Phalangopsidae crickets from the Indian Region ...

Zootaxa 3444: 1–39 (2012) www.mapress.com / zootaxa/ Copyright © 2012 · Magnolia Press

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ZOOTAXA ISSN 1175-5334 (online edition)

Phalangopsidae crickets from the Indian Region (Orthoptera, Grylloidea), with the descriptions of new taxa, diagnoses for genera, and a key to Indian genera LAURE DESUTTER-GRANDCOLAS1 & RANJANA JAISWARA2 1 Muséum national d'Histoire naturelle, Département systématique et évolution, UMR 7205 CNRS, Case postale 50 (Entomologie), 57 rue Cuvier, 75231 Paris cedex 05, France. E-mail: [email protected] 2 Centre for Ecological Sciences, Indian Institute of Science, Bangalore 560012, India

Table of contents Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 List of Phalangopsidae from the Indian region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Key for Phalangopsidae crickets of continental India . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Cacoplistes Brunner von Wattenwyl, 1873 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Meloimorpha Walker, 1870 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Paragryllodes Karny, 1909 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Arachnomimus Saussure, 1897 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Arachnomimus maindroni (Chopard, 1969), n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Kempiola Uvarov, 1940 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Kempiola subalatus (Chopard, 1970), n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Kempiola flavipunctatus Desutter-Grandcolas n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Phalangopsina Chopard, 1933 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Phalangopsina dubia (Bolivar, 1900) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Phalangopsina bolivari Desutter-Grandcolas n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Phalangopsina (?) chopardi Desutter-Grandcolas n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 Phalangopsina (?) gravelyi Desutter-Grandcolas n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Speluncasina Desutter-Grandcolas n. gen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Speluncasina annandalei (Chopard, 1928), n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 Opiliosina Desutter-Grandcolas n. gen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 Opiliosina meridionalis Desutter-Grandcolas n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 Aspidogryllus Uvarov, 1940 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 Larandopsis Chopard, 1924 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 Luzaropsis Chopard, 1925 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 Zacla Gorochov, 2003 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 Acknowledgment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37

Abstract The Indian region is presently the second region after the Neotropics in terms of diversity of phalangopsid crickets. Yet their study is impeded by the lack of necessary taxonomic tools for taxon identification. In the present paper, all generic diagnoses are clarified, using morphological and genitalic characters; female genitalia are described and illustrated for all genera with known females. New taxa are described from southern India: Kempiola flavipunctatus Desutter-Grandcolas n. sp., Opiliosina meridionalis Desutter-Grandcolas n. gen., n. sp., Phalangopsina bolivari Desutter-Grandcolas n. sp., P. chopardi Desutter-Grandcolas n. sp., P . gravelyi Desutter-Grandcolas n. sp., and Speluncasina Desutter-Grandcolas n. gen. The list of phalangopsid crickets from the Indian Region is updated, and a key to phalangopsid genera proposed. Accepted by D. Rentz: 31 May 2012; published: 29 Aug. 2012

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A lectotype and a paralectotype are designated to fix the name of Phalangopsina dubia (Bolivar, 1900). Opilionacris annandalei Chopard, 1928, previously transferred to the African genus Phaeophilacris Walker, 1871, is transferred to the genus Speluncasina Desutter-Grandcolas n. gen., while Larandopsis jharnae Bhowmik, 1981 and L. newguineae Bhowmik, 1981 described from New Guinea are transferred to the eneopterine genus Lebinthus Stål, 1877. Finally Luzaropsis confusa Chopard, 1969 is removed from its synonymy with L. ferruginea Walker, 1871. Key words: New genera, New Species, Taxonomy, India, Sri Lanka, Orthoptera

Introduction The current data on extant crickets show that the Indian Region is the second richest region after the Neotropics in terms of phylogenetic and biological diversity of the cricket Phalangopsidae family. Available data document a potentially huge diversity of behaviour, habitat and way of life, not mentioning unique phenotypes, such as Cacoplistes Brunner von Wattenwyl, 1873, or the supposedly glandular specialization of Aspidogryllus Chopard, 1933b. Yet, studies on Indian Phalangopsidae are still very scarce; apart from few studies mostly devoted to species found in caves (Sinha 1973, Sinha and Agarwal 1977, Agarwal and Sinha 1987, Sahu et al. 2011), most concern taxonomy. In his Fauna of India, Chopard (1969) synthesized all previous works and proposed keys to genera and species. This huge work is now still in use, because it is highly practical, even though it is far from up-to-date in terms of taxonomy. From an opposite point of view, Chandra et al. (2010) published an updated list of Indian crickets, which sums up another perspective of basic taxonomic knowledge (i.e., species list), but should be completed with other practical tools such as diagnoses and identification keys. This unbalanced state of knowledge impedes modern studies on the evolutionary biology and behavioural ecology of Indian phalangopsids for lack of identification tools. The aim of the present paper is then to propose a synthetic view of Indian phalangopsid taxonomy, in order to facilitate further studies of these insects, in taxonomy or in other fields of biology. In the present paper, generic diagnoses are clarified using morphological characters as much as possible in addition to genitalic ones; new Phalangopsid taxa are described from Southern India; and the list of species from the Indian Region is updated accordingly. A key for Indian genera is proposed, taking into account male and female morphological characters, and biological data are given when available. Cricket taxonomy has been deeply modified with the analysis of male genitalic structures (Chopard 1961, Randell 1964). An additional step, initiated by the seminal paper by Alexander and Otte (1967), has been the study of female genitalia, and most specifically the shape of copulatory papilla, which may have a diagnostic value at genus and species levels. Male and female genitalia are now more and more routinely studied in cricket taxonomy, sometimes at the expense of morphological characters, which may result in unpractical taxonomic works for taxa diagnoses and descriptions. Precise descriptions of male and female genitalia are used here as a complement to morphological features in the definition of genera and description of species.

Material and methods Studied material. The specimens studied here come from the MNHN collections or were collected in Southern India during field work for the CEFIPRA project 3009-1 (PIs L. Desutter-Grandcolas, MNHN and R. Balakrishnan, IISc). They were collected by sight only, by day and during the night, to get habitat information for each individually collected specimen. Descriptions. Forewing venation is named after Desutter-Grandcolas (2003) and Robillard and DesutterGrandcolas (2004). Male and female genitalia have been dissected on water relaxed specimens, cleaned in cold KOH, and drawn using a stereomicroscope Leica MZx12 with a camera lucida. They are preserved in glycerin in a small vial pinned with each examined specimen. Male genitalia are named after Desutter (1987), modified in Desutter-Grandcolas (2003). SEM observations were performed at the Plateforme de Microscopie électronique of the MNHN, using a JEOL-JSM 840 electronic microscope (7kV), after a 60sec gold-coating.

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DESUTTER-GRANDCOLAS & JAISWARA

Abbreviations and symbols Dotted areas represent FW coloured parts, and membranous parts in male and female genitalia. Specimen origin: fn, field number. General morphology: a, s, i, o, apical, subapical, inner, outer TIII spurs; d, m, v, dorsal, median and ventral TIII apical spurs; FW, forewings; HW, hindwings; I, II, III, fore, median, hind (leg, femur, tibia, tarsus, basitarsus). Forewing venation: 1-nA, anal veins 1 to n; CuA, CuP, anterior, posterior cubital veins; MA, MP, anterior, posterior media veins; R, radial vein. Male genitalia: arc, ectophallic arc; ec.a., ectophallic apodeme; ec.f., ectophallic fold; en.s., endophallic sclerite; ps. l., pseudepiphallic lateral part; ps.m., pseudepiphallic median part; ps.s., pseudepiphallic sclerite; ps.p., pseudepiphallic paramere; r., rami; v.v., ventral valves. Measurements (in mm, mean in parentheses): LFIII, length of hind femur; LFW, maximal length of forewings; LOvip, length of ovipositor; LPron, median length of pronotum dorsal disc; LTIII, length of hind tibia; LTar, length of hind basitarsus; mwPron, maximal width of pronotum; wFW, width of forewings at mirror anterior angle; wPron, posterior width of pronotum. Institution: IISc, Indian Institute of Sciences, Bangalore; IM, Indian Museum, Calcutta; MNCN, Museo Nacional de Ciencias Naturales, Madrid; MNHN, Muséum national d’Histoire naturelle, Paris; ZSI, Zoological Survey of India, Calcutta.

List of Phalangopsidae from the Indian region (studied taxa in bold) Arachnomimus Saussure, 1897 A. annulicornis Chopard, 1936 (Sri Lanka) A. bicolor Chopard, 1928 (Sri Lanka) A. brevipalpis Chopard, 1969 (Sri Lanka) A. lepidus Chopard, 1969 (Tamil Nadu) A. maindroni (Chopard, 1969), n. comb. (Coromandel coast) A. nietneri (Saussure, 1870) (Sri Lanka) Aspidogryllus Chopard, 1933 A. singularis Chopard, 1933 (Tamil Nadu) Cacoplistes Brunner von Wattenwyl, 1873 C. derelictus Gorochov, 2003 (Assam) C. indicus Chopard, 1935 (Assam) C. proximus Gorochov, 2003 (India) C. rogenhoferi Saussure, 1877 (Kashmir) Kempiola Uvarov, 1940 K. flavipunctatus Desutter-Grandcolas n. sp. (Karnataka) K. longipes (Chopard, 1924) (Garo Hills) K. shankari Sinha and Agarwal, 1977 (India) K. subalatus Chopard, 1970, n. comb. (Basyar state) Larandopsis Chopard, 1924 L. choprai Chopard, 1924 (Assam) Luzaropsis Chopard, 1925 L. confusa Chopard, 1969 (Sri Lanka), n. erect. L. ferruginea (Walker, 1869) (Sri Lanka) L. henryi Chopard, 1928 (Sri Lanka) L. omissa Gorochov, 2003 (Sri Lanka) Meloimorpha Walker, 1870 M. albicornis (Walker, 1869) (N. Hindusthan) M. cincticornis Walker, 1870 (N India; Tamil Nadu; Maharashtra) M. indicus (Agarwal and Sinha, 1988) (Madhya Pradesh) PHALANGOPSIDAE CRICKETS FROM INDIA

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Opiliosina Desutter-Grandcolas n. gen. O. meridionalis Desutter-Grandcolas n. sp. (Karnataka) 0. squamifera Gorochov, 2003 n. comb (Southern India) Paragryllodes Karny, 1909 P. ceylonicus (Chopard, 1936) (Sri Lanka) P. gravelyi (Chopard, 1928) (Kerala) Phalangopsina Chopard, 1933 P. bolivari Desutter-Grandcolas n. sp. (Southeast Inde) P. (?) chopardi Desutter-Grandcolas n. sp. (Tamil Nadu) P. discifera Gorochov, 2003 (Southern India) P. dubia (Bolivar, 1900) (Tamil Nadu) P. (?) gravelyi Desutter-Grandcolas n. sp. (Tamil Nadu) P. palpata Chopard, 1969 (Tamil Nadu) Speluncasina Desutter-Grandcolas n. gen. S. annandalei (Chopard, 1928), n. comb. (Darjeeling) Zacla Gorochov, 2003 Z. pictipes Gorochov, 2003 (Hindustan) Note. Parendacustes pendleburyi Chopard, 1969, P. latifrons Chopard, 1969 and Arachnomimus microphthalmus Chopard, 1929 from Malaysia, Laranda rogenhoferi Saussure, 1877 from Brazil (Desutter-Grandcolas 1994), Paragryllodes anjani Bhowmik, 1970 (Nicobar islands) and Parendacustes sanyali Bhowmik, 1970 (Nicobar islands), listed by Chopard (1969) and/or Chandra et al. (2010), are not considered here because they are distributed out of the geographical scope of the paper. Also, the Indian species listed in the genus Homoeogryllus Guérin-Méneville, 1844 are transferred to the genus Meloimorpha (see infra and Jaiswara et al. in prep.).

Key for Phalangopsidae crickets of continental India 1. 2.

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3.

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4. 5.

6.

