Skip to main content Skip to main navigation menu Skip to site footer
Type: Monograph
Published: 2022-11-22
Page range: 1-119
Abstract views: 533
PDF downloaded: 60

Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision

Instituto de Ecología, A.C., Red Interacciones Multitróficas, Carretera antigua a Coatepec 351, El Haya, 91070 Xalapa, Veracruz, Mexico. 2 Departamento de Biología Evolutiva, Facultad de Ciencias, Universidad Nacional Autónoma de México, Ciudad de México, Mexico
Instituto de Ecología, A.C., Red Interacciones Multitróficas, Carretera antigua a Coatepec 351, El Haya, 91070 Xalapa, Veracruz, Mexico.
Coleoptera Euligyrus Ligyrus Proculigyrus white grubs Pentodontini phylogeny nomenclature keys

Abstract

The phylogeny of Tomarus Erichson, 1847 (Coleoptera: Scarabaeidae: Dynastinae: Pentodontini) was explored by parsimony using morphological characters to approach its taxonomic and nomenclature problems. The genus was not supported as a monophyletic group, and the recovered lineages did not match at all any of the previously known classifications. Three different lineages and six monophyletic groups were recovered, which could be treated as independent genera. However, the classification proposed here is as conservative as possible concerning to the creation of many new names and generic taxa. Thus, four genera are now recognized: Euligyrus Casey, 1915 new status (2 species), Ligyrus Burmeister, 1847 revised status (18 species, including Oxygrylius Casey, 1915 new synonymy), Proculigyrus new genus (1 species), and Tomarus Erichson, 1847 revised status (18 species). Three subgenera are recognized for the genus Ligyrus: Anagrylius Casey, 1915 revised status (2 species), Ligyrodes Casey, 1915 revised status (3 species), and Ligyrus revised status (13 species). A complete taxonomic treatment is presented for each genus, including descriptions, keys, and occurrence maps. A revised status is presented for the following species: Ligyrus (Ligyrus) nasutus (Burmeister, 1847), T. amazonicus (Arrow, 1914), and T. gyas Erichson, 1848. Lectotypes are designated for Bothynus neglectus LeConte, 1847, L. amazonicus Arrow, 1914, L. bryanti Rivers, 1891, Scarabeus fossor Latreille, 1813, S. relictus Say, 1825 T. gyas, T. maimon Erichson, 1847, and some synonyms of L. sallaei Bates, 1888 (Ligyrodes propinquus Casey, 1915 and Ligyrodes aztecus Casey, 1915), S. gibbosus DeGeer, 1774 (Bothynus morio LeConte, 1847, L. californicus Casey, 1909, and L. spissipes Casey, 1909), S. relictus (Ligyrodes parviceps Casey, 1915 and Ligyrodes quadripennis Casey, 1915), and S. bituberculatus Palisot de Beauvois, 1811 (Ligyrus maximus Arrow, 1913 and Ligyrus latus Arrow, 1914). Ligyrus patagonus Steinheil, 1874 is a new synonym for L. rubripes Boheman, 1858. Type species are designated for the synonyms Grylius Casey, 1915 and Ligyrellus Casey, 1915. New combinations (new combination) are presented for: Euligyrus ebenus (De Geer, 1774), E. similis (Endrödi, 1968), Ligyrus (Anagrylius) moroni (López-García & Deloya, 2019), L. (Ligyrus) paranaensis (López-García & Deloya, 2019), L. (Ligyrus) peninsularis (Casey, 1915), L. (Ligyrus) spinipenis (Neita & Ratcliffe 2017), and Tomarus amazonicus. Ligyrus (Ligyrus) allonasutus new species is described to include populations of T. nasutus from western Mexico to Panama. Phylogenetic and taxonomic value of mouthparts, elytro-tergal stridulatory apparatus, and male genitalia is discussed along with a description of the biogeographic patterns of distribution for each genus.

