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Published: 2022-11-29

Distribution, Regionalization, and Diversity of the dung beetle genus Phanaeus MacLeay (Coleoptera: Scarabaeidae) using Species Distribution Models

Museo de Zoología, Facultad de Ciencias, Universidad Nacional Autónoma de México, 04510 Mexico City, Mexico.
Centro Tlaxcala de Biología de la Conducta, Universidad Autónoma de Tlaxcala, 90070 La Loma Xicohtencatl, Tlaxcala de Xicohténcatl, Tlaxcala, Mexico.
Red de Ecoetología, Instituto de Ecología, A. C., 91063 Xalapa, Veracruz, Mexico.
Coleoptera Biogeography Phanaeini Wallacean shortfall Mexican Transition Zone Neotropical beta diversity


The genus Phanaeus is a well-known group whose taxonomy has been described multiple times. Its distribution was previously classified into 11 ecogeographic groups that are equivalent to areas of endemism. Here we use Species Distribution Models to describe species richness patterns. We measured beta-diversity and regionalized its distribution into one region and one transition zone, both with three dominions: Mexican Transition Zone (North American, Mexican, and Mesoamerican dominions) and Neotropical region (Pacific, Brazilian, and Atlantic Forest dominions). We also present a species checklist and updated the distribution maps for 73 of 81 species described so far that reflects all the taxonomical updates. We include a list of all the recorded locations (by country, state, and province), list the recorded habitats and biomes, and describe the modelled environmental conditions for each species.



  1. Alvarado, F., Escobar, F. & Montero-Muñoz, J. (2014) Diversity and biogeographical makeup of the dung beetle communities inhabiting two mountains in the Mexican Transition Zone. Organisms Diversity and Evolution, 4, 105–114.
    Araújo, M.B., Guisan, A. & Arau, M.B. (2006) Five (or so) challenges for species distribution modelling. Journal of Biogeography, 33, 1677–1688.
    Arnaud, P. (2002) Phanaeini. Les coléoptères du monde. Vol. 28. Hillside Books, Canterbury, 151 pp.
    Arnaud, P. (2018) Description d’une nouvelle espèce de Phanaeus d’Équateur et revalidation de l’espèce Coprophanaeus (C.) edmondsi Arnaud (Coleoptera: Scarabaeidae, Scarabaeinae). Besoiro, 26, 3–7.
    Barve, N. & Barve, V. (2019) ENMGadgets: Pre and Post Processing in ENM Workflow. R package Version [program]
    Barve, N., Barve, V., Jiménez-Valverde, A., Lira-Noriega, A., Maher, S.P., Peterson, A.T., Soberón, J. & Villalobos, F. (2011) The crucial role of the accessible area in ecological niche modeling and species distribution modeling. Ecological Modelling, 222, 1810–1819.
    Baselga, A., Orme, D., Villeger, S., De Bortoli, J., Leprieur, J. & Logez, M. (2022) betapart: Partitioning Beta Diversity into Turnover and Nestedness Components. R package. Version 1.5.6. [program]
    Birch, C.P., Oom, S.P. & Beecham, J.A. (2007) Rectangular and hexagonal grids used for observation, experiment and simulation in ecology. Ecological Modelling, 206, 347–359.
    Boyce, M.S., Vernier, P.R., Nielsen, S.E. & Schmiegelow, F.K.A. (2002) Evaluating resource selection functions. Ecological Modelling, 157, 281–300.
    Cardoso, P., Rigal, F. & Carvalho, J.C. (2015) BAT—Biodiversity Assessment Tools, an R package for the measurement and estimation of alpha and beta taxon, phylogenetic and functional diversity. In: Kembel, S. (Ed.), Methods in Ecology and Evolution, 6, pp. 232–236.
    Carvalho, J.C., Cardoso, P. & Gomes, P. (2012) Determining the relative roles of species replacement and species richness differences in generating beta-diversity patterns. Global Ecology and Biogeography, 21, 760–771.
