Skip to main content Skip to main navigation menu Skip to site footer
Responsive image
Issue: Vol. 5722 No. 4: 21 Nov. 2025
Type: Article
Published: 2025-11-21
DOI: 10.11646/zootaxa.5722.4.2
Page range: 485-508
Abstract views: 40
PDF downloaded: 2

New species of Cretaceous Locustopsidae (Orthoptera: Caelifera) from the Crato Formation of Brazil and a taxonomic revision of the family

Leibniz Institute for the Analysis of Biodiversity Change (LIB); Centre for Taxonomy and Morphology (ztm); 20146; Hamburg; Germany
Museu de Paleontologia Plácido Cidade Nuvens; Universidade Regional do Cariri; Santana do Cariri; Ceará; Brazil
Illinois Center for Paleontology; Prairie Research Institute; University of Illinois Urbana-Champaign; Forbes Natural History Building; 1816 South Oak Street; Champaign; Illinois 61820; USA
Museu de Paleontologia Plácido Cidade Nuvens; Universidade Regional do Cariri; Santana do Cariri; Ceará; Brazil
Leibniz Institute for the Analysis of Biodiversity Change (LIB); Centre for Biodiversity Monitoring and Conservation Science (zbm); 20146; Hamburg; Germany
State Museum of Natural History Karlsruhe (SMNK); Karlsruhe; Baden-Württemberg; 76133; Germany
Orthoptera Araripe Basin Lower Cretaceous fossil grasshoppers Konservat-Lagerstätte

Abstract

A new genus and four new species of Locustopsidae Handlirsch, 1906 are described from the Crato Formation of northeastern Brazil. Aestuacrida stereofemoris gen. et sp. nov. differs from previous genera of the family by having four branches of CuA + CuPaα. A new species of Araripelocusta Martins-Neto, 1995, Araripelocusta imperatrix sp. nov., is much larger than previous congeners with a wingspan of ca. 39 mm and features two little leaf-like appendages on the dorsal front part of the head. Parapleurites brasiliensis sp. nov. is the first Cretaceous member of this genus following two Jurassic predecessors. Parapleurites morrisonensis Smith, Gorman, Pardo & Small, 2011 has a wing venation that more closely resembles Mesolocustopsis Hong and Wang, 1990 and is moved to this genus as M. morrisonensis comb. nov. Cratolocustopsis aquila sp. nov. joins C. cretacea (Martins-Neto, 1990) and C. contumax Martins-Neto, 2003 as the third member of this genus, helping to solidify its separate position from other genera of Locustopsidae. A comprehensive taxonomic revision of all known species of Locustopsidae is given to improve our understanding of the diversity of these short-horned Mesozoic grasshoppers. The genus Cratozeunerella Martins-Neto, 1998 is considered a junior synonym of Zeunerella Sharov, 1968. Cratozeunerella godoii Martins-Neto, 2003, C. nervosa Martins-Neto, 2003, C. soaresi Martins-Neto, 2003 and C. titanella Martins-Neto, 2003 are suggested to be included in Locustopsis Handlirsch, 1906. Plesioschwinzia Zessin, 1988 is considered a junior synonym of Locustopsis. Further changes on the species level concern former Zeunerella reticulata (Handlirsch, 1939) which is moved to Cratolocustopsis Martins-Neto, 2003. Mesolocustopsis petruleviciusi (Martins-Neto, 2003) is considered a junior synonym of M. caririensis (Martins-Neto, 1990). In Locustopsis two synonyms are proposed: Locustopsis elongata Handlirsch, 1906 syn. nov. of L. elegans Handlirsch, 1906, and Locustopsis lacera (Zessin, 1983) (former Plesioschwinzia lacera) syn. nov. of L. gyra Zessin, 1983. Additionally, Locustopsis anatolica Sharov, 1968 is moved to Schwinzia Zessin, 1983. The recently described species of the Chinese Sinolocustopsis Huang & Nel, 2024 are moved to the genus Locustopsis: Locustopsis elongatus comb. nov. (Huang & Nel, 2024) and the genus Mesolocustopsis: Mesolocustopsis brevis comb. nov. (Huang & Nel, 2024). Liadolocusta auscultans Handlirsch, 1906 is considered a nomen nudum. Locustopsites gigantea Théobald, 1937 is probably a member of Tettigoniidae Krauss, 1902 and should be excluded from Locustopsidae.

