black, their inner surface with 4 yellow spots above and between spurs; distal part of tarsomeres III1 and III3 and dark brown. Hind tibiae with 5 inner and 10 outer spines above spurs and 4 inner and 7 outer spines between spurs. Tarsomeres III1 with 5 spines on dorsal outer edges and 12 spines at inner bases. Abdomen homogeneously dark brown dorsally; sternites yellowish brown. Cerci orange brown. Male: FW almost not setose, not reaching abdomen midlength. FW colouration (Fig. 2): Cells homogeneously black, not translucent; dorsal field veins brown; Median and R veins black, intermedian area yellow. FW venation (Fig. 1A): 1A angle right (90 ). CuP faint, limited to FW basis. Harp wide, almost flat, with no complete harp vein, without a distinctive rounded area. CuA very faint, its distal part curved inwards, around the median fold. Diagonal vein prolonged posteriorly by a strong transverse vein fused to R. Longitudinal veins strong at apex, transverse veins faint. Mirror (d1) well differentiated but not rounded. Apical field restricted to a few cells in E alignment. Lateral field black, with 5 strong longitudinal veins including MA, R and 3 more ventral veins; laterodorsal angle made by MP; R without bifurcating veins; distal part of MA and MP very weak. Subgenital plate short, not indented (Fig. 4A). Male genitalia (Fig. 5A C): Pseudepiphallic sclerite triangular; anterior apex bisinuate, its lateral part folded dorsally; posterior apex with individualized lophi, setose; pseudepiphallus dorsally convex with a narrow mediodorsal crest starting between bases of lophi (Fig. 5A). Apex of rami little sclerotized. Pseudepiphallic parameres with 3 elongate lobes. Ectophallic arc complete and wide, near base of pseudepiphallic parameres, prolonged ventrally by hooklike expansions. Ectophallic fold short and wide, with paired lateral sclerotizations. Endophallic sclerite long, reaching beyond pseudepiphallus anterior margin, with short lateral arms and a short medioposterior expansion; endophallic apodeme without a median crest. NEW LEBINTHUS OF INDONESIA AND SOLOMON ISLANDS Zootaxa 2386 2010 Magnolia Press 27

FIGURE 1. Male FW venation in dorsal view of: A, Lebinthus ambonensis n. sp.; B, Lebinthus buruensis n. sp.; C, Lebinthus cyclopus n. sp.; D, Lebinthus kolombara n. sp.; E, Lebinthus greensladei n. sp.; F, Lebinthus truncatipennis; G, Lebinthus villemantae n. sp. Abbreviations, see Material and methods. Dotted lines correspond to folds; dotted parts correspond to areas of light colouration. Colours: red, harp area; yellow, c1 cell(s); orange, end of C alignment; green, mirror (d1 cell(s)), blue, d2 cell(s); cream, rest of D alignment. Scale bar: 1 mm. 28 Zootaxa 2386 2010 Magnolia Press ROBILLARD

FIGURE 2. Lebinthus ambonensis n. sp., male habitus, holotype. Scale bar: 5 mm. Female: unknown. Measurements. see Table 1. Habitat and life history traits. unknown. Behaviour. unknown. Lebinthus buruensis n. sp. (Figs 1B, 3B, 4B, 5D F, 6A, 7A, 8A C) Type material. Holotype male: Indonesia: Province Maluku: [Pulau] Buru Is., Station 16, 20/2X1921, leg. L. J. Toxopeus (MZB Orth 1753). Allotype female: same locality as HT, Station 9, 28VI1921, leg. L. J. Toxopeus (MZB Orth 1853), (MNHNENSIF2365). NEW LEBINTHUS OF INDONESIA AND SOLOMON ISLANDS Zootaxa 2386 2010 Magnolia Press 29

FIGURE 3. Cheek, lateral lobe of pronotum and lateral field of male FW of: A, Lebinthus ambonensis n. sp.; B, Lebinthus buruensis n. sp.; C, Lebinthus cyclopus n. sp.; D, Lebinthus greensladei n. sp.; E, Lebinthus kolombara n. sp.; F, Lebinthus truncatipennis; G, Lebinthus villemantae n. sp. Scale bar: 1 mm. Type locality. Indonesia, Pulau Buru Island. Etymology. Species named after the type locality of Buru Island, Indonesia. Other material examined. Indonesia: Province Maluku: [Pulau] Buru Is., Station 16, 20/2X1921, 1 juvenile, leg. L. J. Toxopeus (MZB Orth 1846). Distribution. Indonesia, Pulau Buru Is. Diagnosis. Species of small size, with eyes small and little protruding, differing strikingly from other members of the genus by male genitalia. Description. Size small. Colouration brownish. Head dorsum yellowish brown with 6 wide dark brown longitudinal bands. Eyes little protruding. Fastigium wider than long, setose and slightly carenated posteriorly 30 Zootaxa 2386 2010 Magnolia Press ROBILLARD

