Nisitrus vittatus ( Haan, 1844 )

Tan, Ming Kai, Wahab, Rodzay bin Haji Abdul, Japir, Razy, Chung, Authur Y. C. & Robillard, Tony, 2021, Revision of the cricket genus Nisitrus Saussure (Orthoptera: Gryllidae: Eneopterinae) and descriptions of five new species, European Journal of Taxonomy 761 (1), pp. 1-75 : 64-68

Nisitrus vittatus ( Haan, 1844)

Figs 1 , 3 , 5–6 , 7G , 9E , 10I , 11I , 12I , 13G , 14I , 16G , 17P–Q , 23E–H, 24D–F, 25D–F, 26 ; Tables 1–2 , 11

Gryllus (Platydactylus) vittatus Haan, 1844: 234 .

Nisitra marginata Walker, 1869: 92 . Synonym reinstated.

Nisitrus vittatus – Saussure 1878: 511, 653. — Chopard 1968: 352 (synonymised N. marginata ). — Otte 1994: 67. — Robillard & Desutter-Grandcolas 2004a: 276 (morphological phylogeny). — Robillard & Tan 2013: 707 (redescription and neotype designation). — Tan et al. 2017: 146 (flower-visiting behaviour); 2019: 303. — Tan & Wahab 2018: 126 . — Cigliano et al. 2020 ( Orthoptera Species File online).

Nisitra vittata – Kirby 1906: 87. — Chopard 1940: 199.

Nisitrus marginatus – Otte 1994: 67.

Nisitrus cf. vittatus – Vicente et al. 2017: 2203 (molecular phylogeny).

Revised diagnosis

Species of average size, colouration contrasting with black and yellow, characterised by black vertex, frons yellow, orange brown legs. Male area between M and R yellow, veins black or yellow; cells beneath R partly yellow, otherwise transparent.Anal-most and middle chords distinctly diverging apically, basalmost chord extend from midpoint/after midpoint of middle chord. Female copulatory papilla without truncated lobule at ventro-anterior end. Species very similar to N. malaya sp. nov., from which it differs by male genitalia with pseudepiphallus faintly less robust, narrower, anterior margin less concave; female copulatory papilla without truncated lobule at ventro-anterior end. Juvenile differs strikingly in colour patterns. Male calling song also differs by shorter echeme period.

Material examined

Neotype

INDONESIA – Sumatra • ♂; Wai Lima, “Z. Sum.” [South Sumatra], Lampongs ; 5°25′38.06″ S, 105°01′22.37″ E; Nov.–Dec. 1921; Karny and Siebers leg.; RMNH.

Additional specimens

BRUNEI DARUSSALAM – Temburong • 1 ♂; Kuala Belalong Field Studies Centre ; 4°32′47″ N, 115°9′27″ E; 8 Jan. 2017; M.K. Tan leg.; KB.17.34; N37; MNHN-EO-ENSIF11069 • 1 ♀; Kuala Belalong Field Studies Centre ; 4°32′28.57″ N, 115°09′28.94″ E; 23 Sep. 2016; M.K. Tan leg.; KB.16.10; UBDM • 1 ♀; Kuala Belalong Field Studies Centre ; 4°32′28.57″ N, 115°09′28.94″ E; 23 Sep. 2016; M.K. Tan leg.; KB.16.11; UBDM • 1 ♀; same locality data as for preceding; 25 Jul. 2017; M.K. Tan leg.; KB.17.53; UBDM • 1 juvenile; 4°32′49.2″ N, 115°9′20.3″ E; 7 Jan. 2014; J. Hodecek leg.; iNaturalist.org/observations/46113125 • 1 ♂; 4°32′49.2″ N, 115°9′20.3″ E; 16 Jan. 2014; J. Hodecek leg.; iNaturalist.org/observations/46113093.

