Abstract
We analyzed species richness and ecogeographic patterns of Scarabaeinae (Coleoptera: Scarabaeidae) in Jalisco, Mexico. Occurrence data were compiled from digital databases, literature, and entomological collections, yielding 6,042 records for 86 species. A current richness model was generated using 100 km² grid cells, and species distribution models were constructed using MaxEnt to estimate potential richness. We also performed a principal component analysis based on 23 climatic variables to identify climatic affinity groups. Our results show that the areas with the highest current and potential richness are located in the Sierra Madre del Sur and the Pacific Coast provinces. We identified four main climatic groups that correspond to biogeographic provinces. These findings provide a comprehensive overview of Scarabaeinae diversity in Jalisco and highlight both sampling gaps and priority areas for future studies. The classification of species based on climatic affinities reveals a richness gradient and suggests distinct assemblage structures driven by environmental conditions.
References
Arriaga L., Aguilar C., Espinosa D. & Jiménez R. 1997. Regionalización ecológica y biogeográfica de México. Taller de la Comisión Nacional para el Conocimiento y Uso de la Biodiversidad, México, 123 pp.
Capello V. & Halffter G. 2019. Listado ilustrado de las especies de Scarabaeinae (Coleoptera: Scarabaeidae) de la Reserva de la Biosfera de Calakmul, Campeche, México. Dugesiana 26(2): 103–131. https://doi.org/10.32870/dugesiana.v26i2.7080
Cultid-Medina C. A. & Escobar F. 2020. Dung beetle diversity and functions suggest no major impacts of cattle grazing in the Colombian Andes. Neotropical Entomology 49: 96–107. https://doi.org/10.1007/s13744-019-00731-3
Delgado L. & Kohlmann B. 2007. Revisión de las especies del género Uroxys Westwood de México y Guatemala. Folia Entomológica Mexicana 46(1): 1–35.
Diaz-García J. M., Díaz de la Vega-Pérez A. H., Navarro-Noya Y., Ramírez-Ponce A., Guillén S., Esparza-Carlos J. P. & Arias-Del Razo I. 2025. Biodiversity between volcanoes: A multi-taxa analysis to identify priority conservation areas in a highly fragmented landscape of the Trans-Mexican Volcanic Belt. Journal for Nature Conservation 84: 126852. https://doi.org/10.1016/j.jnc.2024.126852
Edmonds W. D. 1994. Revision of Phanaeus Macleay, a New World genus of Scarabaeinae dung beetles (Coleoptera: Scarabaeidae, Scarabaeinae). Contributions in Science, Natural History Museum of Los Angeles County 443: 1–105.
Escalante T., Rodríguez-Tapia G., Morrone J. J., Linaje M. & Illoldi-Rangel P. 2021. Biogeographic transitions in the Mexican Transition Zone: A multi-taxon approach for biodiversity conservation. Journal of Biogeography 48: 1183–1197. https://doi.org/10.1111/jbi.14072
Favila M. E. & Halffter G. 1997. The use of indicator groups for measuring biodiversity as related to community structure and function. Acta Zoológica Mexicana (nueva serie) 72: 1–25.
García E. 2004. Modificaciones al sistema de clasificación climática de Köppen. Instituto de Geografía, UNAM, México, 91 pp.
GBIF.org 2015. Global Biodiversity Information Facility. Available from: https://www.gbif.org (Accessed 15 July 2015)
Génier F. 1996. A revision of the Neotropical genus Ontherus Erichson (Coleoptera: Scarabaeidae, Scarabaeinae). Memoirs of the Entomological Society of Canada 170: 1–169. https://doi.org/10.4039/entm128170fv
Génier F. 2009. Le genre Eurysternus Dalman, 1824: révision taxonomique et clés de determinación illustrées. Pensoft Series Faunistica 85: 1–430.