    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.
    di Cola, V., Broennimann, O., Petitpierre, B., Breiner, F.T., D’Amen, M., Randin, C., Engler, R., Pottier, J., Pio, D., Dubuis, A., Pellissier, L., Mateo, R.G., Hordijk, W., Salamin, N. & Guisan, A. (2017) ecospat: an R package to support spatial analyses and modeling of species niches and distributions. Ecography, 40, 774–787.
    Conservation Biology Institute (2011) Los insectos del orden Coleoptera para Costa Rica. Data Basin. Available from: (accessed 5 November 2020)
    Cultid-Medina, C.A., Medina-Uribe, C.A., Martínez-Quintero, B.G., Escobar-Villa, A.F., Constantino-Chuaire, L.M. & Betancur-Posada, N.J. (2012) Biota Colombiana. Escarabajos coprófagos (Scarabaeinae) del Eje Cafetero: guía para el estudio ecológico, Villa María, 2012, 1–196.
    de Klerk, H.M., Crowe, T.M., Fjeldså, J. & Burgess, N.D. (2002) Biogeographical patterns of endemic terrestrial Afrotropical birds. Diversity and Distributions, 8, 147–162.
    Dinerstein, E., Olson, D., Joshi, A., Vynne, C., Burgess, N.D., Wikramanayake, E., Hahn, N., Palminteri, S., Hedao, P., Noss, R., Hansen, M., Locke, H., Ellis, E.C., Jones, B., Barber, C.V., Hayes, R., Kormos, C., Martin, V., Crist, E., Sechrest, W., Price, L., Baillie, J.E.M., Weeden, D., Suckling, K., Davis, C., Sizer, N., Moore, R., Thau, D., Birch, T., Potapov, P., Turubanova, S., Tyukavina, A., de Souza, N., Pintea, L., Brito, J.C., Llewellyn, O.A., Miller, A.G., Patzelt, A., Ghazanfar, S.A., Timberlake, J., Klöser, H., Shennan-Farpón, Y., Kindt, R., Lillesø, J.-P.B., van Breugel, P., Graudal, L., Voge, M., Al-Shammari, K.F. & Saleem, M. (2017) An ecoregion-based approach to protecting half the terrestrial realm. BioScience, 67, 534–545.
    Diniz-Filho, J.A.F., de Marco, P. & Hawkins, B.A. (2010) Defying the curse of ignorance: Perspectives in insect macroecology and conservation biogeography. Insect Conservation and Diversity, 3, 172–179.
    Edmonds, W.D. (1994) Revision of Phanaeus MacLeay, a new world genus of scarabaeine dung beetles (Coleoptera: Scarabaeidae, Scarabaeinae). Contributions in Science of the Natural History Museum of Los Angeles County, 443, 1–105.
    Edmonds, W.D. & Zídek, J. (2012) Taxonomy of Phanaeus revisited: Revised keys to and comments on species of the New World dung beetle genus Phanaeus MacLeay, 1819 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). Insecta Mundi, 274, 1–108.
    Elith, J., Graham, C.H., Anderson, R.P., Dudík, M., Ferrier, S., Guisan, A., Hijmans, R.J., Huettmann, F., Leathwick, J.R., Lehmann, A., Li, J., Lohmann, L.G., Loiselle, B.A., Manion, G., Moritz, C., Nakamura, M., Nakazawa, Y., Overton, J.McC.M., Peterson, A.T., Phillips, S.J., Richardson, K., Scachetti-Pereira, R., Schapire, R.E., Soberón, J., Williams, S., Wisz, M.S. & Zimmermann, N.E. (2006) Novel methods improve prediction of species’ distributions from occurrence data. Ecography, 29, 129–151.
    Ennen, J.R., Agha, M., Sweat, S.C., Matamoros, W.A., Lovich, J.E., Rhodin, A.G.J., Iverson, J.B. & Hoagstrom, C.W. (2020) Turtle biogeography: Global regionalization and conservation priorities. Biological Conservation, 241, 108323.
    Escalante, T., Sánchez-Cordero, V., Morrone, J.J. & Linaje, M. (2007) Deforestation affects biogeographical regionalization: A case study contrasting potential and extant distributions of Mexican terrestrial mammals. Journal of Natural History, 41, 965–984.
