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Published: 2021-07-26

Bioacoustics and systematics of Mecopoda (and related forms) from South East Asia and adjacent areas (Orthoptera, Tettigonioidea, Mecopodinae) including some chromosome data

Grillenstieg 18, 39120 Magdeburg, Germany
Department of Electronic Engineering, University of York, York, YO10 5DD, United Kingdom & Department of Life Sciences, Natural History Museum, London, SW7 5BD, United Kingdom.
Zoological Research Museum Alexander Koenig, Adenauerallee 160, D-53113, Bonn, Germany.
Lomonosov Moscow State University; Leninskie Gory, 1, building 12, Moscow, 119234, Russia.
Key Laboratory of the Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing, China.
Zoological Research Museum Alexander Koenig, Adenauerallee 160, D-53113, Bonn, Germany.
Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland.
Neotype Gryllus javanicus new species song stridulation Orthoptera


Bush-crickets (or katydids) of the genus Mecopoda are relatively large insects well-known for their sounds for centuries. Bioacoustic studies in India and China revealed a surprisingly large diversity of sound patterns. We extend these studies into the tropics of South East Asia using integrative taxonomy, combining song analysis, morphology of sound producing organs and male genitalia as well as chromosomes, to get a better understanding of the phylogeny and evolution of this widespread group. Besides the closely related genus Eumecopoda, the genus Mecopoda contains some isolated species and a large group of species which we assign to the Mecopoda elongata group. Some species of this group have broad tegmina and stridulatory files with different tooth spacing patterns and produce continuous, often relatively complicated, trill-like songs. The species of another subgroup with narrower wings have all similar files. Their songs consist of echemes (groups of syllables) which differ in syllable number and syllable repetition rate and also in echeme repetition rate. Our results show that South East Asia harbours a large and certainly not yet fully explored number of Mecopoda species which are most easily and clearly identified by song. Based on the data, five new forms are described: Mecopoda mahindai Heller sp. nov., Mecopoda paucidens Ingrisch, Su & Heller sp. nov., Mecopoda sismondoi Heller sp. nov., Mecopoda niponensis vietnamica Heller & Korsunovskaya subsp. nov., Eumecopoda cyrtoscelis zhantievi Heller subsp. nov. In addition, some taxonomic changes are proposed: Eumecopoda Hebard, 1922 stat. rev., Paramecopoda Gorochov, 2020, syn. nov. of Eumecopoda Hebard, 1922, Mecopoda javana (Johansson, 1763) stat. nov. (neotype selected) with M. javana minahasa Gorochov, 2020 stat. nov., M. javana darevskyi Gorochov, 2020 stat. nov., M. javana buru Gorochov, 2020 stat. nov., Mecopoda macassariensis (Haan, 1843) stat. rev., Mecopoda ampla malayensis Gorochov, 2020 syn. nov., Mecopada ampla javaensis Gorochov, 2020 syn. nov., Mecopoda fallax aequatorialis Gorochov, 2020 syn. nov., the last three are all synonyms of Mecopoda himalaya Liu, 2020, Mecopoda yunnana Liu 2020, stat. nov.


  1. Annandale, N. (1900) Observations on the habits and natural surroundings of insects made during the “Skeat Expedition” to the Malay Peninsula, 1899–1900. Proceedings of the Zoological Society, London, 4, 837–869.

  2. Asana, J.J., Makino, S. & Niyama, H. (1938) A chromosomal survey of some Indian insects. 1. Morphology of the chromosomes in eight species of the Locustidae. Journal of the Faculty of Sciences, Hokkaido University, 6, 221–234.

  3. Aswathanarayana, N.V. & Aswath, S.K. (1994) Karyotypes of two Indian grasshoppers of Mecopodinae (Orthoptera-Tettigoniidae). Cytologia, 59, 285–287.

  4. Ballentes, M.G., Mohagan, A.B., Gapud, V.P., Espallardo, M.C.P., Myrna, O. & Zarcilla, M.O. (2006) Arthropod Faunal Diversity and Relevant Interrelationships of Critical Resources in Mt. Malindang, Misamis Occidental. Southeast Asian Regional Center for Graduate Study and Research in Agriculture (SEARCA), Bangkok, 166 pp.

