Article. New Cardiodactylus species from unsuspected places in Southeast Asia (Orthoptera, Grylloidea, Eneopterinae)

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1 Zootaxa 2909: (2011) Copyright 2011 Magnolia Press Article ISSN (print edition) ZOOTAXA ISSN (online edition) New Cardiodactylus species from unsuspected places in Southeast Asia (Orthoptera, Grylloidea, Eneopterinae) TONY ROBILLARD Muséum national d'histoire naturelle, Département Systématique et Evolution, UMR 7205 CNRS OSEB, CP 50 (Entomologie), Paris Cedex 05, France. Abstract Two new species of Cardiodactylus crickets are described from unsuspected places in Southeast Asia: C. singapura n. sp., from within the islandcity of Singapore, and C. thailandia n. sp. from Thailand. Descriptions focus on male and female genitalia, forewing venation and coloration. Data are provided for C. singapura n. sp. finescaled habitat and calling song. Consequences in terms of distribution of the genus and conservation are discussed. Key words: Eneopterinae, Lebinthini, Cardiodactylus, Southeast Asia, Singapore, Thailand, new species Introduction The genus Cardiodactylus is part of the tribe Lebinthini, a speciose and diverse cricket tribe of the subfamily Eneopterinae, which is widely distributed from Southeast Asia to many archipelagos in the Western Pacific, to Northern Australia and New Caledonia (Robillard and DesutterGrandcolas, 2004a, 2006, 2008; Robillard, 2010). This genus occupies an important part in the acoustic diversity of Eneopterinae (Robillard & DesutterGrandcolas, 2011). Based on the analysis of the frequency spectra of the calling song, previous studies showed that Cardiodactylus species use highfrequency calling songs (Robillard & DesutterGrandcolas 2004b), with a dominant frequency corresponding to the third harmonic of their song spectrum (Robillard et al. 2007). In terms of species richness, Cardiodactylus has recently grown by a factor of about 500% (Otte, 2007: 39 new species; Robillard, 2009: 4 new species; Ma & Zhang, 2010: 1 new species), but the continued fieldwork and museum studies indicate that the diversity of this genus still remains undescribed, while its precise distribution boundaries have to be determined. In this study two new species of Cardiodactylus are described from unsuspected places in Southeast Asia: C. singapura n. sp. from within the islandcity of Singapore, and C. thailandia n. sp. from Thailand, far north from the previously hypothesized northern limits of the genus distribution. Descriptions focus on male and female genitalia and forewing venation. Based on recent fieldwork in Singapore followed by laboratory studies, the life habits and calling song of C. singapura n. sp. are described. Consequences in terms of distribution of the genus and conservation are discussed. Material and methods Material. Fieldwork was made in several localities in Singapore in June and July Specimens were collected by sight only, by night and day, in order to observe their habitat and type of activity. Most specimens of C. singapura were collected as juveniles or subadults and kept in captivity until adulthood. Newly collected specimens are deposited in the collections of the Zoological Reference Collection of Singapore, in Osaka Museum of Natural History and in the Muséum national d Histoire naturelle, Paris. In the lists of material, square brackets are used for additional information not mentioned on specimen labels. 14 Accepted by D. Rentz: 4 May 2011; published: 8 Jun. 2011