    Escobar, F., Halffter, G. & Arellano, L. (2007) From forest to pasture: an evaluation of the influence of environment and biogeography on the structure of beetle (Scarabaeinae) assemblages along three altitudinal gradients in the Neotropical region. Ecography, 30, 193–208.
    Escobar, F., Lobo, J.M. & Halffter, G. (2005) Altitudinal variation of dung beetle (Scarabaeidae: Scarabaeinae) assemblages in the Colombian Andes. Global Ecology and Biogeography, 14, 327–337.
    Fick, S.E. & Hijmans, R.J. (2017) WorldClim 2: new 1-km spatial resolution climate surfaces for global land areas. International Journal of Climatology, 37, 4302–4315.
    Figueroa, L., Edmonds, W.D. & Martínez-Luján, N. (2014) La tribu Phanaeini (Coleoptera, Scarabaeidae, Scarabaeinae) en el Perú. Revista Peruana de Biología, 21, 125–138.
    Global Administrative Areas (2018) GADM database of Global Administrative Areas. Version 3.4. 2018. Available from: (accessed 1 November 2021)
    Gámez, J. & Acconcia, R. (2018) Escarabajos coprófagos y necrófagos (Scarabaeidae: Scarabaeinae: Phanaeini) del flanco andino llanero, piedemonte y llanos, con el reporte de nuevos registros para los estados de Barinas y Mérida (Venezuela). Revista Gaditana de Entomología, 9, 65–75.
    GBIF (2019) GBIF Occurrence Download [Data set].
    Gillett, C.P.D.T. & Toussaint, E.F.A. (2020) Macroevolution and shifts in the feeding biology of the New World scarab beetle tribe Phanaeini (Coleoptera: Scarabaeidae: Scarabaeinae). Biological Journal of the Linnean Society, 130 (4), 661–682.
    Gillett, C., Gillett, M., Gillett, J. & Vaz-de-Mello, F.Z. (2010) Diversity and distribution of the scarab beetle tribe Phanaeini in the northern states of the Brazilian Northeast (Coleoptera: Scarabaeidae: Scarabaeinae). Insecta Mundi, 118, 1–19.
    Gressitt, J.L. (1974) Insect Biogeography. Annual Review of Entomology, 19, 293–321.
    Halffter, G. (1987) Biogeography of the montane entomofauna of Mexico and Central America. Annual Review of Entomology, 32, 95–114.
    Halffter, G. (2017) La zona de transición mexicana y la megadiversidad de México: del marco histórico a la riqueza actual. Dugesiana, 24, 77–89.
    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), 1–46.
    Halffter, G., Favila, M.E. & Arellano, L. (1995) Spatial distribution of three groups of Coleoptera along an altitudinal transect in the Mexican Transition Zone and its biogeographical implications. Elytron, 9, 151–185.
    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, 329–335.
    Hamel-Leigue, A.C. & Herzog, S.K. (2008) Composición y riqueza de una comunidad de escarabajos peloteros (Coleoptera: Scarabaeinae) en los Yungas bajos de la Cordillera Mosetenes, Bolivia. Revista Boliviana de Ecología y Conservación Ambiental, 23, 39–49.
    Hamel-Leigue, A.C., Herzog, S.K., Larsen, T.H., Mann, D.J., Gill, B.D., Edmonds, W.D. & Spector, S. (2013) Biogeographic patterns and conservation priorities for the dung beetle tribe Phanaeini (Coleoptera: Scarabaeidae: Scarabaeinae) in Bolivia. Insect Conservation and Diversity, 6, 276–289.
    Hamel-Leigue, A.C., Herzog, S.K., Mann, D.J., Larsen, T.H., Gill, B.D., Edmonds, W.D. & Spector, S. (2009) Distribución e historia natural de escarabajos coprófagos de la tribu Phanaeini (Coleoptera: Scarabaeidae: Scarabaeinae) en Bolivia. Kempffiana, 5, 43–95.
