Tetrigidae

Tetrigidae
Tetrix species
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Orthoptera
Suborder: Caelifera
Superfamily: Tetrigoidea
Family: Tetrigidae
Rambur, 1838
Subfamilies

Tetrigidae[1] is an ancient family in the order Orthoptera,[2] which also includes similar families such as crickets, grasshoppers, and their allies. Species within the Tetrigidae are variously called groundhoppers,[3] pygmy grasshoppers,[4] pygmy devils[5] or (mostly historical) "grouse locusts".[6]

Diagnostic characteristics

Tetrigidae are typically less than 20 mm (0.79 in) in length and are recognizable by a long pronotum. This pronotum extends over the length of the abdomen, sometimes to the tip of the wings, and ends in a point.[4] In other Orthoptera, the pronotum is short and covers neither the abdomen nor the wings. Tetrigidae are generally cryptic in coloration.[7] Some species have enlarged pronota that mimic leaves, stones or twigs.[8][9]

Other characteristics pygmy grasshoppers exhibit in comparison to other Orthoptera families are the lack of an arolium between the claws, the first thoracic sternite being modified into collar-like structure called sternomentum, a tarsal formula of 2-2-3, scaly fore -wings, and developed hindwings.[10]

General biology

In temperate regions, Tetrigidae are generally found along streams and ponds, where they feed on algae and diatoms.[11] The North American species Paratettix aztecus and Paratettix mexicanus, for example, depend on aquatic primary production for between 80% and 100% of their diet.[12] Detritus, moss and fungal hyphae also dominates the diet in many species.[13] Riparian species are capable of swimming on the surface of the water, and readily leap into the water when alarmed[9] Some species in the tribe Scelimenini are fully aquatic and capable of swimming underwater.[10][11]

The highest biodiversity of Tetrigidae is found in tropical forests.[10] Some tropical species are arboreal and live among mosses and lichens in tree buttresses or in the canopy,[11] while others live on the forest floor.[7]

Like other Orthoptera, Tetrigidae have a hemimetabolous development, in which eggs hatch into nymphs. Unlike other temperate Orthoptera, however, temperate Tetrigidae generally overwinter as adults.[4]

Some subfamilies within the Tetrigidae, such as the Batrachideinae, are sometimes elevated to family rank besides the Tetrigidae.

Arulenus miae is a pygmy grasshopper species from the tropical mountainous rainforests of the Philippines. The species was firstly discovered in Facebook post.[5]

Etymology

Origin of the name of the family is not completely clear as there are different sources on its etymology. The name may be derived from Latin tetricus or taetricus, meaning harsh, sour, severe.[14] The name may also originate from the earlier name 'Tettigidae', based on Tettix (synonym of Tetrix), which was preoccupied by Tettigidae (synonym of Cicadidae).[15] Because of the preoccupation by the cicadas' family name, the second 't' in 'tt' was changed into 'r', resulting in the word Tetrigidae.

Subfamilies, tribes and selected genera

More than 320 genera in 10 subfamilies have been described; according to the Orthoptera Species File[16] the following are included:

Subfamily Batrachideinae[17]

Auth.: Bolívar, 1887; selected genera:

Subfamily Cladonotinae

Auth.: Bolívar, 1887; selected genera:

Tribe Cladonotini Bolívar, 1887[18]
  • Cladonotus Saussure, 1862
  • Deltonotus Hancock, 1904
  • Piezotettix Bolívar, 1887
Tribe Choriphyllini Cadena-Castañeda & Silva, 2019[19]
  1. Choriphyllum Serville, 1838
  2. Phyllotettix Hancock, 1902
Tribe Valalyllini Deranja, Kasalo, Adžić, Franjević & Skejo, 2022[20]
  1. Lepocranus Devriese, 1991
  2. Valalyllum Deranja, Kasalo, Adžić, Franjević & Skejo, 2022
Tribe Xerophyllini Günther, 1979

SE Asia - selected genera:

Tribe Unassigned

Subfamily Criotettiginae[21]

Auth. Kevan, 1966

Criotettigini

Auth. Kevan, 1966

  1. Acanthalobus Hancock, 1904
  2. Criotettix Bolívar, 1887
  3. Dasyleurotettix Rehn, 1904

Thoradontini

Auth. Kevan, 1966

  1. Aryalidonta Subedi & Kasalo, 2023[22]
  2. Eucriotettix Hebard, 1930
  3. Loxilobus Hancock, 1904
  4. Thoradonta Hancock, 1909
  5. Yunnantettix Zheng, 1995