INDONESIA – Sumatra • 1 ♂; same collection data as for neotype; Karny leg.; MNHN-ENSIF1742 • 1 ♀; same collection data as for neotype; MNHN-EO-ENSIF3200 • 1 ♀; same collection data as for neotype; MNHN-ENSIF3198 • 1 ♀; Pahoe; 22 Oct. [1925]; O. Posthumus leg.; MNHN-EO- ENSIF3199 • 1 ♀; same collection data as for preceding; MNHN • 1 ♀; S Sumatra, SW Lampongs, Mt Tanggaamues ; 1°37′15.25″ S, 103°36′23.79″ E; 600 m a.s.l.; Dec. 1934; Giesting, Lieftinck and Toxopeus leg.; MNHN-EO-ENSIF3201 • 1 ♀; S Sumatra, Lalembang, Soeban Djerigi ; 15 Jun. 1933; Soekarno leg.; MNHN-EO-ENSIF3202 • 1 ♀; Damm [?] Muntok Banka ; 2°03′16.62″ S, 105°10′43.11″ E; 25 Nov. [19]23; MNHN-EO-ENSIF3203 • 1 ♀; [ Pidada ] “Pedada-B Lampongs”; 5°27′42.68″ S, 105°19′20.91″ E; MNHN-EO-ENSIF3204 • 6 ♂♂; “southern Sumatra, Bengkulu, environs of town Curup (not far from city Bengkulu)”; 3°28–29′ S, 102°31–38′ E; 24 Apr.–2 May 2009; A. Gorochov, M. Berezin and E. Tkatsheva leg.; ZIN • 1 ♀; same collection data as for preceding; ZIN • 2 ♀♀; “southern Sumatra, Lampung, National Park Way Kambas, ~ 60 km NE of city Tanjung Karang, camp Way Kenam ”; 5°1.678′ S, 105°46572′ E; 20–30 m a.s.l.; 13–14 Apr. 2009; A. Gorochov, M. Berezin and E. Tkatsheva leg.; ZIN • 1 ♂; same collection data as for preceding; ZIN • 7 ♂♂; “ Sumatra, southern Sumatra, environs of village Banding Agung on Lake Ranau Danau ”; 4°48.695′ S, 103°55.289′ E; 600–700 m a.s.l.; 19–29 Apr. 2009; A. Gorochov, M. Berezin and E. Tkatsheva leg.; ZIN • 3 ♀♀; “ Southern Sumtra, prov Lampung, National Park Way Kambas, ~ 60 km NE of city Tanjung Karang, camp Way Kenam ”; 5°1.678′ S, 105°46.572′ E; 20–30 m a.s.l.; 13–14 Apr. 2009; A. Gorochov, M. Berezin and E. Tkatsheva leg.; PII N31; ZIN • 1 juvenile; “ Sumatra, southern Sumatra, environs of village Banding Agung on Lake Ranau Danau ”; 4°48.695′ S, 103°55.289′ E; 600–700 m a.s.l.; 19–29 Apr. 2009; A. Gorochov, M. Berezin and E. Tkatsheva leg.; ZIN • 1 ♂; “southern Sumatra, Lampong, National Park Bukit Barisan Selatan, 15–20 km NW of town Krui, camp Kubu Prahu ”; 5°4.341′ S, 104°3.093′ E; ~ 600 m a.s.l.; 18–19 Apr. 2009; A. Gorochov, M. Berezin and E. Tkatsheva leg.; N33; ZIN . – Java • 1 ♂; “ Java ”; H.D. Saussure leg.; MHN • 1 ♂; Padang ; 0°57′22.4″ S, 100°25′5.36″ E; 21 Apr. 1929; Prince Leopold leg.; RBINS • 1 ♂; Tandjong-Karang ; 0°32′12.4″ S, 105°15′29.88″ E; 21 Apr. 1929; Prince Leopold leg.; RBINS • 1 ♀; Sungai Penuh City , Jambi; 2°7′53.7″ S, 101°15′53.8″ E; 18 Jun. 2019; S. Wollney leg.; iNaturalist.org/observations/27481225 • 1 ♂; Pesawaran Regency , Lampung; 5°26′2.5″ S, 105°9′57.6″ E; 9 Apr. 2019; J.