to yellow median ocellus. Scapes yellowish brown with faint dark patterns on facial side; antennae yellowish brown to dark brown. Cheeks dark brown posterior to eye, except ventral margin yellow (Fig. 3B). Face, front head, mandibles, clypeus and area surrounding median ocellus dorsally black; ventral margins of eyes yellow; faint light brown patterns on epistomal suture, front head and clypeus. Palpi brown. Pronotum: Dorsal disk slightly trapezoidal, straight posteriorly; yellowish brown with dark brown spots, a faint dark brown band medially, lateral margins yellow, anterior and posterior margins with dark brown patterns. Lateral lobes black except a yellow spot on ventral margin (Fig. 3B). Legs: fore and median femora yellowish brown with dark brown spots on dorsal surface, fore and median tibiae brown. Hind femora red brown mottled with yellow, with striated brown patterns on outer face, 3 5 black spots on each ventral edge; hind tibiae brown, distal half of tarsomeres III1 dark brown. Hind tibiae with 4 5 inner (m = 4.5, n = 2) and 7 8 outer (m = 7.5, n = 2) spines above spurs and 3 inner (n = 2) and 5 6 outer (m = 5.5, n = 2) spines between spurs. Tarsomeres III1 with 4 spines on dorsal outer edges (n = 2). Abdomen homogeneously dark brown. Cerci yellowish brown basally, then homogeneously brown. TABLE 1. Measurements (in mm, mean value in brackets). PronL, pronotum length; PronW, pronotum width at midlength; FIIIL, hind femur length; FIIIW, hind femur maximal width; TIIIL, hind tibia length; FWL, forewing length; FWW, forewing width at midlength (at the first bifurcation of CuA1 in males); OvipL, ovipositor length; Strid, number of stridulatory file teeth. PronL PronW FWL FWW FIIIL FIIIW TIIIL Strid OvipL L. ambonensis Male holotype 2.8 3.6 3.3 2.4 10.1 3.7 7.4? L. buruensis Male holotype Female allotype 2.3 2.5 3.3 3.7 3.5 0.8 2.5 1.3 8.1 8.4 3.0 3.2 6.1 6.3? 8.3 L. cyclopus Male holotype 2.2 3.4 2.7 2.4 8.3 3.2 6.2? L. greensladei Male holotype Female allotype Males (n=3) Females (n=3) 2.4 2.1 2.4 2.12.4 (2.2) 3.6 3.3 3.4 3.7 (3.6) 3.1 3.6 (3.3) 3.4 1.2 3 3.4 (3.2) 1.2 1.4 (1.3) 2.1 1.1 2.1 2.2 (2.1) 1.1 1.3 (1.2) 9.9 9.3 9.6 9.9 (9.8) 9.3 9.5 (9.4) 3.6 3.6 3.5 3.9 (3.7) 3.6 3.7 (3.6) 7.3 6.6 7.2 7.8 (7.4) 6.6 7.9 (7.3) 195 190 195 (193, n=2) 7.3 7.3 8.4 (7.9) L. kolombara Male holotype 2.6 3.4 3.0 2.2 10.3 3.8 7.9? L. truncatipennis Male allotype Males (n=6) Female (n=1) 2.6 2.3 2.6 (2.5) 2.5 3.9 3.4 3.9 (3.7) 3.8 3.7 3.6 3.9 (3.8) 1.8 2.7 2.7 3 (2.9) 0.9 12.6 10 12.6 (11.3) 10.8 4.0 3.2 4.0 (3.7) 3.8 10.0 8.3 10 (9.1) 8.2? 223 (n=1) 8.3 L. villemantae Male holotype Female allotype Males (n=5) Females (n=5) 2.4 2.3 2 2.3 (2.2) 1.9 2.2 (2.0) 3.5 3.4 3.1 3.6 (3.4) 3 3.4 (3.2) 3.8 1.9 3.4 4.3 (3.8) 1.8 2.3 (2.0) 2.5 1.7 2.2 2.9 (2.6) 1.7 1.9 (1.8) 10.4 10.0 9.4 11.1 (10.3) 9.2 10.4 (9.7) 3.3 3.4 3 3.6 (3.3) 2.9 3.5 (3.2) 8.5 8.3 8.1 9 (8.5) 7.8 8.6 (8.3)? 198 213 (203, n=4) 7.5 7.5 8.3 (7.9) NEW LEBINTHUS OF INDONESIA AND SOLOMON ISLANDS Zootaxa 2386 2010 Magnolia Press 31

Male: FW setose, not reaching abdomen midlength (Fig. 1B). FW colouration: Cells and veins homogeneously brown, not translucent. FW venation: Most veins very faint. 1A angle wide (>100 ). CuP missing. Harp wide, almost flat, with no harp vein and without a distinctive rounded area. Proximal part of CuA very faint, distal part slightly curved inwards around the median fold. Diagonal vein prolonged posteriorly by a strong elevated vein fused to R. Longitudinal veins very strong at apex, transverse veins very weak or absent, except a faint but wide transverse vein posterior of the undifferentiated mirror (d1). Apical field restricted to 2 cells in E alignment. Lateral field homogeneously brown, except a translucent area on ventral margin; with 5 strong longitudinal veins including MA, R and 3 more ventral veins, the most ventral one bifurcated; laterodorsal angle made by MP; R without strong bifurcating veins. Subgenital plate trapezoidal, slightly indented posteriorly (Fig. 4B). Male genitalia (Fig. 5G I): Pseudepiphallic sclerite as wide as long, convex dorsally; posterior apex with large triangular lophi, separated by a deep Vshaped indentation; anterior apex straight, its lateral margins slightly curved dorsally. Rami short. Pseudepiphallic parameres very large, trilobate, the posterior lobe short and dorsal, the two other lobes ventral, the posterior one very elongate, surrounding outerly the pseudepiphallic lophi. Ectophallic arc complete, narrow, near basis of pseudepiphallic parameres. Ectophallic fold short and wide, membranous. Ectophallic apodemes parallel and short. Endophallic sclerite short, but exceeding anterior margin of pseudepiphallus, convex dorsally, its posterior apex with a median expansion; endophallic apodeme made of lateral lamellas and a narrow median crest. Female: FW very short (Fig. 6A), reaching posterior margin of first tergite, not overlapping; homogeneously dark brown, except a lighter area near lateral margin of dorsal field; dorsal field with 4 strong and 1 weak longitudinal veins. Lateral field with 2 strong dark brown longitudinal veins. Ovipositor as long as hind femora; apex lanceolate, darker brown, denticulate on dorsal edge (Fig. 7A). Female genitalia: Copulatory papilla (Fig. 8A C) concave ventrally, well sclerotized except apex and median dorsal area. Juvenile: Subadults similar to adults in colouration, dark brown. Measurements. see Table 1. Habitat and life history traits. unknown. Behaviour. unknown. Lebinthus cyclopus n. sp. (Figs 1C, 3C, 4C, 5G I) Type material. Holotype male: Indonesia: Province Papua: [Papua Is.], Mt[s] Cyclops, 4000 ft., 12III1936 (L.E. Cheesman) (BMNHB.M.1936271). Type locality. Indonesia, Papua Island, Mounts Cyclops. Etymology. Species named after the type locality. Distribution. Indonesia, Papua Island. Diagnosis. Species of small size, very close in shape, FW venation and male genitalia to L. nattawa Robillard, 2009, from which it differs by colouration and larger size of endophallic sclerite. Description. Size small. Colouration mostly dark brown. Head dorsum with 6 wide but faint brown longitudinal bands; vertex light brown. Eyes well protruding. Fastigium wider than long, setose, darker than vertex and slightly carenated posteriorly to brown median ocellus; apical and lateral margins yellow. Scapes yellowish brown with light brown patterns; antennae yellowish brown. Cheeks dark brown (Fig. 3C). Face wider than high, almost entirely dark brown, with a thin yellow line on epistomal suture; mouthparts dark brown; dorsal part of front head with 2 black spots. Palpi light brown except base and apex, dark brown. Pronotum: Dorsal disk slightly trapezoidal, straight posteriorly; dark brown, lateral margins whitish with dark brown spots. Lateral lobes dark brown with a faint light spot on anteroventral angle. Legs: Fore and median femora yellowish brown mottled with dark brown; tibiae orange brown. Hind legs homogeneously brown, knees dark brown. Hind tibiae with 3 inner and 5 outer spines above spurs and 2 inner and 5 outer spines 32 Zootaxa 2386 2010 Magnolia Press ROBILLARD

