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 Nisitrus insignis Saussure, 1878 Figs 2A , 5 , 7D , 8F , 10E , 11E , 12E , 13D , 14E , 16D , 17H–I , 22; Tables 1–2 , 7 Nisitrus insignis Saussure, 1878: 515 . Crystallomorpha sumatrensis Rehn, 1909: 209 . Syn . nov . Nisitra insignis – Kirby 1906: 87 . — Chopard 1968: 352 . Nisitrus insignis – Chopard 1968: 352 . — Vicente et al. 2017: 2203 (molecular phylogeny). — Hollier et al. 2013: 487 (no types in MHNG). — Cigliano et al. 2020 ( Orthoptera Species File online). Nisitrus sumatrensis – Robillard & Desutter-Grandcolas 2004a: 276 (morphological phylogeny). Nisitrus hyalinus – Chopard 1929: 110 . — Robillard & Desutter-Grandcolas 2004a: 276 (morphological phylogeny). Nisitrus sp. – Preston-Mahfam 2000: 1 (mating behaviour). Diagnosis Of smaller size but stouter habitus than congeners. Vertex coloration ranging from black with creamcoloured margins around eyes and fastigium to cream-coloured with a strong longitudinal black band and one or two faint transverse black band (forming a T- or a cross-shaped black pattern). FIIIs brown, with a brown longitudinal stripe along external ventral half; sometimes dorsal margin with a paler longitudinal stripe. Dorsal field of male FW always with cream-coloured triangular (sometimes rounded) patch, with very distinctly different venation; harp very small, slightly longer than wide, with one distinct straight oblique vein. This species is similar to N. malaya sp. nov. and N. vittatus but differs by legs with lighter colouration and dorsum of head yellow with variable black pattern (instead of entirely black). This species also differs from the sympatric N. hughtani sp. nov. by smaller size, frons yellow (black in N. hughtani sp. nov. ) and vertex colouration pattern; FW lateral field M/R area dark (instead of forming a clearly defined white/yellow stripe), and female FW dorsal field with veins yellow (instead of mostly black). Material examined Lectotype (here designated) INDONESIA – Sumatra • ♀ ; “ Neit Suma ” [ North Sumatra ]; 1853; Mus. Caes.Vindobon; NMW . Additional specimens INDONESIA – Sumatra • 1 ♂ (holotype of N. sumatrensis ); R. Weber leg.; No. 20486; AMNH • 1 ♂ ; “ Mentawei ” [ Siberut ] “Siberoet”; 1°23′17.3″ S , 98°56′7.7″ E ; 22 Sep. 1924 ; H.H. Karny leg.; MNHN • 2 ♂♂ ; same locality data as for preceding; 8 Sep. 1924 ; H.H. Karny leg.; MNHN • 2 ♀♀ ; same locality data as for preceding; 8 Sep. 1924 ; H.H. Karny leg.; identified as Nisitra hyalina by L. Chopard ; MNHN • 3 ♀♀ ; same locality data as for preceding; 12 Sep. 1924 ; H.H. Karny leg.; MNHN • 3 ♀♀ ; same locality data as for preceding; Sep. 1924 ; C.B. K. and N. S. leg.; MNHN • 3 ♂♂ ; same locality data as for preceding; Sep. 1924 ; C.B. K. and N. S. leg.; MNHN ; • 1 ♂ ; “Mentawei” Sipora Island ; 2°13′28.7″ S , 99°40′9.4″ E ; Oct. 1924 ; C.B. K. and N. S. leg.; identifed Nisitrus brunnerianus by T. Robillard , 2004; MNHN-EO-ENSIF1748 • 1 ♀ ; same collection data as for preceding; MNHN • 1 ♀ ; “Mentawei” [ Siberut ] “Siberoet”; 1°23′17.3″ S , 98°56′7.7″ E ; Sep. 1924 ; C.B. K . and N. S. leg.; ZRC • 1 ♂ ; same collection data as for preceding; PII: N55; ZRC • 2 ♀♀ ; Sibolangit ; 3°18′2″ N , 98°34′8″ E ; Sep.