Published March 17, 2025
| Version v1
Journal article
Restricted
A review of the species groups of the Western Hemisphere Onthophagus Latreille (Coleoptera: Scarabaeidae: Scarabaeinae) using COI barcoding and gene trees
Creators
- 1. BioAlfa Barcoding Project Santo Domingo de Heredia, Costa Rica.
Description
Kohlmann, Bert, Solís, Ángel (2025): A review of the species groups of the Western Hemisphere Onthophagus Latreille (Coleoptera: Scarabaeidae: Scarabaeinae) using COI barcoding and gene trees. Zootaxa 5604 (4): 401-447, DOI: 10.11646/zootaxa.5604.4.1, URL: https://doi.org/10.11646/zootaxa.5604.4.1
Files
Linked records
Additional details
Identifiers
- URL
- https://www.checklistbank.org/dataset/309262
- LSID
- urn:lsid:zoobank.org:pub:6A95109D-6F33-4DE7-9D47-6A722DD26918
- LSID
- urn:lsid:plazi.org:pub:FF935B1A1463FF96FFEFAE2EFF91AE3E
- URL
- http://publication.plazi.org/id/FF935B1A1463FF96FFEFAE2EFF91AE3E
References
- 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 of London B: Biological Sciences, 281 (20141470), 1-10. https://doi.org/10.1098/rspb.2014.1470
- Alfaro, A., Gazel, E., White, W.M., Jicha, B. & Rasbury T. (2021) Unravelling the genesis of young continental arc shoshonites in the Talamanca Cordillera, Costa Rica. Lithos, 386-387, 106017. https://doi.org/10.1016/j.lithos.2021.106017
- Almquist, D. (2017) Surveying for gopher tortoise obligate invertebrate commensals. The Tortoise Burrow, 37 (2), 8-13.
- Arriaga-Jimenez, A. (2015) Distribution des coleopteres coprophages (Scarabaeinae, Geotrupinae et Aphodiinae) dans les hautes montagnes de la Zone de Transition Mexicaine: analyse ecologique et biogeographique. Biologie animale. PhD Thesis, Universite Paul Valery - Montpellier III, Montpellier, 292 pp. Available from: https://theses.hal.science/tel- 01231005/file/2015_ARRIAGAJIMENEZ_arch.pdf (accessed 19 November 2024)
- Arriaga-Jimenez, A., Escobar-Hernandez, F., Ros, M. & Kohlmann, B. (2020) Establishment of the Onthophagus anthracinus (Coleoptera: Scarabaeidae: Scarabaeinae) species complex, with a description of a new species. The Canadian Entomologist, 152, 1-17. https://doi.org/10.4039/tce.2019.62
- Arrow, G.J. (1931) The Fauna of British India, including Ceylon and Burma. Coleoptera Lamellicornia. Part III (Coprinae). Taylor & Francis, London, 428 pp.
- Axelrod, D.I. (1975) Evolution and biogeography of Madrean-Tethyan sclerophyll vegetation. Evolution, 62, 280-334. https://doi.org/10.2307/2395199
- Axelrod, D.I. (1979) Age and origin of Sonoran Desert vegetation. Occasional Papers of the California Academy of Sciences, 132, 1-74.
- Azuma, H., Garcia-Franco, J.-G., Rico-Gray, V. & Thien, L.-B. (2001) Molecular phylogeny of the Magnoliaceae: the biogeography of tropical and temperate disjunctions. American Journal of Botany, 88, 2275-2285. https://doi.org/10.2307/3558389
- Barrantes, G. (2009) The role of historical and local factors in determining species composition of the highland avifauna of Costa Rica and Western Panama. Revista de Biologia Tropical, 57, 333-349.
- Bates, H.W. (1887) Biologia Centrali-Americana. Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. Taylor and Francis, London, 432 pp.