between spurs. Tarsomeres III1 with 3 spines on dorsal outer edges. Abdomen and cerci homogeneously dark brown. Male: FW not reaching abdomen midlength. FW colouration: Cells and veins homogeneously brown. FW venation (Fig. 1C): 1A angle wide (>100 ). CuP absent. Harp wide, posterior margin raised, without a distinctive rounded area, with one straight harp vein. CuA faint anteriorly, its distal part curved inwards, around the median fold. Diagonal vein prolonged posteriorly by a strong transverse vein fused to median veins. Longitudinal veins strong at apex, transverse veins faint. Mirror (d1) well differentiated but not rounded. Apical field restricted to one cell in E alignment. Lateral field brown, with 6 strong longitudinal veins including MA, R and 4 more ventral veins; laterodorsal angle made by MP. Subgenital plate short, not indented (Fig. 4C). FIGURE 4. Male subgenital plate of: A, Lebinthus ambonensis n. sp.; B, Lebinthus buruensis n. sp.; C, Lebinthus cyclopus n. sp.; D, Lebinthus greensladei n. sp.; E, Lebinthus kolombara n. sp.; F, Lebinthus truncatipennis; G, Lebinthus villemantae n. sp. Scale bar: 1 mm. NEW LEBINTHUS OF INDONESIA AND SOLOMON ISLANDS Zootaxa 2386 2010 Magnolia Press 33

FIGURE 5. Male genitalia of: A C, Lebinthus ambonensis n. sp.; D F, Lebinthus buruensis n. sp.; G I, Lebinthus cyclopus n. sp.; in dorsal (A, D, G), ventral (B, E, H) and lateral (C, F, I) views. Abbreviations, see Material and methods. Scale bars: 1 mm. 34 Zootaxa 2386 2010 Magnolia Press ROBILLARD

FIGURE 6. Female FW venation of: A, Lebinthus buruensis n. sp.; B, Lebinthus greensladei n. sp.; C, Lebinthus truncatipennis; D, Lebinthus villemantae n. sp. Dotted areas represent patterns of white or yellow areas. Black arrows show the posterior limit of first abdominal tergite. Abbreviations, see Material and methods. Scale bar: 1 mm. NEW LEBINTHUS OF INDONESIA AND SOLOMON ISLANDS Zootaxa 2386 2010 Magnolia Press 35

Male genitalia (Fig. 5G I): Pseudepiphallic sclerite triangular; anterior apex slightly bisinuate; posterior apex with individualized lophi, rounded and setose; pseudepiphallus convex dorsally. Pseudepiphallic parameres large, trilobate. Ectophallic arc complete and wide, near base of pseudepiphallic parameres, prolonged ventrally by hooklike expansions. Ectophallic fold short and wide, with a Wshaped preapical sclerotization. Endophallic sclerite long, reaching beyond pseudepiphallus anterior margin, with short lateral arms and a short posterior expansion; endophallic apodeme made of a thin median crest. Female: unknown. Measurements. see Table 1. Habitat and life history traits. unknown. Behaviour. unknown. Lebinthus greensladei n. sp. (Figs 1E, 3D, 4D, 6B, 7B, 8D F, 9A C) Type material. Holotype male: Solomon Islands: Guadalcanal Province: Guadalcanal [Is.], Mt. Austen 4III 1963, (4605), (P.[J.M.] Greenslade), labelled Lebinthus greensladei by L. Chopard (MNHNENSIF1450). Allotype female: same locality and collector as HT, 9VII1963, (8108) (MNHNENSIF1451). Paratypes: same locality and collector as HT; 1VII1963, 1 (6964), Pres. by P.J.M. Greenslade, labelled Lebinthus greensladei by L. Chopard (BMNHB.M.1966477); 20V1963, 1 (6152), Pres. by P.J.M. Greenslade, labelled Lebinthus greensladei by L. Chopard (BMNHB.M.1966477); 10V1965, 1 (19119), carrion trap (BMNH); I1966, 1 (19973) (BMNH); 20IV/3V1965, 1 (17519), carrion trap (BMNH); V1966, 1 (21533) (BMNH); 22VII1963, 1 (11302) (BMNH). Type locality. Solomon Islands, Guadalcanal Island, Mt. Austen. Etymology. Species named after the collector, P.J.M. Greenslade. Some of the type specimens were separated as a new species by L. Chopard and labelled Lebinthus greensladei. Other material examined. Solomon Islands: Isabel Province: Santa Ysabel [Isabel Is.], Tatamba, 29IX 1965, 1 (Roy. Soc. Exped.), wooded hillside behind resthouse (BMNHB.M.19661). Malaita Province: Malaita [Is.], Tangtalau 200 m, 30IX1957, 1 (J. L. Gressit) (MNHNENSIF1459). MakiraUlawa Province: San Cristoval [Is.], camp 2, confluence of Warachito & Pagato Rivers, 6 7 mls. inland, 24VII 1965, 1 (Roy. Soc. Exped.) (BMNHB.M.19661); camp 2, Warachito Pagato confluence, 3VIII1965, 1 (Roy. Soc. Exped.) (BMNHB.M.19661); camp 3, Huni R., 14VIII1965, 1 (Roy. Soc. Exped.) (BMNH B.M.19661). Papua New Guinea: Autonomous Region of Bougainville: Solomon Is., Bougainville [Is.], Simba Mission, 30VI1966, 1 (E.J. Ford Jr.) (MNHNENSIF1452). Distribution. Isabel Is., Malaita Is., San Cristoval Is., Bougainville Is. Diagnosis. Species of small size, close to L. lifouensis DesutterGrandcolas, 1997 and L. santoensis Robillard, 2009 in shape, colouration, FW venation and male genitalia, differing by shape of pseudepiphallic lophi, less globulose, and shape of ectophallic fold preapical sclerotization. Description. Size small. Head dorsum yellowish brown with 6 irregular brown longitudinal bands as in L. santoensis, 2 median ones, patchy at level of fastigium, 2 lateral ones curved outerly toward eye midlength, 2 most outer ones behind eye and short; dark spots at bases of setae between longitudinal bands. Eyes well protruding, dark brown with a light brown longitudinal band. Fastigium wider than long, setose, yellow, with a dark area posterior to whitish median ocellus; margin between front head and dorsum yellow. Scapes whitish, antennae light brown. Cheeks dark brown posterior to eye, except ventral margin of eyes orange brown (Fig. 3D). Face black except a whitish inverted Tshaped pattern, paired whitish spots between antennae and median part of epistomal suture. Clypeus dark brown; palpi whitish, except basis of 3 rd segment and apex, dark brown. Pronotum: Dorsal disk slightly trapezoidal, straight posteriorly; yellowish brown with dark brown spots, a faint dark brown band medially, lateral margins yellow, posterior margin with dark brown patterns. Lateral lobes black or dark brown except ventral margin, orange brown (Fig. 3D). Legs: Fore and median femora yellowish brown with dark brown spots on dorsal surface, fore and median tibiae brown with a faint darker ring. Hind femora yellowish brown with dark brown striated patterns on dorsal outer faces and 36 Zootaxa 2386 2010 Magnolia Press ROBILLARD