–Oct. 1929 ; D.V. Leeuwen leg.; MNHN • 2 ♂♂ ; same collection data as for preceding; MNHN • 2 ♀♀ ; Sibolangit ; 3°18′2″ N , 98°34′8″ E ; 8 Oct. 1925 ; Fulmek and H. Karny leg. ; • 2 ♀♀ ; 20 km east of Sasak environs of Harau Valley National Park ; 0°4′57″ S , 100°39′12″ E ; 600 m a.s.l. ; 24–26 Nov. 1999 ; A.V. Gorochov leg.; ZIN • 3 ♂♂ ; same collection data as for preceding; ZIN • 1 ♀ ; Pematang Siantar “Pem. Siantar”, Research Centre ? “Prapat Hutan Penelitian”; 10–1100 m a.s.l. ; 10 Mar. 1993 ; S. Ingrisch leg.; SMTD • 1 ♂ ; same collection data as for preceding; SMTD • 2 ♂ ; Alas Valley , Belelutu ; 3°43′ N , 97°38′ E ; ca 320 m a.s.l. ; 3–8 Aug. 1972 ; J. Krikken leg.; RMNH • 1 ♀ ; Alas Valley , vicinity of Gumpang ; 3°48′ N , 97°29′ E ; 13 Jun. 1972 ; J. Krikken leg.; RMNH • 1 ♀ ; Deli Sibolangit Nature Reserve ; 3°19′ N , 98°35′ E ; 500 m a.s.l. ; 29 Jul. 1972 ; J. Krikken leg.; RMNH • 1 ♂ ; Soban Ajam ; Jul. 1916 ; E. Jacobson leg.; RMNH • 2 ♀♀ ; Kuala Simpang , lowland forest ; 4°17′3.7″ N , 98°3′29″ E ; Jan. 1954 ; A. Sollaart leg.; RMNH • 1 ♀ ; Serdang , Tanjong Morawai ; 1882; B. Hagen leg.; RMNH • 2 ♀♀ ; Sumatra Expedition 1877–1878 leg.; RMNH • 1 ♀ ; Ketambe , at Jeb Tenggara ; 3°41′6″ N , 97°44′57″ E ; 350 m a.s.l. ; 18 Nov. 1973 ; H.D. Rijksen leg.; RMNH • 1 ♂ ; Medan ; 3°35′10″ N , 98°40′19″ E ; 20 Jun. 1949 ; Kalshoven leg.; RMNH • 1 ♂ ; 1908; E. Jacobson leg.; RMNH • 1 ♂ ; Padang Panjang , Singalang ; 0°54′9″ S , 100°23′16″ E ; 700 m a.s.l. ; 15 Feb. 1995 ; H. Deumer and M. Schaarschmidt leg.; N5; MNHN-EO-ENSIF4180 • 2 ♀♀ ; 2 km from Padang Panjang ( Taglang ) [Talang?]; “ 3.9.1991 ”; Wolfram Guidatti leg.; MNHN • 2 ♂♂ ; [ Lubu ?] “Lubu Sampin”; 19 Jan. 1885 ; MHN • 1 ♀ ; [ Lubu ?] “Lubu Sampin”; 2 Dec. 1884 ; MHN • 2 ♀♀ ; Padang in Batangproepoe “Pad. Bovenl”; 0°54′9″ S , 100°23′16″ E ; Nov. 1924 ; C.B. K. and N. S. leg.; ZRC • 1 ♂ ; Jambi “ Djambi ”; O. Posthumus leg.; MNHN • 1 ♂ ; “ Habinsaran Tangga ”; 2 Aug. 1928 ; J.C. v. d. MeerMohr leg.; MNHN • 1 ♀ ; same collection data as for preceding; MNHN • 1 ♂ ; Fort de Kock ; 0°18′1″ S , 100°22′8″ E ; 920 m a.s.l. ; 1924; E. Jacobson leg.; MNHN • 1 juvenile ; same collection data as for preceding; MNHN • 1 ♀ ; same collection data as for preceding; MNHN • 1 ♂ ; Soilak Daras , Korinchi Valley ; 1°41′12″ S , 101°13′46″ E ; 3100 ft a.s.l. ; Mar. 1914 ; NHMUK • 1 ♀ ; Lut [“Laut″] Tawar ; 4°37′36″ N , 96°51′4″ E ; 1400 m a.s.l. ; 23 Dec. 1929 ; H.T. Pagdan leg.; NHMUK • 1 ♂ ; Dohrn leg.; identified as Nisitrus hyalinus by L. Chopard; MNHN • 1 ♀ ; Dohrn leg.; MNHN • 1 ♀ ; Padang in Batangproepoe “Pad. Bovenl”; 0°54′9″ S , 100°23′16″ E ; 9 Aug. 1920 “20”; MNHN • 1 ♂ ; same locality data as for preceding; MNHN • 3 ♀♀ ; same locality data as for preceding; MNHN • 1 ♂ ; Medan ; 3°35′10″ N , 98°40′19″ E ; Mjöberg leg.; GNME • 1 ♀ ; Dolok Baros ; 1906; “Maindon” leg.; MNHN • 1 ♀ ; Medan Batu-Seri ; 3°35′10″ N , 98°40′19″ E ; 10 Mar. 1900 ; B. Jachan