Revision of the cricket genus Nisitrus Saussure (Orthoptera: Gryllidae: Eneopterinae) and descriptions of five new species

  • Ming Kai Tan 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
  • Rodzay bin Haji Abdul Wahab Institute for Biodiversity and Environmental Research, Universiti Brunei Darussalam, Jalan Universiti, BE1410, Brunei Darussalam
  • Razy Japir Forest Research Centre (Sepilok), Sabah Forestry Department, P.O. Box 1407, 90715 Sandakan, Sabah, Malaysia
  • Authur Y.C. Chung Forest Research Centre (Sepilok), Sabah Forestry Department, P.O. Box 1407, 90715 Sandakan, Sabah, Malaysia
  • Tony Robillard 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
Keywords: Nisitrini, key, lectotype, redescription, Southeast Asia


Nisitrus Saussure, 1878 is a diurnal and monophyletic group of eneopterine crickets found in parts of Southeast Asia. Species often have vivid colourations which make them charismatic photograph subjects among macro photographers. However, their taxonomy has not been revised since their original descriptions, whereas there are also species awaiting to be discovered and described. A synoptic revision is therefore needed prior to studying the phylogenetic relationships of the genus, and before undertaking more precise studies on biogeography and evolution of traits. By holistically incorporating information about general morphology, male and female genitalia, and bioacoustics, we revise the taxonomy of these crickets and recognise 10 valid species of Nisitrus. Five new species are described: N. crucius Robillard & Tan sp. nov. from Borneo; N. danum Robillard & Tan sp. nov. from Borneo; N. hughtani Robillard & Tan sp. nov. from Sumatra; N. malaya Robillard & Tan sp. nov. from Malay Peninsula; and N. rindu Robillard & Tan sp. nov. from Borneo. We redescribe N. brunnerianus and N. insignis; describe the calls of N. malaya sp. nov., N. musicus and N. vittatus. We consider N. sumatrensis as a junior synonym of N. insignis; and N. marginata as a junior synonym of N. vittatus. Nisitrus hyalinus requires further revision. We also consider N. maculosus as a dubious name.


Baroga-Barbecho J.B., Yap S.A., Tan M.K. & Robillard T. 2019. Taxonomic review of the genus Paranisitra Chopard (Orthoptera: Gryllidae: Eneopterinae: Nisitrini) with description of a new species from Mindanao. Zootaxa 4568 (1): 81–96.

Buys D.W., Witkamp H., Endert F.H., Siebers H.C. & Bosch D.F.K. (1927). Midden-Oost-Borneo Expeditie 1925. Kolff & Co., Weltevreden.

Cigliano M.M., Braun H., Eades D.C. & Otte D. 2020. Orthoptera Species File Online. Version 5 (5.0). Available from [accessed 8 Mar. 2020].

Chopard L. 1929. Spolia Mentawiensia: Gryllidae. Bulletin of the Raffles Museum 2: 98–118.

Chopard L. 1940. Results of the Oxford University expedition to Sarawak (Borneo), 1932. Gryllacridae and Gryllidae (Orthoptera). The Entomologist’s Monthly Magazine 4 76 (8–9): 189–192; 193–204.

Chopard L. 1968. Fam. Gryllidae: Subfam. Mogoplistinae, Myrmecophilinae, Scleropterinae, Cachoplistinae, Pteroplistinae, Pentacentrinae, Phalangopsinae, Trigonidiinae, Eneopterinae; Fam. Oecanthidae, Gryllotalpidae. In: Beier M. (ed.) Orthopterorum Catalogus 12: 213–500. Junk, 's-Gravenhage [The Hague].

Chopard L. 1969. Grylloidea. In: Sewell R.B.S. (ed.) The fauna of India and the adjacent countries. Orthoptera. Vol. 2.: 1–421. Zoological Survey of India and Baptist Mission Press, Calcutta.

Desutter L. 1987. Structure et évolution du complexe phallique des Gryllidea (Orthoptera) et classification des genres néotropicaux de Grylloidea.1ère partie. Annales de la Société entomologique de France (N.S.) 23: 213–239.

Desutter-Grandcolas L. 1990. Étude phylogénétique, biogéographique et écologique des Grylloidea néotropicaux (Insectes, Orthoptères). PhD thesis, Universite de Paris-Sud, Centre d´Orsay, France.