-S. Wang leg.; iNaturalist.org/observations/36723277 • 1 ♂; East Lampung Regency, Lampung; 5°7′3.2″ N, 105°44′16.6″ E; 7 Aug. 2019; pmaaskant leg.; iNaturalist.org/observations/37027091 • 1 ♀; Sungai Penuh City , Jambi; 2°7′56.8″ S, 101°15′53.2″ E; 20 Jun. 2019; S. Wollney leg.;iNaturalist.org/observations/27481235 • 1 ♀; Pesawaran Regency , Lampung; 5°26′4.5″ S, 105°10′29.3″ E; 9Apr. 2019; J.-S. Wang leg.; iNaturalist.org/observations/36723272 • 1 ♀; Eka Jaya, Kec. Jambi Sel. , Kota Jambi , Jambi 36132; 1°37′1.2″ S, 103°40′16.8″ E; 7 Aug. 2020; suhendra leg.; iNaturalist.org/observations/55695657 • 1 ♂; Banyu Asin, Riau Silip , Bangka Regency , Bangka Belitung Islands; 1°53′3″ S, 105°44′55″ E; 20 Oct. 2020; R. Maharani leg.; iNaturalist.org/observations/63092557 . – Kalimantan • 6 ♂; “ Marah ”; 0°58′31.3″ N, 116°33′14.8″ E; 10–28 Nov. 1925; H.C. Siebers leg.; MNHN • 3 ♂; “Midden” [near border of Koetai and Beraoe]; 1°50′14″ N, 116°12′34″ E; 26 Sep. 1925; H.C. Siebers leg.; MNHN • 1 ♂; “Midden” [near border of Koetai and Beraoe]; 1°42′28″ N, 116°15′40″ E; 4 Aug. 1925; H.C. Siebers leg.; MNHN • 1 ♂; “ Midden ”; 1925; H.C. Siebers leg.; MNHN1730 • 1 ♂; “Midden” [near border of Koetai and Beraoe]; 1°50′14″ N, 116°12′34″ E; 27 Sep. 1925; H.C. Siebers leg.; MNHN • 1 ♀; “Midden” [near border of Koetai and Beraoe]; 1°47′35″ N, 116°12′22″ E; 12 Sep. 1925; H.C. Siebers leg.; MNHN • 1 juvenile; “Midden” [near border of Koetai and Beraoe]; 1°50′14″ N, 116°12′34″ E; 30 Sep. 1925; H.C. Siebers leg.; MNHN • 1 ♀; “ Kac. Pujungan Kayan Mentarang Nat. Reserve, lowland dipterocarp forest , WWF station, Lalut Birai Base Camp ”; 2°52′ N, 115°49′ E; 355 m a.s.l.; Jun. 1993; D.C. Darling and U. Rosichon leg.; MZB • 1 ♀; “ Karimata Island Nat. Keptapang Res. , primary rainforest, canopy cover”; 1°25′ N, 106°40′ E; 350 m a.s.l.; Feb. 1991; Sutrisno Djenal leg.; MZB • 1 ♂; Samarinda ; 0°31′54.17″ S, 117°7′35.31″ E; 9 Feb. 1929; Prince Leopold leg.; RBINS • 2 ♀♀; “ Melawi ”; 0°37′11.5″ S, 111°45′4″ E; Nov.–Dec. 1924; Blanchmanche leg.; MNHN • 1 ♂; “Hollandska Ostindien”; Mrs M.-E. Walsh leg.; ZMLU • 2 ♀♀; same collection data as for preceding; ZMLU • 4 ♂♂; “ Samarinda ”; 0°28′59″ S, 117°8′33″ E; Mrs M.-E. Walsh leg.; ZMLU • 3 ♀♀; same collection data as for preceding; ZMLU • 1 ♂; [ Tanggarong ] “Tanggarang”; 0°25′39″ S, 116°59′16″ E; 5 Jul. 1957; Mrs M.-E. Walsh leg.; ZMLU • 1 ♂; same locality data as for preceding; 4 Jul. 1957; Mrs M.-E. Walsh leg.; ZMLU • 1 ♀; [ Kaliorang ] “Kariorang”; 0°52′29″ N, 117°56′31″ E; Apr. 1937; Mrs M.-E. Walsh leg.; ZMLU .