leg.; ZMH • 1 ♀ ; Deli ; 3°19′ N , 98°35′ E ; 3 Jan. 1896 ; Gebr. Siemssen leg.; ZMH • 3 ♂♂ ; Deli [Bindjai] “Bindjey-Estate”; 3°36′44″ N , 98°29′20″ E ; 28 Apr. 1894 ; W. Burchard leg.; ZMH • 1 ♀ ; same collection data as for preceding; ZMH • 1 ♂ ; A. Grubauer leg.; ZMH • 1 ♂ ; “ Anei Kloof ”; 500 m a.s.l. ; 1925; E. Jacobson leg.; ZMH • 1 ♀ ; Fort de Kock ; 0°18′1″ S , 100°22′8″ E ; 920 m a.s.l. ; 1925/1926?; ZMH • 1 ♂ ; “ Deli ”; 3°19′ N , 98°35′ E ; MNHN • 1 ♀ ; same locality data as for preceding; MNHN • 1 ♂ ; v. W. leg.; MNHN • 1 ♀ ; v. W. leg.; MNHN • 1 ♂ ; “ Sumatra ”; MNHN • 1 ♀ ; “ Sumatra ”; MNHN • 3 ♀♀ ; Haran Kloof ; 0°27′14.6″ S , 101°24′53.192″ E ; 23 Apr. 1929 ; Prince Leopold leg.; RBINS • 1 ♂ ; same collection data as for preceding; RBINS • 1 ♂ ; Singalang [Mount Singgalang]; 0°24′36″ S , 100°19′51″ E ; 23 Apr. 1929 ; Prince Leopold leg.; RBINS • 1 ♂ ; Medan S.O.K. [ Deli River ]; 3°35′42.7″ N , 98°40′20.0″ E ; Fulmek L. leg.; NHMW • 1 ♂ ; Labuhan Batu ; 2°0′4.8″ N , 99°53′54.1″ E ; 22 Sep. 2020 ; K.S. Zebua leg.; iNaturalist.org/observations/60403336 • 1 ♂ ; Labuhan Batu ; 2°10′42.9″ N , 99°57′25.3″ E ; 6 Oct. 2020 ; K.S. Zebua leg.; iNaturalist.org/observations/61849044 • 1 ♂ ; Labuhan Batu ; 2°08′7.3″ N , 99°52′59.2″ E ; 6 Oct. 2020 ; K.S. Zebua leg.; iNaturalist.org/observations/61849074 • 1 ♂ ; Kong Paluh , Kutapanjang , Gayo Lues Regency , Aceh ; 3°48′16.0″ N , 97°14′6.9″ E ; 20 Jul. 2018 ; G. Jarasunas leg.; iNaturalist.org/observations/20738568 • 1 ♀ ; Kerinci , Jambi ; 2°14′54.4″ S , 101°32′21.3″ E ; 12 Aug. 2018 ; pmaaskant leg.; iNaturalist.org/observations/37307112 • 1 ♂ ; “Ланкат, СевернаЯ Суматра, ИндонеЗИЯ” [ Langkat , North Sumatra , Indonesia ]; 3°33′6.3″ N , 98°6′37.2″ E ; 3 May 2019 ; reflectitur_photons leg.; iNaturalist.org/observations/25730568 • 1 ♀ ; Lima Puluh Koto , ID-SB, ID; 0°6′39.0″ S , 100°40′15.2″ E ; 6 Jun. 2011 ; D. Nys and M. Vancauwenbergh leg.; iNaturalist.org/observations/34083286 • 1 ♀ ; Lubuk Landua ; 0°6′19.5″ N , 99°53′12.3″ E ; 13 Mar. 2018 ; O. Johnson leg.; iNaturalist.org/observations/11018930 • 1 ♂ ; Gayo Lues Regency , Aceh ; 3°52′51.7″ N , 97°19′23.1″ E ; 21 May 2010 ; ninjawil leg.; iNaturalist.org/observations/9181395 . MALAYSIA • 4 ♀♀ ; “Camp Jor”, border between Perak and Pahang ; 4°9′43″ N , 102°22′20″ E ; 30 Dec. 1902 ; Alb. Grubauer leg.; ZMH . Redescription Slightly smaller than other species ( Fig. 7D ). Vivid yellow, brown and black when alive ( Fig. 22A–C ). Vertex variable: ranging from black with yellow margins around eyes (mostly in individuals from Sumatra) to cream-coloured with a strong longitudinal black band and one or two faint transverse black bands (forming a T- or cross-shaped black pattern; mostly in individuals from Siberut Islands and Malaysia ) ( Fig. 11E ); dark pattern variable between individuals, some have a narrow pattern appearing more like a longitudinal band. Scapes red brown to black. Antennae red brown (individuals from Siberut) to black (individuals from Sumatra), with wide whitish rings. Frons yellow, cream-coloured to yellow, face