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Type: Monograph
Published: 2017-06-08
Page range: 1–86
Abstract views: 182
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Integrative taxonomy reveals cryptic diversity in neotropical grasshoppers: taxonomy, phylogenetics, and evolution of the genus Sphenarium Charpentier, 1842 (Orthoptera: Pyrgomorphidae)

Laboratorio de Ecología, UBIPRO, Facultad de Estudios Superiores Iztacala, Universidad Nacional Autónoma de México (UNAM), A.P. 314, Tlalnepantla, 54090 México, México.
Department of Entomology, Texas A&M University, 2475 TAMU, College Station, TX, U.S.A.
Escuela Nacional de Estudios Superiores Unidad Morelia, UNAM, Morelia, 58190 Michoacán, México.
Instituto de Investigaciones en Ecosistemas y Sustentabilidad, UNAM, Morelia, 58190 Michoacán, México.
Laboratorio de Ecología, UBIPRO, Facultad de Estudios Superiores Iztacala, Universidad Nacional Autónoma de México (UNAM), A.P. 314, Tlalnepantla, 54090 México, México.
Diptera Integrative taxonomy Neotropics Pyrgomorphidae Sphenarium male genitalia morphology molecular phylogenetics species phylogeny

Abstract

The genus Sphenarium Charpentier, 1842 comprises the most diverse group of the New World Pyrgomorphidae. These grasshoppers show an extensive variation in external morphology, and are culturally and economically important for Mexican people since pre-Hispanic times. Nevertheless, the taxonomy of Sphenarium has been chaotic and remained incompletely resolved until now. Following an integrative taxonomic framework, we infer the number of species in this genus by species delimitation based on morphological, phylogenetic, and geographic information. Based on our results, we revise the genus and redefine 9 species and describe 8 new species (S. adelinae sp.n., S. crypticum sp.n., S. infernalis sp.n., S. miztecum sp.n., S. occidentalis sp.n., S. tarascum sp.n., S. totonacum sp.n. and S. zapotecum sp.n.). Moreover, we update the knowledge of the evolutionary relationships and biogeographic patterns of Sphenarium species. Our results also demonstrate the importance of historic geological and climatic events on the lineage diversification of this genus. Different levels of morphological and genetic differentiation among species suggest a complex interplay between evolutionary forces during the evolution of these neotropical grasshoppers.

 

References

  1. Allegrucci, G., Massa, B., Trasatti, A. & Sbordoni, V. (2013) A taxonomic revision of western Eupholidoptera bush crickets (Orthoptera: Tettigoniidae): Testing the discrimination power of DNA barcode. Systematic Entomology, 39 (1), 7–23.
    https://doi.org/10.1111/syen.12031

    Allegrucci, G., Rampini, M., Di Russo, C., Lana, E., Cocchi, S. & Sbordoni, V. (2014) Phylogeography and systematics of the westernmost Italian Dolichopoda species (Orthoptera, Rhaphidophoridae). ZooKeys, 23 (437), 1–23.
    https://doi.org/10.3897/zookeys.437.7917

    Bargues, M.D., Marcilla, A., Ramsey, J.M., Dujardin, J.P., Schofield, C.J. & Mas-Coma, S. (2000) Nuclear rDNA-based Molecular Clock of the Evolution of Triatominae (Hemiptera: Reduviidae), Vectors of Chagas Disease. Memorias do Instituto Oswaldo Cruz, 95 (4), 567–573.
    https://doi.org/10.1590/S0074-02762000000400020

    Barrowclough, G.F. & Zink, R.M. (2009) Funds enough, and time: mtDNA, nuDNA and the discovery of divergence. Molecular Ecology, 18 (14), 2934–2936.
    https://doi.org/10.1590/S0074-02762000000400020

    Bolivar, I. (1884) Monografia de los Pyrgomorfinos. Anales de la Sociedad Española de Historia Natural, 13, 1–500.

    Bolivar, I. (1904) Notas sobre los Prygomórfidos (Pyrgomorphidae). Boletín de la Sociedad Española de Historia Natural, 4, 306–326.

    Bolivar, I. (1909) Orthoptera Fam. Acridiidae Subfam. Pyrgomorphinae. Genera insectorum, 90, 1–59.