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Both males and females with long FWs covering the whole abdomen. TIII with three pairs of subapical spurs (Meloimorpha), or less (Cacoplistes). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Males and females either apterous, or with short, more or less reduced FWs. TIII with four outer subapical spurs, and three (Paragryllodes) or four inner subapical spurs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Large species, with strong body and legs. Pronotum concave dorsally, its margins deeply carinated. TIII thick with strong latero-dorsal margins; subapical spurs very short; apical spurs widely set apart from each other on both inner and outer sides. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cacoplistes Brunner von Wattenwyl, 1873 Small and thin species, with thin legs. Pronotum margins not carinated, but dorsal disc saddle-like. TIII without lateral carina, quadrangulate in section; subapical spurs long and clearly articulated; apical spurs close to each other on each tibia side. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Meloimorpha Walker, 1870 TIII with three inner subapical spurs. Fastigium very narrow, much more narrow than the scape, and furrowed. Male subgenital plate elongate; FWs short; stridulum: harp crossed by many veins, mirror not well defined (Fig. 41 in Chopard 1928). Male genitalia typical for the genus, with a short, transverse pseudepiphallic sclerite and long ectophallic dorsal valves (DesutterGrandcolas 1999). Female winged, FWs with projecting veins (Fig. 40 in Chopard 1928). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paragryllodes Karny, 1909 TIII with four inner subapical spurs. Fastigium wider and not furrowed. Male apterous or winged; when the stridulum is present, harp crossed by no more than two or three veins; mirror present, reticulate or not. Male genitalia without ectophallic dorsal valves. Females apterous when known (except in Luzaropsis). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 TIII apical spurs: median spurs the longest on both inner and outer sides; on outer side, the median either greatly (Arachnomimus, Zacla?, Fig. 1J) or slightly (Kempiola, Fig. 3C) longer than the dorsal spur. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 TIII apical spurs different . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Both sexes apterous. TI without tympanum. Male genitalia: median process resembling those of Zacla (cf. infra). Female genitalia long and horse-foot shaped, with a thinner basis and a wider, split apex, and with a plicated, basal membrane (Fig. 1G, H). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arachnomimus Saussure, 1897 Male FWs more or less reduced in length, but with a complete stridulum; female apterous. TI with an inner tympanum (except in K. flavipunctatus Desutter-Grandcolas n. sp.). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 TIII serrulation very sparse (except in K. flavipunctatus Desutter-Grandcolas n. sp.); TIII median and dorsal outer apical spurs subequal, the median only slightly longer than the dorsal one (Fig. 3C). Male stridulum reticulated. Male genitalia: pseudepiphallus with a long and narrow median process, and elongate lateral lobes (Figs 3F, 5B). Female copulatory papilla elongate and very flat, entirely sclerotized (Fig. 3D, E, G–I). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kempiola Uvarov, 1940 TIII serrulation strong; TIII median outer apical spur clearly longer than the dorsal one. Male stridulum not reticulated (Fig.



7.

8. 9. 10.

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166 in Chopard 1969). Male genitalia with a short and straight median lobe, slightly shorter than lateral sclerites (Fig. 4.1–3 in Gorochov 2003a). Female genitalia unknown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zacla Gorochov, 2003 TIII apical spurs: dorsal spur the longest on outer side, and dorsal spur the longest on inner side (except in P. (?) chopardi Desutter-Grandcolas n. sp., in which the median is slightly longer than the dorsal). TI without a tympanum. Males apterous or with very short FWs. Females apterous, when known . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 TIII apical spurs: median spur the longest on outer side, and dorsal spur slightly longer on inner side. TI with an inner tympanum, or two tympana. Both sexes winged . . . . . . . . . . . . . . . . Larandopsis Chopard, 1924, Luzaropsis Chopard, 1925 TIII not serrulated. Eyes reduced, as long as the scape. Male first abdominal tergite very large and produced above the other tergites in the shape of a triangular shield. Female unknown . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Aspidogryllus Chopard, 1933 TIII serrulated. Eyes not as reduced. Male first abdominal tergite not modified.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Pronotum distinctly transverse (Figs 7C, G, 10E). Fastigium much wider than the scape (Figs 7G, 10E). . . . . . . . . . . . . . . . . 10 Pronotum less transverse (Fig. 10A). Fastigium slightly more narrow than the scape (Fig. 10B). Male unknown. Female copulatory papilla small and flat, flap-like (Fig. 9O) . . . . . . . . . . . . . . . . . . . . . . . . .Speluncasina Desutter-Grandcolas n. gen. TIII subapical spurs: outers much longer than the inners. Male apterous or with flap-like, largely separate FWs; lateral parts of supra anal plate elongate. Male genitalia resembling those of Phalangopsina, but small, rounded and compact (Fig. 12); pseudepiphallic sclerite median lobe flat; pseudepiphallic parameres having the shape of acute hooks (Fig. 12B, C). Female ovipositor shorter than FIII; copulatory papilla flat, concave, sclerotized only distally (Fig. 11H–J). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Opiliosina Desutter-Grandcolas n. gen. TIII subapical spurs: outers only slightly longer than the inners. Male FWs as long as tergite 1, slightly overlapping; supra anal plate simple. Male genitalia: pseudepiphallic sclerite with a wide median part, more or less reversed overhead, and two long lateral sclerites; pseudepiphallic parameres very large, concave, making a kind of wide wide forceps (Fig. 6D–F). Female ovipositor longer than FIII; copulatory papilla large, sclerotized and elongate (Figs 6G–H, 8F, G), with a longer and thinner distal part (shorter in P. (?) chopardi Desutter-Grandcolas n. sp., see Fig. 8K, L) . . . . . . . . . . Phalangopsina Chopard, 1933

Systematics Cacoplistes Brunner von Wattenwyl, 1873 (Fig. 1A) Cacoplistes Brunner von Wattenwyl, 1873: 169. Cachoplistus Saussure 1877: 325.

Type species. Cachoplistus brunnerianus Saussure, 1877 Other species included. Cacoplistes (Cacoplistes) indicus Chopard, 1935, Cacoplistes (Cacoplistes) proximus Gorochov, 2003, Cacoplistes (Laminogryllus) derelictus Gorochov, 2003, Cacoplistes (Laminogryllus) rogenhoferi Saussure, 1877 and Cacoplistes (Laminogryllus) westwoodianus Saussure, 1877. Distribution. Northern India, Nouvelle Hollande (?). Diagnosis. Large species. Pronotum large, concave dorsally; dorsal disc and lateral lobes with distinctly carinated margins. Eyes small, protruding. Fastigium short and narrow, much narrower than the scape; flat, separated from the rounded vertex by a transverse furrow. Antennae thick. Maxillary palpi short; joint 5 concave dorsally, greatly widened in distal half. Legs relatively short; femora higher than wide, FIII only slightly inflated. TI with an inner tympanum; two short and thick ventral apical spurs. TII with two short and thick ventral apical spurs. TIII flattened dorsally, with strong lateral margins; three apical spurs on each side, widely separated from each other: outer spurs very small, the median the longest; inner spurs longer but short, the median and dorsal subequal; subapical spurs very short hardly distinct from the thick spines. Basitarsomeres all higher than wide; basitarsomeres III strongly furrowed dorsally, with both inner and outer thick spines. Males FWs much longer than the body; stridulatory apparatus complete, the harp with several close, parallel veins, the mirror separated into several wide cells by parallel crossing veins, falsely prolonged by the cell between the mirror and the chords, and circled distally by an empty cell. TIII spurs not modified. Male genitalia (see Fig. II (1–14) in Gorochov 2003b): pseudepiphallic sclerite long and thick, variously membranous dorsally, with a small, subapical median process, and more or less prolonged distally; rami thick and long, separated from pseudepiphallus. Pseudepiphallic parameres made of one or two sclerites, more or less hook-like or spiny. Ectophallic apodemes long and thick; arc thick. Ectophallic fold short, its ventral sclerite bifid basally. Endophallic sclerite with three distal prongs, the lateral ones the longest; endophallic apodeme long and flat. No endophallic cavity. Female FWs longer than the body; strong parallel veins separated by many small cells. Subgenital plate wider than long. Ovipositor apex not widened, without ornamentation. Female genitalia: copulatory papilla nested in a thick membrane (Fig. 1A); small, much wider than long. Habitat. Unknown. PHALANGOPSIDAE CRICKETS FROM INDIA


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Meloimorpha Walker, 1870 (Fig. 1B, C) Meloimorpha Walker, 1870: 468.

Type species. Meloimorpha cincticornis Walker, 1870 Other species included. Phalangopsis albicornis Walker, 1870, Gryllus / Phalangopsis japonicus Haan, 1842, Homoeogryllus indicus Agarwal and Sinha, 1988. Remark. Meloimorpha has long been synonymized with Homoeogryllus Guérin-Méneville, 1844. Actually both genera are extremely similar by their general shape and male stridulum, although they differ markedly by their male (Desutter 1985, Gorochov 2003b) and female (compare Fig. 1B, C and 1D–F) genitalia. Both genera can be readily separated by their number of inner subapical spurs on TIII, equal to 3 in Meloimorpha and 2 or less in Homoeogryllus (Desutter 1985, Jaiswara et al. in prep.). Distribution. From Northern India in the West to Japan in the East. Many photos on the web are identified « Homoeogryllus indicus » while they represent an African cricket. These photos clearly show a Homoeogryllus species, which may be H. xanthographus Guérin-Méneville, 1844 or H. orientalis Desutter, 1985, according to locations in Eastern Africa (Desutter 1985). The species described as “Homoeogryllus indicus” by Agarwal and Sinha in 1988 actually belongs to Meloimorpha, which is not present in Africa. Diagnosis. Size small to medium. Eyes small, protruding. Fastigium narrow, as wide as about half scape width; not separate from the vertex by a distinct transverse furrow. Ocelli not distinct. Maxillary palpi moderately long; joint 5 concave dorsally, slightly widened in apical fourth. Pronotum transverse, greatly widened posteriorly, saddle-shaped. TI with inner and outer tympana; two ventral apical spurs, the inner slightly longer than the outer. TII with two ventral apical spurs, the inner slightly longer than the outer. TIII with three pairs of apical spurs; outer spurs small, the dorsal and median subequal in length; inner spurs longer, the median greatly longer than the dorsal; three pairs of subapical spurs, only very slightly alternate, the outers slightly longer than the inners, and located in TIII distal third; TIII strongly serrulated over their whole length. FIII with a distinct filiform apical part. Colouration often contrasted, head and body brown to black, with some times yellow spots on head and pronotum dorsum; tibiae and antennae white, yellow or light ochre. Males with very long FWs greatly widened from their base, but their lateral margins almost parallel. Stridulatory apparatus complete; harp wider than long, crossed by several transverse, parallel veins; mirror wider than long, subdivided into wide cells by crossing veins, and prolonged by wide cells toward the chords; a thin, continuous cell surrounding the mirror distally. Apical field short, truncated. Lateral field: R with many parallel, transverse veins. TIII spurs not modified. Male genitalia: pseudepiphallic sclerite rounded, surrounding long, hook-like pseudepiphallic parameres; median part of pseudepiphallus more or less membranous; rami thin; ectophallic apodemes thick; ectophallic fold small and membranous; endophallic sclerite with one main, median, part, but bordered laterally by a pair of wide sclerites; endophallic apodeme crest-like. Female FWs longer than the body; longitudinal and transverse veins strong, delimiting wide cells. Female genitalia: Copulatory papilla having the shape of a short, sclerotized tube slightly narrowed before apex, and slightly plicated dorsally; aperture of spermathecal duct ventral, sub apical (Fig. 1B, C). Habitat. Species possibly cavicolous, mentioned in termite nests (Bhagava 1982).

Paragryllodes Karny, 1909 Paragryllodes Karny, 1909: 478.

Type species. Paragryllodes borgerti Karny, 1909



FIGURE 1. Female copulatory papilla (A–H) in: A, Cacoplistes sp, female identified Cachoplistus rogenhoferi Saussure, 1877 by L. Chopard (specimen MNHN-ENSIF2960, Tonkin, Dong Daky), distal view; B–C, Meloimorpha japonica (specimen MNHN-ENSIF2963, Japan, Kyoto), ventral (B), lateral (C); D–F, Homoeogryllus xanthographus Guérin-Méneville, 1847 (specimen MNHN-ENSIF2944, Tchad, Amdjamena), dorsal (D), ventral (E), lateral (F); G–H, Arachnomimus maindroni (Chopard, 1969), n.comb., dorsal (G), lateral (H). Hind tibia apical spurs in: I, Arachnomimus maindroni (Chopard, 1969), n.comb. inner spurs; J, Arachnomimus nietneri (Saussure, 1878), outer spurs. Scale 1 mm.