part of fastigium brown to dark brown or black between eyes, lighter for individuals from Siberut than from Sumatra ( Fig. 10E ). Mouthparts dark brown to black in specimens from Sumatra, lighter or sometimes yellow in specimens from Siberut ( Fig. 10E ). Maxillary palpi cream-coloured; apical segment black apically ( Fig. 10E ). Head lateral side yellow to cream-coloured, including gena, with a black band posterior to eyes ( Fig. 12E ). Pronotum dorsal disk black, covered with white setae, rectangular with lateral margin barely widening posteriorly, posterior margin substraight to faintly bisinuous ( Fig. 11E ). Lateral lobes of pronotum dorsal half black, vivid yellow ventrally (cream-coloured in preserved specimens) ( Fig. 12E ). Legs generally light grey brown, sometimes with some small dark spots. FIIIs brown, with a brown longitudinal stripe along ventral half (more distinct and darker in individuals from Sumatra); sometimes dorsal margin with a paler longitudinal stripe; knees dark brown to black. TIIIs brown with black spines and spurs, dark brown to black near distal end; tarsomeres dark brown to black. Hindwings hyaline brown apically; longer than FWs and surpassing cerci, forming a black tail with yellow veins exceeding FWs for less than twice the length of the pronotum. Tergites red brown, with a pale band laterally; sternites pale, sometimes with a dark brown median area. Subgenital plate with a dark median area. Male FOREWINGS. FW colouration ( Fig. 13D ): dorsal field narrow, cells mostly transparent, veins varying from pale yellow to black. Basal area either infumated brown to black (individuals from Sumatra ) or almost transparent (individuals from Siberut), but always with a cream-coloured triangular patch (sometimes more rounded). Chords most often with a white patch, most readily observed in individuals from Sumatra . R black, Sc projections black basally, cream-coloured or transparent apically. Lateral field transparent. FW venation ( Fig. 13D ): 1A curved, slightly angulated. Harp very small, slightly longer than wide, with one distinct straight oblique vein. c1 long and wide, broader basally; c2 diamond-shaped; mirror (d1) small, longer than wide, not rounded, generally separated into two parts by a transverse vein, the anterior part triangular, about as long as posterior part, posterior part rectangular. Cell d2 about as wide as d1, usually subdivided by accessory veins. Apical field short, including one cell alignment posterior to mirror and a narrow apical alignment; its veins cream-coloured. Lateral field transparent, with 5 or 6 projections of Sc. Epiproct black. Subgenital plate with a dark median area. Cerci red brown. GENITALIA ( Fig. 16D ). Pseudepiphallus sclerotized, stout (medium-sized but broad) compared to congeners, anterior margin somewhat broad and straight, posterior margin also straight, lateral margins faintly converging posteriorly. Posterior apex with paired lophi slightly longer than wide (at base), obtuse at apex, moderately spaced apart from one another. Rami swollen preapically, anterior