González-Hernández A. L., Navarrete-Heredia J. L., Quiroz-Rocha G. A. & Deloya C. 2015. Coleópteros necrócolos (Scarabaeidae: Scarabaeinae, Silphidae y Trogidae) del Bosque Los Colomos, Guadalajara, Jalisco, México. Revista Mexicana de Biodiversidad 86(3): 764–770. https://doi.org/10.1016/j.rmb.2015.07.006
Griffiths H. M., Louzada J., Bardgett R. D. & Barlow J. 2015. The value of dung beetles for ecosystem function. Proceedings of the Royal Society B: Biological Sciences 282(1817): 20151299. https://doi.org/10.1098/rspb.2015.1299
Halffter G. & Matthews E. G. 1966. The natural history of dung beetles of the subfamily Scarabaeinae (Coleoptera, Scarabaeidae). Folia Entomológica Mexicana 12–14: 1–312.
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: 1–46. https://doi.org/10.11646/zootaxa.4226.1.1
Halffter G. 1984. Biogeography of the montane entomofauna of Mexico and Central America. Annual Review of Entomology 29: 95–114.
Halffter G. 1987. Biogeography of the montane entomofauna of Mexico and Central America. Annual Review of Entomology 32(1): 95–114. https://doi.org/10.1146/annurev.en.32.010187.000523
Halffter G., Moctezuma V. & Nogueira G. 2022. A new species of Phanaeus MacLeay, 1819 (Coleoptera: Scarabaeidae: Scarabaeinae) from the mountains of Jalisco, Mexico. The Coleopterists Bulletin 76(3): 329–335. https://doi.org/10.1649/0010-065X-76.3.329
Halffter G., Rivera-Cervantes L. E. & Halffter V. 2015. Diversificación del grupo humectus del género Canthon (Coleoptera: Scarabaeidae: Scarabaeinae) en el occidente de México. Acta Zoológica Mexicana (nueva serie) 31(2): 208–220. https://doi.org/10.21829/azm.2015.3121747
Halffter G., Zunino M., Moctezuma V. & Sánchez-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): 1–34. https://doi.org/10.11646/zootaxa.4586.1.1
Hanski I. & Cambefort Y. 1991. Dung Beetle Ecology. Princeton University Press, Princeton, 520 pp.
Hernández B. & Navarrete-Heredia J. L. 2018. Annotated Checklist and Biogeographical Affinities of Scarabaeinae Beetles from Los Altos de Jalisco Region, Mexico. Southwestern Entomologist 43(1): 131–149.
Hernández B., Barragán-Ramírez J. L., Navarrete-Heredia J. L., Quiroz-Rocha G. A. & Vásquez-Bolaños M. 2021. Dung-beetle (Coleoptera: Scarabaeidae: Scarabaeinae) assemblage in two livestock production systems in a southern Mexican High Plateau semiarid ecosystem. The Canadian Entomologist 153(3): 285–300.
Hernández B., Navarrete-Heredia J. L., Vásquez-Bolaños M., Quiroz-Rocha G. A. & Deloya C. 2022. Necrophagous beetles (Coleoptera: Scarabaeoidea and Silphidae) along an elevational gradient from Cerro de García, Jalisco, Mexico. The Coleopterists Bulletin 76(2): 143–156.
Herzog S. K., Hamel-Leigue A. C., Larsen T. H., Mann D. J., Soria-Auza R. W., Gill B. D., Edmonds W. D. & Spector S. 2013. Elevational distribution and conservation biogeography of Phanaeini dung beetles (Coleoptera: Scarabaeinae) in Bolivia. PLOS ONE 8(5): e64963. https://doi.org/10.1371/journal.pone.0064963
Hijmans R. J. & van Etten J. 2020. Raster: Geographic data analysis and modeling. R package.
Hijmans R. J., Cameron S. E., Parra J. L., Jones P. G. & Jarvis A. 2005. Very high resolution interpolated climate surfaces for global land areas. International Journal of Climatology 25: 1965–1978. https://doi.org/10.1002/joc.1276
Howden H. F. & Génier F. 2004. Seven new species of Onthophagus Latreille from Mexico and the United States (Coleoptera: Scarabaeidae, Scarabaeinae). Fabreries 29(1): 53–76.
