Nisitra Walker, 1869: 91 Nisitra Chopard 1940: 199 Nisitrus Saussure 1878: 511 Nisitra Walker Chopard 1968: 352 Desutter-Grandcolas 1990: 239 Otte 1994: 67 Preston-Mafham 2000: 2241 Robillard & Desutter-Grandcolas 2004a: 276 2004b: 578 2004c: 304 2006: 644 2008: 67 Nisitrini Robillard et al. 2007: 1265 Desutter-Grandcolas et al. 2010: 616 Nattier et al. 2011: 2201 Robillard & Tan 2013: 707 Vicente et al. 2017: 2203 Orthoptera Nisitrus brunnerianus N. crucius N. danum N. hughtani N. hyalinus N. insignis N. malaya N. musicus N. rindu N. vittatus Revision of the cricket genus Nisitrus Saussure (Orthoptera: Gryllidae: Eneopterinae) and descriptions of five new species Tan, Ming Kai Wahab, Rodzay bin Haji Abdul Japir, Razy Chung, Authur Y. C. Robillard, Tony European Journal of Taxonomy 2021 2021-07-28 761 1 1 75  Robillard & Tan 2013: 707  8NVZY 323486 Saussure, 1878 Saussure 1878 [694,976,603,629] Insecta Gryllidae Nisitrus Animalia Orthoptera 11 12 Arthropoda genus      Nisitra Walker, 1869: 91.    Nisitra–  Chopard 1940: 199.    Nisitrus–  Saussure 1878: 511, 522 (nomen novum for  NisitraWalker). —  Chopard 1968: 352. —  Desutter-Grandcolas 1990: 239. —  Otte 1994: 67. —  Preston-Mafham 2000: 2241(behaviour). —  Robillard & Desutter-Grandcolas 2004a: 276;  2004b: 578;  2004c: 304;  2006: 644;  2008: 67( Nisitrinitribe); 2011: 637. —  Robillard et al.2007: 1265. —  Desutter-Grandcolas et al.2010: 616. —  Nattier et al.2011: 2201(phylogeny and molecular dating). —  Robillard & Tan 2013: 707(redescription and diagnosis). —  Vicente et al.2017: 2203(phylogeny, historical biogeography). — Cigliano et al.2020( OrthopteraSpecies File Online).     Typespecies   Nisitra vittatus( Haan, 1844).    Remarks on typespecies  Kirby (1906)and Chopard (1968)listed  N. vittataas the typespecies of  Nisitraand treated the original typespecies  N. marginata Walker, 1869as a synonym of  N. vittata. Subsequent authors listed  N. vittatusas the typespecies of the replacement name  Nisitrus, although Otte (1994)reinstated  N. marginatusas a valid species (Braun 2013 in Cigliano et al.2020). The present revision revalidates the synonymy of  N. marginatusunder  N. vittatus, making  N. vittatusthe valid typespecies of the genus.   Diagnosis(emended from Robillard & Tan 2013) Among genera of Eneopterinae,  Nisitrusis characterised by an elegant, wasp-like, slender and colourful body ( Figs 7–9). Size average for subfamily. Colouration vivid, dominantly yellow, whitish and black, with species-specific patterns, particularly in the head ( Figs 10–11), pronotum ( Figs 11–12) and FWs ( Figs 12–14). Characteristic head shape with narrow fastigium and large prominent eyes with thin dorso-ventral black stripes. Pronotum setose, square-shaped to slightly trapezoidal, usually bisinuated posteriorly. Metanotum with large glandular structures in males, made of a wide pit on scutum and a pair of posterior pits on scutellum, with glandular pores mostly organised by groups of 2 or 3 ( Fig. 15). FWs more or less as long as abdomen. Male  Nisitrusshow modified FW venation and characteristic glossy transparency. Hindwings longer than FWs, bicoloured in most species, their base transparent with a wide dark posterior margin. Legs long and thin, FIII with a characteristic narrowed area before knee. Other general traits: TI with two tympana, inner one slit-like, covered by a swollen cuticular expansion; outer tympanum oval, its membrane transversally plicate in dorsal half. TI with three apical spurs; outer dorsal spur missing. TII with four apical spurs, inner longest. TIII with three inner and three outer apical spurs, median longest on each side; four pairs of subapical spurs, inner and outer spurs almost straight, their apex hook-like; TIII serrulate over whole length, slightly furrowed dorsally. Tarsomeres III-1 with one row of dorsal spines on external edge in addition to apical spines. Apical claws of legs slightly indented. Both males and females show characteristic genitalia.  Male genitalia ( Fig. 16) with membranous and setose lophi well individualised; pseudepiphallic parameres including two membranous posterior lobes and a sclerotized anterior lobe oriented anteriorly toward median axis of genitalia and carrying a secondary preapical sclerotized lobe; ectophallic arc transverse, incomplete due to a little-sclerotized median area; ectophallic fold narrowed preapically, its apex narrow, located between anterior apex of pseudepiphallic parameres; dorsal part of fold sclerotized, forming two elongated sclerites connected to the median area of arc; ventral part of ectophallic fold membranous, with two strong rounded lateral sclerites of variable shapes. Membrane of endophallic cavity smooth. Female copulatory papilla ( Fig. 17) conical, apex generally sclerotized and pointed. Ovipositor typically longer than FIII; apex thin with both dorsal and ventral edges smooth.   Phylogenetic relationships Based on previous phylogenetic studies ( Robillard & Desutter-Grandcolas 2004c; Nattier et al.2011; Vicente et al.2017),  Nisitrusis the sister genus of  ParanisitraChopard, 1925(revision in Gorochov 2009and Baroga-Barbecho et al.2019). The two genera form the Nisitrinitribe ( Robillard & Desutter-Grandcolas 2008) and are very close morphologically;  Paranisitrais mostly characterised by a lack of wings in both sexes, but except for this, it shares with  Nisitrusits general body shape, long thin legs with indented claws, colouration with yellow and black, head shape with large prominent eyes and narrow fastigium, and the general structure of male and female genitalia. Nisitriniare less easy to relate to other clades of Eneopterinae.    Natural history Species tend to be found at forest edges and gaps in the forest where the vegetation is more exposed. They are also found among shrubs and herbaceous plants (but not grasses).  Nisitrusare observed to be generally diurnal, during which the males can be heard calling and individuals are found foraging in the vegetation. It seems as  Nisitrusare mostly herbivorous, feeding on leaves, but also on flowers ( Tan et al.2017).  