Nisitrus malaya Robillard & Tan, 2021

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 : 44-55

Nisitrus malaya Robillard & Tan sp. nov.

urn:lsid:zoobank.org:act:AD6647DB-81AE-421B-8433-C4E3D9ACC314

Figs 4– 5 , 7E , 9A , 10F , 11F , 12F , 13E , 14F , 16E , 17J–K , 23A–D, 24A–C, 25A–C, 26 ; Tables 1–2 , 8

Nisitrus vittatus – Robillard & Desutter-Grandcolas 2004a: 276; 2004b: 578; 2004c: 304; 2006: 644; 2008: 67 ( Nisitrini tribe); 2011: 637. — Robillard et al. 2007: 1265; 2013: 2002 (mechanism of stridulation). — Desutter-Grandcolas et al. 2010: 616. — Nattier et al. 2011: 2201 (phylogeny and molecular dating). — Tan 2012: 4; 2017: 4. — Tan et al. 2012: 66; 2013a: 220; 2013b: 99; 2015: 47; 2017: 146 (flower-visiting behaviour). — Robillard & Tan 2013: 707 . — Tan & Robillard 2014: 375. — ter Hofstede et al. 2015: 3245–3252 (communication). — Schneider et al. 2017: 1–12 (tympanum structure).

Diagnosis

Average size, colouration contrasting with black and yellow, characterised by black vertex, yellow frons and orange brown legs. This new species is very close to N. vittatus from which it differs by adult and juvenile colouration, male and female genitalia and calling song. General colouration darker, particularly among FW cells and veins beneath R; R black (instead of cream-coloured as in N. vittatus ); in male venation, anal-most and middle chords narrow and nearly parallel with one another (instead of diverging apically), and basal-most chord extend from basal half of middle chord (instead of at/after midpoint). It differs from N. vittatus by the male genitalia with pseudepiphallus more robust, broad, anterior margin more strongly concave. Female copulatory papilla with truncated lobule at the ventro-anterior end. Juvenile differs strikingly in the body colour pattern. Male calling song also differs by longer echeme period. This species is also similar to N. crucius sp. nov. in size and habitus, but differs by the absence of cross-shaped black pattern on frons and does not occur in Borneo.

Etymology

This specific name is a noun in apposition referring to the speciesʼ distribution in the Malay Peninsula (including Singapore and Peninsular Malaysia).

Material examined

Holotype

SINGAPORE • ♂; Dairy Farm Nature Park ; 1°21′44″ N, 103°46′9″ E; 17 Jul. 2020; M.K. Tan leg.; ZRC.

Paratypes

MALAYSIA – Selangor • 1 ♂; Petaling Jaya, Mount Kiara ; 3°10′17.44″ N, 101°38′42.70″ E; 7 Sep. 2002; T. Robillard leg.; MNHN-EO-ENSIF3132 • 1 ♂; same collection data as for preceding; call recording; MNHN-EO-ENSIF3133 • 1 ♂; same collection data as for preceding; MNHN-EO-ENSIF3134 • 1 ♀; same collection data as for preceding; MNHN-EO-ENSIF11140 • 1 ♀; same locality data as for preceding; Aug. 2002; T. Robillard leg.; MNHN-EO-ENSIF11331 • 1 ♀; same locality data as for preceding; T. Robillard leg.; PIII: Nvith2; MNHN-EO-ENSIF1708 • 1 ♂; 1909; C.J. Brooks leg.; NHMUK .

SINGAPORE • 1 ♂; Bukit Timah , “près de l’entrée de la réserve” [near entrance]; 1°21′6.7″ N, 103°46′45.2″ E; 92 m a.s.l.; 10 Jun. 2011; T. Robillard leg.; call recording; MNHN-EO-ENSIF3136 • 1 ♂; Bukit Timah , “sommet” [summit]; 1°21′16.2″ N, 103°46′35.9″ E; 170 m a.s.l.; 29 Jun. 2009; T. Robillard leg.; call recording MNHN-SO-2016-41 ; TR9; MNHN-EO-ENSIF3102 • 1 ♂; same locality data as for preceding; 10 Jun. 2011; T. Robillard leg.; TR1; MNHN-EO-ENSIF10948 • 1 ♂; Bukit Timah Nature Reserve , Hindhede Trail; 1°20′57.1″ N, 103°46′45.2″ E; 68 m a.s.l.; 20 Jun. 2011; T. Robillard leg.; TR26; MNHN-EO-ENSIF10949 • 1 ♂; MacRitchie Reserve ; 1°21′10.2″ N, 103°46′33.6″ E; 68 m a.s.l.; 2 Jul. 2009; T. Robillard leg.; call recording; MNHN-EO-ENSIF3135 • 1 ♂; MacRitchie Reserve ; 1°21′10.2″ N, 103°46′33.6″ E; 68 m a.s.l.; 2 Jul. 2009; T. Robillard leg.; call recording; MNHN-EO-ENSIF2742 • 1 ♀; Mandai ; 1°24′52.7″ N, 103°47′22.2″ E; 27 Dec. 2015; M.K. Tan and H. Yeo leg.; C2.F.50; MNHN-EO- ENSIF11136 • 1 ♀; same collection data as for preceding; C2.F.51; MNHN-EO-ENSIF11138 • 1 ♀; same collection data as for preceding; C2.F.28; MNHN-EO-ENSIF11137 .