INEGI 2023. Anuario estadístico y geográfico de Jalisco 2023. Instituto Nacional de Estadística y Geografía, Aguascalientes, Mexico.
Kohlmann B. & Solís A. 2006. El género Canthidium (Coleoptera: Scarabaeidae) en Norteamérica. Giornale Italiano di Entomologia 53(11): 235–295.
Lizardo V., Escobar F. & Rojas-Soto O. 2017. Diversity and distribution of Phanaeini (Coleoptera: Scarabaeidae: Scarabaeinae) in Mexico. Zootaxa 4358(2): 271–294. https://doi.org/10.11646/zootaxa.4358.2.3
Lizardo V., Escobar F., Martínez-Meyer E. & Morrone J. J. 2024. Adaptive shifts in Phanaeini dung beetles of the Mexican plateau cenocron in the Mexican transition zone. Zoologica Scripta 53(4): 451–460. https://doi.org/10.1111/zsc.12660
Lizardo V., García-Trejo E. A. & Morrone J. J. 2024. Niche conservatism and convergence in birds of three cenocrons in the Mexican Transition Zone. PeerJ 12: e16664. https://doi.org/10.7717/peerj.16664
Lizardo V., Moctezuma V. & Escobar F. 2022. Distribution, regionalization, and diversity of the dung beetle genus Phanaeus MacLeay (Coleoptera: Scarabaeidae) using species distribution models. Zootaxa 5213(5): 546–568. https://doi.org/10.11646/zootaxa.5213.5.4
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://doi.org/10.1111/jbi.12541
Matthews E. G. & Halffter G. 1968. New data on American Copris with discussion of a fossil species (Col., Scarab.). Ciencia México 26(4): 147–162.
Matthews E. G. 1961. A revision of the genus Copris Müller of the Western Hemisphere (Coleoptera, Scarabaeidae). Entomologica Americana 41(1): 1–139.
Moctezuma V. & Halffter G. 2020a. New species and redescriptions of the Onthophagus chevrolati species complex (Coleoptera: Scarabaeoidea: Scarabaeinae). Annales Zoologici 70(2): 245–261. https://doi.org/10.3161/00034541ANZ2020.70.2.004
Moctezuma V. & Halffter G. 2020b. Three new species of the Onthophagus chevrolati species group (Coleoptera: Scarabaeoidea: Scarabaeinae). Biologia 75(12): 2277–2286. https://doi.org/10.2478/s11756-020-00562-z
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: e30. https://doi.org/10.6620/ZS.2021.60-30
Moctezuma V. 2021a. El género Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae) de México. Dugesiana 28(2): 175–220. https://doi.org/10.32870/dugesiana.v28i2.7166
Moctezuma V. 2021b. Spatial autocorrelation in a Mexican dung beetle ensemble: Implications for biodiversity assessment and monitoring. Ecological Indicators 125: 107548. https://doi.org/10.1016/j.ecolind.2021.107548
Moctezuma V., Espinosa de los Monteros A. & Halffter G. 2024b. Phylogenetic analyses of the subfamily Scarabaeinae (Coleoptera: Scarabaeidae) provide new insights into the Mexican Transition Zone theory. Zootaxa 5415: 501–528. https://doi.org/10.11646/zootaxa.5415.4.1
Moctezuma V., Halffter G. & Arriaga-Jiménez A. 2018. Archipelago reserves, a new option to protect montane entomofauna and beta-diverse ecosystems. Revista Mexicana de Biodiversidad 89(3): 927–937. https://doi.org/10.22201/ib.20078706e.2018.3.2424