 

References

  1. Assine, M.L. (2007) Bacia do Araripe. Boletim de Geociências da Petrobras, 15, 371–389.
  2. Assine, M.L., Perinotto, J.A.J., Neumann, V.H., Custódio, M.A., Varejão, F.G. & Mescolotti, P.C. (2014) Sequências Deposicionais do Andar Alagoas (Aptiano superior) da Bacia do Araripe, Nordeste do Brasil. Boletim de Geociências da Petrobras, 22 (1), 3–28. https://doi.org/10.25249/0375-7536.1992289300
  3. Assine, M.L., Quaglio, F., Warren, L.V. & Simões, M.G. (2016) Comments on paper by M. Arai, “Aptian/Albian (Early Cretaceous) paleogeography of the South Atlantic: a paleontological perspective”. Brazilian Journal of Geology, 46, 3–7. https://doi.org/10.1590/2317-4889201620150046A
  4. Ander, K.E.V. (1939) Vergleichend-anatomische und Phylogenetische studien über die Ensifera (Saltatoria). Vol. 2. Berlingska Boktryckeriet, Lund. [unknown pagination]
  5. Ansorge, J. (1991) Locustopsis africanus n. sp. (Saltatoria, Caelifera) aus der Unterkreide SW-Agyptens. Neues Jahrb. Geol. Palaeontol. Monatsh., 1991 (4), 205–212. https://doi.org/10.1127/njgpm/1991/1991/205
  6. Arai, M. (2014) Aptian/Albian (Early Cretaceous) paleogeography of the South Atlantic: a paleontological perspective. Brazilian Journal of Geology, 44 (2), 339–350. https://doi.org/10.5327/Z2317-4889201400020012
  7. Arai, M. & Assine, M.L. (2020) Chronostratigraphic constraints and paleoenvironmental interpretation of the Romualdo Formation (Santana Group, Araripe Basin, Northeastern Brazil) based on palynology. Cretaceous Research, 116, 104610. https://doi.org/10.1016/j.cretres.2020.104610
  8. Béthoux, O. & Nel, A. (2002) Venation pattern and revision of Orthoptera sensu nov. and sister groups. Phylogeny of Palaeozoic and Mesozoic Orthoptera sensu nov. Zootaxa, 96 (1), 1–88. https://doi.org/10.11646/zootaxa.96.1.1
  9. Bezerra, F.I., da Silva, J.H., Miguel, E.C., Paschoal, A.R., Nascimento Jr., D.R., Freire, P.T.C., Viana, B.C. & Mendes, M. (2020) Chemical and mineral comparison of fossil insect cuticles from Crato Konservat Lagerstätte, Lower Cretaceous of Brazil. Journal of Iberian Geology, 46, 61–76. https://doi.org/10.1007/s41513-020-00119-y
  10. Bezerra, F.I., Solórzano-Kraemer, M.M. & Mendes, M. (2021) Distinct preservational pathways of insects from the Crato Formation, Lower Cretaceous of the Araripe Basin, Brazil. Cretaceous Research, 118, 104631. https://doi.org/10.1016/j.cretres.2020.104631
  11. Bezerra, F.I. & Mendes, M. (2024) A palaeoecological analysis of the Cretaceous (Aptian) insect fauna of the Crato Formation, Brazil. Palaeogeography, Palaeoclimatology, Palaeoecology, 641, 112134. https://doi.org/10.1016/j.palaeo.2024.112134
  12. Bode, A. (1953) Die Insektenfauna des ostniedersächsischen oberen Lias. Palaeontographica. Beitraege zur Naturgeschichte der Vorzeit, 103, 1–375.
  13. Brauer, F.M., Redtenbacher, J. & Ganglbauer, L. (1889) Fossile Insekten aus der Juraformation Ost-Sibiriens. Mémoires de l‘Académie Impériale des Sciences de St. Pétersbourg, 36, 1–22.