dark brown spots on ventral outer faces; knees dark brown; 3 5 black spots on each ventral edge; hind tibiae dark brown, their inner surface with 4 yellow spots above and between spurs; basal and distal parts of tarsomeres III1 dark brown. Hind tibiae with 5 7 inner (m = 6.3, n = 6) and 8 11 outer (m = 9.5, n = 6) spines above spurs and 3 5 inner (m = 4.3, n = 6) and 4 6 outer (m = 5.7, n = 6) spines between spurs. Tarsomeres III1 with 3 6 spines on dorsal outer edges (m = 4, n = 2). Abdomen orange brown mottled with dark brown. Cerci homogeneously brown. FIGURE 7. Apex of ovipositor in lateral view of: A, Lebinthus buruensis n. sp.; B, Lebinthus greensladei n. sp.; C, Lebinthus truncatipennis; D, Lebinthus villemantae n. sp. Scale bar: 1 mm. Male: FW little setose, not reaching abdomen midlength. FW colouration contrasted; cells dark brown, translucent, veins yellowish brown (HT) or brown (male PT); intermedian area strongly sclerotized, whitish. FW venation (Fig. 1E): Veins strong, including transverse veins; 1A angle wide (>100 ); stridulatory file with 190 195 teeth (m = 193, n = 2). CuP missing. Harp wide, almost flat, with one straight harp vein and without a distinctive rounded area. Distal part of CuA slightly curved inwards, around the median fold. Diagonal vein very strong, prolonged posteriorly by a strong yellow vein connected to MA/MP fusion. Mirror (d1) little differentiated. Apical field restricted to a few cells in E alignment. Lateral field homogeneously dark brown, except translucent area on ventral margin, and intermedian area whitish including cells and transverse veins, the median veins orange brown; with 5 strong longitudinal veins including MA, R and 3 more ventral veins, the most ventral one bifurcated; laterodorsal angle made by MP; R without strong bifurcating veins. Subgenital plate trapezoidal (Fig. 4D), quite short compared to L. villemantae (Fig. 4G). Male genitalia (Fig. 9A C): Pseudepiphallic sclerite triangular, slightly convex dorsally; anterior apex slightly indented; posterior apex with individualized lophi, setose, straight and acute at apex. Pseudepiphallic parameres large, trilobate and close together. Ectophallic arc complete, closer to pseudepiphallic parameres than to anterior margin of pseudepiphallus. Ectophallic fold short and wide, with a Ushaped preapical sclerotization. Endophallic sclerite Yshaped, exceeding anterior margin of pseudepiphallus, convex dorsally, its posterior apex with a median expansion; endophallic apodeme made of lateral lamellas and a narrow median crest. Female: FW short (Fig. 6B), exceeding posterior margin of first tergite, not overlapping. FW colouration contrasted dorsally, cells dark brown and veins yellowish brown; dorsal field with 6 7 strong longitudinal veins and sclerotized transverse veins. Lateral field dark brown with 3 4 strong longitudinal veins. Ovipositor slightly shorter than hind femora; apex lanceolate, brown, faintly denticulate on dorsal edge (Fig. 7B). Female genitalia: Copulatory papilla (Fig. 8D F) concave ventrally, not sclerotized except a basal ring. Juvenile: unknown. Variation: Specimens from San Cristoval Is. (2, 1 ) have a more homogeneous dark brown colouration and show very slight differences in male FW venation and male genitalia. Measurements. see Table 1. Habitat and life history traits. unknown. Behaviour. unknown. NEW LEBINTHUS OF INDONESIA AND SOLOMON ISLANDS Zootaxa 2386 2010 Magnolia Press 37