Desutter-Grandcolas L. 2003. Phylogeny and the evolution of acoustic communication in extant Ensifera (Insecta, Orthoptera). Zoologica Scripta 32: 525–561.

Desutter-Grandcolas L., Blanchet E., Robillard T., Magal C., Vannier F. & Dangles O. 2010. Evolution of the cercal sensory system in a tropical cricket clade (Orthoptera: Grylloidea: Eneopterinae): A phylogenetic approach. Biological Journal of the Linnean Society 99: 614–631.

Dong J., Kergoat G.J., Vicente N., Rahmadi C., Xu S. & Robillard T. 2018. Biogeographic patterns and diversification dynamics of the genus Cardiodactylus Saussure (Orthoptera, Grylloidea, Eneopterinae) in Southeast Asia. Molecular Phylogenetics and Evolution 129: 1–14.

Gorochov A.V. 2009. A review of the genus Paranisitra Chopard (Orthoptera: Gryllidae: Eneopterinae). Proceedings of the Zoological Institute of the Russian Academy of Sciences 313 (4): 435–439.

Gorochov A.V. 2015. Evolution and taxonomic significance of the copulatory apparatus in Ensifera (Orthoptera). Part 2: Male genitalia in Grylloidea (Orthoptera: Ensifera). Zoosystematica Rossica 24 (1): 31–41.

Haan W. 1844. Bijdragen tot de kennis der Orthoptera. In: Temminck C.J. (ed.) Verhandelingen over de Natuurlijke Geschiedenis der Nederlandsche Overzeesche Bezittingen 24: 229–248. In commissie bij S. en J. Luchtmans, en C.C. van der Hoek, Leiden.

Hall T.A. 1999. BioEdit: a user‐friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95–98.

Hoang D.T., Chernomor O., von Haeseler A., Minh B.Q. & Vinh L.S. 2017. UFBoot2: improving the ultrafast bootstrap approximation. Molecular Biology and Evolution 35 (2): 518–522.

Hollier J., Bruckner H. & Heads S.W. 2013. An annotated list of the Orthoptera (Insecta) species described by Henri de Saussure, with an account of the primary type material housed in the Muséum d’histoire naturelle de Genève, Part 5: Grylloidea. Revue suisse de Zoologie 120 (3): 445–535.

iNaturalist 2020. iNaturalist. Available from [accessed 17 Jun. 2021].

Ingrisch S. 1987. Neue Grillen von Borneo und aus Thailand (Insecta: Saltatoria: Grylloidea). Senckenbergiana biologica 68 (1–3): 163–185.

Jaiswara R., Dong J. & Robillard T. 2018. Revision of the genus Pseudolebinthus (Orthoptera, Gryllidae, Eneopterinae) with the description of a new species from Southeast Africa. Zootaxa 4521 (2): 265–274.

Jaiswara R., Dong J., Ma L., Yin H. & Robillard T. 2019. Taxonomic revision of the genus Xenogryllus (Orthoptera, Gryllidae, Eneopterinae). Zootaxa 4545 (3): 301–338.

Kalyaanamoorthy S., Minh B.Q., Wong T.F.K., von Haeseler A. & Jermiin L.S. 2017. ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods 14: 587–589.

Katoh K., Rozewicki J. & Yamada K.D. 2017. MAFFT online service: multiple sequence alignment, interactive sequence choice and visualization. Briefings in Bioinformatics 20 (4): 1160–1166.

Kearse M., Moir R., Wilson A., Stones‐Havas S., Cheung M., Sturrock S., Buxton S., Cooper A., Markowitz S., Duran C., Thierer T., Ashton B., Meintjes P. & Drummond A. 2012. Geneious basic: An integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28 (12): 1647–1649.

Kirby W.F. 1906. A Synonymic Catalogue of Orthoptera. Vol. 2. Orthoptera Saltatoria. Part I. (Achetidae et Phasgonuridae). Trustees of the british Museum, London.

Kuraku S., Zmasek C.M., Nishimura O. & Katoh K. 2013. aLeaves facilitates on‐demand exploration of metazoan gene family trees on MAFFT sequence alignment server with enhanced interactivity. Nucleic Acids Research 41: 22–28.