2 Observations and morphological analysis. Direct observations and dissections have been made using a binocular microscope Leica MZ16 at magnifications up to 160, equipped with a camera lucida for the line drawings. Photographs of head, FW venation and male genitalia have been done using a binocular microscope Leica MZ12 and the montage software Leica Application Suite ver (Leica Microsystems). Male tegminal venation. Male tegminal veins and cells (Figs 2D, 7B) are named after DesutterGrandcolas (2003) and Robillard & DesutterGrandcolas (2004a). 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. Photographs of male genitalia have been done after coloration using blue artist acrylic ink (non permanent coloration). Scan electron microscopy. Forewing have been cut off from dry mounted specimens, metalized (20 s, then 30 s) and observed with a JEOL JSM840 electronic microscope (15kV). SEM observations were performed at the Service de Microscopie électronique of the MNHN. 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. C. singapura specimens have been recorded in the laboratory from specimens collected in the field as juveniles or subadults. 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). Correlation between emitted sounds and FW movements were established using an AOS SPri high speed camera (AOS Technologies) at 1250 frames per second. 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. All recording files are deposited in the Sound Library of the Muséum national d Histoire naturelle, Paris. Abbreviations Descriptions General morphology: I, II, III, front, median, hind (femora, legs, tibiae); F, femora; FW, forewing; Tarsomere III1: basal segment of hind leg tarsomere; T, tibiae. Male genitalia: ec arc, ectophallic arc; ec ap, ectophallic apodeme; en ap, endophallic apodeme; en s, endophallic sclerite; ps p, pseudepiphallic paramere; 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; CARDIODACTYLUS FROM SINGAPORE AND THAILAND Zootaxa Magnolia Press 15

3 c13, first to third cells of C alignment; d1 cell (mirror), first cell(s) of D alignment; d2, second cell of D alignment; e1, first cell of E alignment; ha, harp area; Measurements FIIIL, length of hind femora; FIIIW, width of hind femora; FWL, forewing length; FWW, forewing width (at the level of maximal width); HWT, Hind wing tail length (part of the hind wings longer than the FWs); Ias, inner spines on TIII dorsal side, above the spurs; Ibs, inner spines on TIII dorsal side, between the spurs; Oas, outer spines on TIII dorsal side, above the spurs; Obs, outer spines on TIII dorsal side, between the spurs; OL, ovipositor length; PronL, pronotum length; PronW, pronotum width; ST, number of stridulatory teeth; Tt, teeth on transverse section of the file; Lt, teeth on longitudinal section of the file; Bt, teeth on basal section of the file; TIIIL, length of hind tibiae; TaIIIs, Spines on outer edge of third hind tarsomere, not including the apical spine; Institutions MNHN ZRC OMNH Muséum national d'histoire naturelle, Paris, France. Zoological Reference Collection, Department of Zoology, University of Singapore, formerly Raffles Museum. Osaka Museum of Natural History, Osaka, Japan. Systematic part Genus Cardiodactylus Saussure, 1878 Type species: Cardiodactylus novaeguineae (Haan, 1842) Novaeguineae species group Otte, 2007 Cardiodactylus singapura Robillard, sp. nov. (Figs 1 5) Type material. Holotype male: Singapore. Central Catchment Nature Reserve, 01 22'58.3''N '50.3''E, 37 m, 4.VII.2009, nuit, adulte en élevage, enregistrement appel (F0male 4), T. Robillard (ZRC). Allotype female: Singapore. Bukit Timah, 01 21'01.0''N '44.3''E, 95 m, VII.2009, adulte en élevage, T. Robillard (ZRC). Paratypes: Singapore. MacRitchie Reservoir, 01 21'10.2''N '32.2''E, 36 m, 2.VII.2009, nuit, adultes en élevage, T. Robillard: 2, enregistrement appel (F0males 12) (MNHNENSIF ); 1 (MNHN ENSIF2756); 1 (ZRC). Central Catchment Nature Reserve, 01 22'58.3''N '50.3''E, 37 m, 4.VII.2009, nuit, 16 Zootaxa Magnolia Press ROBILLARD