PHALANGOPSIDAE CRICKETS FROM INDIA


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Other species included. Indian species: Pseudendacustes gravelyi Chopard, 1928. West African species: Paragryllodes fuscifrons Chopard, 1934, Paragryllodes optimus Gorochov, 1996. Central African species: Paragryllodes pictus Chopard, 1958, Paragryllodes makandensis Desutter-Grandcolas, 1999, Paragryllodes makokou Desutter-Grandcolas, 1999, Paragryllodes kessala Desutter-Grandcolas, 1999, Paragryllodes orensis Desutter-Grandcolas, 1999, Paragryllodes deleportei Desutter-Grandcolas, 1999, Paragryllodes bipunctatus Desutter-Grandcolas, 1999, Paragryllodes longixiphus Desutter-Grandcolas, 1999, Paragryllodes centralis Desutter-Grandcolas, 1999. East African species: Paragryllodes minor Desutter-Grandcolas, 1999, Paragryllodes dissimilis Desutter-Grandcolas, 1999, Paragryllodes unicolor Desutter-Grandcolas, 1999, Paragryllodes campanella Desutter-Grandcolas, 1998, Paragryllodes kenyanus Kaltenbach, 1982, Paragryllodes pyrrhopterus Kaltenbach, 1982, Montigryllus silvaepluvialis Sjöstedt, 1910, Montigryllus affinis Sjöstedt, 1910. Species from Madagascar: Paragryllodes annulicornis Kaltenbach, 1982, Paragryllodes annulipes Chopard, 1952, Paragryllodes madecassus Chopard, 1927. Species from the Comoros: Paragryllodes milloti Chopard, 1958 (not 1952; contra Eades et al. 2011). Species from Sri Lanka: Seychellesia ceylonicus Chopard, 1936. Species from the Nicobar islands: Paragryllodes anjani Bhowmik, 1970. Distribution. Tropical Africa, Madagascar, the Comoros islands, India (Kerala), Sri Lanka and the Nicobar islands. Diagnosis. According to Desutter-Grandcolas 1999 (p. 498, and figures therein): Size variable. Eyes strongly protruding. Fastigium very narrow, grooved, sharply separated from the vertex. Ocelli set as a very acute triangle. Scapes large, longer than wide. Maxillary palpi moderatly elongate; joint 5 widened and truncated at apex. TI with an inner tympanum, the outer tympanum lacking; two apical spurs. TII with three apical spurs. TIII with four outer, and three inner subapical spurs; three inner apical spurs, the median and dorsal subequal in length; three outer apical spurs, the median the longest; TIII strongly serrulated, except between first subapical spurs. Cerci very long. Colouration black brown mottled with yellow, legs ringed with yellow and brown. Male metanotum and tergites without glandular structures; TIII spurs not modified; FWs covering only part of the abdomen, always overlapping; stridulum: harp crossed by several oblique, parallel veins, not always clearly separated from the chords; mirror more or less separated from apical venation; apical field always short; posterior angles of supra anal plate not elongate. Male genitalia characterized by well-developed dorsal ectophallic valves; pseudepiphallic sclerite transverse or quadrate, not separate from long rami; pseudepiphallic parameres more or less triangular in shape, not hook-like; ectophallic apodemes long and thick; no dorsal cavity. Female FWs short, not covering the body, slightly or not overlapping; longitudinal veins variably separated by transverse veinlets. Copulatory papilla long and tubular, or shorter and more triangular in shape. Remark. Morphologically, P. gravelyi fits the definition of the genus, but male and female genitalia remain to be described. Habitat. Paragryllodes species live in tropical forests, inhabiting only plain forests (Central Africa), or both plain and gallery forests (East Africa) (Desutter-Grandcolas 1998, 1999). They are typically nocturnal, dendrophilous species, which forage at night on dead trunks (lying on the ground or standing), or on standing trees. They hide during the day in cavities, located either near the ground (species from the Nicobar islands, Bhowmik 1970), or above the ground (African species, Desutter-Grandcolas 1999).

Arachnomimus Saussure, 1897 Arachnopsis Saussure, 1878: 582. Arachnomimus Saussure, 1897: 251. Gorochov, 1996: 49 (redescription of type species, lectotype designation).

Type species. Arachnopsis nietneri Saussure, 1878 Other species included. A. annulicornis Chopard, 1936, A. bicolor Chopard, 1928, A. brevipalpis Chopard, 1969, A. lepidus Chopard, 1969, A. microphtalmus Chopard, 1929 (see Chopard 1969), Phalangopsis amboinensis Karsch, 1886 (see Gorochov 1996), Kempiola maindroni Chopard, 1969, A. aureopubescens Wiendl, 1970, A. istrapura Gorochov, 2003c, A. jacobsoni Chopard, 1924. Distribution. Southern India (Coromandel coast, Kerala), Sri Lanka, Java. One species described from Brazil, probably erroneously. Diagnosis. Size small to large. Head not very long . Eyes greatly protruding (Fig. 2B, except in A.



microphthalmus: eyes flat and reduced). Fastigium much narrower than the scape (Fig. 2B). Ocelli set in an acute triangle (lacking in A. microphthalmus), the distance between lateral ocelli much shorter than the distance between one lateral and the median ocelli. Maxillary palpi very elongate, with joint 5 distinctly longer than joint 3. Pronotum transverse (Fig. 2A, D). Mesonotum short, metanotum as long as the pronotum (Fig. 2A, E). All legs long and thin (Fig. 2A). TI without tympanum. TIII with four pairs of alternate subapical spurs (five pairs mentioned by Chopard in A. bicolor), the outers longer than the inners; three inner and three outer apical spurs, the median the longest on both sides: the inner median spur very long (Fig. 1I), the outer median spur much longer than the dorsal spur (Fig. 1J); TIII serrulated on both sides. Tarsi all long, hind basitarsi with few dorsal spines. Male and female apterous. Male metanotum and tergites lacking glandular structures; TIII spurs not modified. Male genitalia resembling those of Zacla Gorochov, 2003a (median process of pseudepiphallus short and straight, slightly shorter than lateral lobes, rami very short, ectophallic fold sclerotized laterally, endophallic sclerite long, bearing a crest-like apodeme). Female genitalia horse-foot shaped, with a plicated basal membrane, and a thick distal sclerite distinctly furrowed at apex (Fig. 1G, H). Habitat. Unknown.

Arachnomimus maindroni (Chopard, 1969), n. comb. (Figs 1G–I, 2) Kempiola maindroni Chopard, 1969: 254.

Type locality. India, Coromandel coast, Gangi. Type material. Holotype: 1 female, India, Coromandel, Gangi, 1901 (M . Maindron) (MNHN-ENSIF2955) (palpi, PI and right PII missing). Diagnosis. Large species (Fig. 2A) characterized by the colouration of its face (Fig. 2C), pronotum (Fig. 2D, E) and FIII (Fig. 2H), ovipositor length (15 mm) and shape of copulatory papilla (long, socket-like, with a distinctly bifid dorsal sclerotization, Fig. 1G, H). Subgenital and supra anal plates as on Fig. 2F, G. TIII apical spurs as on Fig. 1I (inner side) and 1J (outer side, here in Arachnomimus nietneri). Measurements, see Chopard 1969 (p. 254). Remark. Originally described by Chopard in the genus Kempiola, this species differs from K. longipes by its very distinctive copulatory papilla, identical to that of other Arachnomimus species. The other distinctive characters of this last genus (lack of tympanum, male genitalia) cannot be checked with the specimen at hand, but the female genitalia are so characteristic of the genus that generic identity is clear.

Kempiola Uvarov, 1940 Kempiella Chopard, 1924: 87. Kempiola Uvarov 1940: 175.

Type species. Kempiella longipes Chopard, 1924 Other species included. Kempiola shankari Sinha and Agarwal, 1977 (see Figs 4.5–15 in Gorochov 2003a), Arachnomimus subalatus Chopard, 1970, Kempiola flavipunctatus Desutter-Grandcolas n. sp. Known distribution. West coast of India, Eastearn India, Assam. Diagnosis. Fastigium nearly as wide as the scape. Maxillary palpi very long; joint 5 only slightly widened, in apical 5th only, and distinctly longer than joint 3 (Fig. 3A). Pronotum transverse. TI with a small, inner tympanum (except in K. flavipunctatus Desutter-Grandcolas n. sp.). FIII with a long apical, filiform part. TIII with four pairs of short, subapical spurs; serrulation very sparse (except in K. flavipunctatus Desutter-Grandcolas n. sp.); three inner apical spurs, the median the longest (Fig. 3B); three outer apical spurs, the median as long or slightly longer than the dorsal (Fig. 3C). Male. Metanotum and tergites glandular (Fig. 4C); TIII spurs not modified. FWs short, dorsal and lateral fields separate by a sharp angle (except in K. flavipunctatus Desutter-Grandcolas n. sp.); stridulum complete. Supra anal plate with elongate angles (except in K. flavipunctatus Desutter-Grandcolas n. sp.). Male genitalia: median and lateral processes of pseudepiphallus all elongate (Fig. 3F, 5B–D). Female. Apterous. Ovipositor shorter than FIII. Female genitalia: Copulatory papilla elongate and very flat, entirely sclerotized (Fig. 3D–E, G–I). PHALANGOPSIDAE CRICKETS FROM INDIA


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FIGURE 2. Arachnomimus maindroni (Chopard, 1969), n.comb. A, female holotype, dorsal, scale 5 mm; B, head, dorsal; C, face, front view; D, pronotum, dorsal; E, head, pronotum and first tergites, lateral; F, supra anal plate; G, subgenital plate; H, hind femur, outer side.



FIGURE 3. Kempiola Uvarov, 1940. A–E, Kempiola longipes (Chopard, 1924): A, maxillary palpus; B–C, inner (B), outer (C) apical spurs of hind tibia; D–E, copulatory papilla, dorsal (D), lateral (E). F, Kempiola subalatus (Chopard, 1970), n. comb., male genitalia, dorsal. G–J, Kempiola flavipunctatus Desutter-Grandcolas n. sp.: G–I, copulatory papilla, dorsal (G), ventral (H), lateral (I); male forewing (J), dorsal. Abbreviations: see text. Scales 1 mm.



11

Remark. Kempiola resembles Arachnomimus morphologically, from which it differs by the inner tympanum, TIII outer apical spurs, male FWs, and male and female genitalia. Its TIII serrulation is also very sparse, except in K. flavipunctatus Desutter-Grandcolas n. sp. K. flavipunctatus Desutter-Grandcolas n. sp. shows several morphological differences, which may justify separating it in a different genus. Better knowledge of Indian Phalangopsidae is however necessary to ascertain the definition of monophyletic entities. Habitat. Some species have been found in caves, while others are dendrophilous (see Discussion).

Kempiola subalatus (Chopard, 1970), n. comb. (Fig. 3F) Arachnomimus subalatus Chopard, 1970: 122.

Type material. 1 male juvenile, India, Baryar State, cave near Jugdalpur, 1960, Agarwal (MNHN-ENSIF2922). Type locality. India, Baryar State, cave near Jugdalpur. Remark. Described as a male by Chopard (1970), the type specimen revealed a juvenile, according to FWs and male genitalia. The structure of male genitalia (Fig. 3F), the proportion of TIII outer apical spurs (median and dorsal spurs subequal) and TIII serrulation place this species within the genus Kempiola, even though the presence of a tympanum, the venation of male stridulum and the shape of male supra anal plate cannot be checked in the juvenile type.

Kempiola flavipunctatus Desutter-Grandcolas n. sp. (Figs 3G–J, 4, 5, Table 1) Type locality. India, Karnataka, env. 14 km Karkala, Kervasae Reserve forest. Type material. Holotype: 1 male, 18.vii.2009, India, Karnataka, env. 14km Karkala, Kervasae Reserve forest, ZSI. Allotype: 1 female, same data as the holotype, ZSI. Paratypes. 22 males, 18 females. Same locality as holotype, 18.vii.2009, 4 males, 4 females. Karnataka, env. 14 km Karkala, Forêt de Panemburi, 25.vii.2002, KCW20, 1 male, 1 female, in alcohol; 17.vii.2009, 6 males, 4 females. Karnataka, close to Kuderemukh National Park, 18.vii.2009, 7 males, 5 females. Forêt de Horabi, 16.vii.2009, 4 males, 5 females (MNHN, IISc). Diagnosis. In addition to the characters of the genus, large species with very long legs but a short body (Fig. 4A). Colouration dark brown to black, with many yellow dots (Fig. 4A), especially on pronotum disc (four yellow dots) and tergites 2 – 5 (two lateral yellow flecks on each side, and a median half-circled one). Face with a wide longitudinal yellow band from median ocellus to the tip of labrum, narrowed on upper margins of clypeus and labrum (Fig. 4B); base of cerci and supra anal plate yellow. Male tergite 1 glandular (Fig. 4C). Male stridulatory file with about 25 teeth (Fig. 5A) . Male genitalia: median process wide, narrowed only distally; lateral processes slightly longer, club-shaped; pseudepiphallic paramere ventral plate acute distally, as long as pseudepiphallic median process (Fig. 5B–D). Female: ovipositor well shorter than FIII, about 10 mm in length. Female genitalia: copulatory papilla elongate, resembling a horse foot but distinctly split on distal margin dorsally (Fig. 3G–I). Description. In addition to the characters of the genus: Large species with very long legs but a short body (Fig. 4A). Body covered with dense, short setae. Head. Eyes distinctly protruding. Fastigium nearly as wide as the scape, not separated from the vertex. Ocelli small, located as an isoceles triangle; median ocellus subapical. Scapes longer than wide. Maxillary palpi very long and thin; joints 3 and 4 subequal in length; joint 5 longer than joints 3 and 4, slightly widened in distal fourth, distal margin truncate (Fig. 3A). Pronotum. Very large and transverse; dorsal disc convex; lateral lobes raised dorsally. Legs. All long and very thin. TI without tympanum; two long and thin apical spurs, the inner the longest. TII with two long and thin apical spurs, the inner the longest. FI and FII thinner at mid length than in apical third. FIII filiform in apical half. TIII with four pairs of subapical spurs, the outers longer and located higher on tibia than the inners; three pairs of apical spurs, the median spur the longest on both inner and outer sides; TIII serrulated on both inner and outer margins, except between apical and subapical spurs; spines most often thick and bent over the tibia; inner serrulation: no spine between spurs 1 and 2, one to three spines between spurs 2 and 3 (mean 1.3, n=9 in males, mean 1.9, n=10 in females), three to seven spines