MALAYSIA – Sabah • 1 ♂; N R Sandakan ; 5°50′38″ N, 117°57′19″ E; 22 Jul. 1927; H.M. Pendlebury leg.; MNHN • 1 ♂; same collection data as for preceding; MNHN • 1 ♂; Mount Kinabalu, Kiau ; 6°1′40″ N, 116°29′26″ E; 3000 ft a.s.l.; H.M. Pendlebury leg.; MNHN • 1 ♀; “ Crocker Range National Park not far from Keningau Town, secondary–primary forest”; 5°28′07.79″ N, 116°04′22.95″ E; 1000–1300 m a.s.l.; 6 May 2013; A. Gorochov, M. Berezin, V. Gorochova and E. Tkatsheva leg.; N32- Nbo 5; ZIN • 1 ♂; “ Mulu National Park, not far from borders with Brunei and Indonesia ”; 4°01′58.72″ N, 114°49′00.56″ E; 100–300 m a.s.l.; 24–27 Mar. 2012; A.V. Gorochov, M. Berezin, E. Tkatsheva and I. Kamskov leg.; N27- Nbo3; ZIN • 9 ♂♂; Mount Trus Madi , “partly primary/partly secondary forest”; 5°33′53.94″ N, 116°30′11.31″ E; ~ 1000 m a.s.l.; 13–25 May 2007; A.V. Gorochov leg.; ZIN • 10 ♀♀; Mount Trus Madi , “partly primary/partly secondary forest”; 5°33′53.94″ N, 116°30′11.31″ E; ~ 1000 m a.s.l.; 13–25 May 2007; A.V. Gorochov leg.; ZIN • 1 ♂; “ southern part of Kinabalu National Park, primary forest”; 5°59′50.78″ N, 116°33′39.77″ E; ~ 1500–2000 m a.s.l.; 26 Apr.–1 May 2013; A. Gorochov, M. Berezin, V. Gorochova and E. Tkatsheva leg.; PII N25-Nbo1; ZIN • 1 ♂; “ Sandakan, environs of Sukau Village on Kinabatangan River (~ 35 km from sea), secondary–primary forest”; 5°51′21.27″ N, 117°51′49.17″ E; 0 m a.s.l.; 8–13 May 2013; A. Gorochov, M. Berezin and E. Tkatsheva leg.; PII N28-Nbo4; ZIN • 1 ♂; “ Tawau Hills National Park near Tawau City, secondary-primary forest”; 4°23′55.38″ N, 117°53′20.60″ E; 200–400 m a.s.l.; 14–20 Apr. 2013; A.V. Gorochov, M. Berezin, E. Tkatsheva and I. Kamskov leg.; N26- Nbo2; ZIN • 4 ♂♂; [ Long Pasia ] “Long Pa Sia”; 4°24′22.05″ N, 115°43′40.97″ E; ca 1025 m a.s.l.; 1–13 Apr. 1987; C. v. Achterberg leg.; RMNH • 2 ♀♀; same collection data as for preceding; RMNH • 2 ♀♀; Danum Valley Field Studies Centre ; 4°57′52.95″ N, 117°48′18.25″ E; 14–20 Mar. 1987; C. v. Achterberg leg.; RMNH • 1 ♀; “ West of 3.1 km on Road Tenom–Keningau Depleted lowland forest; 5°19′15.2″ N, 116°13′2″ E; 240 m a.s.l.; 25 Jun. 1986; C. v. Achterberg leg.; RMNH • 2 ♂♂; Ranau “ west coast Residency, Ranau , 8 miles north Paring Hot Springs ”; 5°50′47.95″ N, 116°44′02.41″ E; 500 m a.s.l.; 8–11 Oct. 1958; L.W. Quate leg.; BPBM • 1 ♂; Tawau, Quoin Hill ; 4°23′55.38″ N, 117°53′20.60″ E; 8–14 Jul. 1962; H. Holtmann leg.; BPBM • 1 ♂; Sandakan “Sandakan Bay (NW), Sepilok Fore. Res.”; 5°50′38″ N, 117°57′19″ E; 1–10 m a.s.l.; 27 Oct 1957; J.L. Gressitt leg.; BPBM • 1 ♂; [ Crocker Range National Park ] “Croker Range n.p.”; 5°28′07.79″ N, 116°04′22.95″ E; 11–17 Aug. 2003; “Whiting, Svenson, Bybee” leg.; PII: NviSA; MNHN-EO-ENSIF1709 • 2 ♀♀; same collection data as for