Cardiodactylus Saussure, 1878

Robillard, Tony, Gorochov, Andrej V., Poulain, Simon & Suhardjono, Yayuk R., 2014, Revision of the cricket genus Cardiodactylus (Orthoptera, Eneopterinae, Lebinthini): the species from both sides of the Wallace line, with description of 25 new species, Zootaxa 3854 (1), pp. 1-104 : 8-11

Genus Cardiodactylus Saussure, 1878

Type species: Cardiodactylus novaeguineae (Haan, 1842)

Diagnosis. Among Eneopterinae and Lebinthini genera, Cardiodactylus is characterized by its average to large size and its long wings in both sexes, while the other genera of the tribe are brachypterous (Robillard 2011b) ( Fig. 1 ). Eyes large and prominent, fastigium narrower than in Lebinthus and Agnotecous , as in Centuriarus . Face slightly higher than wide. Dorsal disk of pronotum trapezoidal, posterior margin straight or slightly bisinuated, especially in Efordi group. Legs long. TI with 2 tympana; inner tympanum covered by a sclerotized expansion, its membrane visible along a small longitudinal slit only; outer tympanum ellipsoidal, transversally plicate. TI with 2 inner and 1 outer apical spurs. TII with 2 inner and 2 outer spurs. FIII strong basally, filiform at apex. TIII serrulated on their whole length, furrowed longitudinally on their dorsal side and with 4 pairs of subapical spurs and 3 pairs of apical spurs; inner spurs long and curved, outer spurs short and straight. Tarsomeres III-1 with 2 dorso-apical spines and a row of spines on outer dorsal edge; rarely with lateral outer spines. Wings longer than abdomen, hind wings longer than forewings, forming a tail of variable length. Cerci thin, longer than FIII.

Male. Metanotal glands absent. Stridulum complete ( Fig. 2 ): Harp longer than wide, with 1–2 w-shaped harp veins (obliques), delimiting a raised triangular area; posterior area of harp raised along the diagonal vein. In Novaeguineae group, FW coloration pattern usually more contrasted than in Efordi group, posterior angle of harp whitish and anterior region translucent, with different yellow or whitish areas around chords and posterior to mirror. Mirror not rounded but differentiated from other cells of D alignment; d2 cell wide. CuP generally absent. Apical field involving 3 or more cell alignments posterior to the mirror. FW lateral field: Sc with several projections along its length; with variable number of more ventral veins. Subgenital plate clog-shaped; inner side with paired subapical swellings.

Male genitalia. ( Fig. 3 ) Pseudepiphallus [epiphallus according to Randell’s (1964) terminology] setose laterally and ventrally, narrowed at mid-length, forming lateral shoulders delimiting the anterior part, which is generally wider than the posterior part, characterized by the presence of a median process of variable size and shape, with pair longitudinal dorsal ridges developed in Novaeguineae species group, generally independent from each other and sometimes asymmetrical, but fused together in C. novaeguineae and a few closely related species. Pseudepiphallic median process most often terminated posteriorly by a spoon-shaped apex more or less rounded, with a rounded translucent region. Structure including the dorsal ridges and the spoon-like apex always present in the genus, also present, although less developed in most species of Agnotecous (except a few species in Yahoue species group), but lacking in most Lebinthus species and in Centuriarus . Pseudepiphallic parameres [ectoparameres] with characteristic trilobate shape, the posterior lobe narrow and oriented externally. Anterior region of pseudepiphallic sclerite terminated with latero-anterior expansions usually triangular, the membrane between them setose in Novaeguineae species group, except in C. novaeguineae where these expansions delimit a membranous sac ( Robillard & Ichikawa 2009). In Efordi group, the whole anterior region of pseudepiphallus is sclerotized and constitutes a wide dorsal concavity. Rami generally strong, with a wide preapical plate and a thin apical stem. Ectophallic apodemes [endoparameral apodemes] parallel and long, their base sometimes with anterior and posterior expansions. Ectophallic arc [endoparameres] well sclerotized, generally curved. Ectophallic fold [rachis] short, with thin lateral sclerites, apex trilobate, the median lobe sponge-like. Endophallic sclerite [formula] comprising a short median sclerotization and 2 lateral arms of variable size; endophallic apodeme [apodeme of formula] most often with a dorsal crest and lateral lamellas.