- Bergsten, J., Bilton, D.T., Fujisawa, T., Elliott, M.T., Monaghan, M., Balke, L., Hendrich, L., Geijer, J., Herrmann, J., Foster, G. N., Ribera, I., Nilsson, A.N., Barraclough, T.G. & Vogler, A.P. (2012) The effect of geographical scale of sampling on DNA barcoding. Systematic Biology, 61, 851-870. https://doi.org/10.1093/sysbio/sys037
- Beutel, R.G., Xu, C., Jarzembowski, E., Kundrata, R., Boudinot, B.R., McKenna, D.D. & Goczal, J. (2024) The evolutionary history of Coleoptera (Insecta) in the late Palaeozoic and the Mesozoic. Systematic Entomology, 49, 355-388. https://doi.org/10.1111/syen.12623
- Beza-Beza, C.F., Jimenez-Ferbans, L. & McKenna, D.D. (2021) Historical biogeography of American passalid beetles (Coleoptera, Passalidae) reveals Mesoamerican tropical forests as a centre of origin and taxonomic diversification. Journal of Biogeography, 48, 2037-2052. https://doi.org/10.1111/jbi.14134
- Blanco, G., Laiolo, P. & Fargallo, J.A. (2014) Linking environmental stress, feeding-shifts and the 'island syndrome': a nutritional challenge hypothesis. Population Ecology, 56, 203-216. https://doi.org/10.1007/s10144-013-0404-3
- Boucomont, A. (1932) Synopsis des Onthophagus d'Amerique du Sud (Col. Scarab.). Annales de la Societe Entomologique de France, 101, 293-332. https://doi.org/10.1080/21686351.1932.12280217
- Breeschoten, T., Doorenweerd, C., Tarasov, S. & Vogler, A.P. (2016) Phylogenetics and biogeography of the dung beetle genus Onthophagus inferred from mitochondrial genomes. Molecular Phylogenetics and Evolution, 105, 86-95. https://doi.org/10.1016/j.ympev.2016.08.016
- Brower, A.V.Z. & Schuh, R.T. (2021) Biological Systematics. Principles and Applications. Comstock Publishing Associates, Ithaca, New York, 436 pp. https://doi.org/10.7591/cornell/9781501752773.001.0001
- Brown, W.J. (1925) Notes on the Oklahoma species of Onthophagus. Proceedings of the Oklahoma Academy of Sciences, 5, 99-101.
- Brown, W.J. (1927) Four new species of Onthophagus (Coleoptera). The Canadian Entomologist, 59, 128-133. https://doi.org/10.4039/Ent59128-6
- Brown, W.J. (1929) Studies in the Scarabaeidae (II). The Canadian Entomologist, 61, 86-93. https://doi.org/10.4039/Ent6186-4
- Chame-Vazquez, E.R. & Sanchez-Hernandez, G. (2022) Onthophagus tacanensis (Coleoptera: Scarabaeidae: Scarabaeinae), a new species from Chiapas, Mexico. Zootaxa, 5182 (2), 196-200. https://doi.org/10.11646/zootaxa.5182.2.6
- Chamorro, W., Marin-Armijos, D., Asenjo, A. & Vaz-De-Mello, F.Z. (2019) Scarabaeinae dung beetles from Ecuador: a catalog, nomenclatural acts, and distribution records. Zookeys, 826, 1-343. https://doi.org/10.3897/zookeys.826.26488
- Chin, K. & Gill, B. (1996) Dinosaurs, dung beetles, and conifers: participants in a Cretaceous food web. Palaios, 11, 280-285. https://doi.org/10.2307/3515235
- Coates, A.G., Collins, L.S., Aubry, M.-P. & Berggren, W.A. (2004) The geology of the Darien, Panama, and the late Miocene - Pliocene collision of the Panama arc with northwestern South America. Geological Society of America Bulletin, 116, 1327-1344. https://doi.org/10.1130/B25275.1
- Cognato, A.I., Sari, G., Smith, S.M., Beaver, R.A., Li, Y., Hulcr, J., Jordal, B.H., Kajimura, H., Lin, C.-S., Pham, T.H., Singh, S. & Sittichaya, W. (2020) The essential role of taxonomic expertise in the creation of DNA databases for the identification and delimitation of Southeast Asian ambrosia beetle species (Curculionidae: Scolytinae: Xyleborini). Frontiers in Ecology and Evolution, 8 (27), 1-17. https://doi.org/10.3389/fevo.2020.00027
- Colinvaux, P. (1997) The history of forests on the Isthmus from the Ice Age to the Present. In: Coates, A.G. (Ed.), Central America. A Natural and Cultural History. Yale University Press, New Haven, Connecticut, pp. 123-136. https://doi.org/10.2307/j.ctt1xp3sdf.11
- Cupello, M., Silva, F.A.B. & Vaz-de-Mello, F.Z. (2023) The taxonomic revolution of American dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae). Frontiers in Ecology and Evolution, 11 (1168754), 1-42. https://doi.org/10.3389/fevo.2023.1168754