 

References

  1. Ahrens, D. (2006) The phylogeny of Sericini and their position within the Scarabaeidae based on morphological characters (Coleoptera: Scarabaeidae). Systematics Entomology, 31, 113–144. https://doi.org/10.1111/j.1365-3113.2005.00307.x
    Ahrens, D., Schwarzer, J. & Vogler, A.P. (2014) The evolution of scarab beetles tracks the sequential rise of angiosperms and mammals. Proceedings of the Royal Society B: Biological Sciences, 281(1791), 20141470. https://doi.org/10.1098/rspb.2014.1470
    Albuquerque, L.S.C., Grossi, P.C. & Iannuzzi, L. (2016) Flight patterns and sex ratio of beetles of the subfamily Dynastinae (Coleoptera, Melolonthidae). Revista Brasileira de Entomologia, 60, 248–254. https://doi.org/10.1016/j.rbe.2016.03.002
    Arrow, G.J. (1904) Sound-production in the lamellicorn. Transactions of the Entomological Society of London, 1904, 709–751. https://doi.org/10.1111/j.1365-2311.1904.tb02761.x
    Arrow, G.J. (1913) Some new species of lamellicorn beetles from Brazil. The Annals and Magazine of Natural History, Series 11 (8), 456–466. https://doi.org/10.1080/00222931308693338
    Arrow, G.J. (1914) Some further notes on lamellicorn beetles of the subfamily Dynastinae. The Annals and Magazine of Natural History, Series 14 (81), 257–276. https://doi.org/10.1080/00222931408693571
    Ayquipa, G.E. & Cueva, M.A. (1979) Nombres científicos y comunes de los insectos que atacan a la caña de azúcar en el Perú. Revista Peruana de Entomología, 22, 95–97.
    Bates, H.W. (1888) Pectinicornia and Lamellicornia, Family Dynastinae. In: Godman, F.D. & Salvin, O. (Eds.), Biología Centrali-Americana, Insecta, Coleoptera, 2 (Part 2), pp. 296–342.
    Bitar, A. & Morón, M.A. (2014) Revisión y análisis filogenético del género Xyloryctes (Coleoptera: Melolonthidae: Dynastinae: Oryctini). Revista Mexicana de Biodiversidad, 85, 753–796.
    Blatchley, W.S. (1922) Some new and rare Coleoptera from southwestern Florida. The Canadian Entomologist, 54, 27–33. https://doi.org/10.4039/Ent5427-2
    Boheman, C.H. (1858) Coleoptera. Species novas descripsit. In: Kongliga svenska fregatten Eugenies resa omkring jorden under befäl af C.A. Virgin; åren 1851–1853 Vetenskapliga Iakttagelser. II. Zoologi. 1. Insecta. P.A. Norstedt & Söner, Stockholm, pp. 1–112.
    Bottrell, D.C., Brigham, R.D. & Jordan, L.B. (1973) Carrot beetle: pest status and bionomics on cultivated sunflower. Journal of Economic Entomology, 66, 86–90. https://doi.org/10.1093/jee/66.1.86
    Bremer, K. (1994) Branch support and tree stability. Cladistics, 10, 295–304. https://doi.org/10.1111/j.1096-0031.1994.tb00179.x
    Burmeister, H. (1847) Handbuch der Entomologie. Coleoptera Lamellicornia, Xylophila et Pectinicornia. Band 5. Enslin, Berlin, viii + 584 pp.
    Buss, E.A. (2006) Flight activity and relative abundance of phytophagous scarabs (Coleoptera: Scarabaeoidea) from two locations in Florida. Florida Entomologist, 89, 32–40. https://doi.org/10.1653/0015-4040(2006)89[32:FAARAO]2.0.CO;2
    Cano, E.B, Schuster, J.C. & Morrone, J.J. (2018) Phylogenetics of Ogyges Kaup and the biogeography of Nuclear Central America (Coleoptera, Passalidae). Zookeys, 737, 81–111. https://doi.org/10.3897/zookeys.737.20741
    Carrasco, F. (1967) Algunas plagas registradas en Cusco. Revista Peruana de Entomología, 10, 62–66.
    Cartwright, O.L. (1944) New Scarabaeidae from the United States. Annals of the Entomological Society of America, 37, 28–36. https://doi.org/10.1093/aesa/37.1.28
    Cartwright, O.L. (1959) Scarab beetles of the genus Bothynus in the United States (Coleoptera: Sacarabaeidae). Proceedings of the United States National Museum, 108, 515–541.
    Carvajal, V., Villamarín, C. & Ortega, A.M. (2011) Escarabajos del Ecuador. Principales géneros. Serie Entomología. No. 1. Instituto de Ciencias Biológicas, Escuela Politécnica Nacional, Quito, 350 pp.