preceding; MNHN • 1 juvenile; same collection data as for preceding; MNHN • 1 ♂; “ Sipitang ”, “ Mendolong T3/ W5”; 4°54′58″ N, 115°45′60″ E; 25 Apr. 1988; S. Adebratt leg.; ZMLU • 1 ♂; “ Sipitang ”, “ Mendolong T1B/W4”; 4°54′58″ N, 115°45′60″ E; 14 Mar. 1989; S. Adebratt leg.; ZMLU • 4 ♀♀; “ Sipitang ”, “ Mendolong T1B/W4”; 4°54′58″ N, 115°45′60″ E; 14 Mar. 1989; S. Adebratt leg.; ZMLU • 1 ♀; “ Sipitang ”, “ Mendolong T3/W5”; 4°54′58″ N, 115°45′60″ E; 31 Mar. 1989; S. Adebratt leg.; ZMLU • 2 ♀♀; “ Sipitang ”, “ Mendolong T3/W5”; 4°54′58″ N, 115°45′60″ E; 9 Apr. 1988; S.Adebratt leg.; ZMLU • 1 ♀; “ Sipitang ”, “ Mendolong T3/W5”; 4°54′58″ N, 115°45′60″ E; 24 Feb. 1989; S.Adebratt leg.; ZMLU • 2 ♀♀; “ Sipitang ”, “ Mendolong T1B/W4”; 4°54′58″ N, 115°45′60″ E; 17 Mar. 1989; S. Adebratt leg.; ZMLU • 7 ♀♀; “ Sipitang ”, “ Mendolong T2B/W5”; 4°54′58″ N, 115°45′60″ E; 19 Apr. 1988; S. Adebratt leg.; ZMLU • 1 juvenile; “ Sipitang ”, “ Mendolong T5/R”; 4°54′58″ N, 115°45′60″ E; 28 Apr. 1988; S. Adebratt leg.; ZMLU • 1 ♂; “ Sipitang ”, “ Mendolong T5/R”; 4°54′58″ N, 115°45′60″ E; 28 Apr. 1988; S. Adebratt; ZMLU • 1 juvenile; “ Batu Niah 18.mbbbb”; 3°48′14″ N, 113°45′15″ E; Ulf Gärdenfors leg.; ZMLU • 1 ♀; “ Sipitang ” Mendolong; 4°54′58″ N, 115°45′60″ E; S. Adebratt leg.; ZMLU • 1 ♀; Kinabatangan; 5°29′33.2″N, 118°12′23″ E; 30Jul.2018;lhurteau leg.;iNaturalist.org/observations/15437369 • 1 ♂; Kota Kinabatangan; 5°28′20.6″ N, 118°15′28.8″ E; 7 Mar. 2013; rbeunen leg.; iNaturalist.org/observations/18843166 • 1 ♂; Jalan Poring, 89300 Ranau; 6°2′36.6″ N, 116°42′10.6″ E; 6 Aug. 2013; doggy0406 leg.; iNaturalist.org/observations/24398754 • 1 ♂; Lahad Datu District; 4°57′48.7″ N, 117°40′20.3″ E; 17 Oct. 2019; N.K. Thangasamy leg.; iNaturalist.org/observations/35226617 • 1 ♀; Lahad Datu; 4°57′42.6″ N, 117°41′21.1″ E; 27 Sep. 2016; S. Myers leg.; iNaturalist.org/observations/17100119 • 1 ♂; Lahad Datu, Danum Valley Field Centre trails; 4°58′15.8″ N, 117°48′8.63″ E; 17 Jun. 2014; B. Crins leg.; iNaturalist.org/observations/40612983 • 1 ♂; Kinabatangan; 5°29′46.2″ N, 118°16′39.7″ E; 26 Sep. 2018; z_mcgrath leg.; iNaturalist.org/observations/27490441 • 1 ♀; Bahagian Sandakan; 5°22′0.8″ N, 117°25′47.2″ E; 24 May 2018; K. Squires leg.; iNaturalist.org/observations/38619342 • 1 ♀; Penampang; 5°50′50.7″ N, 116°5′27.8″ E; 28 Apr. 2020; tanuki18 leg.; iNaturalist.org/observations/44101645 • 1 juvenile; Penampang; 5°50′50.6″ N, 116°5′28″ E; 23 May 2020; tanuki18 leg.; iNaturalist.org/observations/46949300 • 1 juvenile; Tuaran; 5°49′17.3″ N, 116°20′28.8″ E; 6 Jun. 2020; tanuki18 leg.; iNaturalist.org/observations/48631961 • 1 ♀; Tuaran; 6°0′34.1″ N, 116°14′48.6″ E; 14 Jun. 2020; tanuki18 leg.; iNaturalist.org/observations/49562737 • 1 ♂; 90000 Kinabatangan; 