- Delgado, L. & Deloya, C. (1990) Una nueva especie mexicana de Onthophagus Latreille, 1802, del "grupo clypeatus" (Coleoptera: Scarabaeidae: Scarabaeinae). Annales de la Societe entomologique Francaise, 26, 211-216. https://doi.org/10.1080/21686351.1990.12277659
- Delgado, L. & Mora-Aguilar, E.F. (2019) A new Mexican species of Onthophagus Latreille (Coleoptera: Scarabaeidae: Scarabaeinae), with a revised key to the species of the O. dicranius species complex. Zootaxa, 4695 (6), 586-592. https://doi.org/10.11646/zootaxa.4695.6.9
- Delgado, L., Peraza, L.N. & Deloya, C. (2006) Onthophagus yucatanus, a new species of the clypeatus group from Mexico and Guatemala (Coleoptera: Scarabaeidae). Florida Entomologist, 89, 6-9. https://doi.org/10.1653/0015-4040(2006)89[6:OYANSO]2.0.CO;2
- DeSalle, R. & Goldstein, P. (2019) Review and interpretation of trends in DNA barcoding. Frontiers in Ecology and Evolution, 7 (302), 1-11. https://doi.org/10.3389/fevo.2019.00302
- d'Orbigny, H. (1913) Synopsis des Onthophagides d'Afrique. Annales de la Societe Entomologique de France, 82, 1-742. https://doi.org/10.1080/21686351.1913.12279402
- Eberhard, W.G. (2009) Postcopulatory sexual selection: Darwin's omission and its consequences. Proceedings of the National Academy of Sciences, 106, 10025-10032. https://doi.org/10.1073/pnas.0901217106
- Emlen, D.J., Marangelo, J., Ball, B. & Cunningham, C.W. (2005) Diversity in the weapons of sexual selection: horn evolution in the beetle genus Onthophagus (Coleoptera: Scarabaeidae). Evolution, 59, 1060-1084. https://doi.org/10.1111/j.0014-3820.2005.tb01044.x
- Foster, J.B. (1964) The evolution of mammals on islands. Nature, 202, 234-235. https://doi.org/10.1038/202234a0
- Funk, J., Mann, P. & McIntosh, K. (2009) Cenozoic tectonics of the Nicaraguan depression, Nicaragua, and Median Trough, El Salvador, based on seismic-reflection profiling and remote-sensing data. Geological Society of America Bulletin, 121, 1491-1521. https://doi.org/10.1130/B26428.1
- Gavriilidi, I., De Meester, G., Van Damme, R. & Baeckens, S. (2022) How to behave when marooned: the behavioural component of the island syndrome remains underexplored. Biology Letters, 18 (20220030), 1-23. https://doi.org/10.1098/rsbl.2022.0030
- Ge, Y., Xia, C., Wang, J., Zhang, X., Ma, X. & Zhou, Q. (2021) The efficacy of DNA barcoding in the classification, genetic differentiation, and biodiversity assessment of benthic macroinvertebrates. Ecology and Evolution, 11, 5669-5681. https://doi.org/10.1002/ece3.7470
- Genier, F. (2017) A new Guatemalan cloud forest endemic Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae). The Canadian Entomologist, 149, 574-580. https://doi.org/10.4039/tce.2017.32
- Genier, F. & Howden, H.F. (1999) Two new central American Onthophagus Latreille of the mirabilis species group (Coleoptera: Scarabaeidae, Scarabaeinae). The Coleopterists Bulletin, 53, 130-144.
- Genier, F. & Howden, H.F. (2014) Onthophagus fragosus n. sp. a second endemic species of Onthophagus Latreille from Cuba (Coleoptera : Scarabaeidae, Scarabaeinae). Zootaxa, 3795 (5), 597-599. https://doi.org/10.11646/zootaxa.3795.5.10
- Gillet, C.P.D.T. & Toussaint, E.F.A. (2020) Macroevolution and shifts in the breeding biology of the American scarab beetle tribe Phanaeini (Coleoptera: Scarabaeidae: Scarabaeinae). Biological Journal of the Linnean Society, 20, 1-22.
- Gunter, N.L., Weir, T.A., Slipinksi, A., Bocka, L. & Cameron, 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 (e0153570), 1-47. https://doi.org/10.1371/journal.pone.0153570
- Gunter, N.L., Monteith, G.B., Cameron, S.L. & Weir, T.A. (2019) Evidence from Australian mesic zone dung beetles support their Gondwanan origin and Mesozoic diversification of the Scarabaeinae. Insect Systematics & Evolution, 50, 162-188. https://doi.org/10.1163/1876312X-00002171
- Halffter, G. (1974) Elements ancienes de l'entomofaune Neotropicale: Ses implications biogeographiques. Quaestiones Entomologicae, 10, 231-253.
- Halffter, G. & Halffter, V. (2009) Why and where coprophagous beetles (Coleoptera: Scarabaeinae) eat seeds, fruit, or vegetable detritus. Boletin Sociedad Entomologica Aragonesa, 45, 1-22.