4 adultes en élevage, T. Robillard: 1, enregistrement appel (F0male 3) (MNHNENSIF2757); 1 (MNHN ENSIF2758), 1, 1 (ZRC). Bukit Timah, 01 21'08.4''N '43.8''E, 116 m, VII.2009, nuit (8:30 PM), T. Robillard: 1, adulte en élevage, enregistrement appel (F0male 6) (MNHNENSIF2759); 1 (TR39), sur plante de sousbois (feuille, h=1.5 m), 2, adultes en élevage, (ZRC). Singapore [no precision], 1, Coll. Baker, (MNHNENSIF2753). Singapore [no precision], VII2009, T. Robillard, nuit, adulte en élevage, 1, enregistrement appel (F0male 5) (MNHNENSIF2760) Type locality. Singapore, Central Catchment Nature Reserve, 01 22'58.3''N '50.3''E, 37 m. Etymology. Named after the type locality. Other material examined. Singapore. Bukit Timah, 01 21'08.4''N '43.8''E, 116 m, 1.VII.2009, nuit, T. Robillard: 1 juvenile (MNHN). Central Catchment Nature Reserve, 01 22'58.3''N '50.3''E, 37 m, 4.VII.2009, nuit, T. Robillard: 1 juvenile (MNHN). Singapore [no precision], reared specimens (F1 generation, TR 2010): 5, 2 (MNHN). FIGURE 1. Cardiodactylus singapura n. sp. Male on top of a leaf in Bukit Timah (A, B); juvenile molting in Bukit Timah, showing the transition between the early black coloration to the later brown coloration (C); juvenile (later brown coloration) on top of a leaf in Bukit Timah (D). Distribution. Singapore island. Diagnosis. Species small with contrasted coloration, close to C. pelagus Otte, 2007 from Borneo, from which it differs by the shape of the pseudepiphallus in male genitalia and lighter colour pattern. Description. Size small for the species group. General coloration contrasted, light brown with dark brown patterns. Head dorsum yellow brown with 4 wide dark brown bands, more or less punctuated, bordered by dark brown triangles posterior to eyes and a yellow line more externally (Figs 1A, 2A). Fastigium mottled with dark brown. CARDIODACTYLUS FROM SINGAPORE AND THAILAND Zootaxa Magnolia Press 17

5 FIGURE 2. Cardiodactylus singapura. Dorsal (A) and facial (B) views of head coloration; first instar juvenile with black body coloration (C); male FW venation (D); ventral SEM view of stridulatory file on vein 1A (E), with details of stridulatory teeth in the median region (F). Scale bars: A E, 1 mm; F, 10 μm. 18 Zootaxa Magnolia Press ROBILLARD

6 FIGURE 3. Cardiodactylus singapura. Dorsal (A), ventral (B) and lateral (C) views of male genitalia. Scale bar: 1 mm. Scapes light brown with a transverse dark brown line anteriorly (Figs 1A, 2B). Antennae heterogeneously brown. Face yellowish to light brown, diversely mottled with brown, with 2 parallel lines on the fastigium, 4 dark spots on face, and 2 dark brown spots ventral to eyes. Mouthparts yellow brown. Maxillary palpi yellowish, dark brown apically. Head lateral side with a dark brown area posterior to eyes, progressively lighter ventrally. Pronotum: Dorsal disk trapezoidal, posterior margin straight or slightly bisinuated; light brown mottled with greyish brown and dark brown spots; anterior corners yellowish. Lateral lobes dark brown, ventral edges yellow. Legs light brown to yellowish brown, femora mottled with dark brown; tibiae more or less banded; tarsomeres with dark brown ends. Tarsomeres III1 with 2 3 spines on dorsoexternal edges (m=2.2, n=10). Hind wings longer than FWs, the dark brown tail exceeding the forewings twice longer than the pronotum. Cerci light brown with black rings. Tergites light brown mottled with black. Male: FW coloration (Figs 1B, 2C): Dorsal field cells dark brown, veins mostly orange brown, except for whitish areas in the chords, part of anal veins and in the region posterior to mirror. Base of chords with a brown sclerotization. Harp posterior angle with a whitish sclerotized area. Lateral field dark brown; veins MA, MP, R orange brown, transverse veins and projections of R whitish. FW venation: 1A slightly bisinuated; stridulatory vein (Figs 2C, 2D) with teeth (Fig. 2E) (m=164, n=4), distributed on both transverse ( teeth, m=133) and longitudinal (15 20 teeth, m=19) parts of 1A, plus a group of 9 16 teeth near base of 1A (m=13, n=4). CuP missing. Harp longer than wide, with 2 wshaped harp veins; posterior margin raised along diagonal vein. Mirror area variable, c1 long and narrow, c2 quite large; mirror (d1) longer than wide, not rounded, generally separated in two parts by a transverse vein, the posterior part triangular; e1 cell very elongated, along the mirror. Apical field long and triangular, with 3 wide cell alignments posterior to mirror and a narrow apical alignment. Lateral field: R with 6 7 projections (m=6, n=5). Subgenital plate yellow brown, the median area dark brown. CARDIODACTYLUS FROM SINGAPORE AND THAILAND Zootaxa Magnolia Press 19