Criotettiginae: tribe unassigned

  1. Afrocriotettix Günther, 1938
  2. Amphinotulus Günther, 1939
  3. Apterotettix Hancock, 1904
  4. Bolivaritettix Günther, 1939
  5. Bolotettix Hancock, 1907
  6. Cotysoides Zheng & Jiang, 2000
  7. Hyboella Hancock, 1915
  8. Indomiriatra Tinkham, 1939
  9. Miriatroides Zheng & Jiang, 2002
  10. Probolotettix Günther, 1939
  11. Rhopalina Tinkham, 1939
  12. Rostella Hancock, 1913
  13. Spadotettix Hancock, 1910
  14. Tettitelum Hancock, 1915

Subfamily Guntheritettiginae

Auth.: Cadena-Castañeda, 2025[23] - Madagascar & Indian Ocean islands:

tribe Guntheritettigini Cadena-Castañeda, 2025
  • Guntheritettix Cadena-Castañeda & Tavares, 2025: monotypic G. formidabilis (Günther, 1974)
  • Holocerus Bolívar, 1887: monotypic Holocerus lucifer (Serville, 1838)[24]
tribe Tumbrinckitettigini Cadena-Castañeda, 2025
  • Cryptotettix Hancock, 1900
  • Tumbrinckitettix Cadena-Castañeda & Tavares, 2025
Subfamily Hildegardiinae Cadena-Castañeda, 2025 - monogeneric
  1. Hildegardia Günther, 1974

Subfamily Lophotettiginae[25][26]

Auth.: Hancock, 1909; distribution S. America, all genera:

  1. Lophotettix Hancock, 1909
  2. Phelene Bolívar, 1906

Subfamily Metrodorinae [27]

Auth.: Bolívar, 1887; selected genera:

Tribe Amorphopini Günther, 1939
  • Amorphopus Serville, 1838
Tribe Cleostratini Hancock, 1907
Tribe Clinophaestini Storozhenko, 2013
  • Birmana Brunner von Wattenwyl, 1893
  • Clinophaestus Storozhenko, 2013
Tribe Miriatrini Cadena-Castañeda & Cardona, 2015 (monotypic)
  • Miriatra Bolívar, 1906
Tribe Metrodorini Bolívar, 1887[28]
Tribe Ophiotettigini Tumbrinck & Skejo, 2017
Tribe Unassigned

Subfamily Scelimeninae

Auth.: Hancock, 1907 Tribe Scelimenini Hancock, 1907; selected genera:

incertae sedis

Subfamily Tetriginae

Auth.: Serville, 1838; selected genera

Tribe Dinotettigini Günther, 1979
  • Dinotettix Bolívar, 1905
  • Ibeotettix Rehn, 1930
Tribe Tetrigini Serville, 1838[29]
Tribe unassigned

Subfamily Tripetalocerinae

Auth.: Bolívar, 1887

Tripetalocerinae was originally described by Bolívar in 1887[30] to gather all the Tetrigidae genera of the old world with widened antennae (e.g. Arulenus, Discotettix, Hirrius, Ophiotettix, Tripetalocera). This subfamily today includes only two species in two genera - Tripetalocera (with one species) from India and Borneo and Tripetaloceroides (with one species) from Vietnam and PR China. Members of the subfamily are characteristic within Tetrigidae by massive antennae built up of only eight segments (other Tetrigidae have usually 11-16, Batrachideinae 18-22).[31] Until recently,[31] the subfamily included two tribes - Tripetalocerini and Clinophaestini (including Clinophaestus and Birmana), but the later was moved to the subfamily Metrodorinae due to similarity to Ophiotettigini.[32]

Subfamily unassigned

tribe Echopraxiini Kasalo & Skejo, 2024
  1. Echopraxia (insect) Kasalo & Skejo, 2024
  2. Eurymorphopus Hancock, 1907
  3. Peraxelpa Sjöstedt, 1932
  4. Planotettix Tumbrinck, 2014
  5. Poseidontettix Kasalo & Skejo, 2025
Exanimini Kasalo, Tumbrinck & Skejo, 2024
  1. Exanimus Kasalo, Tumbrinck & Skejo, 2024
  2. Fijixistra Kasalo, Tumbrinck & Skejo, 2024
  3. Ginixistra Kasalo, Tumbrinck & Skejo, 2025
Fijitettigini Kasalo, Tumbrinck & Skejo, 2024
  1. Fijitettix Kasalo, Tumbrinck & Skejo, 2024
  2. Salomonotettix Günther, 1939
Nophthini Kasalo & Skejo, 2025
  1. Dystopia (insect) Kasalo & Skejo, 2025
  2. Nophtha Kasalo & Skejo, 2025
Quasimodini Kasalo & Skejo, 2024
  1. Quasimodo (insect) Kasalo & Skejo, 2024
  2. Seraph (insect) Kasalo & Skejo, 2024
  3. Willemsetettix Tumbrinck, 2014