- Halffter, G., Zunino, M., Moctezuma, V. & Sanchez-Huerta, J.-L. (2019) The integration processes of the distributional patterns in the Mexican Transition Zone: phyletic, paleogeographic and ecological factors of a case study. Zootaxa, 4586 (1), 10-34. https://doi.org/10.11646/zootaxa.4586.1.1
- Han, T., Lee, W., Lee, S., Park, I.G. & Park, H. (2016) Reassessment of species diversity of the subfamily Denticollinae (Coleoptera: Elateridae) through DNA barcoding. PloS One, 11 (e0148602), 1-28. https://doi.org/10.1371/journal.pone.0148602
- Hebert, P.D.N., Ratnasingham, S. & de Waard, J.R. (2003) Barcoding animal life: cytochrome c oxidase subunit 1 divergence among closely related species. Proceedings of the Royal Society of London, Series B, Biological Sciences, 270, S596- S599. https://doi.org/10.1098/rsbl.2003.0025
- Hebert, P.D.N., Stoeckle, M.A., Zemlak, T.S. & Francis, C.M. (2004) Identification of birds through DNA barcodes. Public Library of Science Biology, 2, 1657-1663. https://doi.org/10.1371/journal.pbio.0020312
- Howden, H.F. (1973) Four new species of Onthophagus from Mexico and the United States (Coleoptera: Scarabaeidae). Proceedings of the Entomological Society of Washington, 75, 329-337.
- Howden, H.F. & Cartwright, O.L. (1963) Scarab beetles of the genus Onthophagus Latreille north of Mexico (Coleoptera: Scarabaeidae). Proceedings of the United States National Museum, 114, 1-148. https://doi.org/10.5479/si.00963801.114-3467.1
- Howden, H.F. & Gill, B.D. (1993) Mesoamerican Onthophagus Latreille in the dicranius and mirabilis species groups (Coleoptera: Scarabaeidae). The Canadian Entomologist, 125, 1091-1114. https://doi.org/10.4039/Ent1251091-6
- Howden, H.F. & Young, O.P. (1981) Panamanian Scarabaeidae: taxonomy, distribution, and habits (Coleoptera, Scarabaeidae). Contributions of the American Entomological Institute, 18, 1-204.
- Howden, H.F., Cartwright, O.L. & Halffter, G. (1956) Descripcion de una nueva especie mexicana de Onthophagus con anotaciones ecologicas sobre especies asociadas a nidos de animales y a cuevas. Acta Zoologica Mexicana, 1, 1-16.
- International Commission on Zoological Nomenclature (2020) Opinion 2457 (Case 3722) - Scarabaeus gazella Fabricius, 1787 (currently Digitonthophagus gazella or Onthophagus gazella; Insecta, Coleoptera, Scarabaeidae): neotype designated. The Bulletin of Zoological Nomenclature, 77, 95-96. https://doi.org/10.21805/bzn.v77.a031
- Jeannel, R. (1942) La Genese des Faunes Terrestres. Presses Universitaires de France, Paris, 513 pp.
- Kappelle, M. (2016) Costa Rica´s Ecosystems: Setting the Stage. In: Kappelle, M. (Ed.), Costa Rican Ecosystems. The University of Chicago Press, Chicago, Illinois, pp. 3-16. https://doi.org/10.7208/chicago/9780226121642.001.0001
- Kijimoto, T., Pespeni, M., Beckers, O. & Moczek,A.P. (2012) Beetle horns and horned beetles: emerging models in developmental evolution and ecology. Wiley Interdisciplinary Review in Developmental Biology, 4, 405-418. https://doi.org/10.1002/wdev.81
- Kimura, M.A. (1980) A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution, 16, 111-120. https://doi.org/10.1007/BF01731581
- Kohlmann, B. (2022) Escarabajos estercoleros. In: Comision Nacional para el Conocimiento y Uso de la Biodiversidad (Ed.), La Biodiversidad en Oaxaca. Volumen III. Comision Nacional para el Conocimiento y Uso de la Biodiversidad, Mexico City, pp. 60-72.
- Kohlmann, B. & Solis, A. (2001) El genero Onthophagus (Coleoptera: Scarabaeidae) en Costa Rica. Giornale Italiano di Entomologia, 9, 159-261.
- Kohlmann, B. & Solis, A. (2004) El genero Canthidium (Coleoptera: Scarabaeidae) en Costa Rica. Giornale Italiano di Entomologia, 11, 1 -73.
- Kohlmann, B. & Wilkinson, J. (2007) The Tarcoles Line: biogeographic effects of the Talamanca Range in Lower Central America. Giornale Italiano di Entomologia, 12, 1-30.