    Casey, T.L. (1909) Studies in the Carabaeoidea and Lamellicornia. The Canadian Entomologist, 41, 253–284. https://doi.org/10.4039/Ent41253-8
    Casey, T.L. (1915) A review of the American species of Rutelinae, Dynastinae, and Cetoniinae. Memoirs on the Coleoptera, 6, 1–394.
    Casey, T.L. (1924) Additions to the known Coleoptera of North America. Memoirs on the Coleoptera, 11, 1–347.
    Chalumeau, F. & Gruner, L. (1977) Scarabaeoidea des Antilles françaises, 3me partie: Dynastinae et Cetoniinae. Annales de la Société Entomologique de France, 13, 579–612.
    Cherry, R., McCray, M. & Sandhu, H. (2017) Changes in the relative abundance of soil-dwelling insect pests in sugarcane grown in Florida. Journal of Entomological Science, 52, 169–176.
    Cisternas, A.E. (2013) Insectos plaga de importancia económica asociados al arándano. Boletín Instituto de Investigaciones Agropecuarias, 263, 91–105.
    Clark, D.R. (2011) Phylogenetic analysis of the scarab beetle tribe Cyclocephalini (Coleoptera: Scarabaeidae: Dynastinae) based on adult morphological characters. Masters Thesis, Wichita State University, Wichita, Kansasmerica. Available from: https://soar.wichita.edu/bitstream/handle/10057/5050/t11061_Clark.pdf?sequence=3&isAllowed=y (accessed 10 September 2022)
    Coddington, J.A., Agnarsson, I., Hamilton, C.A. & Bond, J.E. (2019) Spiders did not repeatedly gain, but repeatedly lost, foraging webs. PeerJ, 7, e6703. https://doi.org/10.7717/peerj.6703
    Couturier, G., Inga, H. & Tanchiva, E. (1996) Insectos plaga del arroz en Loreto, Amazonía peruana. Revista Peruana de Entomología, 39, 131–134.
    Davis, J.J. (1916) A progress report on white grub investigations. Journal of Economic Entomology, 9, 261–281.
    Daza, M. & Rincón, H. (1993) Perfil Tecnológico del Camote (Batata) en la Costa Central del Perú. Centro Internacional de la Papa, Lima, 38 pp.
    Dechambre, R.-P. & Lumaret, J.-P. (1985) Un Ligyrus nouveau (Coleoptera, Dynastidae). Description de l’imago, de la lame et indications ethologiques. Revue Francaise d’Entomologie, 7, 107–110.
    De Geer, C. (1774) Memories pour Servir à l´Historie des Insectes. Vol. 4. L.L. Grefing, Stockholm, 456 pp.
    Dellacasa, G., Dellacasa, M. & Mann, D.J. (2010) The morphology of the labrum (epipharynx, ikrioma and aboral surface) of adult Aphodiini (Coleoptera: Scarabaeoidea: Aphodiinae), and its implications for systematics. Insecta Mundi, 132, 1–21.
    De Queiroz, K. (2005) A unified species concept and its consequences for the future taxonomy. Proceedings of the California Academy of Science, 56 (18), 196–215.
    De Queiroz, K. (2007) Species concept and species delimitation. Systematic Biology, 56, 879–886.
    Dupuis, F. (2005) L’abdomen et les genitalia des femelles de coléoptères Scarabaeoidea (Insecta, Coleoptera). Zoosystema, 27, 733–823.
    Dupuis, F. (2014) Tomarus rostratus nouvelle espece du Pérou (Coleoptera, Dynastidae). Coleóptères, 20, 1–4.
    Dupuis, F. (2016) Tomarus maimon fossator Burmeister, 1847, nouvelle sous-spèce du bouclier Guyanais (Coleoptera, Dynastidae). Coléoptères, 22 (4), 30–37.
    Eberhard, W.G. (1977) Fighting behavior of male Golofa porteri beetles (Scarabeidae: Dynastinae). Psyche: A Journal of Entomology, 84, 292–298.
    Eberhard, W.G. (1985) Sexual Selection and Animal Genitalia. Harvard University Press, Princeton. New Jersey, 501 pp.
    Endrödi, S. (1968) Neue Arten der Pentodontini. (Col. Dynastinae). Folia Entomologica Hungarica, 21, 161–177.
    Endrödi, S. (1969) Monographie der Dynastinae. 4. Tribus: Pentodontini (Coleoptera, Lamellicornia). Entomologische Abhandlungen, 37, 1–145.
