Abstract
The genus Endecous (Orthoptera, Grylloidea, Phalangopsidae), commonly found in caves, comprises more than 20 species, grouped into three subgenera: E. (Endecous), E. (Pedroecous) and E. (Notendecous). In this study, from differences observed mainly in the male genitalia, we describe a new subgenus—Endecous (Ramalhoecous) n. sp.—as well as three new species—E. (R.) infernalis n. sp., E. (E.) zin n. sp. and E. (E.) zaum n. sp.—all of them collected in caves in the Serra do Ramalho region, Northeastern, Brazil. We also briefly discuss some troglomorphisms verified in one of the new species: E. (R.) infernalis n. sp.
References
- Alvares, C.A., Stape, J.L., Sentelhas, P.C., Moraes-Gonçalves, J.L. & Sparovek, G. (2013) Köppen’s climate classification map for Brazil. Meteorologische Zeitschrif, 22, 711–728. https://doi.org/10.1127/0941-2948/2013/0507 DOI: https://doi.org/10.1127/0941-2948/2013/0507
- Álvarez-Padilla, F. & Hormiga, G. (2007) A protocol for digesting internal soft tissues and mounting spiders for scanning electron microscopy. The Journal of Arachnology, 35 (3), 538–542. https://doi.org/10.1636/Sh06-55.1 DOI: https://doi.org/10.1636/Sh06-55.1
- Badman, T., Bomhard, B., Fincke, A., Langley, J., Rosabal, P. & Sheppard, D. (2008) Outstanding universal value: Standards for natural world heritage. International Union for Conservation of Nature and Natural Resources (IUCN), Gland, 52 pp.
- Bischof, H.J. (1975) Die keulenförmigen Sensillen auf den Cerci der Grille Gryllus bimaculatus als Schwererezeptoren. Journal of Comparative Physiology, 98, 277–288. https://doi.org/10.1007/BF00656974 DOI: https://doi.org/10.1007/BF00656974
- Bolfarini, M.P. & Bichuette, M.E. (2015) Endecous peruassuensis n. sp. (Orthoptera: Grylloidea: Phalangopsidae) from caves of Eastern Brazil: evidence of isolation in the subterranean realm and discussion about troglomorphisms. Zootaxa, 4032 (3), 297–308. https://doi.org/10.11646/zootaxa.4032.3.5 DOI: https://doi.org/10.11646/zootaxa.4032.3.5
- Campos, L.D., Souza-Dias, P.G.B. & Nihei, S.S. (2017) Taxonomic review of Eidmanacris Chopard, 1956 (Orthoptera: Grylloidea: Phalangopsidae). Zootaxa, 4321 (1), 1–93. https://doi.org/10.11646/zootaxa.4321.1.1 DOI: https://doi.org/10.11646/zootaxa.4321.1.1
- Castro-Souza, R.A., Zefa, E. & Ferreira, R.L. (2020a) New troglobitic and troglophilic syntopic species of Endecous (Orthoptera, Grylloidea, Phalangopsidae) from a Brazilian cave: a case of sympatric speciation?. Zootaxa, 4810 (2), 271–304. https://doi.org/10.11646/zootaxa.4810.2.3 DOI: https://doi.org/10.11646/zootaxa.4810.2.3
- Castro-Souza, R.A., Junta, V.G.P. & Ferreira, R.L. (2020b) Description and ecology of a new species of the cricket genus Endecous (Orthoptera: Grylloidea: Phalangopsidae) in the speleological province of Arcos-Pains-Doresópolis, Southeastern Brazil. Zootaxa, 4821 (2), 305–332. https://doi.org/10.11646/zootaxa.4821.2.4 DOI: https://doi.org/10.11646/zootaxa.4821.2.4
- Castro-Souza, R.A., Zefa, E. & Ferreira, R.L. (2017) Two new species of cave crickets Endecous (Notoendecous) Gorochov, 2014 (Orthoptera: Grylloidea: Phalangopsidae) from northeastern Brazil. Zootaxa, 4318 (3), 474–498. https://doi.org/10.11646/zootaxa.4318.3.3 DOI: https://doi.org/10.11646/zootaxa.4318.3.3
- Cavalcanti, L.F., Lima, M.F.D., Medeiros, R.D.C.S. & Meguerditchian, I. (2012) s.n. In: Plano de ação nacional para a conservação do patrimônio espeleológico nas áreas cársticas da Bacia do Rio São Francisco. Instituto Chico Mendes de Conservação da Biodiversidade, Instituto Chico Mendes, Brasília, pp. 240–256.