    Bouckaert, R., Heled, J., Kühnert, D., Vaughan, T., Wu, C.H., Xie, D., Suchard, M.A., Rambaut, A. & Drummond, A.J. (2014) BEAST 2: A Software Platform for Bayesian Evolutionary Analysis. PLoS Computational Biology, 10 (4), 1–6.
    https://doi.org/10.1371/journal.pcbi.1003537

    Boyle, W.K. (1974) A Revision of the Genus Sphenarium (Orthoptera: Pyrgomorphidae). McGill University, McGill, 214 pp.

    Brower, A.V. (1994) Rapid morphological radiation and convergence among races of the butterfly Heliconius erato inferred from patterns of mitochondrial DNA evolution. Proceedings of the National Academy of Sciences of the United States of America, 91 (14), 6491–6495.
    https://doi.org/10.1073/pnas.91.14.6491

    Bruner, L. (1906) Insecta. Orthoptera. Vol. II. In: Godman, F.D. & Salvin, O. (Eds.), Biologia Centrali-Americana. R. H. Porter, London, pp. 199–207.
    https://doi.org/10.5962/bhl.title.730

    Bryson, R.W., García-Vázquez, U.O. & Riddle, B.R. (2011) Phylogeography of Middle American gophersnakes: mixed responses to biogeographical barriers across the Mexican Transition Zone. Journal of Biogeography, 38 (8), 1570–1584.
    https://doi.org/10.1111/j.1365-2699.2011.02508.x

    Bryson, R.W., García-vázquez, U.O. & Riddle, B.R. (2012a) Relative roles of Neogene vicariance and Quaternary climate change on the historical diversification of bunchgrass lizards ( Sceloporus scalaris group ) in Mexico. Molecular Phylogenetics and Evolution, 62 (1), 447–457.
    https://doi.org/10.1016/j.ympev.2011.10.014

    Bryson, R.W., Nieto-Montes de Oca, A., Jaeger, J.R. & Riddle, B.R. (2010) Elucidation of cryptic diversity in a widespread nearctic treefrog reveals episodes of mitochondrial gene capture as frogs diversified across a dynamic landscape. Evolution, 64 (8), 2315–2330.
    https://doi.org/10.1111/j.1558-5646.2010.01014.x

    Cano-Santana, Z. & Castellanos-Vargas, I. (2009) Historia natural y ecología de Sphenarium purpurascens (Orthoptera : Pyrgomorphidae). In: Lot, A. & Cano-Santana, Z. (Eds.), Biodiversidad del Ecosistema del Pedregal de San Ángel. Universidad Nacional Autonoma de Mexico, Mexico D.F., pp. 337–346.

    Castoe, T.A., Daza, J.M., Smith, E.N., Sasa, M.M., Kuch, U., Campbell, J.A., Chippindale, P.T. & Parkinson, C.L. (2009) Comparative phylogeography of pitvipers suggests a consensus of ancient Middle American highland biogeography. Journal of Biogeography, 36 (1), 88–103.
    https://doi.org/10.1111/j.1365-2699.2008.01991.x

    Cerritos, R. & Cano-Santana, Z. (2008) Harvesting grasshoppers Sphenarium purpurascens in Mexico for human consumption: a comparison with insecticidal control for managing pest outbreaks. Crop Protection, 27, 473–480.
    https://doi.org/10.1016/j.cropro.2007.08.001

    Cueva del Castillo, R. (2003) Body size and multiple copulations in a neotropical grasshopper with an extraordinary mate-guarding duration. Journal of Insect Behavior, 16, 503–522.
    https://doi.org/10.1023/A:1027303323242

    Cueva del Castillo, R. & Cano-Santana, Z. (2001) Variación de la coloración corporal de Sphenarium purpurascens, (Orthoptera: Pyrgomorphidae) en función del sexo y su relación con la formación de parejas en un ambiente heterogéneo. Folia Entomológica Mexicana, 40 (3), 297–309.