  14. Brodie, P.B. (1845) A history of the fossil insects in the secondary rocks of England. J.van Voorst, London, 140 pp. https://doi.org/10.5962/bhl.title.52321
  15. Brullé, G.A. (1835) Cinquième Ordre. Orthoptères. In: Histoire naturelle des insectes: Vol. 9 [1] (5). Société bibliophile, Paris, pp. 1–225. [225–416 in 1836]
  16. Catto, B., Jahnert, R.J., Warren, L.V., Varejão, F.G. & Assine, M.L. (2016) The microbial nature of laminated limestones: lessons from the upper Aptian, Araripe Basin, Brazil. Sedimentary Geology, 341, 304–315. https://doi.org/10.1016/j.sedgeo.2016.05.007
  17. Cigliano, M.M., Braun, H., Eades, D.C. & Otte, D. (2025) Orthoptera Species File. Version July 2025. Available from: http://orthoptera.speciesfile.org/ (accessed 31 July 2025)
  18. Cockerell, T.D.A. (1916) British fossil insects. Proceedings of the United States National Museum, 49, 469–499. https://doi.org/10.5479/si.00963801.49-2119.469
  19. Coimbra, J.C., Arai, M. & Carreño, A.L. (2002) Biostratigraphy of Lower Cretaceous microfossils from the Araripe basin, northeastern Brazil. Geobios, 35 (6), 687–698. https://doi.org/10.1016/S0016-6995(02)00082-7
  20. Coimbra, J.C. & Freire, T.M. (2021) Age of the Post-rift Sequence I from the Araripe Basin, Lower Cretaceous, NE Brazil: Implications for spatio-temporal correlation. Revista Brasileira de Paleontologia, 24 (1), 37–46. https://doi.org/10.4072/rbp.2021.1.03
  21. Corecco, L., Bezerra, F.I., Silva Filho, W.F., Nascimento Jr., D.R., da Silva, J.H. & Felix, J.L. (2022) Petrological meaning of ethnostratigraphic units: Laminated Limestone of the Crato Formation, Araripe Basin, NE Brazil. Pesquisas em Geociências, 49 (1), e121139. https://doi.org/10.22456/1807-9806.121139
  22. de Moura-Júnior, D.A., Scheffler, S.M. & Fernandes, A.C.S. (2018) A paleoentomofauna brasileira: cenário atual. Anuário do Insituto de Geociências, 41, 142–166. https://doi.org/10.11137/2018_1_142_166
  23. Deichmüller, J.V. (1886) Die Insecten aus dem lithographischen Schiefer in Dresdener Museum. Fischer, Basel, 88 pp.
  24. Dias, J.J. & Carvalho, I.D.S. (2022) The role of microbial mats in the exquisite preservation of Aptian insect fossils from the Crato Lagerstätte, Brazil. Cretaceous Research, 130, 105068. https://doi.org/10.1016/j.cretres.2021.105068
  25. Geinitz, F.E. (1880) Der Jura von Dobbertin in Mecklenburg und seine Versteinerungen. Zeitschrift der Deutschen Geologischen Gesellschaft, 32, 510–535.
  26. Germar, E.F. (1842) Beschreibung einiger neuen fossilen Insekten. I. in den lithographischen Schiefern von Bayern. II. Insektenflügel im Schieferthone des Steinkohlegebirges von Wettin. Beiträge zur Petrefactenkunde, 5, 79–94.
  27. The GIMP Team (2025) GIMP. Version 3.0.2-1. Available from: https://www.gimp.org/ (accessed 7 October 2025)
  28. Gorochov, A.V. (1985) Mesozoic crickets (Orthoptera, Grylloidea) of Asia [in Russian]. Paleontologicheskiy Zhurnal, 198 (2), 59–68.
  29. Gorochov, A.V. (1995) System and evolution of the suborder Ensifera (Orthoptera). Part II. Proceedings of the Zoological Institute of the Russian Academy of Sciences, 260, 1–213.