- Kohlmann, B., Solis, A. & Alvarado, G.E. (2019) Description of Onthophagus humboldti and Uroxys bonplandi, two new scarab beetles (Coleoptera, Scarabaeidae, Scarabaeinae) from Costa Rica, with notes on tropical mountain brachyptery and endemicity. ZooKeys, 881, 23-51. https://doi.org/10.3897/zookeys.881.38026
- Kohlmann, B., Salomao, R.P. & Solis, A. (2024) New world dung-beetle (Scarabaeidae: Scarabaeinae) colonization of a recent Miocene insular territory: the case of Costa Rica. Ecology and Evolution, 14 (e11436), 1-17. https://doi.org/10.1002/ece3.11436
- Krell, F.-T. & Moon, A.R. (2019) Dung beetles. Current Biology, 19, R551-R567. https://doi.org/10.1016/j.cub.2019.05.027
- Lavin, M. & Luckow, M. (1993) Origins and relationships of tropical North America in the context of the Boreotropics Hypothesis. American Journal of Botany, 80, 1-14. https://doi.org/10.1002/j.1537-2197.1993.tb13761.x
- Lopes, F., Rossini, M., Losacco, F., Montanaro, G., Gunter, N. & Tarasov, S. (2023) Metagenomics reveals that dung beetles (Coleoptera: Scarabaeinae) broadly feed on reptile dung. Did they also feed on that of dinosaurs? Frontiers in Ecology and Evolution, 11 (1132729), 1-8. https://doi.org/10.3389/fevo.2023.1132729
- Ma, Z., Liu, J., Shen, Y., Fan, Z., Yue, B. & Zhang, X. (2019) Population genetic structure and intraspecific genetic distance of Periplaneta americana (Blattodea: Blattidae) based on mitochondrial and nuclear DNA markers. Ecology and Evolution, 19, 12928-12939. https://doi.org/10.1002/ece3.5777
- Ma, Z., Ren, J. & Zhang, R. (2022) Identifying the genetic distance threshold for Entiminae (Coleoptera: Curcilionidae) species delimitation via COI barcodes. Insects, 13 (261), 1-12. https://doi.org/10.3390/insects13030261
- MacGaughran, A., Hogg, I.D., Stevens, M.I., Chadderton, W.L. & Winterbourn, M.J. (2006) Genetic divergence of three freshwater isopod species from southern New Zealand. Journal of Biogeography, 33, 23-30. https://doi.org/10.1111/j.1365-2699.2005.01338.x
- MacVean, C. & Schuster, J.C. (1981) Altitudinal distribution of passalid beetles (Coleoptera, Passalidae) and Pleistocene dispersal in the volcanic chain of Northern Central America. Biotropica, 13, 29-38. https://doi.org/10.2307/2387868
- Matthews, E. (1966) A taxonomic and zoogeographic survey of the Scarabaeinae of the Antilles (Coleoptera: Scarabaeidae). Memoirs of the American Entomological Society, 21, 1-134.
- Matthews E.G. (1972) A revision of the scarabaeine dung beetles of Australia. I. Tribe Onthophagini. Australian Journal of Zoology, Supplementary Series, 9, 1-330. https://doi.org/10.1071/AJZS009
- Meng, H.-H., Jacques, F.M.B., Su, T., Huang, Y.-J., Zhang, S.-T., Ma, H.-J. & Zhou, Z.-K. (2014) New biogeographic insight into Bauhinia s.l. (Leguminosae): integration from fossil records and molecular analyses. BMC Evolutionary Biology, 14 (181), 1-14. https://doi.org/10.1186/s12862-014-0181-4
- Miller, M.A., Pfeiffer, W. & Schwarz, T. (2010) Creating the CIPRES science gateway for the inference of large phylogenetic trees. Available from: https://doi.org/10.1109/GCE.2010.5676129 (accessed 19 November 2024)
- Moctezuma, V. & Halffter, G. (2019) New biogeographical makeup for colonisation of the Baja California Peninsula, with the description of a new Onthophagus (Coleoptera: Scarabaeidae: Scarabaeinae). Journal of Natural History, 53, 2057-2071. https://doi.org/10.1080/00222933.2019.1685694
- Moctezuma. V. & Halffter, G. (2021) Species redescriptions and new species of the Onthophagus mexicanus species group (Coleoptera: Scarabaeidae), with notes on distribution and rodent-dung beetle associations. Zoological Studies, 60, 1-30.