    Cueva del Castillo, R. & Núnez-Farfán, J. (1999) Sexual Selection on maturation time and body size in Sphenarium purpurascens (Orthoptera: Pyrgomorphidae): correlated response to selection. Evolution, 53 (1), 209–215.
    https://doi.org/10.2307/2640933

    Cueva del Castillo, R., Núñez-Farfán, J. & Cano-Santana, Z. (1999) The role of body size in mating success of Sphenarium purpurascens in Central Mexico. Ecological Entomology, 24 (2), 146–155.
    https://doi.org/10.1046/j.1365-2311.1999.00188.x

    Cueva del Castillo, R. & Núñez‐Farfán, J. (2002) Female mating success and risk of pre‐reproductive death in a protandrous grasshopper. Oikos, 96 (2), 217–224.
    https://doi.org/10.1034/j.1600-0706.2002.960203.x

    Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9 (8), 772–772.
    https://doi.org/10.1038/nmeth.2109

    Dayrat, B. (2005) Towards integrative taxonomy. Biological Journal of the Linnean Society, 85 (3), 407–415.

    Descamps, M. (1975) Etude du peuplement acridien de L’etat de Veracruz (Mexique). Folia Entomológica Mexicana, 31, 3–98.

    Descamps, M. & Wintrebert, D. (1966) Pyrgomorphidae et Acrididae de Madagascar : observations biologiques et diagnoses (Orth. Acridoidea). EOS: Revista Española de Entomología, 42, 41–263.

    Drummond, A.J. & Rambaut, A. (2015) Bayesian Evolutionary Analysis with BEAST. Cambridge University Press, Cambridge, 249 pp.

    Duennes, M.A., Lozier, J.D., Hines, H.M. & Cameron, S.A. (2012) Geographical patterns of genetic divergence in the widespread Mesoamerican bumble bee Bombus ephippiatus (Hymenoptera: Apidae). Molecular Phylogenetics and Evolution, 64, 219–231.
    https://doi.org/10.1016/j.ympev.2012.03.018

    Eberhard, W.G. (1985) Sexual selection and animal genitalia. Harvard University Press, Massachusetts, 244 pp.

    Edgar, R.C. (2004) MUSCLE: a multiple sequence alignment method with reduced time and space complexity. BMC bioinformatics, 5, 113.
    https://doi.org/10.1186/1471-2105-5-113

    Ferrari, L., Orozco-Esquivel, T., Manea, V. & Manea, M. (2012) The dynamic history of the Trans-Mexican Volcanic Belt and the Mexico subduction zone. Tectonophysics, 522–523, 122–149.
    https://doi.org/10.1016/j.tecto.2011.09.018

    Fujisawa, T. & Barraclough, T.G. (2013) Delimiting species using single-locus data and the generalized mixed yule coalescent approach: A revised method and evaluation on simulated data sets. Systematic Biology, 62 (5), 707–724.

    Gerstaecker, A. (1869) Beitrag zur Insekten-Fauna von Zanzibar No. 11 Orthoptera et Neuroptera. Archiv für Naturgeschichte, 35 (1), 201–223.
    https://doi.org/10.5962/bhl.part.12294

    Gerstaecker, A. (1873) Acridiodea nonnulla nova insigniora descropsit. Entomologische Zeitung, 34, 185–197.

    Hare, M.P. & Avise, J.C. (1998) Population structure in the American oyster as inferred by nuclear gene genealogies. Molecular Biology and Evolution, 15 (2), 119–128.
    https://doi.org/10.1093/oxfordjournals.molbev.a025908

    Hebard, M. (1932) New species and records of Mexican Orthoptera. Transactions of the American Entomological Society, 58 (3), 201–371.

    Hebert, P.D.N., Cywinska, A., Ball, S.L. & deWaard, J.R. (2003a) Biological identifications through DNA barcodes. Proceedings of the Royal Society B Biological sciences, 270 (1512), 313–321.
    https://doi.org/10.1098/rspb.2002.2218

    Hebert, P.D.N., Ratnasingham, S. & deWaard, J.R. (2003b) Barcoding animal life: cytochrome c oxidase subunit 1 divergences among closely related species. Proceedings of the Royal Society B Biological sciences, 270 (Supplement l), S96–S99.

    Heled, J. & Drummond, A.J. (2010) Bayesian inference of species trees from multilocus data. Molecular Biology and Evolution, 27 (3), 570–80.
    https://doi.org/10.1093/molbev/msp274

    Hosken, D.J. & Stockley, P. (2004) Sexual selection and genital evolution. Trends in Ecology & Evolution, 19 (2), 87–93.
    https://doi.org/10.1016/j.tree.2003.11.012

    Kevan, D.K.M. (1964) The Pyrgomorphidae (Orthoptera: Acridoidea): Their systematics, tribal divisions and distribution. Canadian Entomologist, 96 (12), 1505–1536.
    https://dx.doi.org/10.4039/Ent961505-12

    Kevan, D.K.M. (1977) The American Pyrgomorphidae (Orthoptera). Revista de la Sociedad Entomológica Argentina, 36 (1–4), 3–28.