    Endrödi, S. (1970) Drei neue Dynastinen (Coleoptera, Melolonthidae) aus Amerika. Mitteilungen aus dem Museum für Naturkunde in Berlin. Zoologisches Museum und Institut für Spezielle Zoologie, Berlin, 46, 105–108. https://doi.org/10.1002/mmnz.19700460113
    Endrödi, S. (1985) The Dynastinae of the World. W. Junk, Dordrecht, 800 pp.
    Erichson, W.F. (1847) Conspectus insectorum coleopterorum quae in Republica Peruana observata sunt. Archiv für Naturgeschichte, 13, 67–185.
    Erichson, W.F. (1848) Die Insekten. In: Schomburg, R. (Ed.), Reisen in British-Guiana in den Jahren 1840–1844. Vol. 3. J.J. Weber, Leipzig, pp. 553–617.
    Escalona, H.E. & Joly, L.J. (2006) El género Ligyrus Burmeister, 1847 en Venezuela (Coleoptera: Scarabaeidae: Dynastinae: Pentodontini). Boletín Sociedad Entomológica Aragonesa, 39, 111–137.
    Fabricius, J.C. (1775) Systema entomologiae, sistens insectorvm classes, ordines, genera, species, adiectis synonymis, locis, descriptionibvs, observationibvs. Korte, Flensburg and Leipzig, [31] + 832 pp. https://doi.org/10.5962/bhl.title.36510
    Fabricius, J.C. (1792) Entomologia Systematica. Vol. 1. Part 2. C.G. Proft et Storch, Copenhagen, Denmark, 538 pp.
    Fabricius, J.C. (1801) Systema eleutheratorum secundum ordines, genera, species adiectis synonymis, locis, observationibus, descriptionibus. Tomus II. Bibliopoli Academici, Kiel, 687 pp.
    Fairmaire, L. (1892) Descriptions de quelques coléoptères argentins. Annales de la Societe Entomologique de Belgique, 36, 242–253.
    Felsenstein, J. (1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution, 39, 783–791. https://doi.org/10.2307/2408678
    Goloboff, P.A. & Catalano, S.A. (2016) TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics, 32, 774–786. https://doi.org/10.1111/cla.12160
    Gordon, R.D. & Anderson, D.M. (1981) The species of Scarabaeidae (Coleoptera) associated with sugarcane in south Florida. Florida Entomologist, 64, 119–138. https://doi.org/10.2307/3494604
    Gunter, N.L., Weir, T.A., Slipinksi, A., Bocak, L. & Cameron, S.L. (2016) If dung beetles (Scarabaeidae: Scarabaeinae) arose in association with dinosaurs, did they also suffer a mass co-extinction at the K-Pg boundary? PloS One, 11 (5), e0153570. https://doi.org/10.1371/journal.pone.0153570
    Gutiérrez, R. (1945) Contribuciones al conocimiento de los Scarabaeidae chilenos. Dynastinae Oryctini. Revista Chilena de Historia Natural, 48, 112–118.
    Halffter, G. (1964) La entomofauna americana, ideas acerca de su origen y distribución. Folia Entomológica Mexicana, 6, 1–108.
    Halffter, G. (1987) Biogeography of the montane entomofauna of Mexico and Central America. Annual Review of Entomology, 32, 95–114. https://doi.org/10.1146/annurev.en.32.010187.000523
    Halffter, G. & Morrone, J.J. (2017) An analytical review of Halffter’s Mexican transition zone, and its relevance for evolutionary biogeography, ecology and biogeographical regionalization. Zootaxa, 4226 (1), 001–1046. https://doi.org/10.11646/zootaxa.4226.1.1
    Hardy, A. (1964) Studies on Oxygrylius Casey (Coleoptera: Scarabaeidae: Dynastinae). The Coleopterists Bulletin, 18, 122–127.
    Hayes, W.P. (1925) A comparative study of the history of certain phytophagous scarabaeid beetles. Kansas Agricultural Experiment Station Technical Bulletin, 16, 1–146.
    Hayes, W.P. (1929) Morphology, taxonomy, and biology of larval Scarabaeoidea. Illinois Biological Monographs, 12 (2), 1–85.
    Hielkema, A.J. & Hielkema, M.A. (2019) An annotated checklist of the Scarabaeoidea (Insecta: Coleoptera) of the Guianas. Insecta Mundi, 732, 1–306.