- Moctezuma, V., Hernandez, B., Sanchez-Huerta, J.L., Lizardo, V., Quiroz-Rocha, G.A. & Navarrte-Heredia, J.L. (2022) Two new species from the Mexican Pacific Slope and new distributional records of Onthophagus Latreille, 1802 (Coleoptera, Scarabaeidae). Journal of Asia-Pacific Entomology, 26 (102015), 1-6. https://doi.org/10.1016/j.aspen.2022.102015
- Moctezuma, V., Halffter, G. & Mora-Aguilar, E. (2023) A new species of the Onthophagus mirabilis species complex (Coleoptera: Scarabaeidae: Scarabaeinae) from the Chimalapas region, Oaxaca, Mexico. Revista Mexicana de Biodiversidad, 94 (e945081), 1-9. https://doi.org/10.22201/ib.20078706e.2023.94.5081
- Moctezuma, V., Espinosa de los Monteros, A. & Halffter, G. (2024) Phylogenetic analyses of the subfamily Scarabaeinae (Coleoptera: Scarabaeidae) provide new insights into the Mexican Transition Zone theory. Zootaxa, 5415 (4), 501-528. https://doi.org/10.11646/zootaxa.5415.4.1
- Monaghan, M.T., Inward, D.J.G., Hunt, T. & Vogler, A.P. (2007) A molecular phylogenetic analysis of the Scarabaeinae (dung beetles). Molecular and Phylogenetic Evolution, 45, 674-692. https://doi.org/10.1016/j.ympev.2007.06.009
- Montes, C., Cardona, A. & Jaramillo, C. (2015) Middle Miocene closure of the Central American Seaway. Science, 348, 226- 228. https://doi.org/10.1126/science.aaa2815
- Muller, P. (1977) Tiergeographie: Struktur, Funktion, Geschichte und Indikatorbedeutung von Arealen. Teubner-Verlag, Stuttgart, 268 pp.
- Nguyen, L.-T., Schmidt, H.A., von Haeseler, A. & Minh, B.Q. (2014) IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution, 32, 268-274. https://doi.org/10.1093/molbev/msu300
- O'Dea, A., Lessios, H.A., Coates A.G., Eytan, R.I., Collins, L.S., Alberto L. Cione, A.L., Queiroz, A.d., Farris, D.W., Norris, R.D., Restrepo-Moreno, S.A., Stallard, R.F., Woodburne, M.O., Aguilera, O., Aubry, M.-P., Berggren, W.A., Budd, A.F., Cozzuol, M.A., Coppard, S.E., Finnegan, S., Gasparini, G.M., Grossman, E.L., Johnson, K.G., Keigwin, L.D., Knowlton, N., Leigh E.G., Leonard-Pingel, J.L., Marko, P.B., Pyenson, N.D., Rachello-Dolmen, P.G., Soibelzon, E., Soibelzon, L., Todd, J.A., Vermeij, G.J. & Jackson, J.B.C. (2016) Formation of the Isthmus of Panama. Science Advances, 2 (e1600883), 1-11. https://doi.org/10.1126/sciadv.1600883
- Palestrini, C. (1985) Problemi filogenetici e biogeografici del popolamento australiano di Onthophagini (Coleoptera Scarabaeidae). Atti XIV Congresso Nazionale Italiano di Entomologia. 28 Maggio -1 Giugno. Erice, Sicily, pp. 249-253.
- Palestrini, C. & Zunino, M. (1986) L'analisi dell'entomofauna nelle Zone di Transizione: prospettive e problemi. Biogeographia, 12, 11-25. https://doi.org/10.21426/B611110243
- Palestrini, C., Simonis, A. & Zunino, M. (1985) Modelli di distribuzione dell´entomofauna della Zona di Transizione Cinesi, nalisis di esempi e ipotese sulle sue origini. Biogeographia, 11, 195-209. https://doi.org/10.21426/B611110243
- Philips, T.K. (2016) Phylogeny of the Oniticellini and Onthophagini dung beetles (Scarabaeidae, Scarabaeinae) from morphological evidence. ZooKeys, 579, 9-57. https://doi.org/10.3897/zookeys.579.6183
- Price, D.L. (2009) Phylogeny and biogeography of the dung beetle genus Phanaeus (Coleoptera: Scarabaeidae). Systematic Entomology, 34, 137-150. https://doi.org/10.1111/j.1365-3113.2008.00443.x
- Ratnasingham, S. & Hebert, P.D.N. (2007) BOLD: The barcode of life data system (www.barcodinglife.org). Molecular Ecology Notes, 7, 355-364. https://doi.org/10.1111/j.1471-8286.2007.01678.x
- Rican, O., Pialek, L., Zardoya, R., Doadrio, I. & Zrzavy, J. (2013) Biogeography of the Mesoamerican Cichlidae (Teleostei: Heroini): colonization through the GAARlandia land bridge and early diversification. Journal of Biogeography, 40, 579- 593. https://doi.org/10.1111/jbi.12023
- Roggero, A., Barbero, E. & Palestrini, C. (2017) Revised classification and phylogeny of an Afrotropical species group based on molecular and morphological data, with the description of a new genus (Coleoptera: Scarabaeidae: Onthophagini). Organisms, Diversity & Evolution, 17, 181-198. https://doi.org/10.1007/s13127-016-0297-z
- Rossini, M. (2021) Additional mislabeling in African Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae): the case of Onthophagus viviensis and Onthophagus laevatus. Zootaxa, 5032 (2), 262-274. https://doi.org/10.11646/zootaxa.5032.2.7
- Rossini, M. & Genier, F. (2024) A new classification for the Onthophagus clypeatus group (Coleoptera: Scarabaeidae: Scarabaeinae) and description of a new species from western Ecuador. The Coleopterists Bulletin, 78, 415-428. https://doi.org/10.1649/0010-065X-78.3.415