7 FIGURE 4. Cardiodactylus singapura. Female FW venation (A); female copulatory papilla in dorsal (B), ventral (C) and lateral (D) views; apex of ovipositor (E). Scale bars: 1 mm. Male genitalia (Fig. 3): Pseudepiphallus very sclerotized, little setose, slightly narrowed at midlength. Dorsal ridges parallel, forming a wide gutter, flattened anteriorly, their external edges carenated. Pseudepiphallic sclerite with lateroanterior expansions, but without a membranous sac as in C. novaeguineae; membrane between the expansions with short strong setae. Apex of pseudepiphallus spoonlike, not rounded, and slightly indented. Rami narrow, with wide preapical plates. Pseudepiphallic parameres trilobate, the posterior lobe wide. Ectophallic arc complete, with a short posterior expansion. Ectophallic apodemes thin, divergent. Apex of ectophallic fold almost hidden by pseudepiphallic parameres, trilobate, membranous. Base of ectophallic apodemes with short sclerotized expansions oriented posteriorly. Endophallic sclerite small. Endophallic apodeme with both lateral lamellas and a short mediodorsal crest. Membrane of endophallic cavity smooth. Female: FW coloration: cells dark brown, veins orange brown, with whitish sclerotization on distal half of CuA and MP, including cells between CuA and MA. MA and R orange brown, R bifurcations whitish. Anterior part of FW with variable number of whitish transverse veins. FW venation (Fig. 4A): 9 11 (m=10, n=5) strong longitudinal veins on dorsal field; lateral field with 9 11 (m=9.5, n=5) longitudinal veins including 5 6 R bifurcations and 3 4 ventral veins. Ovipositor: Shorter than hind femora; apex with both dorsal and ventral edges denticulate (Fig. 4E). Female genitalia (Fig. 4B D): Copulatory papilla triangular with basolateral sclerotizations; apex rounded and folded ventrally. Juvenile: First instar after hatching observed in laboratory, similar in coloration to that of C. novaeguineae, yellow with brown longitudinal bands. Later instars with more contrasted coloration, body black, legs and head yellow brown (Fig. 1C 2C). Black coloration reverses in subadults, characterized by a more homogeneous brown coloration (Fig. 1C D). Measurements. See Table Zootaxa Magnolia Press ROBILLARD

8 TABLE 1. Measurements of Cardiodactylus singapura n. sp. Male holotype Males (n=5) (Mean) Female allotype Females (n=5) (Mean) PronL PronW FWL FWW HWT FIIIL FIIIW TIIIL (2.4) ( (4) (4.3) (13.7) (14.3) (4.2) (4.1) (4.9) (5.1) (14.3) (14.9) (3.9) (4.1) (12.5) (13) continued. Male holotype Males (n=5) (Mean) Female allotype Females (n=5) (Mean) TIIIs TaIIIs ST OL Ias Ibs Oas Obs Tt Lt Bt (8.8) (8) 6 6 (6) (6.4) (11.8) (11.8) 7 7 (7) (6.4) (2.2)??? (10.2) FIGURE 5. Cardiodactylus singapura. Calling song: oscillogram of 5 syllables (A), oscillogram (B) and sonagram (C) of a syllable; power spectrum (D). Symbols: f1, f2, f3 correspond to the first, second and third harmonic frequencies in the spectrum. CARDIODACTYLUS FROM SINGAPORE AND THAILAND Zootaxa Magnolia Press 21