between spurs 3 and 4 (mean 4.1, n=9 in males, mean 4.8, n=10 in females), 10–18 above spurs (mean 12.7, n=9 in males, mean 14.2, n=10 in females); outer serrulation: zero to four spines between spurs 1 and 2 (mean 1.4, n=10 in males, mean 2.2, n=10 in females), five to 11 spines between spurs 2 and 3 (mean 6.7, n=10 in males, mean 7.1, n=10 in females), nine to 14 spines between spurs 3 and 4 (mean 10.8, n=10 in males, mean 12.2, n=10 in females), 14 to 21 spines above spurs (mean 17.2, n=10 in males, mean 17.9, n=10 in females). Tarsomeres 1 very long, longer than half TI and TII; tarsomeres 1–III about one third TIII length; basitarsomeres III with two rows of small spines dorsally: one to three inner spines (mean 2.3, n=9 in males and females), and three to seven outer spines (mean 5.4, n=9 in males, mean 5.5, n=10 in females), in addition to distal spines; tarsomeres 3 about one third tarsomere 1. Terminalia. Cerci very long, more than twice body length (Fig. 4A). Colouration. Black brown with yellow dots and lines. Head: vertex yellow with four brown lines, one close to each eye, and one running from each lateral side of fastigium, through one lateral ocellus, up to occiput; eyes circled with light yellow except on the face; face mottled brown and black brown, with a wide longitudinal yellow line running from median ocellus to labrum tip, a triangular yellow dot under each eye (wider along epistemal suture) and a faint, often hardly distinct, transverse dot below each antennal pit; posterior margin of cheeks brown; maxillari palpi brown; scapes light yellow and brown; antennae brown. Pronotum: black brown, pyriform inscriptions and two rounded dots close to distal margin lighter; two yellow pairs of dots on dorsal disc; anterior and posterior margins sometimes yellow, and the faint median furrow of dorsal disc variously yellow; distal margin of lateral lobes sometimes yellow. Legs yellow and brown, annulated (Fig. 4A); tibiae and tarsomeres light brown, with small yellow dots at the base of TIII subapical spurs; FI and FII yellow or brown at base, with three brown rings more or less prolonged along femur lower margin; FIII with two brown rings on filiform part, two wide brown bands on inner basal half and prolonged obliquely on outer side; a brown longitudinal band bordered with yellow on outer lower basal half. Distal margin of metanotum and tergite 1 yellow, this pattern more clear in females than in males. Tergites black , with a lighter median rounded fleck including a short yellow line on mid line and an elongate yellow spot on lateral side; tergites 1, 2 and 4 with two additional rounded yellow spots on each side; tergites 3, 6 and 7 with the elongate lateral spot very long. Sternites yellowish to light brown; distal margin darker. Terminalia: supra anal plate light brown, the base light yellow; cerci light brown with a light yellow ring at base, more or less prolonged on dorsal and ventral sides. Male. Mesonotum and part of metanotum hidden under the pronotum; metanotum distal margin strongly raised above a large and hairy paired structure on tergite 1; distal margin of tergite 1 strongly bisinuate, with paired hollowed structures overlooked by a small cuticular hook (Fig. 4C). FWs short and not overlapping, FW convex with a surface with strong ultrastructural reliefs; their surface either convex (areas close to inner margin and above the file), or concave (area below the file, ie more distal, between CuA and media veins, and median fan) (Fig. 4D); venation faint, except the veins separating the dorsal and the lateral fields, 1A (file), 3A, diagonal and part of the chords; lateral field coming over the dorsum. Stridulatory apparatus complete but embedded in a network of large, regular cells (Fig. 3J); harp with two veins; diagonal short, oblique; mirror wide, crossed by one irregular, zigzaging vein; chords incomplete (Figs 3J, 4D); stridulatory file with about twenty five broadly aligned teeth (Fig. 5A). Supra anal plate longer than wide; a deep longitudinal, median furrow distally (Fig. 4E); a median triangular concavity in basal third; lateral angles not elongate. Subgenital plate short, quadrate; a deep longitudinal distal furrow (Fig. 4F). Male genitalia. Anterior margin of pseudepiphallus reversed posteriorly, partly covering the pseudepiphallic sclerite (Fig. 5D); pseudepiphallic sclerotization delimiting a strong bottle-shaped median process (wide at base, abruptly narrow before apex; narrow part about one third the total length of the processus), and two long clubshaped lateral lobes longer than the median process (Fig. 5B, C); rami short, thick and curved, connected to pseudepiphallic sclerite; pseudepiphallic parameres with two short plates, one lateral and one dorsal, at the base of a long hook-shaped sclerite nearly as long as pseudepiphallic median process (Fig. 5C, D). Ectophallic apodemes thick, diverging from the arc (Fig. 5B); a thick plate between the pseudepiphallus and the arc (Fig. 5B). Ectophallic fold very short, truncated at apex (Fig. 5C). Endophallic sclerite long and thin, its distal margin with three long prongs (Fig. 5C); no dorsal cavity. Female. Apterous. Distal margin of subgenital plate only slightly sinuate. Ovipositor well shorter than FIII (See Table 1); valves apex not widened and without ornementation. Female genitalia. Copulatory papilla long and well sclerotized, as on Fig. 3G–I. Measurements. See Table 1.



13

FIGURE 4. Kempiola flavipunctatus Desutter-Grandcolas n. sp. A, male holotype, dorsal, scale 3mm; B, face, front view; C, male meso- and metanotum; D, male FWs; E, male supra anal plate; F, male subgenital plate; G–H, hind femur, colouration pattern of inner (G) and outer (H) sides, scale 1mm.



FIGURE 5. Kempiola flavipunctatus Desutter-Grandcolas n. sp. A, male stridulatory file (male fn LD119), SEM image, scale 100 µm; B–D, male genitalia, dorsal (B), ventral (C), lateral (D). Abbreviations, inv, invagination between pseudepiphallic sclerite and epi-ectophallic invagination; others see text. Scale 1 mm.



15

TABLE 1. Measurements (mm) of Kempiola flavipunctatus Desutter-Grandcolas n. sp. Pron-L

Pron-W

Pron-mW

FW-L

FIII-L

TIII-L

Ovip-L

Holotype

3.3

4.5

5.1

2.9

18.6

20.4

----

Males (n=5)

2.8–3.4

3.8–4.9

4.2–5.5

2.1–2.6

15.4–17.9

17.7–19.3

----

Mean (n=6)

3.1

4.3

4.9

2.5

17.2

18.8

----

Allotype

3.3

4.6

5.0

----

17.1

18.7

11.7

Females (n=5)

2.8–3.2

3.9–4.5

4.2–4.7

----

15.4–18.6

16.7–19.1

9.5–11.7

Mean (n=6)

3.0

4.3

4.5

----

16.8

17.9

10.5

Variation. Very dark individuals are almost black and show only the brightest elements of the whole colouration pattern (face dark brown with only the median yellow line and the line under the triangular spot under the eye, pronotum with four yellow spots and a small longitudinal median line, tergites with few yellow dots). Habitat. Kempiola flavipunctatus Desutter-Grandcolas n. sp. has been observed mostly on tree trunks, by day and by night. Contrary to other phalangopsid crickets active on tree trunks at night (Desutter-Grandcolas 1992, 1995), they do not seem to specifically hide in cavities during daytime: 44% were thus found waiting on a trunk, while 56% were found in a kind of shelter, i.e. under raised bark (5%), in a small cavity (7%), in a shallow trough (9%) or most often on the lower side of a lean tree or liana (35%), where several couples were observed without apparent protection. This species was also found at night in small cavities of steep ground slopes along roadsides.

Phalangopsina Chopard, 1933 Phalangopsina Chopard, 1933a: 165.

Type species. Arachnopsis dubius Bolivar, 1900 Other species included. Phalangopsina palpata Chopard, 1969, P. bolivari Desutter-Grandcolas n. sp. and possibly P. discifera Gorochov, 2003a. Phalangopsina (?) chopardi Desutter-Grandcolas n. sp. and Phalangopsina (?) gravelyi Desutter-Grandcolas n. sp. are tentatively described in the genus Phalangopsina. Remark. The type species of the genus has been tentatively described by Bolivar (1900(1899)) in the genus Arachnopsis Saussure, 1878 on a series comprising an unspecified number of males and females, deposited in Bolivar’s and Pantel’s collections (now in MNCN and MNHN respectively), and originating from Maduré, Kodaikanal (collectors: Castets, Décoly). Listing the types of Ignacio Bolivar, Mercedes Paris (1994) identified several putative male and female syntypes of P. dubia: one female identified “Arachnop. Dubius” by Bolivar in Madrid, and one male and 3 females in Paris: 1/ one female labeled “Type // Arachnopsis dubius // Mission Maduré”; 2/ one female labeled “Arachnopsis dubius // Kod[aikanal] Dec[oly]“; 3/ one female labeled “Indes Or., P. Castets”; and 4/ one male, from Kod[aikanal] Dec[oly]. The male specimen revealed a Landrevinae and does not fit Bolivar’s description and measurements: it is not considered here as belonging to the original syntypic series; the male syntype mentioned by Bolivar has not been found yet in Pantel’s collection. The four females are considered here syntypes: in order to fix the species name, two conspecific females are designated here Lectotype (MNCN) and Paralectotype (MNHN) of P. dubia; the other two paralectotype females belong to two new species of Phalangopsina described here, P. bolivari DesutterGrandcolas n. sp. and Phalangopsina (?) chopardi Desutter-Grandcolas n. sp. Known distribution. Southern India. Diagnosis. According to Bolivar (1900), Chopard (1933a) and type observation: Small to very small species with a wide fastigium (wider than scape) and transverse pronotum (less so in P. (?) chopardi Desutter-Grandcolas, n. sp.). Ocelli arranged in a wide triangle, the distance between the lateral ocelli larger than the distance between the median and one lateral ocelli. Scape as long as wide (Figs 6A, 8A), except in P. (?) chopardi DesutterGrandcolas n. sp. (Fig. 8H); TI without tympanum. TIII with four pairs of subapical spurs, the outers slightly longer than the inners and set less distally on TIII than inner spurs; three pairs of apical spurs, the dorsal the longest on both sides (Fig. 8C, D, except in P. (?) chopardi Desutter-Grandcolas n. sp. Fig. 8I). Male with short, slightly



overlapping (P. dubia) or not overlapping (P. discifera) FWs, lacking a stridulatory apparatus. Metanotum glandular (P. discifera) or not (P. dubia). TIII spurs not modified. Male genitalia: pseudepiphallic sclerite with a wide median part, more or less reversed overhead, and two long lateral sclerites, clearly articulated in P. dubia; pseudepiphallic parameres very large, concave, making a kind of wide forceps (Fig. 6D–F); ectophallic apodemes short and wide; ectophallic fold long and narrow, membranous; endophallic sclerite long and thin. Dorsal cavity lacking. Female apterous. Ovipositor longer than FIII. Female genitalia: copulatory papilla large, sclerotized and elongate, with a longer and thinner distal part (except in P. (?) chopardi Desutter-Grandcolas n. sp.) . Habitat. Unknown.

Phalangopsina dubia (Bolivar, 1900) (Figs 6, 7A–D, Table 2) Arachnopsis dubius Bolivar, 1900 (1899): 803. Phalangopsina dubia, Chopard, 1933a: 166.

Type material: Lectotype here designated: 1 female, Maduré, n°1.244, Arachnop. dubius sp. n. [handwritten label by I. Bolivar], identified syntype of Arachnopsis dubius by M. Paris, MNCN (right PIII and antenna missing). Paralectotype, here designated: 1 female, Indes Or. (P . Castets) / Syntype [of Phalangopsina dubia], test. M. Paris, 1991, ex. Coll. Pantel (MNHN-ENSIF2932). Examined material: Female paralectotype. Male genitalia previously observed and figured by L. Chopard (1933a, Fig. a, b). Diagnosis. According to Bolivar (1900), Chopard (1933a) and type observation, and in addition to the characters of the genus: Very small species (see Table 2). Colouration. Head dorsum yellowish (Fig. 7C), with three short black lines at the level of the fastigium; face, cheeks and lateral lobes of pronotum shining black (Fig. 7A, B); scape yellowish; legs I and II yellowish annulated with dark; TIII yellowish with unclear apical and distal brown rings; FIII yellowish striated with brown (Fig. 7D). Meso, metanotum and first tergites lightly coloured on mid dorsum (Fig. 7B, C). Male with small, slightly overlapping FWs (Chopard 1933a, Fig. a). Male genitalia. Pseudepiphallic sclerite with a median distal part returned overhead, narrowed at its base, and wide distally (Fig. 6D, F); pseudepiphallic lateral lobes made of a long hook articulated at its base on a small and narrow lateral sclerite (Fig. 6F); pseudepiphallic parameres made of a wide outer lobe, bearing a long crest on its outer margin and many picots on its inner side (Fig. 6D), and a flap-like, smaller inner lobe (Fig. 6E). Female. Ovipositor longer than FIII and TIII, but about 8–9 mm long; straight, with the apex slightly curved upward. Female genitalia as on Fig. 6G, H. Additional characters to description. Scape as long as wide (Fig. 6A). Maxillary palpi quite short; joint 5 distinctly widened distally (Fig. 6B). TIII inner apical spurs as in P. bolivari Desutter-Grandcolas n. sp. (Fig. 8C); outer spurs as on Fig. 6C; inner serrulation in paralectotype female: no spine between spurs 1 and 2, one spine between spurs 2 and 3, and between spurs 3 and 4, four or five spines above spurs; outer serrulation: two spines between spurs 1 and 2, two to three spines between spurs 2 and 3, three spines between spurs 3 and 4, four to seven spines above spurs. Basitarsomeres III with two rows of small spines dorsally: two inner spines and four outer spines, in addition to distal spines. Colouration. Median part of meso-, metanotum and first tergites lightly coloured, prolonging the clear head dorsum and pronotum dorsal disc (Fig. 7A–C). Distal part of palpi joints darker. FIII as on Fig. 7D. Female. Subgenital plate bisinuate distally. Measurements. See Table 2. TABLE 2. Measurements (mm) of Phalangopsina dubia (Bolivar, 1900). Pron-L

Pron-W

FIII-L

TIII-L

Ovip-L

Lectotype

~ 1.5

-

7.1

6.9

8.8

Paralectotype

1.4

-

6.8

6.4

7.9

Variation. The paralectotype female has a more contrasted colour pattern for the head and pronotum, laterally deep black, and dorsally yellowish. Habitat. Unknown. PHALANGOPSIDAE CRICKETS FROM INDIA


17

FIGURE 6. Phalangopsina dubia (Bolivar, 1900). A, right scape, anterior side; B, maxillary palpus, last joint; C, outer apical spurs of hind tibia; D–F, male genitalia, dorsal (D), ventral (E), lateral (F); G–H, female copulatory papilla, dorsal (G), lateral (H). Abbreviations: see text p. ***. Scales 1 mm.