apex somewhat truncated. Pseudepiphallic parameres narrow. Ectophallic fold rounded lateral sclerites appearing beanshaped to rectangular. Endophallic sclerite wide and rounded laterally, with relatively long lateral arms. Female FOREWINGS. FW colouration ( Fig. 14E ): base of dorsal field with a yellow area; cells black, veins mostly vivid yellow (cream-coloured in preserved specimens). Region of CuA, M and R black, including the veins. Sc projections yellow basally, black apically. Lateral field transparent. FW venation: 6–7 strong longitudinal veins on dorsal field; lateral field with 6 projections of Sc. GENITALIA. Ovipositor slightly longer than FIII. Copulatory papilla conical, smaller and stout; apex folded ventrally, stout, broader than congeners, pointed; dorsal face with a sclerotized area, forming a somewhat triangular rim ( Fig. 17H–I ). Fig. 22. Nisitrus insignis Saussure, 1878 in its natural environment. A–B . Adult ♂♂. C . Adult ♀. D–E . Nymphs. Photograph source and credits: iNaturalist; Michael Kesl (A, D–E), Oscar Johnson (B), Dries Nys and Marion Vancauwenbergh (C). Table 7. Measurements of Nisitrus insignis Saussure, 1878 (in mm, means in brackets).

PronL	PronW	FWL	FWW	HWT	FIIIL	FIIIW	TIIIL
Males (n = 5)	1.7–2.4 (2.0)	2.5–3.2 (2.8)	7.3–9.5 (8.7)	2.5–2.9 (2.8)	3.4–4.6 (3.8)	13.9–16.3 (15)	2.3–2.9 (2.6)	14.3–19.4 (15.8)
Females (n = 5)	2.1–2.5 (2.2)	2.9–3.5 (3.1)	10.2–11.4 (10.7)	3.0–3.7 (3.3)	3.7–4.6 (4.2)	16.3–17.6 (17.0)	2.7–3.2 (3.0)	15.6–18.2 (16.9)

Ias	Dorsal spines of TIII Ibs Oas		Obs	TaIIIs	OL
Males (n = 5)	13–15 (14)	2–4	19–21 (20)	2–6	2–3	–
Females (n = 5)	13–15 (14)	2–5	19–20 (20)	2–5	2	15.7–17.7 (16.7)

Juvenile ( Fig. 22D–E ) Young instar with head nearly completely cream-coloured. Pronotum disk velvety black, lateral lobe with dorsal half velvety black and ventral half cream-coloured. Legs generally cream-coloured with small dark spots; tarsi darker. Abdomen with tergites velvety black dorsally, with sternites cream-coloured ventrally. Epiproct cream-coloured. Later instar resembles adult colouration. Measurements See Table 7 . Ecology Habitat and mating behaviour, including multiple mating, male guarding behaviour, female eating spermatophores and male-male competition for females, were described in detail by Preston-Mahfam (2000). Distribution Sumatra, Malay Peninsula ( Malaysia ). Type locality Indonesia : Sumatra . Calling song Unknown. Remarks One syntype of Saussure from Northern Sumatra found in NMW is designated here as the lectotype of the species. Based on comparisons with the type of N. sumatrensis , the latter name is considered as a junior synonym of N. insignis (new synonymy, see below). The observed specimen series vary greatly in size and colouration, which made us hesitate between defining one or more species, in particular for the lighter specimens from the Mentawai Islands. To remain conservative, we considered these differences as species variation only, but more information about molecular divergence and acoustic parameters may lead to revise this decision in future studies.