9 Habitat and life history traits. Cardiodactylus singapura is a nocturnal species living in the forested areas. Males produce their discreet calling songs at night, from low branches and leaves (Fig. 1). Juveniles are found day and night in the leaf litter and on leaves of low plants. Behavior. Calling song (Fig. 5): High speed video observations at 1250 frames per second confirm that each calling song corresponds to only one FW closure, the gaps within the syllable being made by the jerking movement of the FWs. At 26 C, this monosyllabic calling song (TRmale1F0male: MNHNENSIF2754) lasts for 87.0 ± 11.8 ms with a period of 8.2 ± 2.2 s and a duty cycle of 1.06%. The dominant frequency is 19.0 ± 0.7 khz and corresponds to the third frequency peak of the song. Cardiodactylus thailandia Robillard, sp. nov. (Figs 6 7) Type material. Male holotype: Thailand. Khaoyai [Khao Yai] National Park, about 14.VII.1986 (M. Takeda) (OMNH). Type locality. Thailand. Khao Yai National Park Etymology. Named after the type locality. Distribution. Thailand. Diagnosis. Species of average size for the genus, with very contrasted coloration, close to C. pelagus Otte, 2007 from Borneo and C. guttulus (Matsumura, 1913) from Japan, with differences on colour pattern and the shape of the pseudepiphallic sclerite and parameres and in male genitalia. Description. Average size for the species group. General coloration very contrasted, orange brown, dark brown and whitish brown. Head dorsum yellow brown with 4 wide dark brown bands, median ones punctuated, fused with lateral ones near fastigium; bordered by dark brown triangles posterior to eyes (Figs 6, 7A). Fastigium dark brown. Scapes and antennae homogeneously yellow brown. Face and mouth parts almost homogeneously yellow brown, slightly mottled with brown, with 2 dark spots ventral to antennae. Maxillary palpi yellowish, dark brown apically. Head lateral sides with a wide dark brown area posterior to eyes. Pronotum: Dorsal disk trapezoidal, posterior margin straight; yellow brown laterally, median area mottled with dark brown. Lateral lobes dark brown, ventral edges yellow. Legs homogeneously orange brown. Tarsomeres III1 with 3 spines on dorsoexternal edges (n=1). Hind wings longer than FWs, the dark brown tail exceeding the forewings one and a half longer than the pronotum. Cerci yellow brown basally then gray brown with dark brown spots. Tergites yellow brown mottled with black. Male: FW coloration very contrasted (Figs 6, 7B): Dorsal field cells and veins brown, some areas dark brown, black, yellow or translucent. Yellow areas include: bases of anal veins, basal 3/4 of CuA, part of 1A bearing the stridulatory file, harp veins, chords, part of diagonal vein, mirror accessory veins, large rounded area posterior to mirror, area between CuA and MP. Posterior corner of harp whitish. Translucent areas include anterior region of harp and cells posterior to mirror. MP/R area red brown. Area ventral to R black. Lateral field veins: MP black; MA red brown on first half, then black; R black; R projections yellow, their bases black. FW venation (Fig. 7B): 1A slightly bisinuated; CuP missing. Harp longer than wide, with 2 wshaped harp veins; posterior margin raised along diagonal vein. Mirror area: c1 long and narrow, c2 quite large; mirror (d1) longer than wide, not rounded, separated in two parts by a strong transverse vein, the posterior part almost rectangular; d2 wide, prolonging the mirror, crossed by many accessory veins; e1 very elongated, along the mirror. Apical field short, with 3 cell alignments posterior to mirror. Lateral field with 8 projections of R and 4 more ventral veins. Subgenital plate yellow brown. Male genitalia (Fig. 7C F): Pseudepiphallus very sclerotized, little setose, clearly narrowed at midlength. Two parallel dorsal ridges, forming a narrow gutter, their dorsal surface beanshaped and setose, slightly asymmetrical, the left ridge curved externally, the right one straight. Pseudepiphallic sclerite with lateroanterior expansions, but without a membranous sac as in C. novaeguineae. Apex of pseudepiphallus spoonlike, not indented. Rami strong, with large preapical plates. Pseudepiphallic parameres trilobate, the posterior lobe pointed and divergent. Ectophallic arc complete, with a triangular posterior expansion. Ectophallic apodemes wide, slightly divergent; base of apodemes with sclerotized expansions oriented posteriorly. Apex of ectophallic fold membranous and trilobate. Endophallic sclerite with long posterior arms. Endophallic apodeme with both lateral lamellas and a long mediodorsal crest. Membrane of endophallic cavity smooth. 22 Zootaxa Magnolia Press ROBILLARD