Nattier R., Robillard T., Desutter-Grandcolas L., Couloux A. & Grandcolas P. 2011. Older than New Caledonia emergence? A molecular phylogenetic study of the eneopterine crickets (Orthoptera: Grylloidea). Journal of Biogeography 38 (11): 2195–2209.

Nguyen L.T., Schmidt H.A., von Haeseler A. & Minh B.Q. 2015. IQ‐TREE: a fast and effective stochastic algorithm for estimating maximum‐likelihood phylogenies. Molecular Biology and Evolution 32 (1): 268–274.

Otte D. 1994. Orthoptera Species File. Vol. 1. Crickets (Grylloidea). Orthopterists' Society and the ANSP, Philadelphia.

Preston-Mafham K. 2000. Diurnal mating behaviour of a Nisitrus sp. cricket (Orthoptera: Gryllidae) from Sumatra. Journal of Natural History 34 (12): 2241–2250.

R Core Team. 2018. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available from [accessed 17 Jun. 2021].

Ragge D.R. & Reynolds W.J. 1998. The Songs of the Grasshoppers and Crickets of Western Europe. Harley Books, Colchester, England.

Rehn J.A.G. 1909. A contribution to the knowledge of the Orthoptera of Sumatra. Bulletin of the American Museum of Natural History 26: 177–211.

Robillard R. & Desutter-Grandcolas L. 2004a. Evolution of acoustic communication in crickets: phylogeny of Eneopterinae reveals an adaptive radiation involving high-frequency calling (Orthoptera, Grylloidea, Eneopteridae). Anais da Academia Brasileira de Ciências 76 (2): 297–300.

Robillard R. & Desutter-Grandcolas L. 2004b. High-frequency calling in Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae): adaptive radiation revealed by phylogenetic analysis. Biological Journal of the Linnean Society 83 (4): 577–584.

Robillard R. & Desutter-Grandcolas L. 2004c. Phylogeny and the modalities of acoustic diversification in extant Eneopterinae (Insecta, Orthoptera, Grylloidea, Eneopteridae). Cladistics 20 (3): 271–293.

Robillard T. & Desutter-Grandcolas L. 2006. Phylogeny of the cricket subfamily Eneopterinae (Orthoptera, Grylloidea, Eneopteridae) based on four molecular loci and morphology. Molecular Phylogenetics and Evolution 40: 643–661.

Robillard R. & Desutter-Grandcolas L. 2008. Clarification of the taxonomy of extant crickets of the subfamily Eneopterinae (Orthoptera: Grylloidea; Gryllidae). Zootaxa 1789: 66–68.

Robillard R. & Desutter-Grandcolas L. 2011. The complex stridulatory behavior of the cricket Eneoptera guyanensis Chopard (Orthoptera: Grylloidea: Eneopterinae). Journal of Insect Physiology 57 (6): 694–703.

Robillard T. & Tan M.K. 2013. A taxonomic review of common but little known crickets from Singapore and the Philippines (Insecta: Orthoptera: Eneopterinae). Raffles Bulletin of Zoology 61 (2): 705–725.

Robillard R., Grandcolas P. & Desutter-Grandcolas L. 2007. A shift toward harmonics for high-frequency calling shown with phylogenetic study of frequency spectra in Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae). Canadian Journal of Zoology 85 (12): 1264–1275.

Robillard T., Montealegre-Z. F., Desutter-Grandcolas L., Grandcolas P. & Robert D. 2013. Mechanisms of high-frequency song generation in brachypterous crickets and the role of ghost frequencies. Journal of Experimental Biology 216 (11): 2001–2011.

Robillard T., Gorochov A.V., Poulain S. & Suhardjono Y.R. 2014. Revision of the cricket genus Cardiodactylus (Orthoptera, Eneopterinae, Lebinthini): the species from both sides of the Wallace line, with description of 25 new species. Zootaxa 3854 (1): 1–104.

Saussure H. 1878. Mélanges orthoptérologiques. VI. Fascicule Gryllides. Mémoires de la Société de Physique et d’Histoire naturelle de Genève 25 (2): 369–704 [505–834].