5°24′49.3″ N, 118°2′15.5″ E; 7 Mar. 2020; M. de Groot leg.; iNaturalist.org/observations/40232424 • 1 ♀, 1 ♂; 5°13′0.40″ N, 118°43′5.7″ E; 24 Jul. 2003; C. Shu leg.; iNaturalist.org/observations/39450135. – Sarawak • 1 ♀; Mount Dulit, moss forest; 3°19′58″ N, 114°9′ E; 4000 ft a.s.l.; 22 Oct. 1932; B.M. Hobby and A.W. Moore leg.; MNHN • 1 ♂; Batu Niah; 3°48′ N, 113°45′ E; 4–8 Mar. 1984; S. Ingrisch leg.; SMTD • 2 ♀; “ Gunung Gading National Park , 80–90 km northwest of Kuching City ”; 1°41′25.88″ N, 109°50′44.68″ E; 100–300 m a.s.l.; 8–9 Mar. 2012; A. Gorochov, M. Berezin, E. Tkatsheva and I. Kamskov leg.; ZIN • 3 ♂♂; “ Kuching, environs of Kubah National Park, (situated on Matang Mountain and partly near it)”; 1°33′05.74″ N, 110°18′54.26″ E; 200–500 m a.s.l.; 10–17 Mar. 2012; A.V. Gorochov, M. Berezin, E. Tkatsheva and I. Kamskov leg.; PII: N24 Nku1; ZIN • 1 ♀; “ Kuching, environs of Kubah National Park, (situated on Matang Mountain and partly near it)”; 1°33′05.74″ N, 110°18′54.26″ E; 200–500 m a.s.l.; 10–17 Mar. 2012; A.V. Gorochov, M. Berezin, E. Tkatsheva and I. Kamskov leg.; ZIN • 1 ♂; Nanga Pelagus near Kapit; 1°42′03.32″ N, 113°32′32.38″ E; 180–585 m a.s.l.; 7–14 Oct. 1958; T. Maa leg.; BPBM • 1 ♂; Kapit District, Merirai Valley ; 1°53′52.61″ N, 113°46′13.62″ E; 30–300 m a.s.l.; 1–6 Aug. 1958; T. Maa leg.; BPBM • 2 ♂♂; Tuba ; 1°16′15″ N, 111°5′13″ E; 16–22 Jan. 1979; Gärdenfors, Hall, Hansson and Samuelsson leg.; ZMLU • 2 ♀♀; same collection data as for preceding; ZMLU • 1 ♀; “ 20 km SE Sebuyau, Tuba Forest clearing at longhouse ”; 1°16′15″ N, 111°5′13″ E; 15–22 Jan. 1979; Gärdenfors, Hall, Hansson and Samuelsson leg.; ZMLU • 1 ♀; Mount Matang , clearing; 1°35′34″ N, 110°11′25″ E; 1000 ft a.s.l.; 1 Jun. 1939; N.C.E. Miller leg.; NHMUK 012497573 ; • 1 ♀; Mount Dulit , moss forest; 3°19′58″ N, 114°9′ E; 4000 ft a.s.l.; 22 Oct. 1932; B.M. Hobby and A.W. Moore leg.; NHMUK 012497571 • 1 ♂; 1°45′24.8″ N, 110°19′11.9″ E; 15 May 2017; N. Vladimirov leg.; iNaturalist.org/observations/8840806 • 1 ♂; Siburan ; 1°7′19.3″ N, 110°12′56.8″ E; 21 Jun. 2013; flosker leg.; iNaturalist.org/observations/19469334 • 1 ♀; Bau ; 1°22′52.2″ N, 110°13′15.3″ E; 17 Sep. 2013; ml_ls leg.; iNaturalist.org/observations/28583992 • 1 ♂; Long Lellang Airport, 98008 Marudi ; 3°25′17.5″ N, 115°9′9.5″ E; 2 Oct. 2016; Q. Lu leg.; iNaturalist.org/observations/48562286 • 1 ♂; Gunung Serapi Kuching, 93050, Petra Jaya, 93050 Kuching ; 1°36′16.3″ N, 110°11′19.6″ E; 19 Jul. 2019; plea84 leg.; iNaturalist.org/observations/29106017 • 1 ♀; Kuching ; 1°34′33.2″ N, 110°17′54.1″ E; 18 Jul. 2017; C. Shu leg.; iNaturalist.org/observations/39208711 • 1 ♀; Bahagian Miri ; 3°25′16.8″ N, 115°9′15.8″ E; 30 Sep. 2018; K. Lin leg.; iNaturalist.org/observations/20053192 .