Lebinthus kolombara n. sp. (Figs. 1D, 3E, 4E, 9D F) Type material. Holotype male: Solomon Islands: Western Province: Kolombangara [Is], base camp, 1 mi inland from Kuzi, by Kolombara r [river], 2IX1965 (BMNH). Type locality. Solomon Islands, Kolombangara Island. Etymology. Species named after the Kolombara river near the type locality in Kolombangara Island, Solomon Islands. Distribution. Solomon Islands, Kolombangara Island. Diagnosis. Species of small size, close to L. lifouensis in shape, male FW venation and genitalia; colouration dark brown and black face, and two lateral yellow lines on head dorsum and dorsal disk of pronotum; harp without a distinctive rounded area. Description. Size small. Colouration mostly dark brown and black. Head dorsum brown with 6 wide dark brown longitudinal bands. Eyes well protruding. Fastigium wider than long, setose and slightly carenated posteriorly to yellow median ocellus. Scapes and antennae yellowish brown. Cheeks black, except a yellow line posterior to eyes, and margin below eyes yellow (Fig. 3E). Face, including mouthparts, black except a horizontal yellow line between scapes, 2 yellow spots above epistomal suture and one at base of labrum; clypeus ventral margin yellow. Palpi yellow, apex dark brown. Pronotum: Dorsal disk slightly trapezoidal, straight posteriorly; dark brown, with a very thin median yellow line, lateral margins yellow. Lateral lobes black except for a yellow spot on ventral margin (Fig. 3E). Legs: Median coxae black with a vertical yellow line; fore and median legs orange brown; hind femora orange brown with striated brown patterns on outer face, 3 5 faint dark spots on each ventral edge; hind tibiae dark brown. Both apical and basal parts of tarsomeres III1 dark brown. Hind tibiae with 6 inner and 7 outer spines above spurs and 3 inner and 5 outer spines between spurs. Tarsomeres III1 with 4 spines on dorsal outer edges. Abdomen homogeneously dark brown. Male: FW setose, not reaching abdomen midlength. FW colouration: Cells and veins homogeneously brown, not translucent. FW venation (Fig. 1D): 1A angle wide (>100 ). CuP missing. Harp wide, almost flat, with one transversal harp vein slightly bisinuated, without a distinctive rounded area. Distal part of CuA slightly curved inwards, around the median fold. Diagonal vein prolonged posteriorly by a strong transverse vein fused to MP. Longitudinal veins very strong at apex. Mirror (d1) little differentiated. Posterior venation faintly expressed; apical field restricted to a few cells in E alignment. Lateral field homogeneously brown; with 7 strong longitudinal veins including MA, R and 5 more ventral veins; laterodorsal angle made by MP; R without bifurcating veins; distal part of MA very weak. Subgenital plate short but longer than wide, not indented (Fig. 4E). Male genitalia (Fig. 9D F): close to that of L. lifouensis but wider. Pseudepiphallic sclerite triangular, slightly convex dorsally; anterior apex straight; posterior apex with individualized lophi, setose. Pseudepiphallic parameres large, trilobate and close together. Ectophallic arc complete and wide, at midlength between pseudepiphallic parameres and anterior apex. Ectophallic fold short and wide, with a Mshaped preapical sclerotization. Endophallic sclerite Yshaped with short lateral arms; endophallic apodeme made of a long and wide median crest, not disconnected from membrane. Female: unknown. Measurements. see Table 1. Habitat and life history traits. unknown. Behaviour. unknown. 38 Zootaxa 2386 2010 Magnolia Press ROBILLARD

FIGURE 8. Female copulatory papilla in dorsal, ventral and lateral views of: A C, Lebinthus buruensis n. sp.; D F, Lebinthus greensladei n. sp.; G I, Lebinthus truncatipennis; J L, Lebinthus villemantae n. sp. Scale bar: 1 mm. NEW LEBINTHUS OF INDONESIA AND SOLOMON ISLANDS Zootaxa 2386 2010 Magnolia Press 39

FIGURE 9. Male genitalia of: A C, Lebinthus greensladei n. sp.; D F, Lebinthus kolombara n. sp.; in dorsal (A, D), ventral (B, E) and lateral (C, F) views. Scale bar: 1 mm. Lebinthus truncatipennis Chopard, 1929 (Figs. 1F, 3F, 4F, 6C, 7C, 8G I, 10A C) Chopard, 1929: 111; 1968: 354. Robillard & DesutterGrandcolas 2004a, (morphological phylogeny), 2006 (molecular and morphological phylogeny); Eades et al. 2008. Orthoptera Species File Online. Type material. Female holotype: [Indonesia: West Sumatra Province]: Sipora [Is.], 9X1924 (H. H. K.), not found in the Raffles Museum (ZRC). Male allotype: [Indonesia]: West Sumatra [Province]: Sipora Is., X1924 (C. B. K. & N. S.) (MNHNENSIF1446). Type locality. Indonesia, Sipora Is. Other material examined. [Indonesia: West Sumatra Province], Wai Lima, Lampongs, XIXII1921, 2, 1 (N 120, 218) (Karny), identified L. truncatipennis by T. Robillard, 2004 (Robillard and Desutter Grandcolas 2004a) (MNHNENSIF1447, 2548, 2549). Indonesia: West Java [Province]: [Java Is.], Pangandaran forest, 125m, 15/15VI1925 (T. C. Forest Entomology) 3 (MZB Orth 94809482). 40 Zootaxa 2386 2010 Magnolia Press ROBILLARD

Distribution. Indonesia: Sipora Island, West Java, Sumatra. Diagnosis. Species of small size, brownish, larger than Lebinthus villemantae sp. n.; with a distinctive rounded area on the harp as in L. villemantae n. sp. and L. makilingus Otte, 2007 from which it differs by male genitalia, shorter female FWs and details of colouration. FIGURE 10. Male genitalia of: A C, Lebinthus truncatipennis; D F, Lebinthus villemantae n. sp.; in dorsal (A, D), ventral (B, E) and lateral (C, F) views. Scale bar: 1 mm. NEW LEBINTHUS OF INDONESIA AND SOLOMON ISLANDS Zootaxa 2386 2010 Magnolia Press 41