[33][34]

Xistrellini

Auth: Skejo, Storozhenko, Tumbrinck & Kasalo, 2025; distribution: India, China, Indochina, western Malesia.[35]

  1. Afrosystolederus Devriese & Husemann, 2023
  2. Bannatettix Zheng, 1993
  3. Kanakacris Kasalo & Skejo, 2025
  4. Phaesticus Uvarov, 1940 (synonym Flatocerus Liang & Zheng, 1984)
  5. Pseudoparatettix Günther, 1937
  6. Pseudosystolederus Günther, 1939
  7. Pseudoxistrella Liang, 1991
  8. Synalibas Günther, 1939
  9. Systolederus Bolívar, 1887
  10. Teredorus Hancock, 1907
  11. Xistrella Bolívar, 1909
No tribe assigned
  1. Almacris Skejo, Patano, Škorput & Kasalo, 2025
  2. †Archaeotetrix Sharov, 1968
  3. †Eozaentetrix Zessin, 2017
  4. Euloxilobus Sjöstedt, 1936
  5. Parahirrius Skejo, Patano & Kasalo, 2024
  6. †Prototetrix Sharov, 1968
  7. Spertor Kasalo & Skejo, 2025
  8. Xistra Bolívar, 1887

References

  1. ^ Rambur (1838) Faune entomologique de l'Andalousie 2:64
  2. ^ Song, Hojun; Amédégnato, Christiane; Cigliano, Maria Marta; Desutter-Grandcolas, Laure; Heads, Sam W.; Huang, Yuan; Otte, Daniel; Whiting, Michael F. (2015-03-09). "300 million years of diversification: elucidating the patterns of orthopteran evolution based on comprehensive taxon and gene sampling". Cladistics. 31 (6): 621–651. doi:10.1111/cla.12116. hdl:11336/10677. ISSN 0748-3007. PMID 34753270. S2CID 53702892.
  3. ^ Ragge DR (1965). Grasshoppers, Crickets & Cockroaches of the British Isles. F Warne & Co, London. p. 299.
  4. ^ a b c Borror DJ, Tripplehorn CA, Johnson NF (1989) An Introduction to the Study of Insects, 6th edition. Harcourt Brace College Publishers. New York. pg 213
  5. ^ a b Skejo, Josip; Caballero, Joy Honezza S. (2016-01-21). "A hidden pygmy devil from the Philippines: Arulenus miae sp. nov. — a new species serendipitously discovered in an amateur Facebook post (Tetrigidae: Discotettiginae)". Zootaxa. 4067 (3): 393. doi:10.11646/zootaxa.4067.3.7. ISSN 1175-5334. PMID 27395882.
  6. ^ Imms AD, rev. Richards OW & Davies RG (1970) A General Textbook of Entomology 9th Ed. Methuen 886 pp.
  7. ^ a b Grimaldi D, Engel MS (2005) Evolution of the Insects. Cambridge University Press, Cambridge. pg 211
  8. ^ Skejo, Josip (2017). Taxonomic revision of the pygmy devils (Tetrigidae: Discotettiginae) with online social media as a new tool for discovering hidden diversity. Zagreb: University of Zagreb, Faculty of Science, Department of Biology. pp. 1–246.
  9. ^ a b Preston-Mafham K (1990) Grasshoppers and Mantids of the World. Facts of File, New York. pg 32
  10. ^ a b c Muhammad, Amira Aqilah; Tan, Ming Kai; Abdullah, Nurul Ashikin; Azirun, Mohammad Sofian; Bhaskar, Dhaneesh; Skejo, Josip (2018-09-25). "An annotated catalogue of the pygmy grasshoppers of the tribe Scelimenini Bolívar, 1887 (Orthoptera: Tetrigidae) with two new Scelimena species from the Malay Peninsula and Sumatra". Zootaxa. 4485 (1): 1–70. doi:10.11646/zootaxa.4485.1.1. ISSN 1175-5334. PMID 30313773. S2CID 52975589.
  11. ^ a b c Resh VH, Cardé RT (2003) Encyclopedia of Insects. Academic Press, Amsterdam, pg 839
  12. ^ Bastow, J. L.; Sabo, J. L.; Finlay, J. C.; Power, M. E. (2002). "A basal aquatic-terrestrial trophic link in rivers: algal subsidies via shore-dwelling grasshoppers". Oecologia. 131 (2): 261–268. Bibcode:2002Oecol.131..261B. doi:10.1007/s00442-002-0879-7. PMID 28547694.