  30. Gorochov, A.V., Jarzembowski, E.A. & Coram, R.A. (2006) Grasshoppers and crickets (Insecta: Orthoptera) from the Lower Cretaceous of southern England. Cretaceous Research, 27 (5), 641–662. https://doi.org/10.1016/j.cretres.2006.03.007
  31. Gorochov, A.V. & Coram, R.A. (2023) New and little known taxa of the order Orthoptera (Insecta) from the Upper Triassic and Lower Jurassic of England. Palaeoentomology, 6 (2), 198–204. https://doi.org/10.11646/palaeoentomology.6.2.11
  32. Gu, J.J., Yue, Y., Shi, F.M., Tian, H. & Ren, D. (2016) First Jurassic grasshopper (Insecta, Caelifera) from China. Zootaxa, 4169 (2), 377–380. https://doi.org/10.11646/zootaxa.4169.2.9
  33. Handlirsch, A. (1906) Die fossilen Insekten und die Phylogenie der rezenten Formen: ein Handbuch für Paläontologen und Zoologen. Vol. 1. W. Engelmann, Leipzig, IX + VI + 160 + XX + 672 + II + XII + II + II pp., XXXVI taf. https://doi.org/10.5962/bhl.title.5636
  34. Handlirsch, A. (1939) Neue Untersuchungen über die fossilen Insekten mit Ergänzungen und Nachträgen sowie Ausblicken auf phylogenetische, palaeogeographische und allgemein biologische Probleme. II. Teil. Annalen des naturhistorischen Museums in Wien, 49, 1–240.
  35. Heimhofer, U., Ariztegui, D., Lenniger, M., Hesselbo, S.P., Martill, D.M. & Rios-Neto, A.M. (2010) Deciphering the depositional environment of the laminated Crato fossil beds (Early Cretaceous, Araripe Basin, North-Eastern Brazil). Sedimentology, 57 (2), 677–694. https://doi.org/10.1111/j.1365-3091.2009.01114.x
  36. Heimhofer, U. & Hochuli, P.A. (2010) Early Cretaceous angiosperm pollen from a low-latitude succession (Araripe Basin, NE Brazil). Review of Palaeobotany and Palynology, 161 (3–4), 105–126. https://doi.org/10.1016/j.revpalbo.2010.03.010
  37. Hong Y.C. & Wang W.L. (1990) Fossil Insects from the Laiyang Basin, Shandong Province] In: Regional Geological Surveying Team, Shandong Bureau of Geology and Mineral Resources (Ed.), The stratigraphy and Palaeontology of Laiyang Basin, Shandong Province. Geological Publishing House, Beijing, pp. 44–189.
  38. Kirby, W.F. (1906) Orthoptera Saltatoria. Part I. (Achetidae et Phasgonuridae). In: A Synonymic Catalogue of Orthoptera (Orthoptera Saltatoria, Locustidae vel Acridiidae). Vol. 2. British Museum (Natural History), London, pp. i–viii + 1–562.
  39. Krauss, H.A. (1902) Die Namen der ältesten Dermapteren- (Orthopteren-) Gattungen und ihre Verwendung für Familien- und Unterfamilien-Benennungen auf Grund der jetzigen Nomenclaturregein. Zoologischer Anzeiger, 25 (676), 530–543.
  40. Latreille, P.A. (1810) Considérations générales sur l‘ordre naturel des animaux composant les classes des Crustacés, des Arachnides, et des Insectes; avec un tableau méthodique de leurs genres, disposés en familles. Chez F. Schoell, rue des Fosses S.G.l‘Auxerrois, Paris, 444 pp. https://doi.org/10.5962/bhl.title.39620
  41. Leach, W.E. (1815) Entomology. In: The Edinburgh Encyclopaedia. Vol. 9. William Blackwood, Edinburgh, pp. 57–172.
  42. Lin, Q. (1982) Class Insecta. Shaanxi-Gansu-Ningxia volumn, Mesozoic and Cenozoic. In: Paleontological atlas of Northwest China: Vol. II. Paleontological Atlas of Northwest China. Geological Publishing House, Beijing, pp. 70–83.