Nisitrusare also observed to congregate with one another on the same plant, sometimes making up of more than 20 individuals of both adults and nymphs. They can be considered a pest in some parts of Southeast Asia ( Willemse 2001).    Included species   Nisitrus brunnerianusfrom Borneo and Java;  N. crucius sp. nov.and  N. danum sp. nov.(both from Borneo);  N. hughtani sp. nov.from Sumatra;  N. hyalinusfrom Borneo;  N. insignisfrom Sumatraand Malay Peninsula;  N. malaya sp. nov.from Malay Peninsula;  N. musicusand  N. rindu sp. nov.from Borneo; and  N. vittatusfrom Sumatraand Borneo.    Distribution Borneo ( Sabahand SarawakStates of East Malaysia, Brunei Darussalam, and Kalimantan Stateof Indonesia), Java, Malay Peninsula (Peninsular Malaysiaand Singapore) and Sumatra( Indonesia) ( Fig. 5).    Key to the species of  Nisitrus Saussure, 1878     1. Frons mostly black, or with large dark markings (e.g., cross-shaped pattern); gena usually black, although may have yellow band ventral of eye ................................................................................ 2  – Frons mostly yellow, sometimes with small dots but without large dark markings; gena usually yellow ............................................................................................................................................... 5   2. FW with longitudinal white stripe around region of CuA, M and R with a distinct margin from black colouration on dorsal field; lateral field usually infumated black ventrally. Distribution: Sumatra...................................................................................  N. hughtaniRobillard & Tan sp. nov.  – FW with longitudinal white stripe around region of CuA, M and R darkens progressively towards the dorsal field without a distinct margin from black colouration on dorsal field; lateral field usually light coloured or transparent ventrally. Distribution: Borneo and Java............................................ 3   3. Smaller in size, female FWL typically < 11.3 mm. Frons with a characteristic black cross marking ....................................................................................................  N. cruciusRobillard & Tan sp. nov.  – Larger in size, female FWL typically> 11.8 mm. Frons completely black; without a characteristic black cross marking .......................................................................................................................... 4   4. Dorsum of head completely black. Pronotal lateral lobe either with ventral half yellow or with an oval yellow spot at the anterior end ................................................  N. brunnerianus Saussure, 1878  – Dorsum of head pale with some dark markings. Pronotal lateral lobe with a characteristic dorsal yellow band that extends ventrally ...............................................  N. rinduRobillard & Tan sp. nov.   5. Frons with black band between eyes, mouthparts mostly yellow. Body darker; female FW typically with infumated black cells, with lateral parts of dorsal field usually with an infumated yellow longitudinal stripe. Distribution: Borneo ( Sabah) ......................  N. danumRobillard & Tan sp. nov.  – Frons without black band between eyes, mouthparts black. Body usually with more yellow colouration; male FW with some black and white infumation, but generally transparent. Distribution: Borneo, Malay Peninsula and Sumatra.......................................................................................................... 6   6. Legs usually pale in colouration; FIII with at least one black longitudinal stripe on external face. Male FW with distinct infumated white spots and infumated black pattern near base, sometimes with infumated white patterns around chords, but otherwise transparent, harp very small, with one distinct straight oblique vein; female FW almost always with yellow/white veins. Distribution: Sumatra....................................................................................................  N. insignis Saussure, 1878  – Legs usually brown in colouration; FIII with faint or without black longitudinal stripe on external face. Male FW with infumated black pattern near base, sometimes with infumated white spots, but otherwise transparent, harp larger, with 3 oblique veins; female FW sometimes with yellow/white veins. Distribution: Sumatra, Borneo, Malay Peninsula ................................................................... 7   7. Eyes dark when alive. Frons yellow, face part of fastigium (including scapes) and mouthparts black, maxillary palpi yellow; dorsal part of legs black. Male pseudepiphallus slender. Distribution: Borneo, near Kinabalu area ....................................................................................  N. musicus Ingrisch, 1987  – Eyes green when alive. Frons yellow to brown, mouthparts sometimes yellow or brown; legs typically homogenous brown. Pseudepiphallus more robust, broad. Distribution: Sumatra, Borneo, Malay Peninsula ........................................................................................................................................... 8   8. Lighter in colouration. Lateral field of FWs usually light in colouration. Juveniles of pale white colouration with numerous dark stripes all around the body. Female copulatory papilla without truncated lobule at the ventro-anterior end. Distribution: Borneo, Sumatra...................................... ......................................................................................................................  N. vittatus( Haan, 1844)  – Darker in colouration. Lateral field of FWs usually dark in colouration. Juveniles of darker colouration, usually grey, their pronotum without longitudinal stripes but dorsal disk sometimes with a large velvety black spot. Female copulatory papilla with a truncated lobule at the ventro-anterior end. Distribution: Malay Peninsula ...................................................  N. malayaRobillard & Tan sp. nov.