FIGURE 7. Phalangopsina dubia (Bolivar, 1900). A, Female lectotype, colouration of head and pronotum, lateral; B–D, female paralectotype, colouration, lateral (B), dorsal (C), FIII, outer side (D). Phalangopsina bolivari Desutter-Grandcolas n. sp., E, FIII colouration, outer side. Phalangopsina chopardi Desutter-Grandcolas n. sp. , F, FIII colouration, outer side. Phalangopsina gravelyi Desutter-Grandcolas n. sp., G, body colouration, dorsal; H, FIII colouration, outer side. Scales 1 mm.

Phalangopsina bolivari Desutter-Grandcolas n. sp. (Figs 7E, 8A–G, Table 3) Type material: Holotype, female, Mission Maduré / type / Arachnopsis dubius / Ex. Coll. Pantel (MNHNENSIF2967).



19

Note: As mentioned above, this female belongs to the type series of P. dubia in Pantel Collection (Paris, 1994) and has been designated here as a paralectotype of P. dubia. Examined material: Female holotype, MNHN-ENSIF2967.

FIGURE 8. Phalangopsina bolivari Desutter-Grandcolas n. sp. (A–G) and Phalangopsina chopardi Desutter-Grandcolas n. sp. (H–L). A, H, scape, anterior side; B, maxillary palpus; C, D, I, apical spurs of hind tibia, inner (C, I), outer (D); E, J, female subgenital plate; F, G, K, L, female copulatory papilla, dorsal (F, K), lateral (G, L). Abbreviations, see text p. ***. Scales 1 mm.



Diagnosis. In addition to the characters of the genus: Size larger than P. dubia. Maxillary palpi elongate, joint 5 regularly widened (Fig. 8B). Colouration: face dark brown, with a yellow band under each antennal pit, prolonged on clypeus and joining on labrum; cheeks yellow; pronotum dark brown; tibiae all annulated with brown; FI and II with two small brown rings in distal half, but yellow in basal half. Male unknown. Female. Ovipositor straight, distinctly longer than FIII, about 10 mm long. Female genitalia. Copulatory papilla higher and narrower dorsally than in P. dubia, and with a longer distal part (compare figs 8F, G and 8K, L); apex straight. Description. In addition to the characters of the genus: Size larger than P. dubia (compare measurements). Head. Scape as long as wide (Fig. 8A). Palpi longer; joint 3 smaller than joint 4; joint 5 the longest, widened regularly from the base, apex as on Fig. 8B. Pronotum. Transverse, but more rounded, with raised lateral anterior lobes. Legs. TIII inner and outer apical spurs as on (Fig. 8C, D); inner serrulation in examined female: no spine between spurs 1 and 2, one or two spine(s) between spurs 2 and 3, four spines between spurs 3 and 4, seven spines above spurs; outer serrulation: two or three spines between spurs 1 and 2, four to six spines between spurs 2 and 3, five spines between spurs 3 and 4, eigth to nine spines above spurs. Basitarsomeres III with two rows of small spines dorsally: one inner and four outer spines, in addition to distal spines. FIII setose, with short filiform apical part. Colouration: head brown, face darker, with a yellow band under each antennal pit prolonged dorsally along each eye toward the vertex; facial bands prolonged on the clypeus and joined on the labrum; an additional thin yellow line on vertex mid line; posterior margin of the cheeks yellowish; palpi joints yellowish basally, brown apically; pronotum brown, the pyriform inscriptions, a thin median line and the mid part of anterior margin of dorsal disc yellowish; TI and TII yellowish with three brown rings; FI and FII yellowish, with two brown rings in distal half; TIII yellowish, with six undistinct brownish flecks or rings; FIII with two light brown rings distally, an additional brown fleck on inner side and brown stripes on outer side; basitarsi light brown with yellow base (Fig. 7E). Abdomen mottled with yellowish, light brown and dark brown. Male. Unknown. Female. Subgenital plate distal margin sinuate (Fig. 8E); yellowish, with a brown fleck in median apical part. Ovipositor straight, well longer than FIII. Female genitalia. Copulatory papilla high and clearly narrowed dorsally; distal apex straight (Fig. 8F, G). Measurements. See Table 3. Habitat. Unknown. TABLE 3. Measurements (mm) of Phalangopsina bolivari Desutter-Grandcolas n. sp.

Holotype

Pron-L

W-Pron

Wmax-pron

FIII-L

TIII-L

Ovip-L

2.0

2.8

3.1

8.8

8.3

10.1

Phalangopsina (?) chopardi Desutter-Grandcolas n. sp. (Figs 7F, 8H–L, Table 4) Type material: Holotype, female, Kod[aikanal] Dec[oly]/ Arachnopsis dubius (handwritten label) / Syntype [of Phalangopsina dubia], test. M. Paris, 1991, / Ex. Coll. Pantel (MNHN-ENSIF2966). Note: As mentioned above, this female belongs to the syntype series of P. dubia in Pantel Collection (Paris, 1994) and has been designated here as a paralectotype of P. dubia. Examined material: Female holotype, MNHN-ENSIF2966. Diagnosis. Species resembling P. bolivari Desutter-Grandcolas n. sp. by its general colouration, but differing by its larger size, its less transverse pronotum, its TIII inner apical spurs (median longer than the dorsal) and reduced serrulation, the distinctly filiform apical part of its FIII (related to its larger size?) and its copulatory papilla. Colouration: head as in P. dubia and P. bolivari Desutter-Grandcolas n. sp. but face darker and shining; cheeks less marked with yellow; pronotum lateral lobes dark brown; tibiae and femora I and II with four brown rings each, but only the two apical ones well developed in FI and II; TIII with six brown flecks on dorsal side, yellowish ventrally except for distal ring; FIII with three brown rings in distal half, and brown stripes on outer side (Fig. 7F). Male unknown. Female. Ovipositor straight, distinctly longer than FIII, about 12 mm long. Female



21

genitalia. Copulatory papilla distinctly devoid of an elongate distal part (compare figs 8F, G and 8K, L); apex truncated. Description. Species of medium size. Head. Fastigium wider than the scape, but not as wide as in P. dubia and P. bolivari Desutter-Grandcolas n. sp., the distance between the lateral ocelli slightly shorter than the distance between one lateral and the median ocelli; eyes somewhat protruding. Palpi missing. Scape longer than wide (Fig. 8H). Pronotum. Less transverse than in P. dubia, and anterior angles of lateral lobes more produced dorsally. Legs. PI missing. TII with two apical spurs, the inner longer than the outer. FIII with a long filiform apical part (Fig. 7E). TIII setose; four pairs of subapical spurs, the outers longer than the inners and set less distally on TIII than inner spurs; three pairs of apical spurs: dorsal spur the longest on outer side as in P. bolivari (Fig. 8D), but dorsal inner spur slightly shorter than the median (Fig. 8I); inner serrulation absent, except for one spine sometimes present between spurs 3 and 4, and above spur 4; outer serrulation: zero to two spines between spurs 1 and 2, four to six spines between spurs 2 and 3, five spines between spurs 3 and 4, one or two spines above spurs. Basitarsomeres III with only one row with two small spines on outer dorsal margin, in addition to distal spines. Colouration. Close to that of P. bolivari Desutter-Grandcolas n. sp. but face darker and shining; vertex variegated yellow and brown; cheeks brown, only the posterior angle and margin yellow; pronotum dark brown laterally, variegated brown and yellow dorsally; tibiae and femora I and II with four brown rings each, but the two basal ones not complete in FI and II; TIII with six brown flecks on dorsal side, yellowish ventrally except for distal ring; FIII with three brown rings in distal half, and brown stripes on outer side (Fig. 7E). Tergites brown to black brown, with rows of yellow dots. Sternites lighter. Male. Unknown. Female. Subgenital plate brown; distal margin only slightly sinuate (Fig. 8J). Ovipositor long and straight, its apex neither widened, nor ornemented. Female genitalia. Copulatory papilla resembling that of P. dubia and P. bolivari Desutter-Grandcolas n. sp. by its general shape and its height, but without an elongate distal part; apex truncated (Fig. 8K, L). Measurements. See Table 4. Habitat. Unknown. TABLE 4. Measurements (in mm) of Phalangopsina bolivari Desutter-Grandcolas n. sp.

Holotype

Pron-L

W-Pron

Wmax-pron

FIII-L

TIII-L

Ovip-L

2.4

3.0

3.6

11.7

12.1

12.3

Phalangopsina (?) gravelyi Desutter-Grandcolas n. sp. (Figs 7G, H, 9A–H, Table 5) Type material: Holotype, female, Ootacammund, Nilgiris, identified Speluncacris gravelyi Chopard, cotype, by Lucien Chopard but never described (MNHN-ENSIF2965). Examined material: Female holotype, MNHN-ENSIF2965. Diagnosis. Very small species, fitting best Phalangopsina than other phalangopsid genera by (Fig. 7G) its wide fastigium, ocelli position, transverse pronotum and general characters of its legs, although the dorsal inner apical spur of TIII is only slightly longer than the median inner spur (Fig. 9C). This species can be recognized by the following characters: copulatory papilla long, high and sclerotized dorsally (Fig. 9F–H); maxillary palpi short (Fig. 9B), joint 5 longer than joint 3 but widened regularly from its base; ovipositor dorsal valve slightly widened and lanceolate; ventral valve with a distinct tooth on ventral margin (Fig. 9E). Remark. The generic position of this species is clearly uncertain, especially because of the structure of its copulatory papilla. Description. Size very small. Head dorsum rounded, without a transverse furrow between fastigium and vertex. Eyes « flat », made of wide ommatidia (Fig. 7G). Fastigium much wider and longer than the scape; ocelli arranged as a wide triangle, but the distance between the lateral ocelli slightly larger than the distance between one lateral and the median ocelli. Scape slightly longer than wide (Fig. 9A). Maxillary palpi as on Fig. 9B. Legs. TI without tympanum. TI and TII with two apical spurs, the inner the longest. TIII setose; four pairs of alternate



subapical spurs, the outers longer than the inners and set less distally on TIII than inner spurs; three pairs of apical spurs: dorsal spur the longest on outer side, dorsal inner spur only slightly greater in length than the median (Fig. 9C); TIII serrulation sparse, but made of large spines; inner serrulation: no spine between spurs 1 and 2, one spine between spurs 2 and 3, two to three spines sometimes present between spurs 3 and 4, and two to three spines above spur 4; outer serrulation: one spine between spurs 1 and 2, three spines between spurs 2 and 3, and between spurs 3 and 4, three spines above spurs. Basitarsomeres III with only two rows of dorsal spines, one spine on inner margin and 3 spines on outer margin, in addition to distal spines. Colouration. Light yellow marked with brown (Fig. 7G); numerous thick brown setae, and small yellow ones. Face dark brown shining with a yellow band under each antennal pit; the two yellow bands slightly prolonged on the clypeus, but not fused distally; fastigium sides, area under median ocellus, cheeks (except for the yellow posterior margin) and area along the posterior margin of the eyes also dark brown. Scape and palpi light yellow, joints 3 and distal half of joint 5 of maxillary palpi marked with brown. Antennae light brown with yellow rings. Pronotum light yellow, with a brown elongated fleck along the yellow anterior margin of dorsal disc; lateral lobes brown in posterior half. Legs light yellow annulated with brown; FI and II with two brown rings in basal half; TI and II with a basal brown flack, a small brown ring at mid length, and a wide brown ring in distal half; FIII light yellow with brown dots on knees and inner sides, and brown stripes more or less fused as a line on outer side (Fig. 7H). Tibiae and basitarsomes III yellow. Tergites yellow with brown dots. Sternites and cerci yellow. Male. Unknown. Female. Apterous. Subgenital plate not transverse; distal margin emarginate (Fig. 9D). Ovipositor shorter than FIII, only slightly widened distally; apex without ornamentation; ventral valves with a ventral tooth and slight apical denticles (Fig. 9E). Female genitalia. Copulatory papilla long, straight and well sclerotized; distal margin sinuate (Fig. 9F–H). Measurements. See Table 5. TABLE 5. Measurements (in mm) of Phalangopsina gravelyi Desutter-Grandcolas n. sp.