10 FIGURE 6. Cardiodactylus thailandia. Habitus drawing (male holotype). Scale bar: 5 mm. TABLE 2. Measurements of Cardiodactylus thailandia n. sp. PronL PronW FWL FWW HWT FIIIL FIIIW TIIIL Male holotype continued. TIIIs TaIIIs ST OL Ias Ibs Oas Obs Tt Lt Bt Male holotype ??? CARDIODACTYLUS FROM SINGAPORE AND THAILAND Zootaxa Magnolia Press 23

11 Female: unknown. Juvenile: unknown. Measurements. See Table 2. Habitat and life history traits. Unknown. Behavior. Unknown. FIGURE 7. Cardiodactylus thailandia. Dorsal view of head coloration (A); male FW venation (B); Lateral (C), dorsal (D) and ventral (EF) views of male genitalia. Scale bars: 1 mm. 24 Zootaxa Magnolia Press ROBILLARD

12 Discussion The cricket genus Cardiodactylus proves more and more diverse and widely distributed as the exploration of biodiversity continues in archipelagos of the West Pacific and in Southeast Asia. Several recent studies have accumulated new species and species observations (Otte, 2007; Robillard, 2009; Robillard & Ichikawa, 2009; Ma & Zhang, 2010). The originality of the present study is to describe two new species from places where these crickets were not expected. The species Cardiodactylus thailandia n. sp. from Thailand is interesting as its discovery significantly extends the geographical boundaries of the total distribution of Cardiodactylus on the continental part of Southeast Asia. According to this information, the genus is expected in countries which are still little known for their cricket faunas, such as Vietnam, Laos, Cambodia, Thailand and Malaysia. Cardiodactylus singapura n. sp. is a discreet species from Singapore, inhabiting the most preserved pieces of forests within the city: it seems common at night in Bukit Timah Nature Reserve, in the Central Catchment Nature Reserve and in the forest surrounding McRitchie Reservoir. It was not found in the more anthropized nature areas, where other Eneopterinae species occur, such as in the Labrador Park, or in the islands of Pulau Ubin and Sentosa, despite species such as Nisitrus vittatus Haan, 1842 and/or Lebinthus bitaeniatus Stål, 1877 are common (Tan, 2010; Robillard pers. obs.). C. singapura is currently endemic to Singapore, although it is likely to be found in appropriate preserved habitats in neighbour countries, from where no material could be studied yet. Because of its ecological preferences apparently more restricted than other eneopterine species in Singapore, C. singapura may represent a good candidate as an ecological indicator species of Singapore forests (Noss, 1990), as far as orthopteran insects are concerned. Acknowledgements I thank Akihiko Ichikawa (Japan) for sending me the Cardiodactylus specimen from Thailand (loan from Osaka Museum, OMNH). The field work in Singapore was done in collaboration with Frédéric Legendre (MNHN) and was funded by the UMR 7205 CNRS OSEB and the ANR project BIONEOCAL (resp. P. Grandcolas). I thank Jonathan Goh (National Parks Board) and Janice Yap (AgriFood & Veterinary Authority of Singapore) for permitting field work in Singapore and exportation of material, and Lua Hui Kheng (Raffles Museum of Biodiversity Research, National University of Singapore) for allowing us to examine the cricket specimens in the Raffles Museum of Biodiversity. I also thank Rudolf Meier (University of Singapore) for advising us about the places to prospect in Singapore, and Andie Ang (University of Singapore) for guiding us in Central Catchment Nature Reserve. I thank Guy Lecorvec (MNHN) for preparation specimens, and Laure DesutterGrandcolas and Jérémy Anso for the SEM picture of the file of C. singapura. The habitus of C. thailandia (Fig. 6) was drawn by Gilbert Hodebert (MNHN). References De Haan, W. (1842) Bijdragen tot de Kennis de Orthoptera. In: Temminck, K.J. (Ed.) Verhanlingen over de Natuurlike Geschiedenis der Nederlandsche Overzeesche Bezittingers. Natuurkuundige Commissie in Indie, Leiden, 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, DesutterGrandcolas, L. (2003) Phylogeny and the evolution of acoustic communication in extant Ensifera (Insecta, Orthoptera). Zoologica Scripta, 32, Ma, L. & Zhang, Y. (2010) New record of the cricket genus Cardiodactylus Saussure (Orthoptera, Grylloidae, Eneopterinae) from Hainan Island, China with description of a new species. Transactions of the American Entomological Society, 3+4, Noss, R. (1990) Indicators for monitoring biodiversity. A hierarchical approach. Conservation Biology, 4, Otte, D. (2007) New species of Cardiodactylus from the western Pacific region (Gryllidae: Eneopterinae). Proceedings of the Academy of Natural Sciences of Philadelphia, 156, 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. CARDIODACTYLUS FROM SINGAPORE AND THAILAND Zootaxa Magnolia Press 25