NO COUNTRY • 1 ♂; “ South Borneo ”; H. Fruhstorfer leg.; MHN • 1 ♀; “ South Borneo ”; H. Fruhstorfer leg.; MHN • 1 ♀; “ South Borneo ”; H. Fruhstorfer leg.; MHN • 1 ♀; MHN • 1 ♀; MHN • 1 ♂; MNHN • 1 ♂; “ Borneo ”; Blanchmanche leg.; MNHN • 1 ♂; “ Borneo ”; MNHN • 1 ♀; “ Borneo ”; 1891; Chaper leg.; MNHN • 1 ♀; “ Borneo ”; 1891; Chaper leg.; MNHN • 1 ♂; “ Borneo ”; 1891; Chaper leg.; MNHN • 1 ♀; “ Borneo ”; Bruner leg.; MNHN • 1 ♂; “ Borneo ”; Bruner leg.; MNHN .

Redescription

Size average ( Figs 2 , 23E–H ), similar to N. malaya sp. nov. but differs by slightly lighter colouration, male genitalia, female copulatory papilla and juvenile colouration.

Male

GENITALIA ( Fig. 16G ). Pseudepiphallus relatively slender, slightly longer than broad, anterior margin straight, posterior margin indented in median area to faintly concave, lateral margins with basal half converge posteriorly, apical half somewhat parallel. Pseudepiphallic lophi slightly longer than wide (at base), sclerotized laterally only and covered with strong short setae; their apex slightly folded dorsally and sub-obtuse; moderately spaced apart from each other. Similar to N. malaya sp. nov. but differs by pseudepiphallus faintly more robust and broad, anterior margin more straight. Rami more straight.

Female

GENITALIA. Copulatory papilla conical, smaller and stout; apex folded ventrally, elongated, pointed; dorsal face with a sclerotized area; ventro-anterior end with an oblong rim, without truncated lobule ( Fig. 17P–Q ).

Juvenile ( Fig. 24D–F )

Characterised by pale white or yellow colouration with numerous dark stripes all around body. Dorsum of head with three transverse black stripes that extends to gena. Eyes with numerous dark stripes. Fastigium with margin black; antennal black with pale rings; frons with a few black spots; palpal segments white with apices black. Pronotum with three thick, dark longitudinal stripes on dorsal disk, one very thick velvety black stripe between dorsal disk and lateral lobe; another curved stripe on lateral lobe ventral of the thick stripe; these stripes not reaching margins. Fore and middle legs with longitudinal dark stripes, two on each face, usually one dorsal and one ventral. FIII with longitudinal dark stripes, two dorsally, and two ventrally. Thoracic (excluding pronotum) and abdominal tergites typically dark or greyish, anterior margin dark, their posterior margin whitish forming a homogeneous light/dark striped pattern. Cerci whitish with a black stripe.

Measurements

See Table 11 .

Ecology

This species can be found among shrubs and herbaceous plants (but not grasses) at the forest edge ( Fig. 27A ). Similar to N. malaya sp. nov., the adults and nymphs of this species have been observed in Brunei Darussalam to feed on the flower parts of a varieties of native and exotic plant species ( Tan et al. 2017).

Distribution

Sumatra, Borneo.