Moctezuma V., Halffter G. & Escobar F. 2024a. Multifactorial structuring in a local guild of coprophilic scarab beetles from the Mexican transition zone: Analyses of ecological hypotheses. Ecological Entomology 49(5): 613–623. https://doi.org/10.1111/een.13320
Moctezuma V., Halffter G. & Lizardo V. 2021. The Phanaeus tridens species group (Coleoptera: Scarabaeoidea): A dung beetle group with genital morphological stasis but a changing ecological niche. Acta Entomologica Musei Nationalis Pragae 61(2): 603–616. https://doi.org/10.37520/aemnp.2021.031
Moctezuma V., Hernández B., Sánchez-Huerta J. L., Lizardo V., Quiroz-Rocha G. A. & Navarrete-Heredia J. L. 2023b. Two new species from the Mexican Pacific Slope and new distributional records of Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae). Journal of Asia-Pacific Entomology 26(1): 102015. https://doi.org/10.1016/j.aspen.2022.102015
Moctezuma V., Hernández B., Sánchez-Huerta J. L., Navarrete-Heredia J. L. & Martínez-Rodríguez P. A. 2021. Onthophagus acernorus (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini), a new dung beetle species from Jalisco, Mexico. Zootaxa 5067: 122–128. https://doi.org/10.11646/zootaxa.5067.1.8
Moctezuma V., Lizardo V., Arias-Del Razo I. & Ramírez-Ponce A. 2024c. Overcoming the Wallacean shortfall in sky-islands of central Mexico: the case of copro-necrophagous beetles and two national parks. Journal of Insect Conservation 28(4): 777–785. https://doi.org/10.1007/s10841-024-00561-3
Moctezuma V., Sánchez-Huerta J. L. & Halffter G. 2017. Two new species of the Phanaeus endymion species group (Coleoptera: Scarabaeidae: Scarabaeinae). ZooKeys 702: 113–127. https://doi.org/10.3897/zookeys.702.14791
Moctezuma V., Sánchez-Huerta J. L. & Hernández B. 2023a. A new Mexican species of the genus Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae). Studies on Neotropical Fauna and Environment 58(1): 204–211. https://doi.org/10.1080/01650521.2022.2108391
Montes de Oca E., Anduaga S. & Rivera E. 1994. Presence of the exotic dung beetle Euoniticellus intermedius (Reiche) (Coleoptera: Scarabaeidae) in northern Mexico. The Coleopterists Bulletin 48: 244.
Morrone J. J. 2017. Biogeographic regionalization of the Sierra Madre del Sur province, Mexico. Revista Mexicana de Biodiversidad 88: 710–714. https://doi.org/10.1016/j.rmb.2017.07.012
Morrone J. J. 2020. The Mexican Transition Zone: A Natural Biogeographic Laboratory to Study Biotic Assembly. Springer Nature, Cham, 200 pp. https://doi.org/10.1007/978-3-030-47917-6
Navarrete-Heredia J. L. & Cortés-Aguilar J. 2004. Beetles (Coleoptera) associated with the external debris of Atta mexicana (F. Smith) (Hymenoptera: Formicidae) from Ojuelos, Jalisco, México. Proceedings of the Entomological Society of Washington 106(2): 481–482.
Navarrete-Heredia J. L., Delgado L. & Fierros-López H. E. 2001. Coleoptera Scarabaeoidea de Jalisco, México. Dugesiana 8: 37–93.
Navarrete-Heredia J. L., Sainz-Medina C. I., González-Hernández A. L., Quiroz-Rocha G. A., Hernández A., Vásquez-Bolaños M. & Hernández-Márquez B. 2012. Coleópteros necrócolos del Bosque Los Colomos, Guadalajara, Jalisco, México. Dugesiana 19(2): 157–162.