- Christiansen, K. (1962) Proposition pour la classification des animaux cavernicoles. Spelunca, 2, 75–78.
- Cigliano, M.M., Braun, H., Eades, D.C. & Otte, D. (2022) Orthoptera Species File. Version 5.0/5.0. Available from: http://Orthoptera.SpeciesFile.org (accessed 30 October 2022)
- Cole, M.M. (1960) Cerrado, Caatinga and Pantanal: The Distribution and Origin of the Savanna Vegetation of Brazil. The Geographical Journal, 126, 168–179. https://doi.org/10.2307/1793957 DOI: https://doi.org/10.2307/1793957
- Culver, D.C. & Sket, B. (2000) Hotspots of subterranean biodiversity in caves and wells. Journal of cave and Karst studies, 62 (1), 11–17. [https://digitalcommons.usf.edu/kip_articles/2593]
- Desutter, L. (1987) Structure et évolution du complexe phallique des Gryllidea (Orthoptera) et classification des genres Nétropicaux de Grylloidea, Première Partie. Annales de la Sociét Entomolologique de France, 23 (3), 213–239.
- Desutter-Grandcolas, L. (1995) Nouveaux grillons cavernicoles de la région néotropicale (Orthoptera, Grylloidea, Phalangopsidae). Annales de la Société entomologique de France, 17, 97–106.
- Desutter-Grandcolas, L. (1998a) Comparative morphology of cercal structures in true crickets and their allies (Orthoptera, Ensifera): a phylogenetic perspective. Zoomorphology, 118 (4), 235–243. DOI: https://doi.org/10.1007/s004350050072
- Desutter-Grandcolas, L. (1998b) II—faune souterraine II. 4—Insecta, 56—Ortóptera, B—Grylloidea. In: Juberthie, C. & Decu, V. (Eds.), Encyclopaedia Biospeologica. V. Tome II. Société de Biospéologie, Moulis, pp. 989–1001.
- Desutter-Grandcolas, L. (2003) Phylogeny and the evolution of acoustic communication in extant Ensifera (Insecta, Orthoptera). Zoologica Scripta, 32, 525–561. https://doi.org/10.1046/j.1463-6409.2003.00142.x DOI: https://doi.org/10.1046/j.1463-6409.2003.00142.x
- Ferreira, R.L., Bernard, E., da Cruz Júnior, F.W., Piló, L.B., Calux, A., Souza-Silva, M. & Frick, W.F. (2022) Brazilian cave heritage under siege. Science, 375 (6586), 1238–1239. https://doi.org/10.1126/science.abo1973 DOI: https://doi.org/10.1126/science.abo1973
- Fontanetti, C.S., Zefa, E., Passetti, F. & Mesa, A. (2002) Morphological characterization and comparative analysis of the proventriculus from three species of Endecous Saussure, 1878 (Orthoptera: Gryllidae: Phalangopsinae). Entomotropica, 17 (1), 15–23.