    Kevan, D.K.M., Akbar, S.S. & Chang, Y.-C. (1969) The concealed copulatory structures of the Pyrgomorphidae (Orth. Acridoidea). Part I. General Introduction. EOS: Revista Española de Entomología, 44, 165–266.

    Kevan, D.K.M., Singh, A. & Akbar, S.S. (1964) A Revision of the Mexican Pyrgomorphidae (Orthoptera: Acridoidea) I. Genera Other than Sphenarium. Proceedings of the Academy of Natural Sciences of Philadelphia, 116 (6), 231–298.

    Kirby, W.F. (1910) A synonymic catalogue of Orthoptera. Vol. 1. Orthoptera euplexoptera, cursoria, et gressoria. Trustees of the British Museum, London, 501 pp.

    Knowles, L.L. & Carstens, B.C. (2007) Delimiting species without monophyletic gene trees. Systematic Biology, 56 (6), 887–895.
    https://dx.doi.org/10.1080/10635150701701091

    Knowles, L.L. & Kubatko, L.S. (2010) Estimating species trees: an introduction to concepts and models. In: Knowles, L.L. & Kubatko, L.S. (Eds.), Estimating Species Trees: Practical and Theoretical Aspects. Wiley-Blackwell, New Jersey, pp. 1–14.

    Kubatko, L.S. & Degnan, J.H. (2007) Inconsistency of phylogenetic estimates from concatenated data under coalescence. Systematic Biology, 56 (1), 17–24.
    https://dx.doi.org/10.1080/10635150601146041

    Leaché, A.D. & Mulcahy, D.G. (2007) Phylogeny, divergence times and species limits of spiny lizards (Sceloporus magister species group) in western North American deserts and Baja California. Molecular Ecology, 16 (24), 5216–5233.
    https://dx.doi.org/10.1111/j.1365-294X.2007.03556.x

    López, H., Contreras-Díaz, H.G., Oromí, P. & Juan, C. (2007) Delimiting species boundaries for endangered Canary Island grasshoppers based on DNA sequence data. Conservation Genetics, 8 (3), 587–598.
    https://dx.doi.org/10.1007/s10592-006-9199-5

    Márquez, C. (1962) Estudios de las especies del género Sphenarium basado en sus genitalia (Acrididae; Orthoptera), con la descripción de una nueva especie. Anales de Instituto de Biología UNAM Serie Zoología, 33, 247–258.

    Márquez, C. (1965a) Contribución al estudio de ortópteros de México, III. Estudios ecológicos preliminares de ortópteros del valle de Mezcala, Guerrero. Anales de Instituto de Biología UNAM Serie Zoología, 35, 87–93.

    Márquez, C. (1965b) Contribución al estudio de ortópteros de México, IV. Ortópteros del Pedregal de San Ángel, Villa Orbegón, DF. Anales de Instituto de Biología UNAM Serie Zoología, 39, 107–122.

    Mastretta-Yanes, A., Moreno-Letelier, A., Piñero, D., Jorgensen, T.H. & Emerson, B.C. (2015) Biodiversity in the Mexican highlands and the interaction of geology, geography and climate within the Trans-Mexican Volcanic Belt. Journal of Biogeography, 42 (9), 1586–1600.
    https://dx.doi.org/10.1111/jbi.12546

    McGuire, J.A., Linkem, C.W., Koo, M.S., Hutchison, D.W., Lappin, A.K., Orange, D.I., Lemos-Espinal, J., Riddle, B.R. & Jaeger, J.R. (2007) Mitochondrial introgression and incomplete lineage sorting through space and time: Phylogenetics of crotaphytid lizards. Evolution, 61 (12), 2879–2897.
    https://dx.doi.org/10.1111/j.1558-5646.2007.00239.x

    McKay, B.D. & Zink, R.M. (2010) The causes of mitochondrial DNA gene tree paraphyly in birds. Molecular Phylogenetics and Evolution, 54 (2), 647–650.
    https://dx.doi.org/10.1016/j.ympev.2009.08.024