Additional specimens

MALAYSIA • 6 juveniles; Petaling Jaya, Mount Kiara ; 3°10′17.44″ N, 101°38′42.70″ E; 7 Sep. 2002; T. Robillard leg.; MNHN • 1 juvenile; Kepong FRIM ( Forest Research Insitute of Malaysia ); 3°13′60″ N, 101°37′51″ E; 18 Aug. 2003; Svenson leg.; MNHN • 16 ♂♂; Petaling Jaya, Mount Kiara ; T. Robillard leg.; élevage; MNHN • 11 ♀♀; same collection data as for preceding; MNHN • 1 ♀; PD Villa Country Resort, 11, Jln PDV3, Taman Rakyat, 71050 Teluk Kemang , Negeri Sembilan; 2°27′27.3″ N, 101°52′5.9″ E; 17 Feb. 2019; J. Howes leg.; iNaturalist.org/observations/20414064 • 1 ♀; Bestari Jaya , Selangor; 3°16′7″ N, 101°27′28.6″ E; 27 Apr. 2019; S. Siva Perumal leg.; iNaturalist.org/observations/24568214 • 1 ♂; Kuala Lumpur; 3°7′44.8″ N, 101°39′3.20″ E; 5 Mar. 2019; msone leg.; iNaturalist.org/observations/30373276 • 1 ♂; Kuala Lumpur; 3°7′44.8″ N, 101°39′3.20″ E; 5 Mar. 2019; msone leg.; iNaturalist.org/observations/30373274 • 1 ♂; Kuala Lumpur; 3°7′56.5″ N, 101°39′16.3″ E; 9 Jan. 2019; msone leg.; iNaturalist.org/observations/20052999 • 1 ♀; Kledong Saiong Forest Eco Park, Perak; 4°40′59.7″ N, 101°4′6.5″ E; 25 Jul. 2019; K. Wenyuh leg.; iNaturalist.org/observations/29510836 • 1 ♀; Bukit Gasing, Petaling Jaya , Selangor; 3°5′54.3″ N, 101°39′29.1″ E; 17 Nov. 2006; J. Howes leg.; iNaturalist.org/observations/35667227 • 1 ♂; Taman Tugu PT88, Jalan Sultan Salahuddin, Kuala Lumpur, 50480 Kuala Lumpur Federal Territory of Kuala Lumpur; 3°9′6.28″ N, 101°41′0.900″ E; 17 Nov. 2019; T. Maul leg.;iNaturalist.org/observations/35769550 • 1 ♀; Semenyih , Selangor; 2°56′36.2″ N, 101°54′4″ E; 3 Dec. 2019; B.C.A. Howes leg.; iNaturalist.org/observations/36316633 • 1 ♂; Bukit Persekutuan, Kuala Lumpur, Federal Territory of Kuala Lumpur; 3°8′21.3″ N, 101°40′50.6″ E; 11 Jul. 2007; J. Howes leg.; iNaturalist.org/observations/36663502 • 1 ♀; Lot 240, Jalan Raja Chulan , Kuala Lumpur, 50250 Kuala Lumpur, Wilayah Persekutuan Kuala Lumpur; 3°9′3.28″ N, 101°42′7.5″ E; 6 Jan. 2020; T. Maul leg.; iNaturalist.org/observations/37332417 • 1 ♀; 31600, Perak; 4°27′29.1″ N, 101°12′30.5″ E; 4 Jan. 2020; M. Ng leg.; iNaturalist.org/observations/37357322 • 1 ♀; Hutan Lipur Sungai Tua ; 3°20′1.2″ N, 101°43′26.1″ E; 6 Oct. 2014; T. Kirschey leg.; iNaturalist.org/observations/49430837 • 1 ♀; unnamed road, Kampung Padang, 43100 Hulu Langat , Selangor; 3°11′55.2″ N, 101°51′4″ E; 11 Jan. 2020; F. Cheong leg.; iNaturalist.org/observations/37615383 • 1 ♂; Beranang , Selangor; 2°56′17.8″ N, 101°53′56.2″ E; 19 Jan. 2020; M.A.M. Nasir leg.; iNaturalist.org/observations/37730365 • 1 ♀; Kampung Tanjung Jelai, Jerantut , Pahang; 4°4′11.1″ N, 102°19′0″ E; 20Feb.2019; djhiker leg.;iNaturalist.org/observations/24363726 • 1 ♂; 5°12′53.7″ N, 100°30′53.2″ E; 19 Mar. 2018; R. Ong leg.; iNaturalist.org/observations/29697626 • 1 ♂; 53100 Kuala Lumpur, Federal Territory of Kuala Lumpur; 3°13′53.1″ N, 101°44′25.8″ E; 15 Feb. 2020; T. Maul leg.; iNaturalist.org/observations/38668098 • 1 ♂; Balai Cerap UTM; 1°34′11.8″ N, 103°38′41″ E; 21 Feb. 2020; S. Mierul leg.; iNaturalist.org/observations/38906538 • 1 ♂; Selangor; 3°35′39.6″ N, 101°44′5.9″ E; 28 Jul. 2019; msone leg.; iNaturalist.org/observations/39490018 • 1 ♀; Johor Baharu , MY-JH; 1°32′28.1″ N, 103°40′32.2″ E; 11 Mar. 2020; gonaturelam leg.; iNaturalist.org/observations/39833380 • 1 ♂; 2°50′40.7″ N, 101°56′32.2″ E; 1 May 2019; anukma leg.; iNaturalist.org/observations/24256308 • 1 juvenile; Taman Tugu PT88, Jalan Sultan Salahuddin, Kuala Lumpur, 50480 Kuala Lumpur, Federal Territory of Kuala Lumpur; 3°9′6.28″ N, 101°41′0.900″ E; 17 Nov. 2019; T. Maul leg.; iNaturalist.org/observations/35769588 • 1 ♀; Mersing , Johor; 2°25′7.56″ N, 103°51′22.8″ E; 7 Feb. 2020; msone leg.; iNaturalist.org/observations/38427233 • 1 juvenile; Tamarind Springs Jalan 1, Taman Tun Abdul Razak, 68000 Ampang Jaya , Selangor; 3°10′14.6″ N, 101°46′14.3″ E; 21 Jun. 2020; T. Maul leg.; iNaturalist.org/observations/50422192 • 1 juvenile; Selama ; 5°13′6.5″ N, 100°41′36.5″ E; 5 Jul. 2020; M. Syarifudin leg.; iNaturalist.org/observations/51984928 • 1 juvenile; Tamarind Springs Jalan 1, Taman Tun Abdul Razak, 68000 Ampang Jaya , Selangor; 3°10′14.6″ N, 101°46′14.3″ E; 5 Sep. 2020; T. Maul leg.; iNaturalist.org/observations/58822053 • 1 ♂; Bentong , Pahang, MY; 3°18′47.2″ N, 101°52′49.6″ E; 12 Sep. 2020; J. Marriappan leg.; iNaturalist.org/observations/59301449 • 1 juvenile; Tamarind Springs Jalan 1, Taman Tun Abdul Razak, 68000 Ampang Jaya , Selangor; 3°10′14.6″ N, 101°46′14.3″ E; 19 Sep. 2020; T. Maul leg.; iNaturalist.org/observations/60070623 • 1 ♂; Jalan Melawati 5, Taman Zoo View , Selangor, MY; 3°13′0.400″ N, 101°45′2″ E; 2 Sep. 2020; K.J.M. Anbumani leg.;iNaturalist.org/observations/60166965 • 1 juvenile; Taman Bukit Perdana, 83000 Batu Pahat , Johor; 1°50′53.6″ N, 102°57′37.3″ E; 26 Sep. 2020; J. Neo leg.; iNaturalist.org/observations/60810954 • 1 ♂; Kulaijaya ; 1°34′12.6″ N, 103°38′40.1″ E; 2 Mar. 2020; A. Zaidi leg.; iNaturalist.org/observations/39477877 • 1 juvenile; Tamarind Springs Jalan 1, Taman Tun Abdul Razak, 68000 Ampang Jaya , Selangor; 3°10′14.6″ N, 101°46′14.3″ E; 8 Mar. 2020; T. Maul leg.; iNaturalist.org/observations/39698484 • 1 ♂; UTM Skudai, Johor Bahru ; 1°33′26.3″ N, 103°38′14.1″ E; 29 Apr. 2020; B. Kumpau leg.; iNaturalist.org/observations/44237939 • 1 juvenile; Kuala Lumpur, 50480 Kuala Lumpur, Federal Territory of Kuala Lumpur; 3°9′23.1″ N, 101°41′5.9″ E; 26 May 2020; H. Shanmugam leg.; iNaturalist.org/observations/47737528 • 1 ♀; 2°48′23.6″ N, 101°52′20.3″ E; 3 May 2020; anukma leg.; iNaturalist.org/observations/47829023 • 1 ♂; Klinik Kesihatan Sungai Lembing, Jalan Sungai Lembing, Kampung Bayas , 26200 Sungai Lembing , Pahang; 3°55′31″ N, 103°4′17″ E; 7 Jun. 2020; A. Yusuf leg.; iNaturalist.org/observations/48735148 • 1 ♀; Perdana Botanical Gardens, Kuala Lumpur, Federal Territory of Kuala Lumpur; 3°8′41.7″ N, 101°41′10.6″ E; 19 Jun. 2020; N. Rahman leg.; iNaturalist.org/observations/50133159 • 1 ♂; Tamarind Springs Jalan 1, Taman Tun Abdul Razak, 68000 Ampang Jaya , Selangor; 3°10′14.6″ N, 101°46′14.3″ E; 20 Jun. 2020; T. Maul leg.; iNaturalist.org/observations/50276251 • 1 ♂; Seberang Perai Selatan; 5°17′13.6″ N, 100°30′30″ E; 5 Jul. 2020; alexiuslzl leg.; iNaturalist.org/observations/52006113 • 1 ♀; KJ75 Pertubuhan Peladang, Pekan Batu 14 Hulu Langat , 43100 Hulu Langat , Selangor; 3°6′50.1″ N, 101°48′56.3″ E; 19 Jul. 2020; S. Saphian leg.; iNaturalist.org/observations/53567997 • 1 juvenile; Batu Pahat , Johor; 1°50′28″ N, 102°57′36.8″ E; 31 Jul. 2020; diclim_eschsbp leg.; iNaturalist.org/observations/54914512 • 1 ♂; Taman Bukit Perdana, 83000 Batu Pahat , Johor; 1°50′53.6″ N, 102°57′37.3″ E; 31 Jul. 2020; ziki_eschsbp leg.; iNaturalist.org/observations/54932721 • 1 juvenile; NCP, Taman Tugu , Kuala Lumpur; 3°9′4.4″ N, 101°41′8.13″ E; 9 Aug. 2020; T. Maul leg.; iNaturalist.org/observations/55895941 • 1 ♀; NCP, Taman Tugu , Kuala Lumpur; 3°9′4.4″ N, 101°41′8.13″ E; 9 Aug. 2020; T. Maul leg.; iNaturalist.org/observations/55896227 • 1 juvenile; Jalan Perdana 17, Taman Bukit Perdana, 83000 Batu Pahat , Johor; 1°50′53.7″ N, 102°57′37.7″ E; 16 Aug. 2020; ziki_eschsbp leg.; iNaturalist.org/observations/56605014 • 1 ♂, 1 ♀; Jalan Perdana 17, Taman Bukit Perdana, 83000 Batu Pahat , Johor; 1°50′53.7″ N, 102°57′37.7″ E; 16 Aug. 2020; ziki_eschsbp leg.; iNaturalist.org/observations/56605016 • 1 juvenile; Taman Bukit Indah, 68000 Ampang Jaya , Selangor; 3°9′34.7″ N, 101°46′34.3″ E; 20 Aug. 2020; T. Maul leg.; iNaturalist.org/observations/57008216 • 1 ♂; Taman Bukit Perdana, 83000 Batu Pahat , Johor; 1°50′53.6″ N, 102°57′37.3″ E; 20 Aug. 2020; ziki_ eschsbp leg.; iNaturalist.org/observations/57014573 • 1 ♀; 83200 Senggarang , Johor; 1°46′20″ N, 103°1′33.1″ E; 22 Aug. 2020; ziki_eschsbp leg.; iNaturalist.org/observations/57193924 • 1 ♂; Taman Bukit Perdana 2, 83000 Batu Pahat , Johor; 1°50′32.8″ N, 102°57′15″ E; 20 Aug. 2020; tstan_eschsbp leg.; iNaturalist.org/observations/57307974 • 1 ♀; NCP, Taman Tugu , Kuala Lumpur; 3°9′4.4″ N, 101°41′8.13″ E; 23 Aug. 2020; T. Maul leg.; iNaturalist.org/observations/57311814 • 1 juvenile; Jalan Cenderasari , Kuala Lumpur; 3°8′44.6″ N, 101°41′18.5″ E; 30 Aug. 2020; mengarooo leg.; iNaturalist.org/observations/58026678 • 1 juvenile; Tamarind Springs Jalan 1, Taman Tun Abdul Razak, 68000 Ampang Jaya , Selangor; 3°10′14.6″ N, 101°46′14.3″ E; 31 Aug. 2020; T. Maul leg.; iNaturalist.org/observations/58233139 • 1 ♀; 83200 Senggarang , Johor; 1°44′6.4″ N, 103°4′1.2″ E; 22 Aug. 2020; J. Neo leg.; iNaturalist.org/observations/58578339 • 1 ♂; Choa Chu Kang Park ; 1°23′11.9″ N, 103°44′51″ E; 6 Sep. 2020; mingwei leg.; iNaturalist.org/observations/58695507 • 1 ♂, 1 ♀; Tamarind Springs Jalan 1, Taman Tun Abdul Razak, 68000 Ampang Jaya , Selangor; 3°10′14.6″ N, 101°46′14.3″ E; 5 Sep. 2020; T. Maul leg.; iNaturalist.org/observations/58826217 • 1 ♀; Ampang Jaya , Selangor; 3°13′0.100″ N, 101°45′3.5″ E; 20 Sep. 2020; B. Sg leg.; iNaturalist.org/observations/60147373 • 1 ♂, 1 ♀; 83200 Senggarang , Johor; 1°46′36.8″ N, 103°1′24.6″ E; 4 Oct. 2020; ziki_eschsbp leg.; iNaturalist.org/observations/61669789 • 1 juvenile; 83200 Senggarang , Johor; 1°46′36.8″ N, 103°1′24.6″ E; 4 Oct. 2020; ziki_eschsbp leg.; iNaturalist.org/observations/61669797 • 1 ♂; Garden Avenue, 70300 Seremban , Negeri Sembilan; 2°40′39.7″ N, 101°54′2.70″ E; 7 Oct. 2020; O.J. Seng leg.; iNaturalist.org/observations/61917637 • 1 ♂; Garden Avenue, 70300 Seremban , Negeri Sembilan; 2°40′53″ N, 101°53′56.2″ E; 7 Oct. 2020; O.J. Seng leg.; iNaturalist.org/observations/61920273 • 1 juvenile; Bukit Kiara, 60000 Kuala Lumpur, Federal Territory of Kuala Lumpur; 3°9′12″ N, 101°38′11″ E; 8 Oct. 2020; V. Ganesh leg.; iNaturalist.org/observations/62013556 • 1 juvenile; Kuala Lumpur, Federal Territory of Kuala Lumpur ; 3°8′20.3″ N, 101°41′12.6″ E; 8 Oct. 2020; C.W. Chun leg.; iNaturalist.org/observations/62037651 • 1 ♀; Perdana Botanical Gardens, Kuala Lumpur, Federal Territory of Kuala Lumpur ; 3°8′37.2″ N, 101°41′9.9″ E; 8 Oct. 2020; C.W. Chun leg.; iNaturalist.org/observations/62075913 • 1 ♀; 1°50′33.5″ N, 102°57′38.2″ E; 10 Oct. 2020; joyce_tan leg.; iNaturalist.org/observations/62628458 • 1 juvenile; Petaling Jaya , Selangor; 3°10′33.6″ N, 101°36′31.1″ E; 25 Oct. 2020; msone leg.; iNaturalist.org/observations/63555073 • 1 juvenile; Petaling Jaya , Selangor; 3°10′33.6″N, 101°36′31.1″E; 25Oct.2020; msone leg.;iNaturalist.org/observations/63555074 • 1 juvenile; Perdana Botanical Gardens, Kuala Lumpur, Federal Territory of Kuala Lumpur ; 3°8′37.2″ N, 101°41′9.9″ E; 8 Oct. 2020; C.W. Chun leg.; iNaturalist.org/observations/62078581 • 1 ♂, 1 ♀; Gombak ; 3°15′41.7″ N, 101°38′7″ E; 28Apr. 2019; Y. Joyce leg.; iNaturalist.org/observations/23581051 • 1 juvenile; Gunung Lambak ; 2°1′39″ N, 103°21′27″ E; 2Aug. 2020; howe leg.; iNaturalist.org/observations/55174550 • 1 ♀; Kuala Terengganu; 5°16′55″ N, 103°6′32″ E; 27 Dec. 2013; Lanzz leg.; https://www.projectnoah. org/spottings/130726139?fbclid=IwAR1CsvK9ownDvtCZtRiad5lj9dnh3aKasTEVLyyZGkwfXm1PNj dkKMrq-TM .