- Gorochov, A.V. (2014) Classification of the Phalangopsinae subfamily group, and new taxa from the subfamilies Phalangopsinae and Phaloriinae (Orthoptera: Gryllidae). Zoosystematica Rossica, 23 (1), 7–88. https://doi.org/10.31610/zsr/2014.23.1.7 DOI: https://doi.org/10.31610/zsr/2014.23.1.7
- Hartman, H.B., Bennett, L.P., Moulton, B.A. (1987) Anatomy of equilibrium receptors and cerci of the burrowing desert cockroach Arenivaga (Insecta, Blattodea). Zoomorphology, 107 (2), 81–87. https://doi.org/10.1007/BF00312117 DOI: https://doi.org/10.1007/BF00312117
- Horn, E. & Föller, W. (1985) Tonic and modulatory subsystems of the complex gravity receptor system in crickets, Gryllus bimaculatus. Journal of insect physiology, 31 (12), 937–946. https://doi.org/10.1016/0022-1910(85)90028-9 DOI: https://doi.org/10.1016/0022-1910(85)90028-9
- Kassambara, A. & Mundt, F. (2020) factoextra: Extract and visualize the results of multivariate data analyses. R package. Version 1.0.7. Available from: https://cran.r-project.org/web/packages/factoextra/index.html (accessed 16 January 2023)
- Magrin, G.O., Marengo, J.A., Boulanger, J.-P., Buckeridge, M.S., Castellanos, E., Poveda, G., Scarano, F.R. & Vicuña, S. (2014) Central and South America. In: Barros, V.R., Field, C.B., Dokken, D.J., Mastrandrea, M.D., Mach, K.J., Bilir, T.E.; Chatterjee, M., Ebi, K.L., Estrada, Y.O., Genova, R.C., Girma, B., Kissel, E.S., Levy, A.N., MacCracken, S., Mastrandrea, P.R. & White, L.L. (Eds.), Climate Change 2014: Impacts, Adaptation, and Vulnerability. Part B. Regional Aspects. Cambridge University Press, Cambridge and New York, pp. 1499–1566.
- Mammola, S., Cardoso, P., Culver, D.C., Deharveng, L., Ferreira, R.L., Fišer, C., Galassi, D.M.P., Griebler, C., Halse, S., Humphreys, W.F., Isaia, M., Malard, F., Martinez, A., Moldovan, O.T., Niemiller, M.L., Pavlek, M., Reboleira, A.S.P.S., Souza-Silva, M., Teeling, E.C., Wynne, J.J., Zagmajster, M. (2019) Scientists’ warning on the conservation of subterranean ecosystems. BioScience, 69 (8), 641–650. https://doi.org/10.1093/biosci/biz064 DOI: https://doi.org/10.1093/biosci/biz064
- Mello, F.A.G. & Andrade, M.A.B.S. (2003) Ottedana cercalis: a new genus and species of phalangopsid cricket from the Mantiqueira Range of southeastern Brazil (Orthoptera: Grylloidea). Journal of Orthoptera Research, 12, 141–148. [https://www.jstor.org/stable/3503767] DOI: https://doi.org/10.1665/1082-6467(2003)012[0141:OCANGA]2.0.CO;2
- Mello, F.A.G. & Pellegatti-Franco, F. (1998) A new cave cricket of the genus Endecous from southeastern Brazil and characterization of male and female genitalia of E. itatibensis Rehn, 1918 (Orthoptera: Grylloidea: Phalangopsinae: Luzarinae). Journal of Orthoptera Research, 7, 185–188. https://doi.org/10.2307/3503517 DOI: https://doi.org/10.2307/3503517
- Mesa, A. & Garcia-Novo, P. (1997) Endecous onthophagus: a new combination. Phallic sclerites and karyology of the species (Orthoptera: Grylloidea). Journal of Orthoptera Research, 6, 117–120. https://doi.org/10.2307/3503544 DOI: https://doi.org/10.2307/3503544
- Mews, C.M. & Sperber, C.F. (2008) A new species of Endecous Saussure, 1878 and redescription of Endecous cavernicolus Costa-Lima, 1940 (Orthoptera: Grylloidea: Phalangopsidae). Studies on Neotropical Fauna and Environment, 43 (2), 159–167. https://doi.org/10.1080/01650520701492389 DOI: https://doi.org/10.1080/01650520701492389
- Nicklaus, R. (1969) The function of the club-shaped sensilla on the cerci of crickets. Zool Ges Verh, 1969, 393–398.