    Hofreiter, M. & Stewart, J. (2009) Ecological change, range fluctuations and population dynamics during the Pleistocene. Current Biology, 19 (14), R584–R594. https://doi.org/10.1016/j.cub.2009.06.030
    International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature. 4th Edition. International Trust for Zoological Nomenclature, London, 306 pp.
    Jameson, M.L. (1998) Phylogenetic analysis of the subtribe Rutelina and revision of the Rutela generic groups (Coleoptera: Scarabaeidae: Rutelinae: Rutelini). Bulletin of the University of Nebraska State Museum, 14, 1–184. [dated 1997]
    Jameson, M.L., Micó, E. & Galante, E. (2007) Evolution and phylogeny of the scarab subtribe Anisopliina (Coleoptera: Scarabaeidae: Rutelinae: Anomalini). Systematic Entomology, 32, 429–449. https://doi.org/10.1111/j.1365-3113.2006.00380.x
    King, L. (2003) The evolution of insect mating structures through sexual selection. Florida Entomologists, 86 (2), 124–133. https://doi.org/10.1653/0015-4040(2003)086[0124:TEOIMS]2.0.CO;2
    Kostromytska, O.S. & Buss, E.A. (2008) Seasonal phenology and management of Tomarus subtropicus (Coleoptera: Scarabaeidae) in St. Augustine grass. Journal of Economic Entomology, 101, 1847–1855. https://doi.org/10.1603/0022-0493-101.6.1847
    Kostromytska, O.S. & Buss, E.A. (2011) Tomarus subtropicus (Coleoptera: Scarabaeidae) larval feeding habits. Florida Entomologist, 94, 164–172. https://doi.org/10.1653/024.094.0207
    Krell, F.T. (2000) The fossil record of Mesozoic and Tertiary Scarabaeoidea (Coleoptera: Polyphaga). Invertebrate Systematics, 14, 871–905.
    Kukalová-Peck, J. & Lawrence, J.F. (1993) Evolution of the hind wing in Coleoptera. The Canadian Entomologist, 125, 181–258.
    Lacordaire, J.T. (1855) Histoire naturelle des insectes, genera des coléopterés. Vol 3. Librairic Encyclopédique de Roret, Paris, France, 594 pp.
    Latreille, P.A. (1813) Insectes de l’Amérique équinoxiale recueillis pendant le voyage de Humboldt et Bonpland, Seconde partie. In: Smith, C.J. & Gide, E.C. (Eds.), Voyage de Humboldt et Bonpland, Deuxième Partie. Observations de Zoologie et d’Anatomie Comparée. Schoell et Dufour, Paris, pp. 1–64.
    LeConte, J.L. (1847) Fragmenta entomologica. Journal of the Academy of Natural Sciences of Philadelphia, Series 2, 1, 71–93.
    LeConte, J.L. (1856) Notice of three genera of Scarabaeidae found in the United States. Proceedings of the Academy of Natural Sciences of Philadelphia, 8, 19–25.
    López-García, M.M. & Deloya, C. (2018) Description of the female of Tomarus rostratus Dupuis and new synonyms and records for the genus Tomarus Erichson (Coleoptera: Scarabaeidae: Dynastinae). The Coleopterists Bulletin, 72 (2), 331–338. https://doi.org/10.1649/0010-065X-72.2.331
    López-García, M.M. & Deloya, C. (2019) Five new species of the dynastine genus Tomarus (Coleoptera: Scarabaeidae), with an illustrated key to species. The Coleopterists Bulletin, 73, 127–141. https://doi.org/10.1649/0010-065X-73.1.127
    López-García, M.M., Gasca-Álvarez, H.J. & Amat-García, G. (2014) A new species of Tomarus Erichson, 1847 (Coleoptera: Scarabaeoidea: Dynastinae: Pentodontini), with a key to the species in Colombia. Zootaxa, 3869 (5), 579–584. https://doi.org/10.11646/zootaxa.3869.5.8
    López-García, M.M., Gasca-Álvarez, H.J. & Amat-García, G. (2015) The scarab beetle tribe Pentodontini (Coleoptera: Scarabaeidae: Dynastinae) of Colombia: taxonomy, natural history, and distribution. Zootaxa, 4048 (4), 451–492. https://doi.org/10.11646/zootaxa.4048.4.1
    Maddison, W. & Maddison, D.R. (2015) Mesquite: A Modular System for Evolutionary Analyses. Version 3.01. Available from: https://mesquiteproject.org (accessed 30 August 2022)
    Montesanto, G. (2015) A fast GNU method to draw accurate scientific illustrations for taxonomy. Zookeys, 515, 191–206. https://doi.org/10.3897/zookeys.515.9459
    Moore, M.R., Beza-Beza, C.F., Wickell, D.A., Beck, J.M. & Jameson, M.L. (2015) Molecules, morphology and Mimeoma scarabs: evolutionary and taxonomic implications for a palm-associated scarab group. Systematic Entomology, 40, 891–900. https://doi.org/10.1111/syen.12139
    Morón, M.A. (2010) Observaciones sobre la reproducción y el ciclo de vida de Plusiotis costata Blanchard, 1851 (Coleoptera: Melolonthidae: Rutelinae). Acta Zoológica Mexicana, 26, 705–720.