    Mulcahy, D.G., Morrill, B.H. & Iii, J.R.M. (2006) Historical biogeography of lowland species of toads (Bufo) across the Trans-Mexican Neovolcanic Belt and the Isthmus of Tehuantepec. Journal of Biogeography, 33 (11), 1889–1904.
    https://dx.doi.org/10.1111/j.1365-2699.2006.01546.x

    Nater, A., Burri, R., Kawakami, T., Smeds, L. & Ellegren, H. (2015) Resolving evolutionary relationships in closely related species with whole-genome sequencing data. Systematic Biology, 64 (6), 1000–1017.
    https://dx.doi.org/10.1093/sysbio/syv045

    Ornelas, J.F. & González, C. (2014) Interglacial genetic diversification of Moussonia deppeana (Gesneriaceae), a hummingbird-pollinated, cloud forest shrub in northern Mesoamerica. Molecular Ecology, 23 (16), 4119–4136.
    https://dx.doi.org/10.1111/mec.12841

    Ornelas, J.F., Sosa, V., Soltis, D.E., Daza, J.M., González, C., Soltis, P.S., Gutiérrez-Rodríguez, C., de los Monteros, A.E., Castoe, T.A., Bell, C. & Ruiz-Sanchez, E. (2013) Comparative phylogeographic analyses illustrate the complex evolutionary history of threatened cloud forests of northern Mesoamerica. PloS ONE, 8 (2), 1–11.
    https://dx.doi.org/10.1371/journal.pone.0056283

    Oyama, K., Cano-Santana, Z. & Careaga, S. (1994) Estudios sobre la interacción herbívoro-planta en el Pedregal de San Ángel, México, D. F. In: Coordinación de la Investigación Científica UNAM (Ed.), Reserva ecológica “El Pedregal” de San Ángel: ecología, historia natural y manejo. Universidad Nacional Autónoma de México, Mexico City, pp. 301–311.

    Padial, J.J.M., Miralles, A., De la Riva, I. & Vences, M. (2010) The integrative future of taxonomy. Frontiers in Zoology, 7 (1), 1–16.
    https://dx.doi.org/10.1186/1742-9994-7-16

    Papadopoulou, A., Anastasiou, I. & Vogler, A.P. (2010) Revisiting the insect mitochondrial molecular clock: the mid-Aegean trench calibration. Molecular Biology and Evolution, 27 (7), 1659–1672.
    https://dx.doi.org/10.1093/molbev/msq051

    Pedraza-Lara, C., Barrientos-Lozano, L., Rocha-Sánchez, A.Y. & Zaldívar-Riverón, A. (2015) Montane and coastal species diversification in the economically important Mexican grasshopper genus Sphenarium (Orthoptera: Pyrgomorphidae). Molecular Phylogenetics and Evolution, 84, 220–231.
    https://dx.doi.org/10.1016/j.ympev.2015.01.001

    Pocco, M.E., Minutolo, C., Dinghi, P. a., Lange, C.E., Confalonieri, V. a. & Cigliano, M.M. (2015) Species delimitation in the Andean grasshopper genus Orotettix Ronderos & Carbonell (Orthoptera: Melanoplinae): an integrative approach combining morphological, molecular and biogeographical data. Zoological Journal of the Linnean Society, 174 (4), 733–759.
    https://dx.doi.org/10.1111/zoj.12251

    Pringle, E.G., Ram, S.R., Rez, I., Bonebrake, T.C., Ramírez, S.R., Bonebrake, T.C., Gordon, D.M. & Dirzo, R. (2012) Diversification and phylogeographic structure in widespread Azteca plant-ants from the northern Neotropics. Molecular Ecology, 21 (14), 3576–92.
    https://dx.doi.org/10.1111/j.1365-294X.2012.05618.x

    De Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56 (6), 879–886.
    https://dx.doi.org/10.1080/10635150701701083

    Rambaut, A. & Drummond, A.J. (2013) Tracer V1.6. Available from: http://tree.bio.ed.ac.uk/software/tracer (accessed 5 September 2016)

    Ramos-Elorduy, J. & Moreno, J.M.P. (1989) Los insectos comestibles en el México antiguo: estudio etnoentomológico. AGT, Mexico City, 108 pp.