  5. Beier, M. (1966) Pars 9. Tettigoniidae: subfam, Meconematinae, Mecopodinae, Phyllophorinae. In: Beier, M. (Ed.), Orthopterum catalogus. W. Junk, s’Gravenhage, pp. 249–342.

  6. Bennet-Clark, H.C. (1989) Songs and the physics of sound production. In: Huber, F., Moore, T.E. & Loher, W. (Eds.), Cricket behavior and neurobiology. Cornell University Press, Ithaca & London, pp. 227–261.

  7. Bolívar, I. (1898) Ortópteros nuevos de Borneo y de Nueva Guinea. Actas de la Sociedad Española de Historia Natural. Vol. 2. Serie Real sociedad, Madrid, 27, 137–142.

  8. Brullé, A. (1835) Cinquième Ordre. Orthoptères. In: Orthoptères et Hemiptères. In: Audouin, J.V. & Brullé, A., Histoire naturelle des Insectes, 9 [1], (5), pp. i–iv + 1–225, [225–416 in 1836]

  9. Burmeister, H. (1838) Handbuch der Entomologie, Zweiter Band. Besondere Entomologie. Zweite Abtheilung Kaukerfe, Gymnognatha. Erste Hälfte; vulgo Orthoptera. T.C.F. Enslin, Berlin, pp. 457–756.

  10. Bugrov, A.G., Warchałowska-Śliwa, E., Ito, G., Tchernykh, A. & Maryati, M. (2004) Karyotype and C-banding patterns of the katydid Mecopoda elongata (L.) (Orthoptera, Tettigoniidae, Mecopodinae) from Amami Is. (Japan) and Borneo (Malaysia). Caryologia, 57, 25–29.

  11. Caudell, A.N. (1916) Orthoptera, Fam. Locustidae, Subfam. Mecopodinae. In: Wytsman, P. (Ed.), Genera Insectorum. Vol. 171. V. Verteneuil & L. Desmet, Bruxelles, pp. 1–32.

  12. Chamorro-Rengifo, J., Braun, H. & Lopes-Andrade, C. (2014) The secret stridulatory file under the right tegmen in katydids (Orthoptera, Ensifera, Tettigonioidea). Zootaxa, 3821 (5), 590–596.

  13. Cigliano, M.M., Braun, H., Eades, D.C. & Otte, D. (2021) Orthoptera Species File. Version 5.0/5.0. 17 February 2021. Available from: (accessed 12 December 2021)

  14. Dutta, R., Balakrishnan, R. & Tregenza, T. (2018) Divergence in potential contact pheromones and genital morphology among sympatric song types of the bush cricket Mecopoda elongata. Frontiers in Ecology and Evolution, 6, 158.

  15. Erregger, B., Kovac, H., Stabentheiner, A., Hartbauer, M., Römer, H. & Schmidt, A.K.D. (2017) Cranking up the heat: relationships between energetically costly song features and the increase in thorax temperature in male crickets and katydids. Journal of Experimental Biology, 220 (14), 2635e2644.

  16. Fritze, A. (1900) Orthoptères de l’ Archipel Malais. (Voyage de MM. M. Bedot et C. Pictet dans l’ Archipel Malais). Revue Suisse de Zoologie, 7 (1899), 335–340.

  17. Griffini, A. (1908) Sopra alcuni Stenopelmatidi e sopra alcune Mecopodidi Malesi ed Austro-Malesi. Atti della Societa Italiana di Scienze Naturali, 46, 271–288.

  18. Gorochov, A.V. (2020) A review of the genus Mecopoda (Orthoptera: Tettigoniidae: Mecopodinae) from Indo-Malayan and Papuan Regions. Proceedings of the Zoological Institute RAS, 324 (4), 564–586.