    Morón, M.A. & Grossi, P.C. (2015) Revision of Philoscaptus Bréthes, 1919 and description of a new genus (Coleoptera, Melolonthidae, Dynastinae). Entomotropica, 30, 1–11.
    Neita-Moreno, J.C. (2011) Escarabajos (Coleoptera, Scarabaeoidea) del departamento de Chocó, Colombia. Revista Biodiversidad Neotropical, 1, 17–27.
    Neita-Moreno, J.C. & Ratcliffe, B.C. (2017) The genus Tomarus Erichson (Coleoptera: Scarabaeidae: Dynastinae: Pentodontini) in Argentina, Chile, and Uruguay. Insecta Mundi, 547, 1–36.
    Nel, A. & De Villiers, W.M. (1988) Mouthpart structure in adult scarab beetles (Coleoptera: Scarabaeoidea). Entomologia Generalis, 13, 95–114. https://doi.org/10.1127/entom.gen/13/1988/95
    Nel, A. & Scholtz, C.H. (1990) Comparative morphology of the mouthparts of adult Scarabaeoidea (Coleoptera). Entomology Memoir, Republic of South Africa, Department of Agricultural Development, 80, 1–84.
    Nixon, K.C. (2004) WinClada-Asado. Version 1.61. Computer software and documentation. Available from: www.diversityoflife.org/winclada (accessed 22 January 2019)
    Palisot de Beauvois, A.M.F.J. (1811) Insectes Recueillis en Afrique et em Amérique, dans les Royaumes d´Oware et de Benin, à Saint-Dominigue et dans les États-Unis, pendant les Anées 1786-1797. Livraison 7. Levrault, Schoell, et Cie, Paris, pp. 101–120, pls. 3c + 15.
    Pamplona A.M.S.R., García, M.V.B., Fonseca, C.R.V., Teles, B.R. & Andreazze, R. (1994) Nova praga da bananeira no Amazonas: Ligyrus similis Endrodi. Technical report. Empresa Brasileira de Pesquisa Agropecuária, Manaus, 4 pp. Available from: https://ainfo.cnptia.embrapa.br/digital/bitstream/item/60527/1/intrucoes-tecnicas-02-ano-19940001.pdf (accessed 10 September 2022)
    Paucar-Cabrera, A. & Moore, M.R. (2018) New tribal placement and review of Parapucaya Prell and Pucaya Ohaus (Coleoptera, Scarabaeidae, Dynastinae). Zookeys, 805, 127–158. https://doi.org/10.3897/zookeys.805.28524
    Perty, J.A.M. (1830) Delectus animalium articulatorum, quae in iterene per Brasilian annis MDCCCXVIIMDCCCXX jussu et auspiciis Maximiliani Josephi I. Bavariae Regis Augustissimi Peracto Collegerunt Dr. J. B. de Spix et Dr. C. F. Ph. De Martins. Fascicle 1. Impensis editoris, Munich, 60 pp.
    Piedrahíta, R., Pardo-Locarno, L.C. & Montoya-Lerma, J. (2007) Registro de Ligyrus ebenus De Geer (Coleoptera: Melolonthidae) en cultivos de papachina (Colocasia esculenta) en los departamentos de Chocó y Valle del Cauca. Boletín del Museo de Entomología de la Universidad del Valle, 8 (2), 30–31.
    Prell, H. (1937) Beiträge zur Kenntnis der Dynastiden. XV. Neue Arten und Rassen. Entomologische Zeitschrift, 51, 89–90.