- Rossini, M., Vaz-de-Mello, F.Z. & Zunino, M. (2018a) Toward a comprehensive taxonomic revision of the "hirculus" group of American Onthophagus Latreille, 1802 (Coleoptera, Scarabaeidae, Scarabaeinae). European Journal of Taxonomy, 432, 1-21. https://doi.org/10.5852/ejt.2018.432
- Rossini, M., Vaz-de-Mello, F.Z. & Zunino, M. (2018b) A taxonomic revision of the American Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae) of the osculatii species-complex, with description of two new species from South America. Journal of Natural History, 52, 541-586. https://doi.org/10.1080/00222933.2018.1437230
- Rowland, J.M., Qualls, C.R. & Buzatto, B.A. (2017) A model for conditional male trimorphisms. Journal of Theoretical Biology, 419, 184-192. https://doi.org/10.1016/j.jtbi.2017.02.006
- Sanchez-Huerta, J.L., Zunino, M. & Halffter, G. (2018) A new species of American Onthophagus Latreille (Coleoptera: Scarabaeidae: Scarabaeinae) associated with rodent (Geomydae) burrows. The Coleopterist Bulletin, 72, 407-416. https://doi.org/10.1649/0010-065X-72.3.407
- Schoolmeesters, P. (2024) World Scarabaeidae Database. Catalogue of Life Checklist. Version 5 August 2024. Available from: https://doi.org/10.48580/dgdwl-38g (accessed 19 November 2024)
- Schwery, O. & O'Meara, B.C. (2021) Age, origin, and biogeography: unveiling the factors behind the diversification of dung beetles. Available from: https://doi.org/10.1101/2021.01.26.428346 (accessed 19 November 2024)
- Slay, M.E., Skelley, P.E. & Taylor, S.J. (2012) New Records of Onthophagus cavernicollis Howden and Cartwright (Coleoptera: Scarabaeidae) from Ozark caves, with a review of scarabaeoids reported from North American caves. The Coleopterists Bulletin, 66, 187-199. https://doi.org/10.1649/072.066.0302
- Smith, S.A. & Bermingham, E. (2005) The biogeography of lower Mesoamerican freshwater fishes. Journal of Biogeography, 32, 1835-1854. https://doi.org/10.1111/j.1365-2699.2005.01317.x
- Solis, A. & Kohlmann, B. (2012) A checklist and distribution atlas of the Scarabaeinae (Coleoptera: Scarabaeidae) of Costa Rica. Zootaxa, 3482 (1), 21-32. https://doi.org/10.11646/zootaxa.3482.1.1
- Solis, A. & Kohlmann, B. (2023) New synonyms and registers of Costa Rican and Panamanian dung beetles (Coleoptera: Scarabaeinae). Insecta Mundi, 972, 1-21.
- Solis, A., Kohlmann, B. & Alvarado, G. (2024) A Costa Rican biological puzzle⸻the geological, landscape, and vicariance history as a conditioning of the past and present biogeography. In: Quesada-Roman, A. (Ed.), Landscapes and Landforms of Costa Rica. Springer, Cham, pp. 1-455. https://doi.org/10.1007/978-3-031-64940-0_10
- Stamatakis, A. (2014) RAxML Version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30, 1312-1313. https://doi.org/10.1093/bioinformatics/btu033
- Stegmann, B. (1930) Die Vogel des dauro-mandschurischen Ubergangsgebietes. Journal fur Ornithologie, 78, 389-471. https://doi.org/10.1007/BF01953148
- Taborsky, M. & Brockmann, H.J. (2010) Alternative reproductive tactics and life history phenotypes. In: Kappeler, E.M. (Ed.), Animal Behavior: Evolution and Mechanisms. Springer-Verlag, Berlin, pp. 537-586. https://doi.org/10.1007/978-3-642-02624-9_18
- Tarasov, S.I. & Dimitrov, D. (2016) Multigene phylogenetic analysis redefines dung beetles relationships and classification (Coleoptera: Scarabaeidae: Scarabaeinae). BMC Evolutionary Biology, 16, 1-19. https://doi.org/10.1186/s12862-016-0822-x
- Tarasov, S.I. & Genier, F. (2015) Innovative Bayesian and parsimony phylogeny of dung beetles (Coleoptera, Scarabaeidae, Scarabaeinae) enhanced by ontology-based partitioning of morphological characters. PLoS One, 10, 1-86. https://doi.org/10.1371/journal.pone.0116671
- Tarasov, S.I. & Solodovnikov, A.Y. (2011) Phylogenetic analyses reveal reliable morphological markers to classify megadiversity in Onthophagini dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae). Cladistics, 27, 1-39. https://doi.org/10.1111/j.1096-0031.2011.00351.x
- Tarasov, S., Vaz-de-Mello, F.Z., Krell, F.T. & Dimitrov, D. (2016) A review and phylogeny of Scarabaeine dung beetle fossils (Coleoptera: Scarabaeidae: Scarabaeinae), with the descriptions of two Canthochilum species from Dominican amber. Peerj, 4 (e1988), 1-35. https://doi.org/10.7717/peerj.1988
- Toledo, V.M. (1982) Pleistocene vegetation changes in tropical Mexico. In: Prance, G.T. (Ed.), Biological Diversification in the Tropics: Proceedings of the Fifth International Symposium of the Association for Tropical Biology. Columbia University Press, New York, New York, pp. 93-111.