  19. Haan, W. de (1843) Bijdragen tot de kennis der Orthoptera. In: Temminck, C.J. (Ed.), Verhandelingen over de Natuurlijke Geschiedenis der Nederlansche Overzeesche Bezittingen, de Leden der Natuurkundige Commissie in Indië en andere Schrijvers, Leiden, 19/20, pp. 165–228.

  20. Hareyama, S. (1932) On the chromosomes of some insects belonging to Locustidae. Zoological Magazine, Japan, 44, 83–84.

  21. Hartbauer, M., Kratzer, S. & Römer, H. (2006) Chirp rate is independent of male condition in a synchronising bushcricket. Journal of Insect Physiology, 52, 221–230.

  22. Hartbauer, M, Siegert, M.E., Fertschai, I. & Römer, H. (2012) Acoustic signal perception in a noisy habitat: lessons from synchronising insects. Journal of Comparative Physiology, A, 198, 397–409.

  23. Hebard, M. (1922) Studies in Malayan, Melanesian and Australian Tettigoniidae (Orthoptera). Proceedings of the Academy of Natural Sciences, Philadelphia, 74, 121–299.

  24. Helfert, B. & Sänger, K. (2007) Variability of diagnostic features in Eumecopoda cyrtoscelis cyrtoscelis (Karsch, 1888) from the Raja Ampat Islands (Indonesia) (Tettigoniidae, Mecopodinae). Zeitschrift der Arbeitsgemeinschaft Oesterreichischer Entomologen, 59 (3–4), 71–83.

  25. Heller, K.-G. (1988) Bioakustik der europäischen Laubheuschrecken. Josef Margraf, Weikersheim, 358 pp.

  26. Helversen, O. von & Elsner, N. (1977) The stridulatory movements of acridid grasshoppers recorded with an opto-electronic device. Journal of Comparative Physiology, 122, 53–64.

  27. Hemp, C., Heller, K.-G., Warchałowska-Śliwa, E. & Hemp, A. (2015) A new species of Philoscirtus (Orthoptera: Phaneropteridae: Mecopodinae) from the West Usambara Mountains of Tanzania and its conservation status. Zootaxa, 3905 (2), 273–282.

  28. Hewitt, G.M. (1979) Grasshoppers and crickets. Animal Cytogenetics, 3. Insecta I. Orthoptera. Borntraeger, Berlin and Stuttgart, 170 pp.

  29. Ichikawa, A., Ito, F., Kanô, Y., Kawai, M., Tominaga, O. & Murai, T. (Eds.) (2006) Orthoptera of the Japanese Archipelago in color. s.n., Sapporo, 688 pp. + 2 CD.

  30. ICZN (1999) International Code of Zoological Nomenclature. 4th Edition. International Trust for Zoological Nomenclature, London, 306 pp.

  31. Johansson, B. [not Johannson, B.] (1763) Centurio insectorum rariorum. In: Linné, C. von. (Ed.), Amoenitates Academicae seu dissertationes variae Physicae, Medicae, Botanicae anthehac seorsim editae. Vol 6. 2nd Edition. 1789. s.n., Holmiae [Stockholm], pp. 384–415.

  32. Karny, H.H. (1920) Zur Deutung der de Haan’schen Laubheuschrecken. Zoologische Mededeelingen, 5, 139–210.

  33. Karny, H.H. (1921) Katydids (Tettigonioidea) of the Philipine Islands, collected by C.F. Baker. Philippine Journal of Science, 18, 607–617.

  34. Karny, H.H. (1924) Beiträge zur malayischen Orthopterenfauna. VIII. Die Mecopodinen des Buitenzorger Museums. Treubia, 5, 137–160.

  35. Karsch, F. (1888) Zwei neue Mecopoda-Arten (Orthoptera). Entomologische Nachrichten, Berlin, 14, 145–148.

  36. Kevan, D.C.M. (1978) The Land of the Locusts: Being Some Further Verses on Grigs and Cicadas Before 450 A.D. Lyman Entomological Museum and Research Laboratory. Memoir No. 6. Lyman Entomological Museum and Research Laboratory, Quebec, 530 pp.