- Rabelo, L.M., Souza-Silva, M. & Ferreira, R.L. (2018) Priority caves for biodiversity conservation in a key karst area of Brazil: comparing the applicability of cave conservation indices. Biodiversity and conservation, 27 (9), 2097–2129. https://doi.org/10.1007/s10531-018-1554-6 DOI: https://doi.org/10.1007/s10531-018-1554-6
- R Core Team (2022) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Available from: https://www.R-project.org/ (accessed 16 January 2023)
- Silva, R.C., Berbert-Born, M., Bustamante, D.E., Santoro, T.N., Sedor, F. & dos Santos Avilla, L. (2019) Diversity and preservation of Pleistocene tetrapods from caves of southwestern Bahia, Brazil. Journal of South American Earth Sciences, 90, 233–254. https://doi.org/10.1016/j.jsames.2018.12.004 DOI: https://doi.org/10.1016/j.jsames.2018.12.004
- Souza-Dias, P.G.B., Bolfarini M.P., Nihei, S.S. & Mello, F.A. (2014) Endecous apterus: A new species of cave cricket from northeast Brazil, with comments on the use of subterranean habitats by Luzarinae crickets (Orthoptera: Grylloidea: Phalangopsidae: Luzarinae). Zootaxa, 3784 (2), 120–130. https://doi.org/10.11646/zootaxa.3784.2.2 DOI: https://doi.org/10.11646/zootaxa.3784.2.2
- Souza-Dias, P.G.B. (2015) Análise cladística e morfologia do complexo fálico de Phalangopsidae, com ênfase em Luzarinae (Orthoptera, Ensifera, Grylloidea). Doctoral thesis, Instituto de Biociências da Universidade de São Paulo, São Paulo, 183 pp.
- Souza-Dias, P.G.B., Mello, F.A.G; Vieira, L.B. (2016) A new genus and species of cricket from the Chapada Diamantina National Park, northeastern Brazil (Grylloidea: Phalangopsidae; Luzarinae). Zootaxa, 4121 (3), 258–266. http://doi.org/10.11646/zootaxa.4121.3.2 DOI: https://doi.org/10.11646/zootaxa.4121.3.2
- Souza-Dias, P.G.B., Szinwelski, N., Fianco M., De Oliveira, E.C., Mello, F.A.G. & Zefa, E. (2017) New species of Endecous (Grylloidea, Phalangopsidae, Luzarinae) from the Iguaçu National Park (Brazil), including bioacoustics, cytogenetic and distribution data. Zootaxa, 4237 (3), 454–470. https://doi.org/10.11646/zootaxa.4237.3.2 DOI: https://doi.org/10.11646/zootaxa.4237.3.2
- Souza-Silva, M., Cerqueira, R.F.V., Pellegrini, T.G. & Ferreira, R.L. (2021) Habitat selection of cave-restricted fauna in a new hotspot of subterranean biodiversity in Neotropics. Biodiversity and Conservation, 30 (14), 4223–4250. https://doi.org/10.1007/s10531-021-02302-8 DOI: https://doi.org/10.1007/s10531-021-02302-8
- Watson, J., Hamilton-Smith, E., Gillieson, D. & Kiernan, K. (1997) Guidelines for cave and karst protection. International Union for Conservation of Nature and Natural Resources (IUCN) Gland and Cambridge, 63 pp.
- Williams, P. (2008) World heritage caves and karst. International Union for Conservation of Nature and Natural Resources (IUCN) World Heritage Studies, Gland, 57 pp.
- Zefa, E., Mesa, A. & Martins, L.D.P. (2010) New Brazilian species of Endecous Saussure, 1878: Phallic sclerites, calling song and tegmen morphometry (Orthoptera: Grylloidea: Phalangopsinae). Entomological science, 13 (1), 150–155. https://doi.org/10.1111/j.1479-8298.2010.00371.x DOI: https://doi.org/10.1111/j.1479-8298.2010.00371.x
- Zefa, E., Redu, D.R., Costa, M.K.M., Gottschalk, M.S., Padilha, G.B., Silva, A.F. & Martins, L.P. (2014) A new species of Endecous Saussure, 1878 (Orthoptera, Gryllidae) from northeast Brazil with the first X¹X²0 chromosomal sex system in Gryllidae. Zootaxa, 3847 (1), 125–132. https://doi.org/10.11646/zootaxa.3847.1.7 DOI: https://doi.org/10.11646/zootaxa.3847.1.7