  43. Linnaeus, C. (1758) Systema naturae. Vol. 1. Laurentii Salvii, Stockholm, 824 pp.
  44. Lúcio, T., Neto, J.A.S. & Selby, D. (2020) Late Barremian/early Aptian Re-Os age of the Ipubi Formation black shales: Stratigraphic and paleoenvironmental implications for Araripe Basin, northeastern Brazil. Journal of South American Earth Sciences, 102, 102699. https://doi.org/10.1016/j.jsames.2020.102699
  45. MacLeay, W.S. (1821) Horae Entomologicae or Essays on the Annulose Animals. Vol. 2. S. Bagster, London. [unknown pagination]
  46. Martill, D.M., Loveridge, R. & Heimhofer, U. (2007) Halite pseudomorphs in the Crato Formation (Early Cretaceous, Late Aptian–Early Albian), Araripe Basin, northeast Brazil: further evidence for hypersalinity. Cretaceous Research, 28, 613–620. https://doi.org/10.1016/j.cretres.2006.10.003
  47. Martins-Neto, R.G. (1990) The family Locustopsidae (Insects, Caelifera) in the Santana Formation (Lower Cretaceous, Northeast Brazil. I. Description of two new species of the genus Locustopsis Handlirsch and three new species of the genus Zessinia n. gen. In: Atas do I Simposio sobre a Bacia do Araripe e bacias interiores do Nordeste. DNPM, Crato, pp. 227–291.
  48. Martins-Neto, R.G. (1995) Araripelocustidae, fam. n. uma familia de gafanhotos (Insecta, Caelifera) da Formação Santana, Cretáceo Inferior do nordeste do Brasil. Revista Brasileira de Entomologia, 39 (2), 311–319, illustr.
  49. Martins-Neto, R.G. (1998) A new genus of the family Locustopsidae (Insecta, Caelifera) in the Santana Formation (Lower Cretaceous, northeastern Brazil). Revista Española de Paleontologia, 13 (2), 133–138. https://doi.org/10.7203/sjp.24046
  50. Martins-Neto, R.G. (2001) Review of some insects from Mesozoic and Cenozoic Brazilian deposits, with descriptions of new taxa. Acta Geologica Leopoldensia, 24 (52/53), 115–124.
  51. Martins-Neto, R.G. (2003) Systematics of the Caelifera (Insecta, Orthopteroidea) from the Santana Formation, Araripe Basin (Lower Cretaceous, northeast Brazil), with a description of new genera and species. Acta Zoologica Cracoviensia, 46, 205–228.
  52. Martynov, A.V. (1937) Liasovyye nasekomyye Shuraba i Kizil-Kii (Liassic Insects of Shurab and Kizyl-Kiya). Trudy Paleontologicheskogo Instituta, Akademiia Nauk SSSR [Transactions of the Institute of Paleontology, USSR Academy of Sciences], 7 (1), 5–232.
  53. Melo, R.M., Guzmán, J., Almeida-Lima, D., Piovesan, E.K., Neumann, V.H.D.M.L. & Sousa, A.D.J.E. (2020) New marine data and age accuracy of the Romualdo Formation, Araripe Basin, Brazil. Scientific Reports, 10 (1), 15779. https://doi.org/10.1038/s41598-020-72789-8
  54. Mendes, M., Vasconcelos, S.M.O.L. de & Oliveira, F.I.B. de(2019) A new fossil stick grasshopper (Proscopioidea: Proscopiidae) from the Crato formation of Brazil. Anuário do Instituto de Geociências, UFRJ, 42 (2), 437–443. https://doi.org/10.11137/2019_2_437_443
  55. Mendes, M., Bezerra, F.I. & Adami, K. (2020) Ecosystem Structure and Trophic Network in the Late Early Cretaceous Crato Biome. In: Iannuzzi, R., Rößler, R. & Kunzmann, L. (Eds.), Brazilian Paleofloras: from Paleozoic to Holocene. Springer, Cham, pp. 1–19. https://doi.org/10.1007/978-3-319-90913-4_33-1
  56. The Inkscape Team (2025) Inkcape. Version 1.4.2. Available from: https://inkscape.org/de/ (accessed 31 July 2025)
  57. Nel, A. & Jouault, C. (2022) New grasshoppers (Orthoptera: Elcanidae, Locustopsidae) from the Lower Cretaceous Crato formation suggest a biome homogeneity in Central Gondwana. Historical Biology, 34 (10), 2070–2078. https://doi.org/10.1080/08912963.2021.2000602
  58. Nel, A. & Huang, D.Y. (2024) New locustopsid fossils from the Middle Jurassic Yan’an Formation (Orthoptera: Caelifera). Journal of Insect Biodiversity, 60 (1), 16–23.