  37. Kim, T.W. (2009) Taxonomy and acoustic signals of Korean Tettigonioidea (Orthoptera : Ensifera) II. National Institut of Biological Reserves, Incheon. [+CD]

  38. Kirby, W.F. (1891) Notes on the Orthopterous family Mecopodidae. Transactions of the Royal Entomological Society of London, 1891, 405–412.

  39. Korsunovskaya, O.S. (2008) Acoustic signals in katydids (Orthoptera, Tettigoniidae). Communication I. Zoologicheskii Zhurnal, 87, 1453–1471. [in Russian; English version: Entomological Review, 88, 1032–1050]

  40. Kostarakos, K. & Römer, H. (2015) Neural mechanisms for acoustic signal detection under strong masking in an insect. Journal of Neuroscience, 35, 10562–10571.

  41. Krobath, I. (2013) Complex signalling, song interaction, and mate choice in a trilling species of the Mecopoda complex (Orthoptera: Tettigoniidae). Master thesis, University of Graz, 59 pp.

  42. Krobath, I., Römer, H. & Hartbauer, M. (2017) Plasticity of signaling and mate choice in a trilling species of the Mecopoda complex (Orthoptera: Tettigoniidae). Behavioral Ecology and Sociobiology, 71, 164.

  43. Levan, A., Fredga, K. & Sanberg, A. (1964) Nomenclature for centromeric position on chromosomes. Hereditas, 52, 201–220.

  44. Linnaeus, C. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima, reformata. Laurentii Salvii., Holmiae [Stockholm], 824 pp.

  45. Liu, C.-X., Heller, K.-G., Wang, X.-S., Yang, Z., Wu, C., Liu, F. & Zhang, T. (2020) Taxonomy of a katydid genus Mecopoda Serville (Orthoptera: Tettigoniidae, Mecopodinae) from East Asia. Zootaxa, 4758 (2), 296–310.

  46. Liu, Y.-F., Shen, C-Z., Gong, P., Zhang, L. & He, Z-Q. (2019) Three new species of genus Mecopoda Serville, 1831 from China (Orthoptera: Tettigoniidae: Mecopodinae). Zootaxa, 4585 (3), 561–572.

  47. Luanpirom, A., Bunnag, S., Kritpetcharat, O. & Kritpetcharat, C. (1999) Cytogenetic research of some animals in Phu-Phan National Park. In: Research reports on Biodiversity in Thailand. Biodiversity Research and Training Program (BRT), Bangkok, pp. 563–570. [in Thai]

  48. Machácková, L. & Fikácek, M. (2014) Catalogue of the type specimens deposited in the Department of Entomology, National Museum, Prague, Czech Republic. Acta Entomologica Musei Nationalis Pragae, 54, 399–450.

  49. Marshall, J.A. (1983) The orthopteroid insects described by Linnaeus, with notes on the Linnean collection. Zoological Journal of the Linnean Society, 78, 375–396.

  50. Nityananda, V. & Balakrishnan, R. (2006) A diversity of songs among morphologically indistinguishable katydids of the genus Mecopoda (Orthoptera: Tettigoniidae) from southern India. Bioacoustics, 15, 223–250.

  51. Nityananda, V. & Balakrishnan, R. (2021) Synchrony of complex signals in an acoustically communicating katydid. Journal of Experimental Biology, 224 (9), jeb241877.

  52. OSF (Orthoptera Species File) (2021) Cigliano et al. 2021.

  53. Panhwar, W.A., Sultana, R., Wagan, M.S. & Khatri, I. (2016) New records of Mecopodinae (Orthoptera: Tettigonioidea: Tettigoniidae) from Pakistan. Arquivos Entomolóxicos, 15, 269–274.

  54. Redtenbacher, J. (1892) Monographische Uebersicht der Mecopodiden. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien, 42, 183–224.