Redescription. In addition to the characters mentioned by Chopard (1929): Colouration brown with more or less contrasted coloured patterns. Head dorsum yellowish with 4 wide brown longitudinal bands more or less distinct, and with a yellow band posterior to eye prolonging eye colouration and underlined ventrally by a black band. Eyes well protruding, black with a red brown dorsal longitudinal band. Fastigium wider than long, setose, brown with a dark spot behind yellowish median ocellus; apex yellow and slightly carenated posteriorly to median ocellus, at margin between front head and dorsum. Scapes yellowish brown with brown patterns on face; antennae light brown. Face yellowish brown with paired spots between scapes; 2 dark brown horizontal bands, one below scapes; 2 yellow spots dorsal to epistomal suture, and one on mandible bases and labrum; clypeus yellowish brown except ventral margin; cheeks yellowish brown except for a dark brown pattern posterior to eyes (Fig. 3F). Palpi yellowish brown, with one dark brown ring per segment. Pronotum: Dorsal disk slightly trapezoidal, slightly sinuate posteriorly; yellowish brown with few dark spots, slightly lighter on lateral margins. Lateral lobes black dorsally, ventral margin yellow with a faint curved brown pattern (Fig. 3F). Legs: Fore and median legs light brown, femora with dark brown spots, tibiae with dark rings. Hind femora orange brown, with striated dark patterns on outer faces, 3 5 black spots on each ventral edge, knees dark brown; hind tibiae with dark brown rings, distal half of tarsomeres III1 and III3 dark brown. Hind tibiae with 5 8 inner (m = 6.4, n = 7) and 8 12 outer (m = 9.7, n = 7) spines above spurs and 3 5 inner (m = 4.8, n = 7) and 5 6 outer (m = 5.7, n = 7) spines between spurs. Tarsomeres III1 with 2 3 spines on dorsoouter edges (m = 2.6, n = 7) and 0 1 (m = 0.7, n = 7) on outer faces. Abdomen homogeneously dark brown, with sometimes a median dark brown band ventrally. Cerci brown basally, yellowish brown with dark brown rings toward apex. Male: FW not reaching abdomen midlength. FW colouration: Cells and veins brown, not translucent; base of CuA and angle of 1A with a yellowish brown sclerotization; intermedian area yellowish brown; lateral field brown, its dorsal margin (MA/R area) dark brown. FW venation (Fig. 1F): 1A angle wide (>100 ); stridulatory file with 223 teeth (n = 1), located on transverse part of 1A only. CuP visible at FW basis only. Harp wide, including a false mirror, i.e. a distinctive rounded area delimited by the strong harp vein, polyfurcated anteriorly, and fused with both diagonal vein and CuA posteriorly. Distal part of CuA weak and curved inwards, surrounding the median fold. Distal part of diagonal vein very weak. Longitudinal veins very strong at apex, transverse veins very weak. Mirror (d1) not differentiated. Apical field absent, with no bifurcation of CuA posterior to diagonal vein. Lateral field with 5 longitudinal veins including MA, R and 3 more ventral veins; laterodorsal angle made by MP; R without strong bifurcating veins. Subgenital plate clogshaped (Fig. 4F), less elongate than in L. villemantae (Fig. 4G). Male genitalia (Fig. 10A C): Pseudepiphallic sclerite elongate, widened anteriorly, straight dorsally; posterior apex with short triangular lophi curved dorsally, setose, separated by a Vshaped indentation; anterior apex straight, its lateral margins slightly curved dorsally. Pseudepiphallic parameres trilobate, the posterior lobe dorsal, the two other lobes ventral. Ectophallic arc complete and narrow, well separate from basis of pseudepiphallic parameres. Ectophallic fold wide, with paired preapical sclerotization. Ectophallic apodemes wide, parallel and exceeding anterior margin of pseudepiphallus. Endophallic sclerite long, exceeding anterior margin of pseudepiphallus, its posterior apex with a median triangular expansion and with short lateral arms; endophallic apodeme made of a narrow median crest. Female: FW very short (Fig. 6C), not reaching posterior margin of second tergite, not overlapping; dorsal field with 6 strong brown longitudinal veins; yellowish brown sclerotization between brown transverse veins. Lateral field darker than dorsal field, with 4 strong dark brown longitudinal veins. Ovipositor shorter than hind femora; apex lanceolate, slightly denticulate on dorsal edge (Fig. 7C). Female genitalia: Copulatory papilla (Fig. 8G I) very small, almost completely membranous. Juvenile: unknown. Measurements. see Table 1. Habitat and life history traits. unknown. Behaviour. unknown. 42 Zootaxa 2386 2010 Magnolia Press ROBILLARD