    Rehn, J.A.G. & Hebard, M. (1912) Fixation of the single type (Lectotype) specimens of species of American Orthoptera. Proceedings of the Academy of Natural Sciences of Philadelphia, 64 (1), 60–183.

    Ronquist, F., Teslenko, M., Van Der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) Mrbayes 3.2: Efficient bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61 (3), 539–542.
    https://dx.doi.org/10.1093/sysbio/sys029

    Ruiz-Sanchez, E. & Specht, C.D. (2013) Influence of the geological history of the Trans-Mexican Volcanic Belt on the diversification of Nolina parviflora (Asparagaceae: Nolinoideae) P. Ladiges (Ed). Journal of Biogeography, 40 (7), 1336–1347.
    https://dx.doi.org/10.1111/jbi.12073

    Sanabria-Urbán, S., Song, H., Oyama, K., González-Rodríguez, A., Serrano-Meneses, M.A. & Cueva del Castillo, R. (2015) Body size adaptations to altitudinal climatic variation in neotropical grasshoppers of the genus Sphenarium (Orthoptera: Pyrgomorphidae). Plos ONE, 10 (12), 1–24.
    https://dx.doi.org/10.1371/journal.pone.0145248

    Saussure, H. (1859) Orthoptera nova americana (Diagnoses praeliminares). Revue et magasin de zoologie pure et appliquée, 2 (11), 390–394.

    Schlick-Steiner, B.C., Steiner, F.M., Seifert, B., Stauffer, C., Christian, E. & Crozier, R.H. (2010) Integrative taxonomy: a multisource approach to exploring biodiversity. Annual Review of Entomology, 55, 421–438.
    https://dx.doi.org/10.1146/annurev-ento-112408-085432

    SENASICA (2017) Chapulín (Plaga endémica). Available from: https://www.gob.mx/senasica/acciones-y-programas/chapulin-plaga-endemica (accessed 9 February 2017)

    Simmons, L.W. (2014) Sexual selection and genital evolution: sexual selection and genital evolution. Austral Entomology, 53 (1), 1–17.
    https://dx.doi.org/10.1111/aen.12053

    Song, H. (2009) Species-specificity of male genitalia is characterized by shape, size, and complexity. Insect Systematics & Evolution, 40 (2), 159–170.
    https://dx.doi.org/10.1163/187631209X424571

    Song, H., Buhay, J.E., Whiting, M.F. & Crandall, K.A. (2008) Many species in one: DNA barcoding overestimates the number of species when nuclear mitochondrial pseudogenes are coamplified. Proceedings of the National Academy of Sciences of the United States of America, 105 (36), 13486–13491.
    https://dx.doi.org/10.1073/pnas.0803076105

    Talavera, G., Lukhtanov, V.A., Rieppel, L., Pierce, N.E. & Vila, R. (2013) In the shadow of phylogenetic uncertainty: the recent diversification of Lysandra butterflies through chromosomal change. Molecular Phylogenetics and Evolution, 69 (3), 469–478.
    https://dx.doi.org/10.1016/j.ympev.2013.08.004

    Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular Biology and Evolution, 30 (12), 2725–2729.
    https://dx.doi.org/10.1093/molbev/mst197

    Uvarov, B.P. (1977) Grasshoppers and locusts: a handbook of general acridology. Vol.2. Centre for Overseas Pest Research, London, 613 pp.

    Walker, F. (1870) Catalogue of the specimens of Dermaptera Saltatoria and supplement of the Blattariæ in the collection of the British Museum. Part III. Trustees of the British Museum, London, 196 pp.

    Wickler, W. & Seibt, U. (1985) Reproductive behaviour in Zonocerus elegans (Orthoptera: Pyrgomorphidae) with special reference to nuptial gift guarding. Zeitschrift für Tierpsychologie, 69 (3), 203–223.
    https://dx.doi.org/10.1111/j.1439-0310.1985.tb00147.x

    Woller, D.A., Fontana, P., Mariño-Pérez, R. & Song, H. (2014) Studies in Mexican grasshoppers: Liladownsia fraile, a new genus and species of dactylotini (acrididae: Melanoplinae) and an updated molecular phylogeny of melanoplinae. Zootaxa, 3793 (4), 475–495.
    https://dx.doi.org/10.11646/zootaxa.3793.4.6