SINGAPORE • 1 juvenile; Central Catchment Nature Reserve , “sentier forestier”; 1°22′49″ N, 103°49′6″ E; 79 m a.s.l.; 16 Jun. 2011; T. Robillard leg.; MNHN • 1 ♂; Elevage F1; T. Robillard leg.; MNHN • 3 ♂♂; Elevage F2; T. Robillard leg.; MNHN • 2 ♂♂; Elevage F3; T. Robillard leg.; MNHN • 1 ♂; Elevage F1; T. Robillard leg.; MNHN • 1 ♂; Elevage F4; T. Robillard leg.; MNHN • 1 ♂; Elevage F5; T. Robillard leg.; MNHN • 1 ♂; Elevage F6; T. Robillard leg.; MNHN • 1 ♂; Elevage F9; T. Robillard leg.; MNHN-EO-ENSIF3102 • 4 ♂♂; Elevage ; T. Robillard leg.; MNHN • 1 ♀; Elevage F2; T. Robillard leg.; MNHN • 1 ♀; Elevage F3; T. Robillard leg.; MNHN • 1 ♀; Elevage F5; T. Robillard leg.; MNHN • 3 ♀♀; Elevage ; T. Robillard leg.; MNHN • 1 ♀; Lim Chu Kang ; 1°26′46.1″ N, 103°43′45.7″ E; 26 Dec. 2019; E.C. Maxwell leg.; iNaturalist.org/observations/39013226 • 1 ♂; Blackmore Drive , Holland Village; 1°19′59.8″ N, 103°46′53.1″ E; 27 Oct. 2020; stefaniek leg.; iNaturalist.org/observations/63613982 • 1 ♂; Central Water Catchment ; 1°20′48.2″ N, 103°50′8.5″ E; 10 Apr. 2020; M. Finlay leg.; iNaturalist.org/observations/41811938 • 1 ♂; Blk 70A; 1°16′35.6″ N, 103°48′42″ E; 23 May 2020; lcwoo leg.; iNaturalist.org/observations/46927932 • 1 ♀; Woodlands (SLE) Park Connector, Woodlands; 1°25′36.7″ N, 103°47′14.3″ E; 29 May 2020; looiec leg.; iNaturalist.org/observations/47701690 • 1 ♀; Champions Way , Woodlands; 1°25′40.1″ N, 103°47′16.8″ E; 29 May 2020; looiec leg.; iNaturalist.org/observations/47701715 • 1 juvenile; 36C Lor Halwa 298637; 1°20′10″ N, 103°49′22.6″ E; 9 May 2020; flyingswallow123 leg.; iNaturalist.org/observations/48998890 • 1 ♀; Tanglin ; 1°18′2.5″ N, 103°48′41.1″ E; 15 Jun. 2020; D. Rham leg.; iNaturalist.org/observations/49656881 • 1 ♂; Yishun ; 1°24′51.7″ N, 103°50′34.2″ E; 20 Jun. 2020; M. Ng leg.; iNaturalist.org/observations/50287104 • 1 ♂, 1 ♀; Rambutan Trail , Thomson; 1°23′6.28″ N, 103°49′13.8″ E; 28 Jun. 2020; simchoonheok leg.; iNaturalist.org/observations/51206798 • 1 juvenile; Tengah ; 1°21′58″ N, 103°43′47.2″ E; 23 Jun. 2020; C.W. Gan leg.; iNaturalist.org/observations/51352950 • 1 ♀; Bishan ; 1°20′57.1″ N, 103°49′59.8″ E; 27 Jun. 2020; M. Finlay leg.; iNaturalist.org/observations/51885537 • 1 ♂; Bukit Timah Nature Reserve ; 1°21′38.8″ N, 103°46′38.1″ E; 5 Jul. 2020; ramesh-birding-butterflying leg.; iNaturalist.org/observations/52003328 • 1 ♂; Kent Ridge Park , Pasir Panjang; 1°17′7.28″ N, 103°47′28.3″ E; 2 Jul. 2020; skm3355 leg.; iNaturalist.org/observations/52035813 • 1 ♀; Bef Dairy Farm Road ; 1°22′0″ N, 103°46′32″ E; 10 Jul. 2020; N. Sanjai leg.; iNaturalist.org/observations/52529111 • 1 ♀; Bukit Panjang ; 1°21′50.8″ N, 103°46′34.2″ E; 10 Jul. 2020; N. Sanjai leg.; iNaturalist.org/observations/52536392 • 1 ♀; Bef Old Upp Thomson Road ; 1°23′12.3″ N, 103°49′12.9″ E; 10 Jul. 2020; Y.H. Koh leg.; iNaturalist.org/observations/52542825 • 1 ♂; MacRitchie Reservoir ; 1°20′41.3″ N, 103°49′20.5″ E; 10 Jul. 2020; falseamoebae leg.; iNaturalist.org/observations/52574945 • 1 ♀; Novena ; 1°20′0.598″ N, 103°49′39.3″ E; 10 Jul. 2020; M. Finlay leg.; iNaturalist.org/observations/52644701 • 1 ♀; Upper Seletar Reservoir Park ; 1°23′58.7″ N, 103°48′29.3″ E; 10 Jul. 2020; K. Well leg.;iNaturalist.org/observations/52703749 • 1 juvenile; Bukit Panjang ; 1°20′52.7″ N, 103°46′30.3″ E; 14 Jul. 2020; M. Nasry leg.; iNaturalist.org/observations/53034819 • 1 ♀; Rifle Range Link ; 1°21′22.6″ N, 103°48′6″ E; 12 Jul. 2020; K. Well leg.; iNaturalist.org/observations/53036305 • 1 ♀; 100 Dairy Farm Road , 679057; 1°21′42.7″ N, 103°46′31.1″ E; 17 Jul. 2020; M. Ng leg.; iNaturalist.org/observations/53356890 • 1 ♀; 100 Dairy Farm Road , 679057; 1°21′42.7″ N, 103°46′31.1″ E; 17 Jul. 2020; M. Ng leg.; iNaturalist.org/observations/53356981 • 1 ♂; Central Water Catchment ; 1°20′48.7″ N, 103°50′8″ E; 12 Jul. 2020; M. Finlay leg.; iNaturalist.org/observations/53451611 • 1 ♂; Central Water Catchment ; 1°20′46.7″ N, 103°50′9.3″ E; 12 Jul. 2020; M. Finlay leg.; iNaturalist.org/observations/53451612 • 1 ♀; Bishan ; 1°20′51.5″ N, 103°50′7.70″ E; 12 Jul. 2020; M. Finlay leg.; iNaturalist.org/observations/53452345 • 1 ♂; Novena ; 1°20′8.13″ N, 103°49′40.6″ E; 18 Jul. 2020; M. Finlay leg.; iNaturalist.org/observations/53454664 • 1 ♀; Upper Thomson Road ; 1°23′7.4″ N, 103°49′15.6″ E; 19 Jul. 2020; M. Ng leg.; iNaturalist.org/observations/53596352 • 1 ♀; Wallace Centre , Dairy Farm Nature Park; 1°21′35.3″ N, 103°46′39″ E; 19 Jul. 2020; K. Well leg.; iNaturalist.org/observations/53622733 • 1 ♂, 1 ♀; Bukit Panjang ; 1°21′42.7″ N, 103°46′31.1″ E; 19 Jul. 2020; lcwoo leg.; iNaturalist.org/observations/53668007 • 1 juvenile; Dairy Farm Nature Park ; 1°21′42.7″ N, 103°46′31.1″ E; 19 Jul. 2020; lcwoo leg.; iNaturalist.org/observations/53673593 • 1 ♂; Bukit Timah Nature Reserve , Bukit Timah; 1°21′22.1″ N, 103°46′8.56″ E; 18 Jul. 2020; gcynature leg.; iNaturalist.org/observations/53775496 • 1 ♂; 1°22′22.5″ N, 103°49′26.6″ E; 24 Jul. 2020; noahartpradid leg.; iNaturalist.org/observations/54131093 • 1 juvenile; Windsor Park ; 1°21′41.3″ N, 103°49′35.5″ E; 22 Jul. 2020; C.W. Gan leg.; iNaturalist.org/observations/54154877 • 1 ♂; Neo Tiew Lane 2 at Kranji Marsh; 1°25′0.80″ N, 103°43′34.2″ E; 25 Jul. 2020; K. Well leg.; iNaturalist.org/observations/54259065 • 1 ♀; Central Water Catchment ; 1°22′26.3″ N, 103°49′13.6″ E; 26 Jul. 2020; gracelsy leg.; iNaturalist.org/observations/54379190 • 1 ♂; Thomson Nature Park ; 1°22′56″ N, 103°49′19.5″ E; 30 Jul. 2020; K. Well leg.; iNaturalist.org/observations/54847762 • 1 ♂, 1 ♀; Thomson Nature Park ; 1°23′7.4″ N, 103°49′15.6″ E; 31 Jul. 2020; S. Chan leg.; iNaturalist.org/observations/54945933 • 1 ♂; Thomson Nature Park ; 1°23′7.4″ N, 103°49′15.6″ E; 31 Jul. 2020; S. Chan leg.; iNaturalist.org/observations/54946444 • 1 juvenile; Lorong Pelita , Thomson; 1°22′58.8″ N, 103°49′17.3″ E; 31 Jul. 2020; yeehwa leg.; iNaturalist.org/observations/55005657 • 1 juvenile; North ; 1°22′5.9″ N, 103°46′52.1″ E; 1 Aug. 2020; S.G. Wen leg.; iNaturalist.org/observations/55005954 • 1 ♂, 1 ♀; Central Water Catchment ; 1°23′49.1″ N, 103°48′13″ E; 2 Aug. 2020; S.K. Yung leg.; iNaturalist.org/observations/55122889 • 1 juvenile; Sungei Buloh Wetland Reserve ; 1°26′46.2″ N, 103°43′39.7″ E; 2 Aug. 2020; K. Well leg.; iNaturalist.org/observations/55167429 • 1 ♂; Botanic Gardens , Holland Village; 1°18′28.8″ N, 103°49′3″ E; 3 Aug. 2020; pb1785 leg.; iNaturalist.org/observations/55235146 • 1 ♂; Aft Tagore Drive ; 1°23′5.78″ N, 103°49′14.9″ E; 8 Aug. 2020; lcwoo leg.; iNaturalist.org/observations/55762965 • 1 juvenile; Bukit Drive , Bukit Timah; 1°20′54.5″ N, 103°46′29.8″ E; 2 Aug. 2020; gcynature leg.; iNaturalist.org/observations/56017697 • 1 ♂; Central Water Catchment ; 1°20′44.7″ N, 103°50′10.1″ E; 8 Aug. 2020; T.Z. Wan leg.; iNaturalist.org/observations/56103224 • 1 juvenile; Andrews Avenue , Sembawang; 1°27′26.8″N, 103°50′21.1″E; 11Aug.2020; K.Well leg.;iNaturalist.org/observations/56116568 • 1 ♀; Mandai ; 1°25′3.5″ N, 103°47′27.8″ E; 16 Aug. 2020; S.K. Yung leg.; iNaturalist.org/observations/56602046 • 1 ♀; Central Water Catchment ; 1°23′50.7″ N, 103°48′20.3″ E; 16 Aug. 2020; kangmin leg.; iNaturalist.org/observations/56625952 • 1 juvenile; Bukit Panjang ; 1°21′2.5″ N, 103°46′44.2″ E; 3 Aug. 2020; T.Z. Wan leg.; iNaturalist.org/observations/56827799 • 1 juvenile; Admiralty West Park Connector , Woodlands; 1°27′4.4″ N, 103°46′48″ E; 18 Aug. 2020; M. Farook leg.; iNaturalist.org/observations/56946793 • 1 ♀; Lower Peirce Reservoir ; 1°22′12.3″ N, 103°49′23.1″ E; 21 Aug. 2020; G. Lim leg.; iNaturalist.org/observations/57110204 • 1 ♀; Bukit Batok East Avenue 2, Bukit Batok Nature Park Playground; 1°20′59.3″ N, 103°45′49″ E; 22 Aug. 2020; V. Lim leg.; iNaturalist.org/observations/57189865 • 1 juvenile; Bukit Batok ; 1°20′58.6″ N, 103°45′51.6″ E; 22 Aug. 2020; V. Lim leg.; iNaturalist.org/observations/57191688 • 1 ♂; Lim Chu Kang ; 1°25′7.5″ N, 103°43′15.6″ E; 23 Aug. 2020; S.K. Yung leg.; iNaturalist.org/observations/57315540 • 1 ♀; Lower Peirce Reservoir Park ; 1°22′23″ N, 103°49′26.8″ E; 23 Aug. 2020; darkendaltrax leg.; iNaturalist.org/observations/57315659 • 1 ♀; Upper Seletar Reservoir Park ; 1°24′3.20″ N, 103°48′25.3″ E; 23 Aug. 2020; S. Chan leg.; iNaturalist.org/observations/57317179 • 1 juvenile; 1°20′46.7″ N, 103°53′9.63″ E; 23 Aug. 2020; lowch leg.; iNaturalist.org/observations/57318298 • 1 ♀; Bukit Timah Nature Reserve ; 1°21′51.1″ N, 103°46′33.2″ E; 24 Aug. 2020; noahartpradid leg.; iNaturalist.org/observations/57429535 • 1 ♂; Lower Peirce Reservoir ; 1°22′12.3″ N, 103°49′23.1″ E; 29 Aug. 2020; G. Lim leg.; iNaturalist.org/observations/57916937 • 1 ♀; Lim Chu Kang ; 1°25′1.2″ N, 103°43′32.3″ E; 3 Aug. 2020; S.K. Yung leg.; iNaturalist.org/observations/58007762 • 1 ♂, 1 ♀; Upper Thomson Road ; 1°23′7.4″N, 103°49′15.6″E; 10Jul.2020; C. Tong leg.;iNaturalist.org/observations/58179315 • 1 ♂; Sungei Kadut ; 1°25′24.8″ N, 103°45′19.8″ E; 18 Aug. 2020; S. Hui leg.; iNaturalist.org/observations/58230167 • 1 ♂, 1 ♀; Champions Golf Course , Bukit Timah; 1°20′24.1″ N, 103°47′54.6″ E; 28 Aug. 2020; noahartpradid leg.; iNaturalist.org/observations/58407499 • 1 ♀; Ang Mo Kio ; 1°22′52.5″ N, 103°49′16.8″ E; 6 Sep. 2020; lowch leg.; iNaturalist.org/observations/58680372 • 1 juvenile; Zhenghua Park ; 1°22′36.2″ N, 103°46′34.8″ E; 6 Sep. 2020; S. Chan leg.; iNaturalist.org/observations/58696289 • 1 ♀; 1°21′49.2″ N, 103°46′19.8″ E; 7 Sep. 2020; kei_y leg.; iNaturalist.org/observations/58931194 • 1 ♀; 1°21′50.2″ N, 103°46′36.1″ E; 1 Sep. 2020; kei_y leg.; iNaturalist.org/observations/58931653 • 1 juvenile; Bef Mandai Road ; 1°24′37.2″ N, 103°46′41.8″ E; 4 Sep. 2020; T.Z. Wan leg.; iNaturalist.org/observations/58957627 • 1 ♂, 1 ♀; Dairy Farm Nature Park ; 1°21′42.7″ N, 103°46′31.1″ E; 9 Sep. 2020; S. Chan leg.; iNaturalist.org/observations/59036868 • 1 ♀; Central Water Catchment ; 1°22′16.5″ N, 103°46′46.2″ E; 9 Sep. 2020; S. Chan leg.; iNaturalist.org/observations/59038833 • 1 juvenile; Upper Seletar Reservoir Park ; 1°24′3.20″ N, 103°48′25.3″ E; 11 Sep. 2020; S. Chan leg.; iNaturalist.org/observations/59209460 • 1 ♀; Novena ; 1°20′13.9″ N, 103°49′36.8″ E; 29 Aug. 2020; M. Finlay leg.; iNaturalist.org/observations/59402675 • 1 ♂; Central Water Catchment ; 1°23′47.7″ N, 103°48′12.1″ E; 13 Sep. 2020; S.K. Yung leg.; iNaturalist.org/observations/59419512 • 1 ♀; Central Water Catchment ; 1°23′48.8″ N, 103°48′13.1″ E; 13 Sep. 2020; S.K. Yung leg.; iNaturalist.org/observations/59419728 • 1 ♀; 1°23′5.28″ N, 103°49′21.3″ E; 15 Sep. 2020; kei_y leg.; iNaturalist.org/observations/59666269 • 1 ♂; 1°22′56.7″ N, 103°49′16.8″ E; 15 Sep. 2020; kei_y leg.; iNaturalist.org/observations/59666275 • 1 ♀; Champions Golf Course , Bukit Timah; 1°20′28.3″N, 103°47′53.7″E; 16Sep.2020; noahartpradid leg.;iNaturalist.org/observations/59759002 • 1 ♀; Bukit Panjang ; 1°21′44.3″ N, 103°46′36.1″ E; 20 Sep. 2020; M. Nasry leg.; iNaturalist.org/observations/60171530 • 1 ♂; Upper Seletar Reservoir Park ; 1°24′3.20″ N, 103°48′25.3″ E; 19 Sep. 2020; S. Chan leg.; iNaturalist.org/observations/60189103 • 1 ♀; Kranji Marshes , Neo Tiew Lane 2; 1°25′0.80″ N, 103°43′35.7″ E; 20 Sep. 2020; mingwei leg.; iNaturalist.org/observations/60196972 • 1 ♀; 1°19′14.3″ N, 103°40′42.2″ E; 20 Sep. 2020; prakashbr leg.; iNaturalist.org/observations/60402098 • 1 ♂; Dairy Farm Nature Park ; 1°21′42.7″ N, 103°46′31.1″ E; 24 Sep. 2020; S. Chan leg.; iNaturalist.org/observations/60596693 • 1 ♂; Block 265; 1°20′59.7″ N, 103°45′35.1″ E; 25 Sep. 2020; yq_1810 leg.; iNaturalist.org/observations/60696154 • 1 ♂; Tanglin ; 1°19′11.3″ N, 103°48′54.3″ E; 26 Sep. 2020; scarlett26 leg.; iNaturalist.org/observations/60807200 • 1 juvenile; Bukit Panjang ; 1°21′22.6″ N, 103°46′56.7″ E; 26 Sep. 2020; S. Chan leg.; iNaturalist.org/observations/60811750 • 1 ♀; Dairy Farm Nature Park ; 1°21′42.7″ N, 103°46′31.1″ E; 26 Sep. 2020; S. Chan leg.; iNaturalist.org/observations/60817045 • 1 juvenile; Bukit Panjang ; 1°21′14.6″ N, 103°46′37″ E; 27 Sep. 2020; S. Chan leg.; iNaturalist.org/observations/60972069 • 1 ♀; Thomson Nature Park ; 1°23′7.5″ N, 103°49′15.6″ E; 1 Oct. 2020; G. Lim leg.; iNaturalist.org/observations/61387731 • 1 ♂; Queenstown ; 1°17′6.70″ N, 103°47′29″ E; 6 Oct. 2020; S.K. Yung leg.; iNaturalist.org/observations/61843628 • 1 ♀; Thomson Nature Park ; 1°22′53.3″ N, 103°49′20.1″ E; 4 Oct. 2020; patricialorenz leg.; iNaturalist.org/observations/61869845 • 1 ♂; 100 Dairy Farm Road , 679057; 1°21′42.7″ N, 103°46′31.1″ E; 9 Oct. 2020; C. Tong leg.;iNaturalist.org/observations/62099010 • 1 ♂; Tanglin , Swan Lake; 1°18′28.5″ N, 103°48′55.2″ E; 11 Oct. 2020; L.J. Yi leg.; iNaturalist.org/observations/62263904 • 1 ♂, 1 ♀; Botanic Gardens ; 1°18′49.7″ N, 103°48′57.2″ E; 11 Oct. 2020; lcwoo leg.; iNaturalist.org/observations/62384745 • 1 ♂; Holland Village ; 1°19′50.5″ N, 103°46′52.7″ E; 13 Oct. 2020; geledek leg.; iNaturalist.org/observations/62453318 • 1 juvenile; Unnamed Road ; 1°26′31.8″ N, 103°44′9″ E; 13 Oct. 2020; S.K. Yung leg.;iNaturalist.org/observations/62472505 • 1 ♂; Central Water Catchment ; 1°22′24.8″ N, 103°49′27.8″ E; 16 Oct. 2020; Y. Jun leg.; iNaturalist.org/observations/62719526 • 1 ♂; Yio Chu Kang ; 1°24′7.06″ N, 103°49′4.78″ E; 17 Oct. 2020; M. Ng leg.; iNaturalist.org/observations/62797963 • 1 ♀; Central Water Catchment ; 1°23′58.7″ N, 103°48′28.6″ E; 18 Oct. 2020; S.K. Yung leg.; iNaturalist.org/observations/62877098 • 1 ♂; HortPark ; 1°16′43.5″ N, 103°48′6.56″ E; 18 Oct. 2020; S. Chan leg.; iNaturalist.org/observations/62896486 • 1 ♀; Kent Ridge Park ; 1°17′2″ N, 103°47′28.6″ E; 18 Oct. 2020; S. Chan leg.; iNaturalist.org/observations/62898852 • 1 ♂; Woodlands ; 1°26′59.6″ N, 103°46′43.2″ E; 19 Oct. 2020; Y. Jun leg.; iNaturalist.org/observations/63007200 • 1 ♀; Holland Village ; 1°19′49.8″ N, 103°46′53″ E; 20 Oct. 2020; irwin88 leg.; iNaturalist.org/observations/63062077 • 1 ♀; Holland Village ; 1°20′2.78″ N, 103°46′53.7″ E; 20 Oct. 2020; rocherussel leg.; iNaturalist.org/observations/63062082 • 1 ♀; Holland Village ; 1°19′50.3″ N, 103°46′52.8″ E; 20 Oct. 2020; khim99 leg.; iNaturalist.org/observations/63062122 • 1 ♀; Bukit Timah Road , Holland Village; 1°20′8″ N, 103°46′53.2″ E; 20 Oct. 2020; i328026 leg.; iNaturalist.org/observations/63062126 • 1 ♀; BukitTimah ; 1°19′33.5″N, 103°46′51.5″E; 20Oct.2020; kangmin leg.;iNaturalist.org/observations/63076249 • 1 ♂; Central Water Catchment ; 1°21′18.5″ N, 103°47′49.8″ E; 23 Oct. 2020; C. Tong leg.; iNaturalist.org/observations/63332336 • 1 ♀; Central Water Catchment ; 1°21′18.5″ N, 103°47′49.8″ E; 23 Oct. 2020; C. Tong leg.; iNaturalist.org/observations/63332337 • 1 ♀; Novena ; 1°20′7.20″ N, 103°49′43.2″ E; 22 Oct. 2020; M. Finlay leg.; iNaturalist.org/observations/63354764 • 1 ♂, 2 ♀♀; Novena ; 1°20′7.78″ N, 103°49′43.2″ E; 22 Oct. 2020; M. Finlay leg.; iNaturalist.org/observations/63354765 • 1 ♀; Novena ; 1°20′7.70″ N, 103°49′41.1″ E; 18 Oct. 2020; M. Finlay leg.; iNaturalist.org/observations/63357376 • 1 ♂; Novena ; 1°20′8.8″ N, 103°49′44.2″ E; 18 Oct. 2020; M. Finlay leg.;iNaturalist.org/observations/63357384 • 1 ♂, 1 ♀; Central Water Catchment ; 1°23′55.5″ N, 103°46′40.1″ E; 23 Oct. 2020; M. Finlay leg.; iNaturalist.org/observations/63371589 • 1 juvenile; Kent Ridge Park ; 1°17′2″ N, 103°47′28.6″ E; 24 Oct. 2020; S. Chan leg.; iNaturalist.org/observations/63383652 • 1 ♂; Kent Ridge Park , Pasir Panjang; 1°16′54.7″ N, 103°47′33.7″ E; 14 Oct. 2020; skm3355 leg.; iNaturalist.org/observations/63400835 • 1 juvenile; Hindhede Nature Park ; 1°20′56.1″ N, 103°46′32.6″ E; 25 Oct. 2020; lcwoo leg.; iNaturalist.org/observations/63466017 • 1 ♂; Aft WʼLands Waterfront Park ; 1°27′2.89″ N, 103°46′47″ E; 29 Oct. 2020; S.G. Wen leg.; iNaturalist.org/observations/63750085 • 1?; Bukit Panjang ; 1°21′37.7″ N, 103°46′37.3″ E; 30 Oct. 2020; M. Nasry leg.; iNaturalist.org/observations/63878925 • 1 ♂, 1 ♀; Thomson Nature Park ; 1°23′7.5″ N, 103°49′15.6″ E; 12 Jul. 2020; G. Lim leg.; iNaturalist.org/observations/52933922 • 1 ♂; Sungei Buloh Wetland Reserve ; 1°26′53.1″ N, 103°43′28.5″ E; 28 Dec. 2013; T.K. Hui leg.; iNaturalist.org/observations/ 19011702 .