    Puker, A., Rodrigues, S.R., Tiago, E.F., Ide, S. & Fuhrmann, J. (2011) Notes on biology and behavior of rhinoceros beetle Enema pan (Coleoptera: Scarabaeidae: Dynastinae). Annals of the Entomological Society of America, 104, 919–927. https://doi.org/10.1603/AN10197
    QGIS Development Team (2017) Quantum GIS geographic information system. Open Source Geospatial Foundation Project. Version 2.18.14. Available from: https://www.qgis.org/en/site/ (accessed 10 September 2022)
    Ramírez-Salinas, C., Morón, M.A. & Castro-Ramírez, A.E. (2004) Descripción de los estados inmaduros de tres especies de Anomala, Ancognatha y Ligyrus (Coleoptera: Melolonthidae: Rutelinae y Dynastinae) con observaciones de su biología. Acta Zoológica Mexicana, 20 (3), 67–82.
    Ramírez-Salinas, C., Pacheco-Flores, C., Castro-Ramírez, A.E. & Morón, M.A. (2010) Descripción de los estados inmaduros de Ligyrus nasutus (Burmeister, 1847) (Coleoptera: Melolonthidae: Dynastinae) con información sobre su biología. Acta Zoológica Mexicana, New Series, 26, 617–625.
    Ratcliffe, B.C. (2003) The dynastine scarab beetles of Costa Rica and Panama. Bulletin of the University of Nebraska State Museum, 16, 1–506.
    Ratcliffe, B.C. (2009) Xyloryctes (Coleoptera: Scarabaeidae: Dynastinae: Oryctini) in the United States. Qui es et ubi fuisti et quo vadis? Insecta Mundi, 100, 1–11.
    Ratcliffe, B.C. (2014) A new genus and species of Dynastinae (Coleoptera: Scarabaeidae) from the Galápagos Islands, Ecuador, other new species of Cyclocephalini, Pentodontini, and Phileurini from South America, and a revised key to the genera of New World Pentodontini. The Coleopterists Bulletin, 68, 663–1680. https://doi.org/10.1649/0010-065X-68.4.663
    Ratcliffe, B.C. & Cave, R.D. (2006) The dynastine scarab beetles of Honduras, Nicaragua, and El Salvador. Bulletin of the University of Nebraska State Museum, 21, 1–424.
    Ratcliffe, B.C. & Cave, R.D. (2010) The Dynastinae (Coleoptera: Scarabaeidae) of the Cayman Islands (West Indies), with descriptions of Tomarus adoceteus, new species (Pentodontini) and Caymania nitidissima, new genus and species (Phileurini). Insecta Mundi, 139, 1–15.
    Ratcliffe, B.C. & Cave, R.D. (2015) The dynastine scarab beetles of the West Indies (Coleoptera: Scarabaeidae: Dynastinae). Bulletin of the University of Nebraska State Museum, 28, 1–346.
    Ratcliffe, B.C. & Cave, R.D. (2017) The dynastine scarab beetles of the United States and Canada (Coleoptera: Scarabaeidae). Bulletin of the University of Nebraska State Museum, 30, 1–298.
    Ratcliffe, B.C. & Morón, M.A. (1997) Dynastinae. In: Morón, M.A., Ratcliffe, B.C. & Deloya, C. (Eds.), Atlas de los escarabajos de México. Coleoptera: Lamellicornia. Vol. I. Familia Melolonthidae. Subfamilias Rutelinae, Dynastinae, Cetoniinae, Trichinae, Valginae y Melolonthinae. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad
    (CONABIO), Mexico City, pp. 53–98.
    Ratcliffe, B.C., Smith, D.M. & Erwin, D. (2005) Oryctoantiquus borealis, new genus and species from the Eocene of Oregon, USA, the world’s oldest fossil dynastine and largest fossil scarabaeid (Coleoptera: Scarabaeidae: Dynastinae). The Coleopterists Bulletin, 59, 127–136. https://doi.org/10.1649/0010-065X(2005)059[0127:OBNGAS]2.0.CO;2
    Ratcliffe, B.C., Cave, R.D. & Cano, E.B. (2013) The dynastine scarab beetles of Mexico, Guatemala, and Belize (Coleoptera: Scarabaeidae: Dynastinae). Bulletin of the University of Nebraska State Museum, 27, 1–666.
    Ritcher, P.O. (1958) Biology of Scarabaeidae. Annual Review of Entomology, 3, 311–334.