Nichols E., Spector S., Louzada J., Larsen T., Amezquita S. & Favila M. E. 2019. Ecological functions and ecosystem services provided by Scarabaeinae dung beetles. Biological Conservation 229: 1–10. https://doi.org/10.1016/j.biocon.2018.11.001
Noriega J. A., Delgado O., Blanco J. I., Gámez J. & Clavijo J. 2017. Introduction, establishment, and invasion of Digitonthophagus gazella (Fabricius, 1787) (Coleoptera: Scarabaeinae) in the savannas of Venezuela and Colombia. Natural Resources 8: 370–381. https://doi.org/10.4236/nr.2017.85023
Noriega J. A., Hortal J., Azcárate F. M., Berg M. P., Bonada N., Briones M. J. I. & Santos A. M. C. 2018. Research trends in ecosystem services provided by dung beetles. Basic and Applied Ecology 26: 1–16. https://doi.org/10.1016/j.baae.2017.09.006
Noriega J. A., Hortal J., de Castro-Arrazola I., Menéndez R. & Hortal J. 2023. Dung removal increases under higher dung beetle functional diversity regardless of grazing intensification. Nature Communications 14: 8070. https://doi.org/10.1038/s41467-023-43760-8
Ojeda M., Trejo I., Challenger A., Dirzo R., López E. & Maass M. 2021. Biodiversity hotspots and conservation gaps in Mexico. Nature Conservation 44: 123–145. https://doi.org/10.3897/natureconservation.44.63377
Pablo-Cea J. D., Velado-Cano M. A., Fuentes R., Cruz M. & Noriega J. A. 2017. First report of Digitonthophagus gazella (Fabricius, 1787) and new records for Euoniticellus intermedius (Reiche, 1849) (Coleoptera: Scarabaeidae Latreille, 1802) in El Salvador. Acta Zoológica Mexicana (nueva serie) 33(3): 527–531.
Pessôa M. B., Alves-Martins F., De Marco Junior P. & Hortal J. 2021. Unveiling the drivers of local dung beetle species richness in the Neotropics. Journal of Biogeography 48(4): 861–871. https://doi.org/10.1111/jbi.14041
Peterson A. T., Soberón J., Pearson R. G., Anderson R. P., Martínez-Meyer E., Nakamura M. & Araújo M. B. 2018. Ecological Niches and Geographic Distributions. Princeton University Press, Princeton, 328 pp.
Peterson A. T., Soberón J., Pearson R. G., Anderson R. P., Martínez-Meyer E., Nakamura M. & Araújo M. B. 2011. Ecological Niches and Geographic Distributions. Princeton University Press, Princeton, 314 pp.
Phillips S. J., Anderson R. P. & Schapire R. E. 2006. Maximum entropy modeling of species geographic distributions. Ecological Modelling 190: 231–259. https://doi.org/10.1016/j.ecolmodel.2005.03.026
QGIS Development Team 2023. QGIS Geographic Information System. Open Source Geospatial Foundation Project.
Quiroz-Rocha G. A., Navarrete-Heredia J. L. & Martínez-Rodríguez P. A. 2008. Especies de Scarabaeinae (Coleoptera: Scarabaeidae) y Silphidae (Coleoptera) necrófilas de bosque de pino-encino y bosque mesófilo de montaña en el municipio de Mascota, Jalisco, México. Dugesiana 15(1): 27–37.
R Core Team 2023. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available from: https://www.R-project.org/ (Accessed 2 March 2026)
Rivera J. D. & Favila M. E. 2022. Dung beetle diversity and functional groups in a landscape with different degrees of anthropogenic disturbance in the Mexican Neotropics. Journal of Insect Conservation 26(1): 75–87. https://doi.org/10.1007/s10841-021-00361-9
Rivera-Cervantes L. E. & Halffter G. 1999. Monografía de las especies mexicanas de Canthon del subgénero Glaphyrocanthon (Coleoptera: Scarabaeidae; Scarabaeinae). Acta Zoológica Mexicana (nueva serie) 77: 23–150. https://doi.org/10.21829/azm.1999.77771693
Rivera-Gasperín S. L., Escobar-Hernández F. & Arellano L. 2025. A new species in the Canthon indigaceus species group (Coleoptera: Scarabaeidae: Scarabaeinae) from the Mexican Pacific Coast. Taxonomy 5(1): 11. https://doi.org/10.3390/taxonomy5010011
Rodríguez A. 2017. Ecorregiones terrestres, pp. 27–60. In: La Biodiversidad en Jalisco. Estudio de Estado. Vol. II (CONABIO, editor). CONABIO, México.