Holotype

Pron-L

W-Pron

Wmax-pron

FIII-L

TIII-L

Ovip-L

1.5

2.2

2.4

6.7

6.7

6.2

Habitat. Unknown. Remark. The specimen observed here cannot be the female of Aspidogryllus singularis Chopard, 1933b also described from Ootacamund in the Nilgiris: although it resembles this species by its very small size and TIII apical and subapical spurs, it is different by TIII serrulation (absent in A. singularis), the size of its eyes (equal to scape length in A. singularis), and the colouration of its face, pronotum and legs (see Chopard 1933b, p. 119).

Speluncasina Desutter-Grandcolas n. gen. Type species. Speluncacris (?) annandalei Chopard, 1928: 26 Distribution. Known from Darjeeling only. Diagnosis. Small species characterized by the following combination of morphological characters: Eyes protruding. Fastigium narrow, not as wide as the scape (Fig. 10B); scape longer than wide. Pronotum not particularly transverse (Fig. 10A); lateral lobes raised dorsally. TI without tympanum. TIII with four pairs of subapical spurs, the outers the longest; median apical spurs the longest on both inner and outer sides (Fig. 9K, L); serrulation sparse. Male unknown. Female apterous. Female genitalia: Copulatory papilla small and flat, flap-like (Fig. 9O). Remark. Speluncasina Desutter-Grandcolas n. gen. resembles Phalangopsina and Opiliosina DesutterGrandcolas n. gen. by the relative size of its TIII apical spurs and the lack of a tympanum. Its differs from these genera by its less transverse pronotum, narrow fastigium and female genitalia. Habitat. Found in caves, but natural habitat not precisely assessed. Description. Size small (Fig. 10A). Head. Eyes slightly protruding anteriorly and laterally (Fig. 10B). Fastigium long and narrow, not as wide as the scape (Fig. 10B). Scape longer than wide, rectangular (Fig. 9I) Ocelli protruding; arranged as a wide triangle, the distance between the lateral ocelli longer than the distance between one PHALANGOPSIDAE CRICKETS FROM INDIA


23

lateral and the median ocelli; median ocellus subapical. Maxillary palpi not very elongate; joint 3 shorter than joint 4, and joint 5 the longest; joint 5 slightly widened apically (Fig. 9J). Pronotum. Wider than long, but not particularly transverse; greatly widened posteriorly (Fig. 10A). Dorsal disc: anterior and posterior margins respectively slightly concave and convex; a median longitudinal furrow. Lateral lobes raised dorsally. Legs. Long and very thin. TI without tympanum; two long and thin apical spurs, the inner the longest. TII with two long and thin apical spurs, the inner the longest. FI and FII thinner at mid length than in apical third. FIII filiform in apical third. TIII with four pairs of subapical spurs, the outers longer and located higher on tibia than the inners, the outer subapical spurs 2 and 3 the longest; three pairs of apical spurs, the dorsal spur the longest on both inner and outer sides (Fig. 9K, L); serrulation sparse with small spines both between and above subapical spurs. Tarsomeres 1-III less than one fourth TIII length, with two sparse rows of dorsal spines. FIII filiform in distal third (Fig. 10D). Male. Unknown. Female. Apterous. Ovipositor widened at apex, lanceolate (Fig. 9N); valves without ornementation apically. Female genitalia. Copulatory papilla very short and flap-like (Fig. 9O).

Speluncasina annandalei (Chopard, 1928), n. comb. (Figs 9I–O, 10A–C, Table 6) Speluncacris (?) annandalei Chopard, 1928: 26; Chopard 1968: 299. Opilionacris (?) annandalei, Chopard, 1969: 263. Phaeophilacris annandalei, Otte, 1994: 53. Phalangopsina (?) annandalei, Gorochov, 2003a: 717.

Remark. In his revision of the African genus Phaeophilacris Walker, 1870, Kaltenbach (1983) synonymized Speluncacris Sjöstedt, 1910 with Phaeophilacris, and transferred the African species of Speluncacris to this genus; he considered however the placement of Speluncacris annandalei doubful (« mit zweifelhafter Zugehörigkeit »), and did not formally transfer it to Phaeophilacris. Type material: 1 female type, India, Sureil, alt . 5 000 ft ., Darjeeling district, E. Himalayas, 11–31.X.17 (N. Annandale and F.H. Gravely), IM. Material examined: 1 female labelled cotype, Sureil, Darjeeling, identified by L. Chopard (MNHNENSIF2964). Diagnosis. In addition to the characters of the genus: Legs. TIII slightly longer than FIII. TIII serrulated on both inner and outer margins, except between apical and subapical spurs, with rather small spines; inner serrulation: no spine between spurs 1 and 2, one spine between spurs 2 and 3, five spines between spurs 3 and 4, five spines above spurs; outer serrulation: no spine between spurs 1 and 2, seven spines between spurs 2 and 3, six spines between spurs 3 and 4, three spines above spurs. Basitarsomeres III with two inner and four outer spines, in addition to distal spines. Colouration. Head dorsum light brown, with three main longitudinal light yellow lines (one from median ocellus, and one along the inner margin of each eye), and two shorter, undistinct ones, between the lateral and the median lines (Fig. 10A). Face yellowish brown; a yellow median line from the median ocellus to the epistemal suture, widening on clypeus and labrum (Fig. 10B). Cheeks brown, with a yellow line under each eye. Palpi brown, the upper and lower margins of joints 3 and 4 yellowish. Scapes marked with yellow and brown, with a distinct brown ring apically. Pronotum dorsal disc brown, marked with yellow, the anterior and posterior margins yellow (Fig. 10A); lateral lobes brown, with a yellow spot in anterior angle. Legs I and II yellow with brown rings: FI and FII with two rings in apical half, and an additional, faint one more basally; TI and TII with four brown rings; FIII with two brown rings in apical, filiform part, many brown stripes in outer side, and an additional brown fleck on inner side (Fig. 10C); TIII yellow ventrally and with six more or less distinct brown flecks dorsally. Tarsi light brown. Tergites light brown, darker along distal margin; a thin, median longitudinal yellow line. Male. Unknown. Female. Apterous. Subgenital plate deeply emarginate (Fig. 9M). Ovipositor as on Fig. 9N; shorter than FIII. Female genitalia. As on Fig. 9O. Measurements. See Table 6.



TABLE 6. Measurements (in mm) of Speluncasina annandalei (Chopard, 1928), n. comb.

Cotype

Pron-L

W-Pron

Wmax-pron

FIII-L

TIII-L

Ovip-L

2.7

3.2

3.4

10.2

10.8

9.5

FIGURE 9. Phalangopsina gravelyi Desutter-Grandcolas n. sp. (A–H) and Speluncasina annandalei (Chopard, 1928) n. comb. (I–O). A, I, scape, anterior side; B, J, maxillary palpus; C, K, L, apical spurs of hind tibia, inner (C, L), outer (K); D, M, female subgenital plate; E, N, female ovipositor apex, lateral; F, G, H, O, female copulatory papilla, dorsal (F), ventral (G, O), lateral (H). Abbreviations, see text p. ***. Scales 1 mm.



25

FIGURE 10. Speluncasina annandalei (Chopard, 1928), n. comb. (A–C) and Opiliosina meridionalis Desutter-Grandcolas n. sp. (D–G). A, D, E, Head, pronotum and body shape; B, F, face; C, G, hind femur. Scales 1 mm.



Opiliosina Desutter-Grandcolas n. gen. Type species. Opiliosina meridionalis n. sp. Other species included. Phalangopsina squamifera Gorochov, 2003a (according to male genitalia, see Fig. 5.13–15 in Gorochov 2003a). Etymology. Named after the names Opilionacris Sjöstedt, synonymized with Phaeophilacris Walker, 1870 by Kaltenbach (1983), and Phalangopsina Chopard, 1933a. Distribution. South India. Diagnosis. Size small. Fastigium wider than scape and rounded; eyes protruding (Fig. 10E). Maxillary palpi not very elongate, but thin; joint 5 greatly widened in apical third, truncated (Fig. 11B). Pronotum transverse (Fig. 10E). TI without tympanum. Legs I and II very thin; TI and TII with two long apical spurs. FIII with a short apical part. TIII inner apical spurs: median and dorsal spurs greatly longer than the ventral one and subequal, the dorsal the longest; TIII outer apical spurs short; dorsal spur clearly longer than the median. TIII with four pairs of subapical spurs, the outer twice as long as the inner, and set more basally on TIII. Metanotum and tergite 1 welldeveloped, clearly larger than abdominal tergites (Fig. 10E). Cerci slightly longer than the body. Male. FWs present and lobiform (O. squamifera: see Gorochov 2003a, Fig . 5.10), or absent (O. meridionalis DesutterGrandcolas n. sp.). Metanotum (O. squamifera: see Fig . 5.10 in Gorochov 2003a) or tergite 1 (O. meridionalis Desutter-Grandcolas n. sp.) glandular; supra anal plate with elongate lateral parts. Male genitalia: Resembling those of Phalangopsina, but small, rounded and compact (Fig. 12); pseudepiphallic sclerite median lobe flat (Fig. 12D); no rami; pseudepiphallic parameres having the shape of acute hooks (Fig. 12B, C); epi-ectophallic invagination short dorsally; ectophallic apodemes short, divergent and high (Fig. 12A, D); ectophallic fold very close to the pseudepiphallic sclerite, largely membranous, and deeply subdivided apically (Fig. 12B); dorsal cavity lacking. Female. Apterous. Subgenital plate emarginate distally. Ovipositor short. Female genitalia: Copulatory papilla flat, concave, sclerotized only distally (Fig. 11H–J). Relationships. Morphologically, this genus is very similar to Phalangopsina Chopard, 1933a (size, transverse pronotum, no tympanum, wide fastigium, FIII shape, TIII apical and subapical spurs). Male genitalia are also similar to those of Phalangopsina, by the pseudepiphallic sclerite separate in a dorsal median lobe and two lateral parts, and the general location of pseudepiphallic parameres. Opiliosina n. gen. differs however by its short and flat pseudepiphallic median lobe (raised dorsally in Phalangopsina), the lack of an invagination between the pseudepiphallus and the epi-ectophallic invagination (present in Phalangopsina), the size and shape of pseudepiphallic parameres, the shape of ectophallic fold (long and thin in Phalangopsina), and the shape of epiectophallic invagination, ectophallic apodemes, endophallic sclerite and endophallic apodeme. Habitat. Unknown. Description. Small species with thin legs and a short body (Fig. 10D). Head. Fastigium wider than the scape; ocelli settled as a wide triangle, the distance between median ocellus and one lateral ocellus slightly longer than the distance between the lateral ocelli. Eyes protruding (Fig. 10E). Maxillary palpi not particularly elongate, but thin; joint 5 greatly widened in apical third (Fig. 11B), well longer than joint 3. Scape longer than wide (Fig. 11A). Pronotum. Transverse, the anterior angles of lateral lobes only slightly raised dorsally (Fig. 10E). Mesonotum mostly hidden under pronotum. Metanotum and tergite 1 elongate. Legs. All very thin. TI without tympanum; two long and thin apical spurs, the inner the longest. TII with two long and thin apical spurs, the inner the longest. FIII filiform on a various apical length. TIII with four pairs of subapical spurs, the outers longer and located higher on tibia, than the inners; outer subapical spur 1 the shortest; three pairs of apical spurs, the dorsal spur the longest on both sides (Fig. 11C, D); inner median and dorsal spurs very long; outer dorsal spur about twice as long as the median; TIII serrulated on both inner and outer margins, but with few spines. Basitarsomeres all very long; basitarsomeres III with two rows of dorsal spines, the inner row reduced or lacking. Male. Metanotum (O. squamifera, see Fig. 5.10 in Gorochov 2003a) or tergite 1 (O. meridionalis DesutterGrandcolas n. sp.) glandular. FWs short and not overlapping (O. squamifera) or lacking. Supra anal plate wider than long; lateral sides more or less elongate (Fig. 11E). Subgenital plate not very short; a shallow longitudinal distal furrow (Fig. 11F). Male genitalia. Small, rounded and compact. Pseudepiphallic sclerite with a short, rounded and flat median part, almost transverse in O. squamifera, and two short and wide lateral sclerites; rami lacking; pseudepiphallic parameres having the shape of acute, curved hooks (Fig. 12). Epi-ectophallic invagination very short. Ectophallic



27

apodemes short and divergent. Ectophallic fold very close to the pseudepiphallic sclerite, membranous, except laterally close to its base. Endophallic sclerite U-shaped and endophallic sclerite crest-like (in O. meridionalis Desutter-Grandcolas n. sp. only?). Dorsal cavity lacking. Female. Apterous. Ovipositor very slightly widened at apex; valves without ornementation apically. Female genitalia. Copulatory papilla more or less flat and sclerotized (Fig. 11I–J).