  13. ^ Feeding patterns in tropical groundhoppers (Tetrigidae): a case of phylogenetic dietary conservatism in a basal group of Caelifera
  14. ^ "Family Tetrigidae - Pygmy Grasshoppers".
  15. ^ Buckton, George Bowdler (1890). Monograph of the British Cicadae or Tettigidae. London: Macmillan & Co.
  16. ^ Orthoptera Species File: family Tetrigidae Rambur, 1838 (retrieved 14 February 2026)
  17. ^ Kasalo, Niko; Brouste, Damien; Desutter-Grandcolas, Laure; Hervé, Christophe; Bogić, Domagoj; Skejo, Josip (2025-06-27). "Atlas of New Caledonian Tetrigidae with preliminary suprageneric classification of Batrachideinae". Zoosystema. 47 (18). doi:10.5252/zoosystema2025v47a18. ISSN 1280-9551.
  18. ^ Bhaskar, Dhaneesh; Stermšek, Sara; Easa, P. S.; Franjević, Damjan; Skejo, Josip (2020-12-10). "Wide-nosed pygmy grasshoppers (Cladonotinae: Cladonotini, Xerophyllini) of India and Sri Lanka: catalogue with an identification key and description of a new species of the genus Tettilobus". Zootaxa. 4894 (3): 474–500. doi:10.11646/zootaxa.4894.3.12. ISSN 1175-5334. PMID 33311079. S2CID 229173782.
  19. ^ Silva, Daniela Santos Martins; Cadena-Castañeda, Oscar J.; Pereira, Marcelo Ribeiro; De Domenico, Fernando Campos; Sperber, Carlos Frankl (2019-10-16). "Review of Lophotettix, the sole member of the subfamily Lophotettiginae Hancock, 1909 (Insecta: Orthoptera: Caelifera: Tetrigidae)". Zootaxa. 4686 (3): 346–360. doi:10.11646/zootaxa.4686.3.2. ISSN 1175-5334. PMID 31719477. S2CID 207937067.
  20. ^ Deranja, Maks; Kasalo, Niko; Adžić, Karmela; Franjević, Damjan; Skejo, Josip (2022-07-01). "Lepocranus and Valalyllum gen. nov. (Orthoptera, Tetrigidae, Cladonotinae), endangered Malagasy dead-leaf-like grasshoppers". ZooKeys (1109): 1–15. Bibcode:2022ZooK.1109....1D. doi:10.3897/zookeys.1109.85565. ISSN 1313-2970. PMC 9848646. PMID 36762343.
  21. ^ Subedi, Madan; Kasalo, Niko; Skejo, Josip (2025-09). "Definition of the pygmy grasshopper subfamily Criotettiginae (Orthoptera: Tetrigidae) with a preliminary catalogue of genera". Zoologischer Anzeiger. 318: 133–151. doi:10.1016/j.jcz.2025.08.003. ISSN 0044-5231. {{cite journal}}: Check date values in: |date= (help)
  22. ^ Subedi, Madan; Kasalo, Niko (2023-04-18). "Aryalidonta itishreea, a new genus and species of Thoradontini (Orthoptera, Tetrigidae) from Nepal honors the Emperor of Laughter". Journal of Orthoptera Research. 32 (1): 63–80. doi:10.3897/jor.32.94918. ISSN 1937-2426.
  23. ^ Cadena-Castañeda OJ, Quintana-Arias RF, Infante IC, Silva DSM, Tavares GC (2025) Studies on pygmy grasshoppers: On the current Metrodorinae sensu lato classification (Orthoptera: Tetrigidae) with emphasis on American and Malagasy taxa. Zootaxa, 5597(1), 1–265. DOI
  24. ^ Skejo, Josip; Medak, Kristian; Pavlović, Marko; Kitonić, Davorka; Miko, Rafanomezanjanahary Jean Christian; Franjević, Damjan (2020-08-10). "Figure 1 from: Skejo J, Medak K, Pavlović M, Kitonić D, Miko RJC, Franjević D (2020) The story of the Malagasy devils (Orthoptera, Tetrigidae): Holocerus lucifer in the north and H. devriesei sp. nov. in the south? ZooKeys 957: 1-15. https://doi.org/10.3897/zookeys.957.52565". doi.org. Retrieved 2026-03-20. {{cite web}}: External link in |title= (help)