Female. FWs with 7–10 strong longitudinal veins. Sc with several projections along its length; with variable number of more ventral veins. FW coloration variable across species, usually brown with patterns of yellow patches. Ovipositor laterally flattened, apex lanceolate (see Fig. 13 ), often denticulate on dorsal edge as in Agnotecous and Lebinthus species ; denticulation also on ventral edge of ovipositor apex in most species of Novaeguineae group.

Female genitalia (see Fig. 14 ). Copulatory papilla with a wide ovoid basal part with pair basal sclerites and a narrowed apex generally rounded.

Classification

Species group Novaeguineae Otte, 2007a

Diagnosis. Species of average to large sizes, usually with more contrasted and vivid coloration patterns on FWs and pronotum, and shorter hind wings in both sexes than in Efordi group. Male genitalia with dorsal pseudepiphallic longitudinal ridges variable amongst species, but generally forming a wide gutter with a rounded spoon-shaped posterior apex; anterior apex of pseudepiphallus generally with pair triangular expansions surrounding a setose membrane.

Species group Efordi Otte, 2007a

Diagnosis. Species of small to average sizes, more slender and usually with little contrasted brownish coloration, and longer hind wings in both sexes than in Novaeguineae group. Male genitalia with a very narrowed posterior part variable amongst species; anterior apex of pseudepiphallus sclerotized and constitutes a wide dorsal concavity.

List and distribution of Cardiodactylus species per island [country] in the region under study ( Fig. 4 )

All the species in the region around the Wallace line (west of New Guinea Island) belong to the Novaeguineae species group. Species in the studied area but not directly considered here are mentioned in brackets, see corresponding references for more details.

Figures

FIGURE 1. Habitus of Cardiodactylus oeroe Robillard, n. sp. Scale bar: 5 mm.

FIGURE 2. Male FW venation. Male forewing venation of Cardiodactylus and terminologies used in the systematic part. The list of abbreviations for veins and cells is given in section materials and methods. Dorsal field (A) with corresponding lateral field (B) of C. enkraussi Otte, 2007 exemplifies the Efordi species group; dorsal field (C) of C. tankara Robillard, 2009 examplifies the Novaeguineae species group (figures modified after Robillard 2009).

FIGURE 3. Male genitalia: Typical structure of male genitalia of Cardiodactylus (species group Novaeguineae) in dorsal (A), ventral (B) and lateral (C) views, showing the terminology used in the descriptions below. Terminology used by TR is mentioned first and terminology by AG is mentioned in brackets. Abbreviations, see section Material and methods.

FIGURE 4. Distributions of Cardiodactylus species around the Wallace line. The species’ distributions are hypothesized broadly according to the material examined and type localities. The Walace line is represented in red.

FIGURE 13. Ovipositor of Cardiodactylus species: Cardiodactylus borneoe Robillard & Gorochov, n. sp. (A); C. buru Gorochov & Robillard, n. sp. (B); C. erniae Robillard & Gorochov, n. sp. (C); C. floresiensis Robillard, n. sp. (D); C. fruhstorferi Gorochov & Robillard, n. sp. (E); C. halmahera Gorochov & Robillard, n. sp. (F); C. kondoi Otte, 2007 (G); C. kotandora Robillard, n. sp. (H); C. lampongsi Robillard & Gorochov, n. sp. (I); C. loboe Robillard, n. sp. (J); C. lombrinjani Robillard, n. sp. (K); C. muiri Otte, 2007 (L); C. muria Robillard, n. sp. (M); C. obi Gorochov & Robillard, n. sp. (N); C. oeroe Robillard, n. sp. (O); C. pelagus Otte, 2007 (P); C. sumba Robillard, n. sp. (Q); C. tello Robillard, n. sp. (R); C. variegatus Gorochov & Robillard, n. sp. (S). Scale bar = 1 mm.

FIGURE 14. Female copulatory papilla of Cardiodactylus species in ventral (left), dorsal (middle) and lateral (right) views: C. borneoe Robillard & Gorochov, n. sp. (A); C. buru Gorochov & Robillard, n. sp. (B); C. erniae Robillard & Gorochov, n. sp. (C); C. floresiensis Robillard, n. sp. (D); C. fruhstorferi Gorochov & Robillard, n. sp. (E); C. halmahera Gorochov & Robillard, n. sp. (F); C. kondoi Otte, 2007 (G); C. kotandora Robillard, n sp.(H); C. lampongsi Robillard & Gorochov, n. sp. (I); Cardiodactylus loboe Robillard, n. sp. (J). Scale bar: 1 mm.