Ruiz-Corral J. A., Medina-García G. & García-Romero G. E. 2018. Sistema de información agroclimático para México-Centroamérica. Revista Mexicana de Ciencias Agrícolas 9(1): 1–10. https://doi.org/10.29312/remexca.v9i1.841
Rzedowski J. 2006. Vegetación de México, 1ra. edición digital. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad, México, 504 pp.
Sánchez-Hernández G. & Gómez B. 2018. First precise locality data for Onthophagus atriglabrus Howden and Gill and new state record for Onthophagus anewtoni Howden and Génier (Coleoptera: Scarabaeidae: Scarabaeinae) in Mexico. The Coleopterists Bulletin 72: 873–876. https://doi.org/10.1649/0010-065X-72.4.873
SNIB-CONABIO 2015. Sistema Nacional de Información sobre Biodiversidad. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad. Available from: http://www.conabio.gob.mx (Accessed 15 July 2015).
Soberón J. & Peterson A. T. 2020. What is the shape of the species richness–environment relationship? Ecography 43: 1131–1140. https://doi.org/10.1111/ecog.05159
Spector S. 2006. Scarabaeine dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae): An invertebrate focal taxon for biodiversity research and conservation. The Coleopterists Bulletin 60: 71–83.
Stanbrook R., Wheater C. P., Harris W. E. & Jones M. 2021. Habitat type and altitude work in tandem to drive the community structure of dung beetles in Afromontane forest. Journal of Insect Conservation 25: 397–410. https://doi.org/10.1007/s10841-020-00289-1
UNEP 1992. World Atlas of Desertification. United Nations Environment Programme, London, 69 pp.
Verdú J. R., Arellano L., Numa C. & Micó E. 2007. Roles of endothermy in niche differentiation for ball-rolling dung beetles (Coleoptera: Scarabaeidae) along an altitudinal gradient. Ecological Entomology 32: 544–551. https://doi.org/10.1111/j.1365-2311.2007.00907.x
Vernes K., Pope L. C., Hill C. J. & Bärlocher F. 2005. Seasonality, dung specificity and competition in dung beetle assemblages in the Australian Wet Tropics, north-eastern Australia. Journal of Tropical Ecology 21: 1–8. https://doi.org/10.1017/S026646740400224X
Villalobos F., Carvajal-Hernández C. I., Meza-Joya F. L., Guevara R. & Sánchez-Ramos G. 2022. Rethinking biodiversity patterns and processes in Mexico: Integrating biogeography and macroecology. Frontiers in Ecology and Evolution 10: 789532. https://doi.org/10.3389/fevo.2022.789532
Villamarín-Cortez S., Hankin L. E., Coronado S., Macdonald J. L. & Noriega J. A. 2022. Diversity and distribution patterns of Ecuador’s dung beetles (Coleoptera: Scarabaeinae). Frontiers in Ecology and Evolution 10: 1008477. https://doi.org/10.3389/fevo.2022.1008477
Wisz M. S., Hijmans R. J., Li J., Peterson A. T., Graham C. H., Guisan A. & NCEAS Predicting Species Distributions Working Group 2008. Effects of sample size on the performance of species distribution models. Diversity and Distributions 14: 763–773. https://doi.org/10.1111/j.1472-4642.2008.00482.x
Zunino M. & Halffter G. 1988. Análisis taxonómico ecológico y biogeográfico de un grupo americano de Onthophagus. Museo Regionale di Scienze Naturali Torino Monografia 9: 1–211.
Zunino M. 2003. Tribu Onthophagini, pp. 66–74. In: Atlas de los Escarabajos de México. Coleoptera: Lamellicornia. Vol. II (M. A. Morón and B. C. Ratcliffe, editors). Argania Editio, Barcelona.