FIGURE 11. Opiliosina meridionalis Desutter-Grandcolas n. sp. A, scape, anterior side; B, maxillary palpus; C, D, apical spurs of hind tibia, inner (C), outer (D); E, male supra anal plate; F, male subgenital plate, lateral; G, female subgenital plate; H–J, female copulatory papilla, dorsal (H), ventral (I), lateral (J). Abbreviations, see text p. ***. Scales 1 mm.

Opiliosina meridionalis Desutter-Grandcolas n. sp. (Figs 10D–G, 11, 12, Table 7) Type locality. India, Karnataka, route de Kadari, 800–900 m. Holotype: 1 male, India, Karnataka, route de Kadari, 800–900 m, forêt dégradée, cavités dans paroi de terre (MNHN-ENSIF2945). Allotype: 1 female, same data as the holotype (MNHN-2947). Paratypes. Same data as holotype, 2 males (MNHN-ENSIF2946, IISc) and 8 females (MNHN-ENSIF2948-2951, ENSIF2961-2962, IISc).



FIGURE 12. Opiliosina meridionalis Desutter-Grandcolas n. sp. male genitalia, dorsal (A), ventral (B), dorso apical (C), lateral (D). Abbreviations, see text p. **. Scale 0,5 mm.

Other material examined. Same data as the holotype, 2 juveniles (MNHN). Diagnosis. In addition to the characters of the genus: Colouration. Face yellowish brown, bordered by a dark band extending between each eye and the mandibula, as on Fig. 10F, and including a dark, thin triangular spot close to epistemal suture bordered by yellow, a brown rounded spot and a transverse fleck under each eye, and a dark round spot circled with black under median ocellus; cheeks with a light-yellow line under the eye lower angle; palpi yellow and brown; pronotum brown to dark brown with a transverse yellow band, the dorsal margins yellow (Fig. 10E), the lateral lobes entirely dark; legs annulated (Fig. 10D); tergites brown, mottled with yellowish and dark brown (Fig. 10E); cerci brown with a basal yellowish ring. Male. Apterous. Metanotum not glandular, but PHALANGOPSIDAE CRICKETS FROM INDIA


29

tergite 1 elongate, covering tergite 2, and with a thick distal margin (Fig. 10E). Male genitalia (Fig. 12): median lobe of pseudepiphallus rounded, only slightly longer than the lateral parts (Fig. 12A); lateral lobes sclerotized on inner and outer sides; pseudepiphallic parameres hook like (Fig. 12B, C); ectophallic fold deeply subdivided apically (Fig. 12C), slightly asymmetrical laterally, and with sclerotized lateral margins; ectophallic arc lacking (Fig. 12A); endophallic sclerite having the shape of a wide U, with a crest-like apodeme (Fig. 12A, B). Female. Subgenital plate distal margin deeply sinuate (Fig. 11G). Ovipositor very small, about half femur length. Female genitalia. As on Fig. 11H–J. Description. In addition to the characters of the genus: Legs. TIII inner and outer apical spurs as on (Fig. 11C, D); inner serrulation: no spine between spurs 1 and 2, zero to four spines between spurs 2 and 3 (mean 0.6, n=5 in males, mean 1.9, n=8 in females), zero to five spines between spurs 3 and 4 (mean 1.8, n=5 in males, mean 2.6, n=8 in females), zero to six spines above spurs (mean 1.8, n=5 in males, mean 2.8 n=8 in females); outer serrulation: one to four spines between spurs 1 and 2 (mean 3.0, n=5 in males, mean 3.0, n=8 in females), four to eight spines between spurs 2 and 3 (mean 4.0, n=5 in males, mean 5.3, n=8 in females), four to seven spines between spurs 3 and 4 (mean 5.6, n=5 in males, mean 6.0, n=8 in females), three to seven spines above spurs (mean 5.6, n=5 in males, mean 5.6, n=8 in females). Basitarsomeres III with two rows of small spines dorsally: zero to one inner spine (mean 0.2, n=9 in males, mean 0.1, n=8 in females), and two to six outer spines (mean 2.8, n=5 in males, mean 4.5, n=8 in females), in addition to distal spines. Terminalia. Cerci long, longer than body length. Colouration. Body dark brown, legs yellow annulated with brown. Face, buccal parts and cheeks yellow, with dark brown spots (Fig. 10F): a black acute triangular close to epistemal suture, a rounded dark spot under each eye, lower margin of each eye brown, connected to a wide brown band extended down to mandibula basal third; a dark round spot circled with black under median ocelli. Palpi yellow and brown; joint 5 yellow, brown in distal third. Antennae brown. Scapes yellow and brown. Pronotum brown to dark brown, the pyriform inscriptions dark ochre, a transverse spot in anterior half light ochre; lateral lobes entirely and distinctly dark. Legs yellow, annulated and marked with brown: FI, FII with three brown rings and a small basal brown spot; TI, TII with three brown rings, the basal one reaching almost tibia mid length; FIII with three brown rings in apical half, and additional brown spots on basal inner and outer sides (Fig. 10G). Body brown; distal margin of meso-, metanotum and tergite 1 yellow (Fig. 10E); a rounded yellow spot more or less distinct on each side. Cerci brown with a basal yellowish ring. Male. Apterous. Metanotum and tergites not glandular. Supra anal plate lateral lobes slightly elongate (Fig. 11E); brown, the base yellow. Subgenital plate long and relatively flat (Fig. 11F). Male genitalia. Median lobe of pseudepiphallus rounded, only slightly longer than the lateral parts, and sclerotized on its margins only (Fig. 12). Lateral lobes higher than wide or long, sclerotized on inner and outer sides. Pseudepiphallic parameres comprising a very thin apical, semi-circular hook and a more basal, not sclerotized plate (Fig. 12B). Ectophallic fold very close to pseudepiphallic sclerite dorsally, deeply subdivided apically (Fig. 12B, C); left lateral margin larger than the right one; membranous, except for sclerotized lateral margins. Epi-ectophallic invagination narrow dorsally (ectophallic arc lacking), but deeper laterally with sclerotized margins (Fig. 12A); ectophallic apodemes short and divergent, higher than wide (Fig. 12A, D). Endophallic sclerite having the shape of a wide, rounded U (Fig. 12A, B); endophallic apodeme short, crest-like (Fig. 12A, B). Female. Apterous. Subgenital plate emarginate distally (Fig. 11G). Ovipositor very short, about two thirds of TIII length. Female genitalia. Distal margin of copulatory papilla emarginate; sclerotization apical only (Fig. 11H–J). Measurements. See Table 7. TABLE 7. Measurements (in mm) of Opiliosina meridionalis Desutter-Grandcolas n. sp. Pron-L

Pron-W

FIII-L

TIII-L

Ovip-L

Holotype

1.9

2.8

10.3

10.9

----

Males (n=2)

1.9

2.8–3.0

9.7–10.3

10.0–10.9

----

Mean (n=3)

1.9

2.9

10.0

10.4

----

Allotype

2.5

3.6

12.0

12.6

5.8

Females (n=4)

2.1-2.5

3.5-3.8

10.9-12.5

11.2-12.9

5.7-7.0

Mean (n=5)

2.3

3.6

12.2

12.2

6.0



Aspidogryllus Uvarov, 1940 Type species. Aspidogryllus singularis Chopard, 1933b Distribution. Southern India (Madras). Diagnosis. According to Chopard (1933b, 1969): Small species characterized by the development and particular shape of the metanotum (hairy cup) and tergite I (produced other other tergites as a triangular shield) in males. Fastigium wide, much wider than the scape. Eyes small, as long as the scape. Ocelli scarcely visible. Maxillary palpi moderately elongate; joints 3 and 4 subequal; joint 5 a little longer, slightly concave dorsally and widened at apex. TI without tympanum. FIII with a short apical part. TIII with four pairs of subapical spurs, the outers slightly longer than the inners; three pairs of apical spurs on each side, the dorsal spur the longest on both side. TIII not serrulated dorsally. Hind metatarsi very long, rounded. Male genitalia « projecting, partly membranous » (Chopard, 1969, p. 266). Female unknown. Habitat. Unknown.

Larandopsis Chopard, 1924 Larandopsis Chopard, 1924: 86.

Type species. Larandopsis choprai Chopard, 1924 Other species included. Two additional species were described from New Guinea, Larandopsis jharnae Bhowmik, 1981 and L. newguineae Bhowmik, 1981. According to Bhowmik’s (1981) descriptions and illustration, these species do not belong to Larandopsis, but to the eneopterine genus Lebinthus Stål, 1877 (fastigium more than twice as wide as the scape, outer tympanum « prominent, elongated, oval », inner tympanum very narrow, elongate; FW venation as in Fig. 1B in Bhowmik 1981). Distribution. Assam. Diagnosis. According to Chopard (1924, 1969): Size small. Fastigium as wide as the scape. Maxillary palpi with very long joint 5. Legs rather long. TI with inner tympanum. FIII rather short. TIII with four pairs of subapical spurs, the inners longer than the outers; three inner apical spurs, the median and dorsal spurs very long, the dorsal the longest, as long as metatarsus; three outer apical spurs, very short, the median about twice as long as the other two. Cerci as long as the body. FWs and HWs present in both sexes. Colouration (juvenile specimens, with a yellow line on brown tergites): Face yellowish with a median brownish band; pronotum dorsal disc brownish with an irregular whitish band and a large, more or less rounded, yellowish spot on each side; lateral lobes fuscous with a small yellow spot; legs yellowish with brown bands. Remark. Chopard (1969) considered that Larandopsis, which has been defined using juvenile specimens, should be close to Luzaropsis Chopard, 1925 from Sri Lanka (see below). Habitat. Found in caves (Chopard 1969).

Luzaropsis Chopard, 1925 (Figs 13, 14) Luzaropsis Chopard, 1925: 521.

Type species. Luzara ferruginea Walker, 1869 Other species included. Luzaropsis confusa Chopard, 1969 n. erect., Luzaropsis henryi Chopard, 1928 and Luzaropsis omissa Gorochov, 2003c. Luzaropsis mjöbergi Chopard, 1930 described for one female from Sarawak resembles these species by its legs (inner tympanum, TIII apical and subapical spurs), but has a subgenital plate which is short, transverse and with a straight apical margin: it certainly belongs to another genus, as suggested by Gorochov (2003c, p. 725). Distribution. Sri Lanka. Diagnosis. Size medium, legs relatively short and (for PIII) thick, body and legs highly setose. Head flattened dorsally (Fig. 14C). Eyes large, but not protruding (Fig. 14A, B). Ocelli all wide; distance between the lateral ocelli PHALANGOPSIDAE CRICKETS FROM INDIA


31

FIGURE 13. Luzaropsis henryi Chopard, 1928 (A, E, F), Luzaropsis confusa Chopard, 1969 (B, C, D, G, H, K, L, M) and Luzaropsis sp. affinis henryi Chopard, 1928 (I, J). A, B, maxillary palpus; C, D, TIII apical spurs, outer (C), inner (D); E–H, male FWs, dorsal (E, G) and lateral (F, H) fields; I, J, Female FWs, dorsal (I) and lateral (J) fields; K, female subgenital plate; L, female ovipositor; M, female copulatory papilla, lateral. Scales 1 mm.



FIGURE 14. Luzaropsis henryi Chopard, 1928 (A, C, E), Luzaropsis confusa Chopard, 1969 (B, D, F). A, B, Head and pronotum, dorsal; C, D, head, profile; E, F, male supra anal plate, dorsal. Scales 1 mm.