- Waltari, E., Hijmans, R.J., Peterson, A.T., Nyari, A.S., Perkins, S.L. & Guralnick, R.P. (2007) Locating Pleistocene refugia: comparing phylogeographic and ecological niche model predictions. PlosOne, 7 (e563), 1-11. https://doi.org/10.1371/journal.pone.0000563
- Wang, C.W. (1961) The Forests of China with a Survey of Grassland and Desert Vegetation. Maria Moors Cabot Foundation/ Harvard University, Cambridge, Massachusetts, United States of America, 313 pp.
- Whittaker, R.J, Fernandez-Palacios, J.M. & Matthews, T.J. (2023) Island Biogeography. Geo-environmental dynamics, ecology, evolution, human impact, and conservation. Oxford University Press, Oxford, 475 pp. https://doi.org/10.1093/oso/9780198868569.001.0001
- Willmore, K.E. (2012) The body plan concept and its centrality in evo-devo. Evolution: Education and Outreach, 5, 219-230. https://doi.org/10.1007/s12052-012-0424-z
- Wolfe, J.A. (1975) Some aspects of plant geography in the Northern Hemisphere during the late Cretaceous and Tertiary. Annals of the Missouri Botanical Garden, 62, 264-279. https://doi.org/10.2307/2395198
- Zhang, Q., Ree, R.H., Salamin, N., Xing, Y. & Silvestro, D. (2021) Fossil-informed models reveal a boreotropical origin and divergent evolutionary trajectories in the walnut family (Juglandaceae). Systematic Biology, 71, 242-258. https://doi.org/10.1093/sysbio/syab030
- Zunino, M. (1979) Gruppi artificiali e gruppi naturali negli Onthophagus (Coleoptera Scarabaeoidea). Bollettino del Museo di Zoologia dell'Universita di Torino, 1, 1-18.
- Zunino, M. (2003) Tribu Onthophagini. In: Moron, M.-A. (Ed.), Atlas de los Escarabajos de Mexico. Vol. II. Familias Scarabaeidae, Trogidae, Passalidae y Lucanidae. Argania Editio, Barcelona, pp. 66-74.
- Zunino, M. & Halffter, G. (1981) Descrizione di Onthophagus micropterus n. sp. (Coleoptera: Scarabaeidae), note sulla sua distribuzione geografica e sulla riduzione alare nel genere. Bollettino del Museo di Zoologia dell'Universita di Torino, 8, 95-110.
- Zunino, M. & Halffter, G. (1988) Analisis taxonomico, ecologico y biogeografico de un grupo americano de Onthophagus. Museo Regionale di Scienze Naturale Torino, Monografia, 9, 1-211.
- Zunino, M. & Halffter, G. (1997) Sobre Onthophagus Latreille, 1802 americanos (Coleoptera: Scarabaeidae: Scarabaeinae). Elytron, 11, 157-178.
- Zunino, M. & Halffter, G. (2007) The association of Onthophagus Latreille, 1802 beetles (Coleoptera: Scarabaeinae) with vertebrate burrows and caves. Elytron, 21, 17-55.
- Zunino, M. & Halffter, G. (2019) About the origin of American Onthophagus (Coleoptera: Scarabaeidae). A critical appraisal of a recent paper by Breeschoten et al. (2016) on phylogenetics and biogeography. Molecular Phylogenetics and Evolution, 133, 1-3. https://doi.org/10.1016/j.ympev.2019.01.001