  59. Neumann, V.H., Borrego, A.G., Cabrera, L. & Dino, R. (2003) Organic matter composition and distribution through the Aptian–Albian lacustrine sequences of the Araripe Basin, northeastern Brazil. International Journal of Coal Geology, 54 (1–2), 21–40. https://doi.org/10.1016/S0166-5162(03)00018-1
  60. Ponte, F.C. & Ponte-Filho, F.C. (1996) Estrutura geológica e evolução tectônica da Bacia do Araripe. Departamento Nacional de Produção Mineral, Recife, 68 pp.
  61. Santos, F.H., Azevedo, J.M., Nascimento Jr., D.R., Souza, A.C.B., Mendes, M., Bezerra, F.I. & Limaverde, S. (2017) Análise de fácies e petrografia de uma seção do Membro Crato em Nova Olinda (CE): contribuições à história deposicional e diagenética do neoaptiano na Bacia do Araripe. Geologia, Série Científica, 17 (1), 3–18. https://doi.org/10.11606/issn.2316-9095.v17-319
  62. Serville, J.G.A. (1838) Histoire naturelle des insectes. Orthoptères. Librairie Encyclopédique de Roret, Paris, xviii + 776 pp., 14 pls. [1839]
  63. Sharov, A.G. (1968) Filogniya orthopteroidnykh nasekomykh [1971 English translation: Phylogeny of the Orthopteroidea]. Trudy Paleontologicheskogo Instituta, Akademiia Nauk SSSR [Transactions of the Institute of Paleontology, USSR Academy of Sciences], 118, 1–216.
  64. Smith, D.M., Gorman, M.A., Pardo, J.D. & Small, B.J. (2011) First fossil Orthoptera from the Jurassic of North America. Journal of Paleontology, 85 (1), 102–105. https://doi.org/10.1666/10-096.1
  65. Strand, E. (1926) Miscellanea nomenclatorica zoologica et palaeontologica. Archiv für Naturgeschichte, 92 (8), 30–75. [1928]
  66. The Paleobiology Database (2025) The Paleobiology Database. Available from: https://paleobiodb.org/#/ (accessed 15 August 2025)
  67. Théobald, N. (1937) Les Insectes Fossiles des terrains oligocenes de France. Imprimerie Georges Thomas, Angle des rues de Solignac et Henri-Lepage, Nancy, 473 pp.
  68. von Laicharting, J.N.E. (1781) Verzeichnis und Beschreibung der Tyroler Insecten. Vol. 1. bey J.C. Füessly, Zürich, XII + 176 pp.
  69. Whalley, P.E.S. (1985) The systematics and palaeogeography of the Lower Jurassic insects of Dorset, England. Bulletin of the British Museum (Natural History) Geology, 39 (3), 107–189.
  70. Zessin, W. (1983) Revision der mesozoischen Familie Locustopsidae unter Berücksichtigung neuer Funde (Orthopteroida, Caelifera). Deutsche Entomologische Zeitschrift, 30 (1–3), 173–237. https://doi.org/10.1002/mmnd.19830300115
  71. Zessin, W. (1983) Locustopsis kruegeri n. sp. (Orthopteroida, Caelifera) aus dem oberen Lias von Schandelah bei Braunschweig (BRD). Zeitschrift für geologische Wissenschaften, 11 (7), 905–910.
  72. Zessin, W. (1988) Neue Saltatoria (Insecta) aus dem Oberlias Mitteleuropas. Freiberger Forschungshefte, Reihe C, 419, 107–121.
  73. Zeuner, F.E. (1942) The Locustopseidae and the phylogeny of the Acridodea (Orthoptera). Proceedings of the Royal Entomological Society of London, Series B, 11 (1), 1–19. https://doi.org/10.1111/j.1365-3113.1942.tb00713.x
  74. Zherikhin, V.V. (1985) Jurassic orthopterans of south Siberia and western Mongolia. Trudy Paleontologicheskogo Instituta, Akademiia Nauk SSSR [Transactions of the Institute of Paleontology, USSR Academy of Sciences], 211, 171–184.

How to Cite

Schall, O.-K.O., Lima, D., Heads, S.W., Pinheiro, A.P., Kotthoff, U. & Husemann, M. (2025) New species of Cretaceous Locustopsidae (Orthoptera: Caelifera) from the Crato Formation of Brazil and a taxonomic revision of the family. Zootaxa, 5722 (4), 485–508. https://doi.org/10.11646/zootaxa.5722.4.2