33

almost equal or greater than the distance between one lateral and the median ocelli; median ocellus subapical. Fastigium nearly as wide as the scape. Scape slightly longer than wide. Maxillary palpi short; joint 4 longer than joint 3; joint 5 slightly shorter than joint 4, distinctly widened from its base and concave dorsally (Fig. 13A, B). Pronotum transverse, but not particularly short; wider posteriorly, with raised antero-lateral angles (Fig. 14A, B). FWs short and coriaceous in both males and females. TI with one inner, or two tympana, the outer smaller than the inner; two apical, ventral spurs. TII with four apical spurs, the inner ventral the longest, the outer dorsal the smallest. FIII short and thick, without a filiform apical part. TIII shorter than FIII, flat, wide and slightly furrowed dorsally; serrulation strong and regular through whole length, except near the knee; four pairs of short subapical spurs, the inners slightly longer than the outers and set slightly more distally than the outers; three pairs of apical spurs, all short, the median the longest on outer side (Fig. 13C), the dorsal spur the longest on inner side (Fig. 13D); TIII spurs not modified. Tarsomeres not elongated; hind basitarsomeres flattened dorsally, with two rows of thick spines. Colouration. Part of the cheeks and lateral lobes of the pronotum black brown; FW media vein yellow, prolonged in males on the dorsal disc of pronotum and behind the eyes (Fig. 14C, D). Males. Metanotum not glandular (at least in the species devoid of a stridulum). FWs not reaching the distal margin of tergites 2 or 3, truncated apically. FWs of two different types, either slightly overlapping and with a venation similar to that of females (L. henryi, Fig. 13E), or overlapping and with a reduced stridulatory apparatus (Fig. 13G, file short, veins of the harp parallel to the chords and not clearly separated from them, no distinct mirror: L. ferruginea, L. confusa, L. omissa); lateral field with few longitudinal parallel veins, but either narrow and regularly narrowed toward apex (Fig. 13F), or very wide basally and narrowed abruptly in distal fourth (Fig. 13H). Tergites not glandular. Distal part of supra anal plate with thick margins bearing bunch of long setae in latero apical corners (Fig. 14E, F). Subgenital plate short and truncate. Male genitalia. Pseudepiphallus comprising two different part, a dorsal transverse sclerite connected to the rami and a distal, median, elongate sclerite. Pseudepiphallic parameres long and high. Epi-ectophallic invagination very deep, without a sclerotized arc; ectophallic apodemes short; ectophallic fold long and narrow, originating from deep inside the genitalia, with a pair of ventral sclerites from which originates a pair of small apodemes. Dorsal cavity present, more or less regularly inflated. Endophallic sclerite transverse at the base of the dorsal cavity. Females. FW slightly overlapping over nearly their whole length; venation with thick, protruding, parallel, longitudinal veins, and faint transverse veinlets (Fig. 13I, J). Subgenital plate transverse, distal margin deeply concave (Fig. 13K). Ovipositor as long or, most often, longer than FIII; dorsal valves with a distinctive dorsal notch and ventral concavity before apex (Fig. 13L). Female genitalia. Copulatory papilla short and flat, hardly sclerotized (Fig. 13M); spermathecal duct widened before aperture. Remarks. The present-day Luzaropsis genus is clearly a homogeneous assemblage, which presents however a wide diversity for a large amount of characters. There are on one hand species with a wider head and pronotum (Fig. 14B), with a stridulatory apparatus (Fig. 13G) and a wide FW lateral field (Fig. 13H) in males, and on the other hand, species with thinner head and pronotum (Fig. 14A), no stridulum (Fig. 13E) and a thinner lateral field (Fig. 13F) in males. For male genitalia, some species have a pseudepiphallus which is clearly composed of two different sclerites (L. confusa, to a lesser extent L. henryi), while in L. ferruginea the two parts are almost continuous. Chopard (1969) correctly represented this variation in his Figures 155, 157, 158, which were misinterpreted by Gorochov (2003c, p.725): this invalidates the synonymy proposed by Gorochov (2003c) between L. ferruginea and L. confusa, which differ thus by their genitalia, colouration, venation and size. Traditionally classified within Phalangopsidae crickets, Luzaropsis presents a whole panel of characters, which cast some doubt on its familial affinity. This is most particularly the case of its male genitalia, which share some characters with Gryllomorpha Fieber, 1853 (two-part pseudepiphallus, ectophallic fold, dorsal cavity and endophallus). This problem will be further addressed in a forthcoming paper (Desutter-Grandcolas in prep.). Habitat. Unknown.

Zacla Gorochov, 2003 Zacla Gorochov, 2003a: 714.

Type species. Phalangopsis pictipes Walker, 1869



Remark. The only species known in the genus has been successively transferred to the genera Arachnopsis Saussure, 1878 ( = Arachnomimus Saussure, 1897) and Parendacustes Chopard, 1924 (Chopard 1969: 248). Distribution. Hindustan. Diagnosis. According to Chopard (1969): Size rather large. Fastigium as wide as the scape, convex dorsally. Eyes rounded, strongly protruding. Lateral ocelli rather large. Maxillary palpi long. Pronotum very strongly transverse, widening in front; dorsal disc feebly convex; lateral lobes almost triangular. Legs very long. TI with a small, inner tympanum. FIII filiform in nearly apical third. TIII very long; with four pairs of subapical spurs, strongly serrulated above and between subapical spurs; three inner and three outer apical spurs, the median the longest on both sides. Metatarsi very long; two rows of dorsal spines, three to four on inner side, six to seven on outer side. Male. FWs covering half of the abdomen. Stridulum complete: harp with oblique, parallel veins, mirror about as long as wide, crossed by three parallel, nearly horizontal veins. Other venation: chords elongate, close to each other, the chords 1 and 2 joining more distally; diagonal vein very short, very oblique; apical field very short. HWs lacking. Male genitalia (see Fig. 4.1–3 in Gorochov 2003a): Pseudepiphallus without extended median process; largelly indented distally, on both sides of ectophallic fold; pseudepiphallic parameres distal, massive; ectophallic apodemes thick, slightly diverging; endophallic sclerite with three distal prongs, the median greatly longer than the laterals; endophallic apodemes as a pair of lateral laminas. Female unknown. Habitat. Unknown.

Discussion Phylogenetic relationships. The cricket family Phalangopsidae has not yet been delimited by an extensive and robust phylogenetic analysis. Its definition is thus highly variable from one author to another. According to the classification adopted by Eades et al. (2011), based on the molecular phylogeny of Jost and Shaw (2006) (but see Legendre et al. 2010 for a critical reanalysis of the data), phalangopsid crickets would include five subfamilies, none of which are properly attested as monophyletic. Only two of them would be present in the Indian Region, the Cachoplistinae, with Cacoplistes, Meloimorpha, Larandopsis, Luzaropsis and Paragryllodes, and the Phalangopsinae, with Phalangopsina, Zacla, Kempiola, Arachnomimus and Aspidogryllus. To these should be added now Opiliosina Desutter-Grandcolas, n.gen. and Speluncasina Desutter-Grandcolas, n. gen. Morphological and anatomical characters reveal clear relationships among some of these genera. This is the case for example of Cacoplistes and Meloimorpha, close to the African genus Homoeogryllus: These genera show similar structures of male genitalia, and resemble each other by the venation of male stridulatory apparatus and the low number of subapical spurs on hind tibia (Jaiswara et al. in prep.). Saussure (1877) placed Cacoplistes in his legion Cachoplistites, together with the genus Pteroplistus Brunner von Wattenwyl, 1874, even though he considered it close to the phalangopsid genus Homoeogryllus. Chopard (1949) separated both genera into two different families, the Cachoplistidae and Pteroplistidae, because of the unique morphology of each genus. Finally, Gorochov (1986) gathered Cacoplistes, Homoeogryllus and Meloimorpha within two different tribes of the Cachoplistinae, where they are now classified, together with some additional phalangopsid genera (Otte 1994, Eades et al. 2011), among which the Luzaropsini (Larandopsis, Luzaropsis) on one hand, and Paragryllodes on the other. None of these taxa looks however close to Cacoplistes, Meloimorpha and Homoeogryllus from the point of view of genitalic structures. As mentioned above, Luzaropsini present unique structures in their male genitalia,that have no equivalent in cachoplistid or phalangopsid crickets, where they have been classified at one time or another. Their phylogenetic position will have to be tested on a wider scale than just within Phalangopsidae. In the same way, Paragryllodes shows in male genitalia the apomorphic structures associated with the development of dorsal ectophallic valves, that are present in the Neoaclini, Strogulomorphini, and part of the genera presently classified in the Endacustini (Desutter-Grandcolas 2002). In the same way, the genera now classified within the Phalangopsinae may not constitute a natural group. Even if one sets apart Aspidogryllus, which is very badly defined, and Speluncasina Desutter-Grandcolas, n. gen. known by females only, two distinct groups can be drawn according to male genitalic structures: Phalangopsina, Kempiola, Arachnomimus and Opiliosina Desutter-Grandcolas, n.gen. on one hand, and Zacla on the other. Distributions. Figure 15 shows the distribution of Indian Phalangopsidae as documented today. Apart from Sri Lanka, Southern India and the Northeastern part of the Indian Region, there exists large geographic gaps which



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have never been prospected for phalangopsid crickets. It is to be expected that the diversity of this group will increase with additional sampling, but present data could also be correlated with environmental variables, such as mean annual precipitations, as shown by our Figure 15.

FIGURE 15. Distribution of Phalangopsidae crickets in the Indian region, with regards to annual precipitation (in mm), taken from Worldclim, version 1.2 (oct. 2004), 10’ resolution. Symbols: , Arachnomimus Saussure, 1897; ●, Aspidogryllus Chopard, 1933; , Cacoplistes Brunner von Wattenwyl, 1873; , Kempiola Uvarov, 1940; √, Larandopsis Chopard, 1924; , Luzaropsis Chopard, 1925; , Meloimorpha Walker, 1870; Δ, Opiliosina Desutter-Grandcolas, n. gen.; +, Paragryllodes Karny, 1909; , Phalangopsina Chopard, 1933; Θ, Speluncasina Desutter-Grandcolas, n. gen. Zacla Gorochov, 2003 is known from “Hindusthan”.

‫ڎ‬

Habitats and biological attributes. If phylogenetic relationships and distribution of Indian Phalangopsidae are poorly known, the situation is still worse for their biology. Apart from taxa found in caves, very few data exist in the litterature describing how and where Indian Phalangopsidae forage and communicate. And if acoustic communication can logically be inferred for taxa with a stridulum, to our knowledge, no Indian Phalangopsidae has ever been recorded. In addition, their courtships have never been described.



Observations in tropical Africa and South America show that phalangopsid crickets usually present locally a certain amount of ecological diversity, which include several ecological niches for habitats (Desutter-Grandcolas 1995, 1997): straminicolous species (foraging and refuging in leaf litter), straminicolous—cavicolous species (foraging in leaf litter, but hiding in cavities located near the ground such as hollow trees or burrows), dendrophilous—cavicolous species (foraging on tree trunks, but hiding in cavities located near the ground, such as burrows, or high on trees), dendrophilous—subcorticolous species (foraging on tree trunks, but hiding under raised barks) and troglobitic species (foraging and refuging in caves, without getting out the cave at night). In the Indian region, some species have been repeatedly observed in caves and could be classified as potentially troglobitic, provided they don’t use caves as diurnal shelters only. Such is the case of Kempiola shankari (Sinha 1973, Biswas 2010, Biswas and Shrotriya 2011), K. longipes (Harries et al. 2008) and K. subalatus (Skalski 1990, Biswas 2010). Others have been collected in caves only, such as Larandopsis choprai (Chopard 1969) and Meloimorpha indicus (Agarwal and Sinha 1987). Some other species have been observed active on tree trunks at night, while during daytime they either hide in cavities near the ground during daytime (Paragryllodes species, Bhowmik 1970), or stay on tree trunks, looking for shallow shelters (K. flavipunctatus DesutterGrandcolas, n. sp. see above). However, most Indian taxa have never been observed in the field. They have been described on few collection specimens, most often without precise geographic localities. The habitats of these species can, however, be hypothesized by comparing their morphology with that of other phalangopsid species from other regions. For example, it can then be hypothesized that Cacoplistes and Meloimorpha could be dendrophilous and cavicolous, as Homoeogryllus species in Africa (Desutter-Grandcolas 1995). Bhargava (1982) mentions the presence of Meloimorpha cincticornis in termite nests, and in the same way one of us observed in tropical Africa that Homooegryllus species often hide in termite nests during the day in savanna environments and get out at night to forage and sing (LDG, pers. obs.). Most Arachnomimus species could be straminicolous and cavicolous, as Phalangopsis Serville, 1839 in South America (Desutter-Grandcolas 1992), while Phalangopsina and Opiliosina could be dendrophilous, as K. flavipunctatus Desutter-Grandcolas, n. sp. Finally, Aspidogryllus and Luzaropsis could represent the only straminicolous phalangopsids found so far in the Indian region, as they have the same short body and legs, and same small size, as many straminicolous Phalangopsidae; L. omissa has been explicitly collected in a rain forest. Clearly Phalangopsidae from the Indian region are largely unknown, and much remains to be studied in the field to document their biology. As phalangopsids are living preferentially in forested areas and look very sensitive to environmental degradation, survey of phalangopsid diversity could give cues to assess habitat preservation.

Acknowledgment This work was part of the CEFIPRA project 3009-1, « Phylogeny and acoustic evolution of crickets ». Additional financial support were provided by CNRS (UMR 7205 « Origine et structure de la Biodiversité », Dir. L. Deharveng) and MNHN (ATM « formes possibles - formes réalisées », Dir. V. Bels and P.H. Gouyon). We thank M. Paris (MNCN) for pictures and measurements of the Lectotype specimen of Phalangopsina dubia, S. Poulain (CNRS) and Guy Lecorvec (MNHN) for their help in specimen preparation and pictures, S. Hugel (Strasbourg) for reading a first draft of the manuscript and for Worldclim maps, and D. Rentz (Australia) for corrections in the manuscript. We are greatly indebted to Khartik Rao (IISc), for his help during field work and his good mood.

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