13 Robillard, T. (2009) Eneopterinae crickets (Orthoptera, Grylloidea) from Vanuatu. Zoosystema, 31, Robillard, T. (2010) New species of the genus Lebinthus (Orthoptera, Grylloidea, Eneopterinae, Lebinthini) from Indonesia and the Solomon Islands Zootaxa, 2386, Robillard, T. & DesutterGrandcolas, L. (2004a) Phylogeny and the modalities of acoustic diversification in extant Eneopterinae (Insecta, Orthoptera, Grylloidea, Eneopteridae). Cladistics, 20, Robillard, T. & DesutterGrandcolas, L. (2004b) Highfrequency calling in Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae): an adaptive radiation revealed by phylogenetic analysis. Biological Journal of the Linnean Society, 83, Robillard, T. & DesutterGrandcolas, L. (2006) Phylogeny of the cricket subfamily Eneopterinae (Insecta, Orthoptera, Grylloidea, Eneopteridae) based on four molecular loci and morphology. Molecular Phylogenetics and Evolution, 40, Robillard, T. & DesutterGrandcolas, L. (2008) Clarification of the taxonomy of extant crickets of the subfamily Eneopterinae (Orthoptera: Grylloidea; Gryllidae). Zootaxa, 1789, Robillard, T. & DesutterGrandcolas, L. (2011) Evolution of calling songs as multicomponent signals in crickets (Orthoptera: Grylloidea: Eneopterinae). Behaviour, in press. Robillard, T., Grandcolas, P. & DesutterGrandcolas, L. (2007) A shift toward harmonics for highfrequency calling shown with phylogenetic study of frequency spectra in Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae). Canadian Journal of Zoology, 85, Robillard, T. & Ichikawa, A. (2009) Redescription of two Cardiodactylus species (Orthoptera, Grylloidea, Eneopterinae): the supposedly wellknown C. novaeguineae (Haan, 1842), and the semiforgotten C. guttulus (Matsumura, 1913) from Japan. Zoological Science, 26, Specht, R. (2008) AvisoftSASLab version copyright , Avisoft Bioacoustics, Berlin. Available from Stål C. (1877) Orthoptera nova ex Insulis Philippinis. Öfversigt af Kongl. VetenskapsAkademiens Förhandlingar 34, Zootaxa Magnolia Press ROBILLARD