    Ritcher, P.O. (1966) White Grubs and their Allies. Oregon State University Press, Corvallis, Oregon, 219 pp.
    Rivers, J.J. (1891) New species of Scarabaeidae. Proceedings of the California Academy of Sciences, 3, 97–98.
    Rosen, D.E. (1975) A vicariance model of Caribbean biogeography. Systematic Biology, 24, 431–464. https://doi.org/10.2307/2412905
    Say, T. (1825) Descriptions of new species of coleopterous insects inhabiting the United States. Journal of the Academy of Natural Sciences of Philadelphia, 5, 160–204.
    Sereno, P.C. (2007) Logical basis for morphological characters in phylogenetics. Cladistics, 23, 565–587. https://doi.org/10.1111/j.1096-0031.2007.00161.x
    Solier, A.J.J. (1851) Orden III. Coleopteros. In: Gay, C. (Ed.), Historia Física y Política de Chile. Zoología. Vol. 5. C. Gay, Paris, pp. 5–285
    Squire, F.A. (1972) Entomological problems in Bolivia. Pest Articles & News Summaries, 18, 249–268. https://doi.org/10.1080/09670877209411802
    Steinheil, E. (1874) Symbolae ad historiam Coleopterorum Argentinae meridionalis, ossia enumerazione dei coleotteri raccolti dal Prof. Strobel, durante il suo soggiorno in Buenos Aires e nei viaggi de la intrapresi a Mendoza e nal Chili, indi a Bahia Blanca ed al Carmen de los Patagones; e descrizione delle specie nuove. Atti della Società Italiana di Scienze Naturali, 15, 554–578. [1872]
    Strong, E.E. & Lipscomb, D. (1999) Character coding and inapplicable data. Cladistics, 15, 363–371. https://doi.org/10.1111/j.1096-0031.1999.tb00272.x
    Tarasov, S. & Génier, F. (2015) Innovative Bayesian and parsimony phylogeny of dung beetles (Coleoptera, Scarabaeidae, Scarabaeinae) enhanced by ontology-based partitioning of morphological characters. PLoS One, 10 (3), e0116671. https://doi.org/10.1371/journal.pone.0116671
    Tarasov, S.I. & Solodovnikov, A.Y. (2011) Phylogenetic analyses reveal reliable morphological markers to classify mega-diversity in Onthophagini dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae). Cladistics, 27, 490–528. https://doi.org/10.1111/j.1096-0031.2011.00351.x
    Walsh, H.E., Kidd, M.G., Moum, T. & Friesen, V.L. (1999) Polytomies and the power of phylogenetic inference. Evolution, 53, 932–937. https://doi.org/10.1111/j.1558-5646.1999.tb05386.x
    Wheeler, Q.D. & Platnick, N.I. (2000) The phylogenetic species concept. In: Wheeler, Q.D. & Meier, R. (Eds.), Species Concepts and Phylogenetic Theory. Columbia University Press, New York, New York, pp. 55–69.
    Wickham, H.F. (1911) Fossil Coleoptera from Florissant, with descriptions of several new species. Bulletin of the American Museum of Natural History, 30, 53–69.
    Wickham, H.F. (1914) New Miocene Coleoptera from Florissant. Bulletin of the Museum of Comparative Zoology, 58, 421–494.
    Wiens, J.J., Chippindale, P.T. & Hillis, D.M. (2003) When are phylogenetic analyses misled by convergence? A case study in Texas cave salamanders. Systematic Biology, 52, 501–514.
    Wiley, E.O. (1981) Phylogenetics: The Theory and Practice of Phylogenetic Systematics. Wiley, New York, New York, 439 pp.
    Zunino, M. (1972) Revisione delle specie paleartiche del genere Onthophagus Latr. (Coleoptera, Scarabeoidea) I.II sottogenere Euonthophagus Balth. Bollettino del Museo di Zoologia dell’Università di Torino, 1, 1–28.
    Zunino, M. (2014) About dung beetle (Coleoptera: Scarabaeoidea) genitalia: some remarks to a recent paper. Acta Zoológica Mexicana, New Series, 30, 439–443.
    Zunino, M. & Halffter, G. (1988) Análisis taxonómico, ecológico y biogeográfico de un grupo americano de Onthophagus (Coleoptera: Scarabaeidae). Monografie del Museo Regionale di Scienze Naturali, Torino, 9, 1–211.