Revision of the cricket genus Nisitrus Saussure (Orthoptera: Gryllidae: Eneopterinae) and descriptions of five new species Author Tan, Ming Kai 2F544BE6-7986-4905-AE51-CF16B44D5251 Institut de Systématique, Evolution et Biodiversité (ISYEB), Muséum national dʼhistoire naturelle, CNRS, SU, EPHE, UA, 57 rue Cuvier, CP 50, 75231 Paris Cedex 05, France. mingkai@gmail.com Author Wahab, Rodzay bin Haji Abdul 50868E6A-34E4-471A-B941-C504CC9E895A Institute for Biodiversity and Environmental Research, Universiti Brunei Darussalam, Jalan Universiti, BE 1410, Brunei Darussalam. rodzay.wahab@ubd.edu.bn Author Japir, Razy B05E507D-FBA4-437F-AB03-94E349912EA5 Forest Research Centre (Sepilok), Sabah Forestry Department, P. O. Box 1407, 90715 Sandakan, Sabah, Malaysia. razy.Japir@sabah.gov.my Author Chung, Authur Y. C. 6E7371D6-6FF9-42DF-8745-4A8304CB0978 Forest Research Centre (Sepilok), Sabah Forestry Department, P. O. Box 1407, 90715 Sandakan, Sabah, Malaysia. arthur.Chung@sabah.gov.my Author Robillard, Tony 19BBE007-85D2-4325-8396-A1F4BAF5775C Institut de Systématique, Evolution et Biodiversité (ISYEB), Muséum national dʼhistoire naturelle, CNRS, SU, EPHE, UA, 57 rue Cuvier, CP 50, 75231 Paris Cedex 05, France. tony.robillard@mnhn.fr text European Journal of Taxonomy 2021 2021-07-28 761 1 1 75 http://dx.doi.org/10.5852/ejt.2021.761.1449 journal article 10.5852/ejt.2021.761.1449 bf24ff1f-569a-4222-afe1-8527c83d8003 2118-9773 5156259 519E2F56-40E1-4431-BC90-8484E308D16E Genus Nisitrus Saussure, 1878 Nisitra Walker, 1869: 91 . Nisitra Chopard 1940: 199 . Nisitrus Saussure 1878: 511 , 522 (nomen novum for Nisitra Walker ). — 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 ( Nisitrini tribe); 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 ( Orthoptera Species File Online). Type species Nisitra vittatus ( Haan, 1844 ) . Remarks on type species Kirby (1906) and Chopard (1968) listed N. vittata as the type species of Nisitra and treated the original type species N. marginata Walker, 1869 as a synonym of N. vittata . Subsequent authors listed N. vittatus as the type species of the replacement name Nisitrus , although Otte (1994) reinstated N. marginatus as a valid species (Braun 2013 in Cigliano et al. 2020 ). The present revision revalidates the synonymy of N. marginatus under N. vittatus , making N. vittatus the valid type species of the genus. Diagnosis (emended from Robillard & Tan 2013 ) Among genera of Eneopterinae , Nisitrus is 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 Nisitrus show 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. Fig. 7. Habitus of ♂♂ in dorsal view. A . Nisitrus brunnerianus Saussure, 1878 . B . N. crucius Robillard & Tan sp. nov. C . N. hughtani Robillard & Tan sp. nov. D . N. insignis Saussure, 1878 . E . N. malaya Robillard & Tan sp. nov. F . N. musicus Ingrisch, 1987 . G . N. vittatus ( Haan, 1844 ) . Scale bar = 10 mm. Fig. 8. Habitus of ♀♀ in dorsal view. A–B . Nisitrus brunnerianus Saussure, 1878 . C . N. crucius Robillard & Tan sp. nov. D . N. danum Robillard & Tan sp. nov. E . N. hughtani Robillard & Tan sp. nov. F . N. insignis Saussure, 1878 . Scale bar = 10 mm. Fig. 9. Habitus of ♀♀ in dorsal view. A . Nisitrus malaya Robillard & Tan sp. nov. B . N. musicus Ingrisch, 1987 . C–D . N. rindu Robillard & Tan sp. nov. E . N. vittatus ( Haan, 1844 ) . Scale bar = 10 mm. Fig. 10. Face in anterior view. A . Nisitrus brunnerianus Saussure, 1878 . B . N. crucius Robillard & Tan sp. nov. C . N. danum Robillard & Tan sp. nov. D . N. hughtani Robillard & Tan sp. nov. E . N. insignis Saussure, 1878 . F . N. malaya Robillard & Tan sp. nov. G . N. musicus Ingrisch, 1987 . H . N. rindu Robillard & Tan sp. nov. I . N. vittatu s ( Haan, 1844 ). Scale bar = 2 mm. Fig. 11. Head and pronotum in dorsal view. A . Nisitrus brunnerianus Saussure, 1878 . B . N. crucius Robillard & Tan sp. nov. C . N. danum Robillard & Tan sp. nov. D . N. hughtani Robillard & Tan sp. nov. E . N. insignis Saussure, 1878 . F . N. malaya Robillard & Tan sp. nov. G . N. musicus Ingrisch, 1987 . H . N. rindu Robillard & Tan sp. nov. I . N. vittatus ( Haan, 1844 ) . Scale bar = 2 mm. Fig. 12. Body in lateral view. A . Nisitrus brunnerianus Saussure, 1878 . B . N. crucius Robillard & Tan sp. nov. C . N. danum Robillard & Tan sp. nov. D . N. hughtani Robillard & Tan sp. nov. E . N. insignis Saussure, 1878 . F . N. malaya Robillard & Tan sp. nov. G . N. musicus Ingrisch, 1987 . H . N. rindu Robillard & Tan sp. nov. I . N. vittatus ( Haan, 1844 ) . Scale bar = 5 mm. Fig. 13. Male tegmina in dorsal view. A . Nisitrus brunnerianus Saussure, 1878 . B . N. crucius Robillard & Tan sp. nov. C . N. hughtani Robillard & Tan sp. nov. D . N. insignis Saussure, 1878 . E . N. malaya Robillard & Tan sp. nov. F . N. musicus Ingrisch, 1987 . G . N. vittatus ( Haan, 1844 ) . Scale bar = 2 mm. Fig. 14. Female tegmina in dorsal view. A . Nisitrus brunnerianus Saussure, 1878 . B . N. crucius Robillard & Tan sp. nov. C . N. danum Robillard & Tan sp. nov. D . N. hughtani Robillard & Tan sp. nov. E . N. insignis Saussure, 1878 . F . N. malaya Robillard & Tan sp. nov. G . N. musicus Ingrisch, 1987 . H . N. rindu Robillard & Tan sp. nov. I . N. vittatus ( Haan, 1844 ) . Scale bar = 2 mm. 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 ), Nisitrus is the sister genus of Paranisitra Chopard, 1925 (revision in Gorochov 2009 and Baroga-Barbecho et al. 2019 ). The two genera form the Nisitrini tribe ( Robillard & Desutter-Grandcolas 2008 ) and are very close morphologically; Paranisitra is mostly characterised by a lack of wings in both sexes, but except for this, it shares with Nisitrus its 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. Nisitrini are 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). Nisitrus are observed to be generally diurnal, during which the males can be heard calling and individuals are found foraging in the vegetation. It seems as Nisitrus are mostly herbivorous, feeding on leaves, but also on flowers ( Tan et al. 2017 ). Nisitrus are 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 brunnerianus from Borneo and Java ; N. crucius sp. nov. and N. danum sp. nov. (both from Borneo); N. hughtani sp. nov. from Sumatra ; N. hyalinus from Borneo; N. insignis from Sumatra and Malay Peninsula; N. malaya sp. nov. from Malay Peninsula; N. musicus and N. rindu sp. nov. from Borneo; and N. vittatus from Sumatra and Borneo. Fig. 15. Metanotal gland. A . Nisitrus hughtani Robillard & Tan sp. nov. B . N. brunnerianus Saussure, 1878 . Scale bar = 2 mm. Fig. 16. Male genitalia in ventral view. A . Nisitrus brunnerianus Saussure, 1878 . B . N. cruciu s Robillard & Tan sp. nov. C . N. hughtani Robillard & Tan sp. nov. D . N. insignis Saussure, 1878 . E . N. malaya Robillard & Tan sp. nov. F . N. musicus Ingrisch, 1987 . G . N. vittatus ( Haan, 1844 ) . Scale bar = 1 mm. Fig. 17. Female genitalia in lateral (A, C, E, F, H, J, L, N, P) and dorsal (B, D, G, I, K, M, O, Q) views. A–B . Nisitrus brunnerianus Saussure, 1878 . C–D . N. crucius Robillard & Tan sp. nov. E . N. danum Robillard & Tan sp. nov. F–G . N. hughtani Robillard & Tan sp. nov. H–I . N. insignis Saussure, 1878 . J–K . N. malaya Robillard & Tan sp. nov. L–M . N. musicus Ingrisch, 1987 . N–O . N. rindu Robillard & Tan sp. nov. P–Q . N. vittatus ( Haan, 1844 ) . Scale bar = 0.5 mm . Distribution Borneo ( Sabah and Sarawak States of East Malaysia , Brunei Darussalam , and Kalimantan State of Indonesia ), Java , Malay Peninsula (Peninsular Malaysia and 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. hughtani Robillard & 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. crucius Robillard & 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. rindu Robillard & 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. danum Robillard & 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. malaya Robillard & Tan sp. nov.