Schneider E.S., Römer H., Robillard T. & Schmidt A.K.D. 2017. Hearing with exceptionally thin tympana: Ear morphology and tympanal membrane vibrations in eneopterine crickets. Scientific Reports 7 (15266): 1–12.

Sueur J., Aubin T. & Simonis C. 2008. Seewave, a free modular tool for sound analysis and synthesis. Bioacoustics 18 (2): 213–226.

Tan M.K. 2012. Orthoptera in the Bukit Timah and Central Catchment Nature Reserves (Part 2): Suborder Ensifera. Raffles Museum of Biodiversity Research, National University Singapore, Singapore. Available from [accessed 16 Jun. 2021].

Tan M.K. 2017. Orthoptera in the Bukit Timah and Central Catchment Nature Reserves (Part 2): Suborder Ensifera. 2nd Edition. Lee Kong Chian Natural History Museum, National University of Singapore, Singapore. Available from [accessed 16 Jun. 2021].

Tan M.K. & Robillard T. 2014. A new species of Cardiodactylus (Orthoptera: Grylloidea: Eneopterinae) from Singapore. Zootaxa 3764 (3): 364–376.

Tan M.K. & Wahab R.A. 2018. Preliminary study on the diversity of Orthoptera from Kuala Belalong Field Studies Centre, Brunei Darussalam, Borneo. Journal of Orthoptera Research 27 (2): 119–142.

Tan M.K., Ngiam R.W.J. & Ismail M.R.B. 2012. A checklist of Orthoptera in Singapore parks. Nature in Singapore 5: 61–67.

Tan M.K. Ngiam R.W.J., Ismail M.R.B. & Ibrahim H. 2013a. Diversity of Orthoptera from Neo Tiew Lane Two, Singapore. Nature in Singapore 6: 211–222.

Tan M.K., Yeo H., Hasnan S., Woon S.Z.S. & Wu B. 2013b. A rapid comparison of the Orthoptera communities between Bukit Brown Cemetery and Lornie Trail of Central Catchment Nature Reserve, Singapore. Nature in Singapore 6: 97–103.

Tan M.K., Yeo H. & Lee J.X.Q. 2015. Diversity of entomofauna (Orthoptera, Reduviidae and Aculeata) in the Mandai-Lake Road area, Singapore. Nature in Singapore 8: 37–51.

Tan M.K., Artchawakom T., Wahab R.A., Lee C.-Y., Belabut D.M. & Tan H.T.W. 2017. Overlooked flower visiting Orthoptera in Southeast Asia. Journal of Orthoptera Research 26 (2): 143–153.

Tan M.K., Japir R. & Chung A.Y.C. 2019. Uncovering the Grylloidea and Tettigonioidea (Orthoptera: Ensifera) in the Forest Research Centre (Sepilok) Entomological Collection. Zootaxa 4701 (4): 301–349.

ter Hofstede H., Schöneich S., Robillard T. & Hedwig B. 2015. Evolution of a communication system by sensory exploitation of startle behavior. Current Biology 25: 3245–3252.

Trifinopoulos J., Nguyen L.‐T., von Haeseler A. & Minh B.Q. 2016. W‐IQ‐TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nuclei Acids Research 44: 232–235.

Vicente N., Kergoat G.J., Dong J., Yotoko K., Legendre F., Nattier R. & Robillard T. 2017. In and out of the Neotropics: historical biogeography of Eneopterinae crickets. Journal of Biogeography 44: 2199–2210.

Walker F. 1869. Catalogue of the Specimens of Dermaptera Saltatoria in the Collection of the British Museum Part 2. Trustees of the British Museum, London.

Walker F. 1871. Catalogue of the Specimens of Dermaptera Saltatoria in the Collection of the British Museum Part 5. Trustees of the British Museum, London.

Willemse L.P.M. 2001. Fauna Malesiana: Guide to the Pest Orthoptera of the Indo-Malayan Region. Backhuys Publishers, Leiden.

How to Cite
Tan, M. K., Wahab, R. bin H. A., Japir, R., Chung, A. Y., & Robillard, T. (2021). Revision of the cricket genus Nisitrus Saussure (Orthoptera: Gryllidae: Eneopterinae) and descriptions of five new species. European Journal of Taxonomy, 761(1), 1-75.
Research article