Description

Habitus similar to N. vittatus ( Figs 7E , 23A–D ): vertex black with cream-coloured margin around eyes ( Fig. 11F ). Fastigium velvety black with cream-coloured margins. Scapes dark red brown to black (particularly on dorso-inner margins). Antennae black with wide whitish rings. Frons and mouthparts most often yellow with a few dark spots ( Fig. 10F ). Maxillary palpi cream-coloured, apical segment black apically. Head lateral side yellow, with a wide black band posterior to eye ( Fig. 12F ). Pronotum covered with white setae; dorsal disk dark red brown to black ( Fig. 11F ); lateral lobes of pronotum dorsal third black, vivid yellow ventrally (cream-coloured in preserved specimens) ( Fig. 12F ). Legs dark orange brown. FIIIs brown, 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, transparent basally; longer than FWs, forming a dark brown tail exceeding FWs for more than twice the length of pronotum. Tergites red brown, with a pale band laterally; sternites pale, with a red brown median area. Cerci red brown, lighter dorsally.

Male

FOREWINGS. FW colouration ( Fig. 13E ): dorsal field cells mostly transparent, veins mostly dark brown to black. Basal FW area black (instead of cream-coloured or brown as in N. vittatus ), with a relatively small cream-coloured patch basally. R black (instead of cream-coloured as in N. vittatus ), Sc projections cream-coloured basally, black apically. Apex of dorsal field somewhat truncated. Lateral field dark. FW venation ( Fig. 13E ): 1A curved, slightly angulated. Harp slightly longer than wide, with 3 oblique veins, basal one bifurcated distally, distal one bifurcated basally. c1 long and wide, c2 diamond-shaped; mirror (d1) longer than wide, not rounded, generally separated into two parts by a distinct transverse vein, the posterior part nearly rectangular, shorter than anterior part. Cell d2 narrower than d1. Anal-most and middle chords narrow and nearly parallel; basal-most chord extending from basal half of middle chord. Apical field short and rounded, with 3 wide cell alignments posterior to mirror and a narrow apical alignment; veins cream-coloured. Lateral field with 5 projections of Sc. Epiproct black. Subgenital plate pale cream-coloured with median area and anterior and posterior margins black.

GENITALIA. Very similar to N. vittatus ( Fig. 16E ) but differ by pseudepiphallus faintly less robust and broad, anterior margin more strongly concave. Rami more strongly curved.

Female

FOREWINGS. FW colouration ( Fig. 14F ): cells black, veins generally cream-coloured. Without distinct longitudinal white stripe around region of CuA, M and R with distinct margin from black colouration on dorsal field; Sc projections cream-coloured basally, black apically. Lateral field dark. FW venation: 6–7 strong longitudinal veins on dorsal field; lateral field with 6 projections of Sc.

GENITALIA. Ovipositor about as long as FIII. Copulatory papilla conical, small and stout; apex folded ventrally, short, pointed; dorsal face with a sclerotized area; ventro-anterior end forming a truncated lobule ( Fig. 17J–K ).

Juvenile ( Fig. 24 A–C )

Differing from that of N. vittatus by darker colouration, usually grey, never as white as N. vittatus . Stripes on head similar to N. vittatus but sometimes very faint; pronotum dorsal disk homogeneously dark grey, without longitudinal stripes; sometimes with a large velvety black spot. Legs grey with dark stripes, but stripes not as distinct as in N. vittatus . Abdominal tergites dark grey, their posterior margins thinly lined with yellow, not forming a pattern of transverse stripes. In some individuals, first and second abdominal tergites with a particularly dark (black), velvety band.

Measurements

See Table 8 .

Ecology

This species can be found among shrubs and herbaceous plants (but not grasses) ( Fig. 27 ). The adults and nymphs from Singapore and Malaysia (previously reported as N. vittatus ) have been reported to feed on the flower parts of a varieties of native (e.g., Dillenia suffruticosa (Griff.) Martelli ) and exotic weedy (e.g., Lantana camara L.) plant species ( Tan et al. 2017). They also feed on the foliages of understory plants such as Piper sarmentosum Roxb. ( Fig. 27 ). In Singapore, N. malaya sp. nov. can be found abundantly in the understory where Piper sarmentosum dominates.

Distribution

Malay Peninsula ( Singapore and Peninsular Malaysia [northern limit at the state of Penang]) ( Fig. 5 ).

Type locality

Singapore.

Calling song ( Singapore, n = 4 males, 20 echemes; MNHN-SO-2851–2854) ( Fig. 25A–C )

At 27.3°C, the calling song consists of bouts of highly variable length, ranging from a few seconds to several minutes. Each bout is composed of short echemes made of triplets of syllables; echeme duration is 32.3 ± 2.8 ms (29.0–37.56 ms); echeme period is 64.5± 4.3 ms (59.5–75.5 ms); downtime between consecutive echemes is 32.1 ±4.4 ms (24.3–41.3 ms). Durations for the first to third syllables of the triplet are as follows: 6.1 ±1.5 ms (3.7–8.0 ms), 6.7 ± 2.7 ms (2.5–10.6 ms) and 5.8± 2.9 ms (3.0–11.2 ms). Call spectrum consists of a series of harmonics with three distinct peaks in the first three harmonics: first harmonic (fundamental frequency) at 7.35± 1.28 kHz (6.201–9.39 kHz), which is also the dominant frequency; second harmonic at 13.33± 1.01 kHz (12.23–15.07 kHz); third harmonic at 18.88 ±0.77 kHz (17.23–20.07 kHz).

Remarks

We split the species previously recognised as N. vittatus into two distinct species after integrating evidence from general and nymphal colouration, female genitalia morphology, acoustics, DNA (12S sequence) and distribution. In terms of adult morphology, the new species is very similar to N. vittatus in habitus, other than being darker in colouration. The male genitalia are not always distinguishable between these species; but the sclerotization of the female genitalia can be differentiated. However, the colour patterns in the nymphs are drastically different. Likewise, a multivariate analysis of the call properties also showed distinct calling songs. The 12S gene tree also demonstrates that the two species form distinct clades. Lastly, the new species occurs in Malay Peninsula whereas N. vittatus occurs in Borneo and Sumatra. Integrating these pieces of evidence indicates that N. vittatus and N. malaya sp. nov. are indeed separate species having probably diverged recently, despite the similarities in adult habitus and male genitalia.

Figures

Fig. 4. Drawing of male genitalia ofN. malaya Robillard &Tan sp. nov. (with labels of parts).Abbreviations: see Material and methods; names of structures follow Desutter (1987), modified after Desutter-Grandcolas (2003) and Robillard & Desutter-Grandcolas (2004a). Terminology used by Gorochov (2015) in square brackets. Scale bar = 1 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. 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. 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.

Tables

Table 8. Measurements of Nisitrus malaya Robillard & Tan sp. nov. (in mm, means in brackets).

	PronL	PronW	FWL	FWW	HWT	FIIIL	FIIIW	TIIIL
Male holotype	1.8	2.7	10.2	3.4	6.6	13.6	2.5	12.9
Males (n = 5)	1.7–2.1 (1.9)	2.7–3.4 (3)	7.9–9.9 (8.9)	3.1–3.7 (3.4)	3.6–6.3 (5.2)	11.3–13.3 (12.6)	1.9–2.7 (2.4)	9.7–13.1 (11.8)
Females (n = 5)	1.7–2.2 (2.0)	2.5–3.1 (2.8)	9.2–10.6 (9.9)	3.2–3.4 (3.3)	5.3–6.8 (6.2)	12.6–14.2 (13.3)	2.2–3.0 (2.6)	11.3–13.6 (12.6)