    Hattab, T., Albouy, C., Lasram, F.B.R., Loc’h, F.L., Guilhaumon, F. & Leprieur, F. (2015) A biogeographical regionalization of coastal Mediterranean fishes. Journal of Biogeography, 42 (7), 1336–1348.
    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.
    Hijmans, R.J., Phillips, S., Leathwick, J. & Elith, J. (2017) Package “dismo’’—Specie Distribution Modeling”. CRAN Repository. [program]
    Hirzel, A.H., Le Lay, G., Helfer, V., Randin, C. & Guisan, A. (2006) Evaluating the ability of habitat suitability models to predict species presences. Ecological Modelling, 199, 142–152.
    Holt, B.G., Lessard, J.P., Borregaard, M.K., Fritz, S.A., Araújo, M.B., Dimitrov, D., Fabre, P.H., Graham, C.H., Graves, G.R., Jønsson, K.A., Nogués-Bravo, D., Wang, Z., Whittaker, R.J., Fjeldså, J. & Rahbek, C. (2013) An update of Wallace’s zoogeographic regions of the world. Science, 339, 74–78.
    Hortal, J., Roura-Pascual, N., Sanders, N.J. & Rahbek, C. (2010) Understanding (insect) species distributions across spatial scales. Ecography, 33, 51–53.
    Hughes, C. & Eastwood, R. (2006) Island radiation on a continental scale: Exceptional rates of plant diversification after uplift of the Andes. Proceedings of the National Academy of Sciences, 106, 9322–9327.
    Jablonski, D., Roy, K. & Valentine, J.W. (2006) Out of the tropics: Evolutionary dynamics of the latitudinal diversity gradient. Science, 314, 102–106.
    Kelley, L.A., Gardner, S.P. & Sutcliffe, M.J. (1996) An automated approach for clustering an ensemble of NMR-derived protein structures into conformationally related subfamilies. Protein Engineering, Design and Selection, 9, 1063–1065.
    Kohlmann, B., Arriaga-Jiménez, A. & Rös, M. (2018) Dung beetle vicariant speciation in the mountains of Oaxaca, Mexico, with a description of a new species of Phanaeus (Coleoptera, Geotrupidae, Scarabaeidae). ZooKeys, 743, 67–93.
    Kreft, H. & Jetz, W. (2010) A framework for delineating biogeographical regions based on species distributions. Journal of Biogeography, 37, 2029–2053.
    Lizardo, V., Escobar, F. & Rojas-Soto, O. (2017) Diversity and distribution of Phanaeini (Coleoptera: Scarabaeidae: Scarabaeinae) in Mexico. Zootaxa, 4358 (2), 271–294.
    Lobo, J.M. & Halffter, G. (2000) Biogeographical and ecological factors affecting the altitudinal variation of mountainous communities of coprophagous beetles (Coleoptera: Scarabaeoidea): A comparative study. Annals of the Entomological Society of America, 93, 115–26.[0115:BAEFAT]2.0.CO;2
    Löwenberg-Neto, P. (2014) Neotropical region: A shapefile of Morrone’s (2014) biogeographical regionalisation. Zootaxa, 3802 (2), 300–300.
    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, 1586–1600.
    Matamoros, W.A., Kreiser, B.R. & Schaefer, J.F. (2012) A delineation of Nuclear Middle America biogeographical provinces based on river basin faunistic similarities. Reviews in Fish Biology and Fisheries, 22, 351–365.
    Mateo, R.G., Vanderpoorten, A., Muñoz, J., Laenen, B. & Désamoré, A. (2013) Modeling species distributions from heterogeneous data for the biogeographic regionalization of the European bryophyte flora. PLoS ONE, 8, e55648.
    Moctezuma, V. & Halffter, G. (2017) A new Species of Phanaeus MacLeay (Coleoptera: Scarabaeidae: Scarabaeinae) from Los Chimalapas, Oaxaca, Mexico. The Coleopterists Bulletin, 71, 47–56.
    Moctezuma, V. & Halffter, G. (2021) Taxonomic revision of the Phanaeus endymion species group (Coleoptera: Scarabaeidae), with the description of five new species. European Journal of Taxonomy, 747, 1–71.