  25. ^ Kasalo, Niko; Husemann, Martin; van de Kamp, Thomas; Skejo, Josip (2024-05-28). "Description of Phelene reinschmidti from Ecuador with notes on the subfamily Lophotettiginae (Orthoptera, Tetrigidae)". Evolutionary Systematics. 8 (1): 119–125. doi:10.3897/evolsyst.8.124285. ISSN 2535-0730.
  26. ^ Kasalo, Niko; Bertner, Paul; González de Rueda, Javier Aznar; Skejo, Josip (2022-06-30). "The True Nature of the Genus Lophotettix Hancock, 1909 (Orthoptera: Tetrigidae): A Brief Taxonomic Revision". Annales Zoologici. 72 (2). doi:10.3161/00034541anz2022.72.2.006. ISSN 0003-4541.
  27. ^ Kasalo, Niko; Yong, Sheyla; Rebrina, Fran; Skejo, Josip (2023-07-28). "Definition of the tribe Metrodorini (Orthoptera: Tetrigidae) with notes on biogeography and evolution of Metrodorinae and Cladonotinae". Acta Entomologica Musei Nationalis Pragae: 187–193. doi:10.37520/aemnp.2023.010. ISSN 1804-6487.
  28. ^ Kasalo, Niko; Yong, Sheyla; Rebrina, Fran; Skejo, Josip (2023-07-28). "Definition of the tribe Metrodorini (Orthoptera: Tetrigidae) with notes on biogeography and evolution of Metrodorinae and Cladonotinae". Acta Entomologica Musei Nationalis Pragae: 187–193. doi:10.37520/aemnp.2023.010. ISSN 1804-6487.
  29. ^ DEVRIESE, HENDRIK; NGUYEN, EILEEN; HUSEMANN, MARTIN (2023-05-17). "An identification key to the genera and species of Afrotropical Tetrigini (genera Paratettix, Leptacrydium, Hedotettix, Rectitettix nov. gen., and Alienitettix nov.gen.) with general remarks on the taxonomy of Tetrigini (Orthoptera, Tetrigidae)". Zootaxa. 5285 (3): 511–556. doi:10.11646/zootaxa.5285.3.4. ISSN 1175-5334.
  30. ^ Bolívar & Urrutia, Ignacio (1887). Essai sur les acridiens de la tribu des Tettigidae. Imprimerie C. Annoot-Braeckman. OCLC 42198873.
  31. ^ a b Storozhenko, Sergey Yu. (2013-10-03). "Review of the subfamily Tripetalocerinae Bolívar, 1887 (Orthoptera: Tetrigidae)". Zootaxa. 3718 (2): 158–170. doi:10.11646/zootaxa.3718.2.4. ISSN 1175-5334. PMID 26258216.
  32. ^ Tumbrinck, Josef; Skejo, Josip (2017). "Taxonomic and biogeographic revision of the New Guinean genus Ophiotettix Walker, 1871 (Tetrigidae: Metrodorinae: Ophiotettigini trib. nov.), with the descriptions of 33 new species" (PDF). Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea. Riga: Entomological Society of Latvia. pp. 525–580, figures 104–124.
  33. ^ Kasalo, Niko; Skejo, Josip (2024-08-19). "The smallest Australian Tetrigidae (Orthoptera): taxonomic revision of Peraxelpa Sjöstedt, 1932 with the descriptions of three new genera and eleven new species". Annales de la Société entomologique de France (N.S.). 60 (5): 515–546. doi:10.1080/00379271.2024.2366405. ISSN 0037-9271.
  34. ^ Kasalo, Niko; Tumbrinck, Josef; Pavlović, Marko; Skejo, Josip (2024-03-30). "Atlas of Fijian Pygmy Grasshoppers (Orthoptera: Tetrigidae) with New Taxa Descriptions and an Identification Key". Annales Zoologici. 74 (1). doi:10.3161/00034541anz2024.74.1.003. ISSN 0003-4541.
  35. ^ Orthoptera Species File: tribe Xistrellini Skejo, Storozhenko, Tumbrinck & Kasalo, 2025.
  • Media related to Tetrigidae at Wikimedia Commons
  • Pygmy locusts
  • Pygmy grasshoppers
  • BugGuide.net
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