Type locality

Indonesia: Sumatra: Lampongs, Wai Lima.

Calling song (Kuala Belalong, n = 2 males, 20 echemes; MNHN-SO-2020-2859 to MNHN-SO-2020-2860) ( Fig. 25D–F )

At 27.0°C, the calling song consists of long homogeneous bouts made of short echemes consisting of triplets of syllables; echeme duration is 32.47± 3.79 ms (27.00–38.65 ms); echeme period is 55.20±5.90 ms (50.52–74.27 ms); downtime between consecutive echemes is 22.73 ± 7.26 ms (14.16–41.82 ms). Durations for the first to third syllables of the triplet are as follows: 6.93 ± 1.73 ms (4.57–9.95 ms), 7.51 ± 2.03 ms (4.80–11.67 ms) and 7.22 ± 2.82 ms (3.24–12.44 ms). The call spectrum consists of a series of harmonics with three distinct peaks corresponding to the first three harmonics: first harmonic (fundamental frequency) at 6.86 ± 0.28 kHz (6.46–7.15 kHz), is usually the dominant frequency; second harmonic peaking at 14.08 ± 0.39 kHz (13.35–14.64 kHz); third harmonic peaking at 19.96 ± 0.35 kHz (19.12–20.67 kHz).

Remarks

See N. malaya sp. nov. for arguments about considering two distinct species.

About synonymy of Nisitrus marginatus ( Walker, 1869) under N. vittatus , considering that N. vittatus is variable in colouration and widely distributed in Borneo and Sumatra, and that the only known syntype of N. marginatus from Sumatra resembles N. vittatus , we consider the species described by Walker (1869) as a junior synonym of N. vittatus .

Figures

Fig. 1. Habitus of Nisitrus vittatus (Haan, 1844) in dorsal (A, C) and lateral (B, D) views. A–B. ♂. C–D. ♀. Scale bar = 5 mm.

Fig. 3. Forewing venations (with labels of the veins and areas) of Nisitrus vittatus (Haan, 1844). A. ♂. B. ♀. Scale bar = 5 mm.

Fig. 5. Map of Southeast Asia showing the distribution of species of Nisitrus Saussure, 1878. Each coloured symbol represents one sampling or observation locality listed in the material in the text.

Fig. 6. Maximum likelihood phylogenetic tree based on the 12S marker of six species ofNisitrus Saussure, 1878. Values under branches indicate bootstrap values.

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. 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. vittatus (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.

Fig. 16. Male genitalia in ventral 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 = 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.

Fig. 26. Principal Coordinates Analysis (PCoA) of the acoustic properties of the calling songs of N. vittatus (Haan, 1844), N. malaya Robillard & Tan sp. nov. and N. musicus Ingrisch, 1987.

Fig. 2. A. Nisitrus insignis Saussure, 1878, adult ♀. B. Natural habitat of a typical Nisitrus Saussure, 1878, Lima Puluh Koto, Sumatra, Indonesia. Photograph source and credits: iNaturalist; Dries Nys and Marion Vancauwenbergh.

Fig. 23. A–D. Nisitrus malaya Robillard & Tan sp. nov. E–H. N. vittatus (Haan, 1844). Adult ♂♂ (A–B, E–F) and ♀♀ (C–D, G–H) in their natural environment. Photograph credit: Ming Kai Tan.

Fig. 24. Nymphs in their natural environment. A–C. Nisitrus malaya Robillard & Tan sp. nov. D–F. N. vittatus (Haan, 1844). Photograph credit: Ming Kai Tan.

Fig. 27. A. Habitat where Nisitrus vittatus (Haan, 1844) can be found in Sungai Belalong in Brunei. B. Habitat where Nisitrus malaya Robillard & Tan sp. nov. can be found in Singapore. The arrows point at the type of plants (Macaranga sp. in A, Piper sarmentosum Roxb. in B) on which Nisitrus can be found in relatively large numbers. Photograph credits: Yale-NUS students (A), Ming Kai Tan (B).

Fig. 25. Acoustics ofNisitrus malaya Robillard & Tan sp. nov. (A–C) and N. vittatus (Haan, 1844) (D–F). A, D. Series of echemes. B, F. One echeme. C, F. Spectral plot of one echeme.