    Moctezuma, V. & Nogueira, G. (2019) Distribución geográfica de Phanaeus furiosus Bates, 1887 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini) y primeros registros para Zacatecas, México. Dugesiana, 26, 69–70.
    Moctezuma, V., Deloya, C., Sánchez-Huerta, J.L. & Halffter, G. (2019) A new species of the Phanaeus endymion species group (Coleoptera: Scarabaeidae: Scarabaeinae), with comments on ecology and distribution. Annales de la Société Entomologique de France, New Series, 55, 249–254.
    Moctezuma, V., Halffter, G. & Escobar, F. (2016) Response of copronecrophagous beetle communities to habitat disturbance in two mountains of the Mexican Transition Zone: influence of historical and ecological factors. Journal of Insect Conservation, 20, 945–956.
    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 Nationailis Pragae, 61, 447–482.
    Moctezuma, V., Nogueira, G. & Halffter, G. (2020) A revalidation and a new species in the genus Phanaeus (Coleoptera: Scarabaeoidea: Scarabaeidae: Scarabaeinae). Besoiro, 30, 3–11.
    Moctezuma, V., Sánchez-Huerta, J.L. & Halffter, G. (2017) Two new species of the Phanaeus endymion species group (Coleoptera, Scarabaeidae, Scarabaeinae). ZooKeys, 702, 113–135.
    Morrone, J.J. (2004) La zona de transición Sudamericana: caracterización y relevancia evolutiva. Acta Entomológica Chilena, 28, 41–50.
    Morrone, J.J. (2005) Hacia una síntesis biogeográfica de México. Revista Mexicana de Biodiversidad, 76, 207–252.
    Morrone, J.J. (2006) Biogeographic areas and transition zones of Latin America and the Caribbean islands based on panbiogeographic and cladistic analyses of the entomofauna. Annual Review of Entomology, 51, 467–494.
    Morrone, J.J. (2007) Hacia una biogeografía evolutiva. Revista Chilena de Historia Natural, 80, 509–520. https://doi.org10.4067/S0716-078X2007000400011
    Morrone, J.J. (2009) Evolutionary biogeography: An integrative approach with case studies. Columbia University Press, New York, New York, 304 pp.
    Morrone, J.J. (2014) Biogeographical regionalisation of the Neotropical region. Zootaxa, 3782 (1), 1–110.
    Morrone, J.J. (2017) Neotropical Biogeography: Regionalization and Evolution. CRC Press, Boca Raton, Florida, 282 pp.
    Morrone, J.J. (2019) Regionalización biogeográfica y evolución biótica de México: encrucijada de la biodiversidad del Nuevo Mundo. Revista Mexicana de Biodiversidad, 90, e902980.
    Morrone, J.J. (2020) The Mexican Transition Zone: A natural biogeographic laboratory to study biotic assembly. Springer International Publishing, Cham, 191pp.
    Morrone, J.J. & Crisci, J.V. (1995) Historical biogeography: Introduction to methods. Annual Review of Ecology and Systematics, 26, 373–401.
    Naimi, B., Hamm, N.A.S., Groen, T.A., Skidmore, A.K. & Toxopeus, A.G. (2014) Where is positional uncertainty a problem for species distribution modelling? Ecography, 37, 191–203.
    Pessôa, M.B., Alves-Martins, F., de Marco Júnior, P. & Hortal, J. (2020) Unveiling the drivers of local dung beetle species richness in the Neotropics. Journal of Biogeography, 48, 861–871.
    Peterson, A.T., Ball, L.G. & Cohoon, K.P. (2002) Predicting distributions of Mexican birds using ecological niche modelling methods. Ibis, 144, E27–E32.
    Peterson, A.T., Papeş, M. & Soberón, J. (2008) Rethinking receiver operating characteristic analysis applications in ecological niche modeling. Ecological Modelling, 213, 63–72.