  55. Rentz, D.C.F., Su, Y.N. & Ueshima, N. (2006) Studies in Australian Tettigoniidae: the mecopodine katydids (Orthoptera: Tettigoniidae; Mecopodinae; Mecopodini). Transactions of the American Entomological Society, 132 (1+2), 1–24.[1:SIATTM]2.0.CO;2

  56. Serville, J.G.A. (1831) Revue méthodique des insectes de l‘ordre des Orthoptères. Annales des Sciences Naturelles Zoologie et Biologie Animale, 22, 28–65 + 134–167 + 262–292.

  57. Shorthouse, D.P. (2010) SimpleMappr, an online tool to produce publication-quality point maps. Available from: http:// (accessed 05 October 2020)

  58. Siegert, M.E., Römer, H. & Hartbauer, M. (2013) Maintaining acoustic communication at a cocktail party: heterospecific masking noise improves signal detection through frequency separation. Journal of Experimental Biology, 216, 4655–4665.

  59. Sismondo, E (1990) Synchronous, alternating, and phase-locked stridulation by a tropical katydid. Science, 249, 55–58.

  60. Stoll, C. (1813) Natuurlijke en naar het leven nauwkeurig gekleurde afbeeldingen en beschrijvingen der spoken, wandelende bladen, zabel-springhanen, krekels, trek-springhanen en kakkerlakken. J.C.Sepp & sohn, Amsterdam. [unpaginated, in Dutch and French]

  61. Storozhenko, S.Y. & Paik, J.C. (2007) Orthoptera of Korea. Dalnauka, Vladivostok, 231 pp.

  62. Storozhenko, S.Y., Kim, T.W. & Jeon, M.J. (2015) Monograph of Korean Orthoptera. National Institute of Biological Resources, Incheon, 377 pp.

  63. Sumner, S.G. (1972) A simple technique for demonstrating centromere heterochromatin. Experimental Cell Research, 75, 304–306.

  64. Tan, M.K., Baroga-Barbecho, J.B. & Yap, S.A. (2019) An account on the Orthoptera from Siargao Island (Southeast Asia: Philippines: Mindanao). Zootaxa, 4609 (1), 1–30.

  65. Tan, M.K. & Kamaruddin, K. M(2016) A contribution to the knowledge of Orthoptera diversity from Peninsular Malaysia: Bukit Larut, Perak. Zootaxa, 4111 (1), 21–40.

  66. 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.

  67. Thunberg, C.P. (1815) Hemipterorum maxillosorum genera illustrata plurimisque novis speciebus ditata ac descripta. Mémoires de l’ Académie Impériale des Sciences de St. Pétersbourg, 5, 211–301.

  68. Uvarov, B.P. (1927) Some Orthoptera of the families Mantidae, Tettigoniidae and Acrididae from Ceylon. Spolia Zeylanica, 14, 85–114.

  69. Walker, F. (1869) Catalogue of the specimens of Dermaptera Saltatoria in the collection of the British Museum. Part II. Catalogue of Locustidae. British Museum of Natural History, London, pp. 225–423.

  70. Walker, F. (1870) Catalogue of the Specimens of Dermaptera Saltatoria in the Collection of the British Museum. Part III. British Museum of Natural History, London, pp. 425–604.

  71. Warchałowska-Śliwa, E. (1998) Karyotype characteristics of katydid orthopterans (Ensifera, Tettigoniidae) and remarks on their evolution at different taxonomic levels. Folia Biologica, Kraków, 46, 143–176.

  72. Willemse, C. (1933) Résultats scientifiques du voyage aux Indes-Orientales Néerlandaises de LL.AA.RR. le Prince et la Princesse Léopold de Belgique. Orthoptera II: fam. Tettigoniidae and Gryllacridae. Mémoires du Musée royal d’histoire naturelle de Belgique, Hors Série 4 (8), 1–15.

  73. Willemse, F. (1977) Classification and distribution of the Sexavae of the Melanesian subregion (Orthoptera, Tettigonioidea, Mecopodinae). Tijdschrift voor Entomologie, 120, 213–277.

  74. Zhantiev, R.D. (1981) Bioacoustics of Insects. Moscow State University Press, Moscow, 256 pp. [in Russian]