Lebinthus villemantae n. sp. (Figs 1G, 3G, 4G, 6D, 7D, 8J L, 10D F, 11, 12, 13) Type material. Holotype male: Indonesia: Province Sulawesi Selatan: Sulawesi Selatan, Mont Bulu Saraung, Pos 5, forêt de pente, espace dégagé en bord de piste, litière, 20VIII2007, jour (T. Robillard), adulte en élevage (MZB Orth). Allotype female: same locality as HT, 31VIII2007, jour (C. Villemant), adulte en élevage (MZB). Paratypes (18, 5 ): Indonesia: Province Sulawesi Selatan: same locality, date and collector as AT, 7, 2, adultes en élevage (MZB); 9 (recorded), 2, adultes en élevage (MNHNENSIF25722582). Sulawesi Selatan, région de Malawa, forêt humide audessus de Boto Siri, sentier longeant ruisseau, litière, 25VIII2007, jour (T. Robillard), 1, 1 (MZB); 1 (MNHNENSIF2583). Type locality. Indonesia, Sulawesi Selatan, Mount Bulu Saraung. Etymology. Species dedicated to the French hymenopterologist Dr. Claire Villemant (MNHN). Other material examined. Indonesia: Province Sulawesi Selatan: Sulawesi Selatan, Mont Bulu Saraung, Pos 5, forêt de pente, espace dégagé en bord de piste, litière, 20VIII2007, jour (T. Robillard), 12 juveniles (7 in alcohol) (MNHNENSIF25912596); 30 juveniles (MZB); 31VIII2007, jour (C. Villemant), 8, 4 (MNHNENSIF25672571, 25842590), 6 juveniles (MZB). Distribution. Indonesia, South Sulawesi. Diagnosis. Species of small size, slightly smaller and darker than Lebinthus truncatipennis, and differing from L. truncatipennis and L. makilingus by male genitalia and details of colouration. Description. Size small. Colouration variable, most often dark brown with more or less contrasted colour patterns (Fig. 11). Head dorsum with 4 6 wide dark brown longitudinal bands more or less distinct, with a yellow band posterior to eyes prolonging their colouration and underlined by black pattern. Eyes well protruding, black with a yellow longitudinal band dorsally. Fastigium wider than long, setose, brown with a dark spot behind median ocellus, yellow. Scapes yellow; antennae yellow at base then progressively dark brown. Upper part of front head yellow. Black mask including lower part of front head, region below antennae and eyes, base of mandibles and upper part of clypeus; epistomal suture yellowish brown; yellow to whitish colouration around the mask, including parts of jaws and lower parts of mandibles and clypeus. Palpi whitish with dark brown rings. Pronotum: Dorsal disk slightly trapezoidal, straight posteriorly; generally dark brown, lighter laterally. Lateral lobes dorsally black, ventral margin yellow with a black line (Fig. 3G). Legs: Fore and mid legs light brown, femora with dark brown spots, tibiae with dark rings. Hind femora brown, with striated dark patterns on outer faces, 3 5 black spots on each ventral edge, knees dark brown; hind tibiae with black rings, distal half of tarsomeres III1 black. Hind tibiae with 6 12 inner (m = 7.9, n = 10) and 10 13 outer (m = 11.3, n = 10) spines above spurs and 4 5 inner (m = 4.3, n = 10) and 4 7 outer (m = 5.4, n = 10) spines between spurs. Tarsomeres III1 with 3 4 spines on dorsal outer edges (m = 3.4, n = 10) and 0 2 (m = 1.3, n=8) on outer faces. Abdomen homogeneously brown dorsally, yellowish brown ventrally with sometimes a median dark brown band. Cerci yellowish brown basally, with black rings near apex. Male: FW not reaching abdomen midlength. FW colouration: Cells and veins brown, not translucent; intermedian area whitish, except for dark brown veins; lateral field brown, its dorsal margin (MA/R area) dark brown. FW venation (Fig. 1G): 1A angle wide (>100 ); stridulatory file with 198 213 teeth (m = 203, n = 4), located on transverse part of 1A only. CuP visible at FW basis only. Harp wide, including a false mirror, i.e. a distinctive rounded area delimited by the strong harp vein, polyfurcated anteriorly, and fused with both diagonal vein and CuA posteriorly. Distal part of CuA weak and curved inwards, surrounding the median fold. Distal part of diagonal vein very weak. Longitudinal veins very strong at apex, transverse veins very weak. Mirror (d1) not differentiated. Apical field absent, with no bifurcation of CuA posterior to diagonal vein. Lateral field with 5 strong longitudinal veins including MA, R and 3 more ventral veins; laterodorsal angle made by MP; R without strong bifurcating veins. Subgenital plate elongate, clogshaped (Fig. 4G). Male genitalia (Fig. 10D F): Pseudepiphallic sclerite triangular, convex dorsally; posterior apex with short individualized lophi, setose, parallel and separated by a Vshaped indentation; anterior margin slightly indented, its lateral margins slightly curved dorsally. Rami short. Pseudepiphallic parameres trilobate, the posterior lobe dorsal, the two other lobes ventral, the median one rectangular. Ectophallic arc complete, NEW LEBINTHUS OF INDONESIA AND SOLOMON ISLANDS Zootaxa 2386 2010 Magnolia Press 43

narrow, very close to basis of pseudepiphallic parameres. Ectophallic fold short and wide, with a wide preapical sclerotization including a rounded part and two anterior expansions laterally. Ectophallic apodemes parallel and very long, exceeding anterior margin of pseudepiphallus. Endophallic sclerite very long, exceeding anterior margin of pseudepiphallus, dorsally convex, its posterior apex with a median triangular expansion and with short lateral arms; endophallic apodeme made of a narrow median crest. FIGURE 11. Lebinthus villemantae n. sp., male habitus, holotype. Scale bar: 5 mm. 44 Zootaxa 2386 2010 Magnolia Press ROBILLARD

Female: FW short (Fig. 6D), exceeding posterior margin of second tergite, not overlapping but close together; dorsal field with 5 strong longitudinal veins, brown. Lateral field darker than dorsal field, with 3 4 strong dark brown longitudinal veins, intermedian area whitish as in male. Ovipositor shorter than hind femora; apex lanceolate, denticulate on dorsal edge (Fig. 7D). Female genitalia: Copulatory papilla (Fig. 8J L) with small basal sclerotized ring; apical part narrowed and long, slightly plicate ventrally and sclerotized at apex. Juvenile: Similar to adults in colouration, dark brown (Fig. 12). Measurements. see Table 1. Habitat and life history traits. L. villemantae lives in the leaf litter of forested areas (Fig. 12). It is found in dense populations in wet leaf litter, generally on or under large dead leaves. Behaviour. Only 3 adult specimens were observed in the field, by day in leaf litter in Malawa region, South Sulawesi. Singing activity was recorded in laboratory mostly during night period, but sexual activity seems to occur during both day and night. Mating behaviour has been observed in laboratory; multiple copulations have been recorded and will be described in a forthcoming paper (Robillard et al. in prep.). FIGURE 12. Lebinthus villemantae n. sp.: A, B, habitat, in Bulu Saraung mount, Sulawesi Selatan, Indonesia; C, male, in Malawa region, Sulawesi Selatan, Indonesia; D, E, juvenile specimens, in Bulu Saraung mount, Sulawesi Selatan, Indonesia. NEW LEBINTHUS OF INDONESIA AND SOLOMON ISLANDS Zootaxa 2386 2010 Magnolia Press 45