    Peterson, A.T., Navarro-Sigüenza, A.G. & Benítez-Díaz, H. (1998) The need for continued scientific collecting; a geographic analysis of Mexican bird specimens. Ibis, 140, 288–294.
    Philips, T.K., Pretorius, E. & Scholtz, C.H. (2004) A phylogenetic analysis of dung beetles (Scarabaeinae: Scarabaeidae): unrolling an evolutionary history. Invertebrate Systematics, 18 (1), 53–88.
    Podani, J. & Schmera, D. (2011) A new conceptual and methodological framework for exploring and explaining pattern in presence-absence data. Oikos, 120, 1625–1638.
    Price, D.L. (2007) A phylogenetic analysis of the dung beetle genus Phanaeus (Coleoptera: Scarabaeidae) based on morphological data. Insect Systematics and Evolution, 38, 1–18.
    Price, D.L. (2009) Phylogeny and biogeography of the dung beetle genus Phanaeus (Coleoptera: Scarabaeidae). Systematic Entomology, 34, 137–150.
    Procheş, Ş. & Ramdhani, S. (2012) The world’s zoogeographical regions confirmed by cross-taxon analyses. BioScience, 62, 260–70.
    Qian, H. & Ricklefs, R.E. (2016) Out of the tropical lowlands: Latitude versus elevation. Trends in Ecology and Evolution, 31, 738–741.
    Qian, H., Sandel, B., Deng, T. & Vetaas, O. R. (2019) Geophysical, evolutionary and ecological processes interact to drive phylogenetic dispersion in angiosperm assemblages along the longest elevational gradient in the world. Botanical Journal of the Linnean Society, 190, 333–344.
    R Core Team (2021) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Available from: (accessed 8 November 2022)
    SiBBr (2020) Phanaeus occurrence data. Available from: (accessed 5 November 2020)
    Sigrist, M.S. & de Carvalho, C.J.B. (2009) Historical relationships among areas of endemism in the tropical South America using Brooks Parsimony Analysis (BPA). Biota Neotropica, 9, 79–90.
    da Silva, J.L. & Vaz-de-Mello, F.Z. (2020) Areas of endemism in the Brazilian Atlantic Forest based on the distribution of dung beetles (Coleoptera, Scarabaeidae, Sscarabaeinae). Iheringia, Série Zoologia, 110, 1–10.
    Soberón, J. & Nakamura, M. (2009) Niches and distributional areas: concepts, methods, and assumptions. Proceedings of the National Academy of Sciences, 106, 19644–19650.
    Soberón, J. & Peterson, A.T. (2005) Interpretation of Models of Fundamental Ecological Niches and Species’ Distributional Areas. Biodiversity Informatics, 2, 1–10.
    Solís, Á. & Kohlmann, B. (2012) Checklist and distribution atlas of the Scarabaeinae (Coleoptera: Scarabaeidae) of Costa Rica. Zootaxa, 3482 (1), 1–32.
    U.S. EPA (2006) Level II Ecoregions of North America Metadata [Data set]. U.S. EPA Office of Research & Development (ORD)—National Health and Environmental Effects Research Laboratory (NHEERL). Corvallis, Oegon. Available from: (accessed 5 June 2020)
    USGS (2017) Digital Elevation - Shuttle Radar Topography Mission (SRTM) 1 Arc-Second Global [Data set]. USGS Earth Resources Observation and Science (EROS).
    Wiens, J.J. & Donoghue, M.J. (2004) Historical biogeography, ecology and species richness. Trends in Ecology and Evolution, 19, 639–644.
    Whittaker, R.H. (1960) Vegetation of the Siskiyou Mountains, Oregon and California. Ecological Monographs, 30, 280–338.
    White, D. & Gramacy, R. (2012) Maptree: Mapping, pruning, and graphing tree models. R package. Version 1.4-7. Available from: (accessed 8 November 2022)
    Yackulic, C.B., Chandler, R., Zipkin, E.F., Royle, J.A., Nichols, J.D., Campbell Grant, E.H. & Veran, S. (2013) Presence-only modelling using MAXENT: when can we trust the inferences? Methods in Ecology and Evolution, 4, 236–243.