FIGURE 13. Calling song of Lebinthus villemantae n. sp.: A, oscillogram showing 3 successive echemes; B, spectrogram (relative amplitude); C, detailed oscillogram of an echeme; D, detailed oscillogram of a syllable; E, detailed sonogram of an echeme. Symbols: f1 f4, first (fundamental) to fourth harmonic frequencies. Calling song (Fig. 13; Table 2): Five males recorded in laboratory at 18 26 C are analysed here. The calling song consists of a succession of short echemes of 1.8 ± 0.1 s emitted every 47.1 ± 19 s, with a duty cycle of 3.8%. At 20 C each echeme is made of 99.5 ± 10.6 syllables, with the following characteristics: syllable rate = 58.7 ± 3.8 /s; syllable duration = 7.2 ± 1.9 ms; syllable period = 17.5 ± 4.1 ms; syllable duty cycle = 41.1%; the dominant frequency is 20.1 ± 2.3 khz and corresponds to the fundamental frequency of the song. Courtship song: Less intense and shorter than calling song. Aggressive song: Not observed despite many observations of male interactions. This song is probably absent from the species repertoire. 46 Zootaxa 2386 2010 Magnolia Press ROBILLARD

TABLE 2. Calling song parameters of Lebinthus villemantae n. sp. Specimens T C Echeme Syllable Dominant frequency (khz) Duration (s) Period (s) Syllable number Duration (ms) Period (ms) TREl4 (MNHNENSIF2574) 23.6 17.7± 0 1.6±0.0 159.9±96.5 108±7 7.8±1.5 14.7±1.7 66.5±3.2 TREl7 (MNHNENSIF2578) 25.8 18.7±0.3 1.5±0.1 49.6±30.9 116±9 7.8±1.3 13.1±1.9 76.3±1.5 TREl9 (MNHNENSIF2586) 25 18.5±0.5 0.9±0.4 101.3±51.1 67±33 6.0±1.7 12.5±1.1 77.7±0.9 TREl11 (MNHNENSIF2590) 24.2 18.2±0.2 1.6±0.1 30.1±19.2 118±4 7.5±1.2 13.1±1.9 75.5±2.9 TREl18 (MNHNENSIF2584) 18.8 20.4±0.2 2.1±0.1 43.2±4.9 114±6 7.3±1.8 19.0±4.8 55.3±0.9 Rate (Hz) Acknowledgements The field work in South Sulawesi was organised by Louis Deharveng (MNHN), Anne Bedos (MNHN) and Yayuk Suhardjono (LIPI), with a grant from the PPF "État et structure phylogénétique de la biodiversité actuelle et fossile", MNHN (P. Janvier). I thank Claire Villemant (MNHN) for helping collecting crickets in Sulawesi. I also thank Simon Poulain (CNRS) for preparation of the specimens, Constance Boitard for specimen measurements, Gilbert Hodebert (MNHN) for habitus drawings and Laure DesutterGrandcolas for helpful comments on the manuscript. I also thank: Judith Marshall and George Beccaloni (BMNH), for their help during the study of Eneopterinae crickets in the Natural History Museum, London, funded by the SYNTHESYS European program (GBTAF531); Rob de Vries and Caroline Pepermans (RMNH) for their help during the study in Leiden collections, funded by a grant from the PPF "État et structure phylogénétique de la biodiversité actuelle et fossile", MNHN (P. Janvier). References Chopard, L. (1929) Spolia mentawiensia, Gryllidae. Bull. Raffles Mus., 2, 98 118. Chopard, L. (1968) Pars 12. Fam Gryllidae: Subfam. Mogoplistinae, Myrmecophilinae, Scleropterinae, Cachoplistinae, Pteroplistinae, Pentacentrinae, Phalangopsinae, Trigonidiinae, Eneopterinae. Fam. Oecanthidae, Gryllotalpidae. Dr.W.Junk N.V., 's Gravenhage, 215 500 pp. Desutter, L. (1987) Structure et évolution du complexe phallique des Gryllidea (Orthoptera) et classification des genres néotropicaux de Grylloidea.1ère partie. Annales de la Société Entomologique de France (N.S.), 23, 213 239. DesutterGrandcolas, L. (1997) Les grillons de Nouvelle Calédonie (Orthoptères, Grylloidea): espèces et données nouvelles. In: Najt, J. & Matile, L. (Eds.) Zoologia Neocaledonica, 4. Mémoires du Muséum National d'histoire Naturelle, Paris, pp. 165 177. DesutterGrandcolas, L. (2003) Phylogeny and the evolution of acoustic communication in extant Ensifera (Insecta, Orthoptera). Zoologica Scripta, 32, 525 561. DesutterGrandcolas, L. & Robillard, T. (2006) Phylogenetic systematics and evolution of Agnotecous in New Caledonia (Orthoptera: Grylloidea, Eneopteridae). Systematic Entomology, 31, 65 92. Eades, D.C., Otte, D. & Naskrecki, P. (2008) Orthoptera Species File Online Vers 2.0/3.4 [20/04/2008]. http://orthoptera. SpeciesFile.org. Otte, D. (2007) New species of Cardiodactylus from the western Pacific region (Gryllidae: Eneopterinae). Proceedings of the Academy of Natural Sciences of Philadelphia, 156, 341 400. Ragge, D.R. & Reynolds, W.J. (1998) The songs of the grasshoppers and crickets of Western Europe. Harley Books, Colchester, England, x + 591, 1 CD pp. Robillard, T. & DesutterGrandcolas, L. (2004) Phylogeny and the modalities of acoustic diversification in extant Eneopterinae (Insecta, Orthoptera, Grylloidea, Eneopteridae). Cladistics, 20, 271 293. Robillard, T. & DesutterGrandcolas, L. (2004) Highfrequency calling in Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae): an adaptive radiation revealed by phylogenetic analysis. Biological Journal of the Linnean Society, 83, 577 584. Robillard, T. & DesutterGrandcolas, L. (2006) Phylogeny of the cricket subfamily Eneopterinae (Insecta, Orthoptera, NEW LEBINTHUS OF INDONESIA AND SOLOMON ISLANDS Zootaxa 2386 2010 Magnolia Press 47