Abstract
The ant genus Pheidole is a highly diverse lineage of ants that are dominant elements of tropical and subtropical ecosystems throughout the world. Biodiversity inventory projects in Middle American wet forests (southern Mexico to Costa Rica, lowland rainforest to high montane forest) allow an improved taxonomy of the genus in this region. An identification guide to 234 species in the region is provided, using a "bird guide" approach. Species are arranged in order of minor worker head width and scape length, and each species has a fixed layout that includes habitat and microhabitat information, standard views (minor lateral, minor face, major face), and a distribution map. The following new synonyms are proposed: biconstricta Mayr 1870 (= inermis Mayr 1870), carapuna Mann 1916 (= tristicula Wilson 2003), fimbriata Roger 1863 (= soesilae Makhan 2007), insipida Forel 1899 (= fariasana Wilson 2003, mooreorum Wilson 2003), longiscapa Forel 1901 (= cocciphaga Borgmeier 1934), plebecula Forel 1899 (= perdiligens Wilson 2003, texticeps Wilson 2003), nitidicollis Emery 1896 (= chalcoides Wilson 2003), nubicola Wilson 2003 (= cielana Wilson 2003, petrensis Wilson 2003), simonsi Wilson 2003 (= arctos Wilson 2003, gangamon Wilson 2003, thrasys Wilson 2003), striaticeps Mayr 1870 (= chloe Forel 1908), transversostriata Mayr 1887 (= scalaris Wilson 2003), vafra Santschi 1923 (= laticornis Wilson 2003). The following 57 new species are described: ajaxigibba, andersoni, angustinigra, atitlana, balatro, belonorte, besalon, bicornisculpta, brownampla, cahui, caliginosa, carinitida, corniclypeus, costaricensis, cusuco, depressinoda, eosimilis, familiaparra, fincanaranjo, hansoni, hectornitida, hitoy, huarache, imbrilis, indagarama, kasparii, kelainos, lagunculiminor, lamancha, laselvoides, lineafrons, luteagossamer, machaquila, marmor, moskitia, muralla, musacolor, musinermis, natalie, nephele, obturaculum, passivaeferox, perissothrix, platyscapa, probolonotum, rima, rogeripolita, savegre, sensipelada, sepultura, tapanti, tikal, tinamu, tsontekonwei, tuculutan, xiloa, zannia.
References
Borgmeier, T. (1934) Contribuição para o conhecimento da fauna mirmecológica dos cafezais de Paramaribo, Guiana Holandesa (Hym. Formicidae). Archivos do Instituto de Biologia Vegetal (Rio de Janeiro), 1, 93–111, estampa 111–112.
Economo, E.P., Huang, J.-P., Fischer, G., Sarnat, E. M., Narula, N., Janda, M., Guénard, B., Longino, J.T. & Knowles, L.L. (2019) Evolution of the latitudinal diversity gradient in the hyperdiverse ant genus Pheidole. Global Ecology and Biogeography, 28, 456–470.
https://doi.org/10.1111/geb.12867
Economo, E.P., Klimov, P., Sarnat, E.M., Guénard, B., Weiser, M.D., Lecroq, B. & Knowles, L.L. (2015) Global phylogenetic structure of the hyperdiverse ant genus Pheidole reveals the repeated evolution of macroecological patterns. Proceedings of the Royal Society B: Biological Sciences, 282, 20141416.
https://doi.org/10.1098/rspb.2014.1416
Economo, E.P., Sarnat, E.M., Janda, M., Clouse, R., Klimov, P.B., Fischer, G., Blanchard, B.D., Ramirez, L.N., Andersen, A.N., Berman, M., Guénard, B., Lucky, A., Rabeling, C., Wilson, E.O. & Knowles, L.L. (2015) Breaking out of biogeographical modules: range expansion and taxon cycles in the hyperdiverse ant genus Pheidole. Journal of Biogeography, 42, 2289–2301.
https://doi.org/10.1111/jbi.12592
Emery, C. (1896) Studi sulle formiche della fauna neotropica. XVII-XXV. Bullettino della Società Entomologica Italiana, 28, 33–107, Tav. I.
Forel, A. (1899) Formicidae. [part c]. In: Insecta. Hymenoptera. Formicidae. R.H. Porter, Dulau & Co., London, pp. 57–80.
Forel, A. (1901) Variétés myrmécologiques. Annales de la Société Entomologique de Belgique, 45, 334–382.
Forel, A. (1908) Fourmis de Costa-Rica récoltées par M. Paul Biolley. Bulletin de la Société Vaudoise des Sciences Naturelles, 44, 35–72.
Forel, A. (1914) Einige amerikanische Ameisen. Deutsche Entomologische Zeitschrift, 1914, 615–620.
Gauld, I.D. & Bolton, B. (1988) The Hymenoptera. Oxford, Oxford University Press. 332 pp.
Kempf, W.W. (1972) Catálogo abreviado das formigas da Região Neotropical. Studia Entomologica, 15, 3–344.
Letourneau, D.K. (1983) Passive aggression: an alternative hypothesis for the Piper-Pheidole association. Oecologia, 60, 122–126.
https://doi.org/10.1007/BF00379331
Lillico-Ouachour, A. & Abouheif, E. (2016) Regulation, development, and evolution of caste ratios in the hyperdiverse ant genus Pheidole. Current Opinion in Insect Science, 19, 43–51.
https://doi.org/10.1016/j.cois.2016.11.003
Longino, J.T. (2009) Additions to the taxonomy of New World Pheidole (Hymenoptera: Formicidae). Zootaxa, 2181, 1–90.
Longino, J.T. & Colwell, R.K. (2011) Density compensation, species composition, and richness of ants on a neotropical elevational gradient. Ecosphere, 2, art29.
https://doi.org/10.1890/ES10-00200.1
Makhan, D. (2007) Pheidole soesilae sp. nov. from Suriname (Hymenoptera: Formicidae). Calodema Supplementary Paper, 59, 1–2.
Mann, W.M. (1916) The ants of Brazil. (The Stanford Expedition to Brazil, 1911, John C. Branner, Director). Bulletin of the Museum of Comparative Zoology, 60, 399–490, pls. 391–397.
Mayr, G. (1870a) Formicidae novogranadenses. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften in Wien. Mathematisch-Naturwissenschaftliche Klasse Abteilung I, 61, 370–417, Tafel.
Mayr, G. (1870b) Neue Formiciden. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien, 20, 939–996.
Mayr, G. (1887) Südamerikanische Formiciden. Verhandlungen der Kaiserlich-Königlichen Zoologisch-botanischen Gesellschaft in Wein, 37, 511–632.
Mertl, A., Sorenson, M. & Traniello, J. (2010) Community-level interactions and functional ecology of major workers in the hyperdiverse ground-foraging Pheidole (Hymenoptera, Formicidae) of Amazonian Ecuador. Insectes Sociaux, 57, 441–452.
https://doi.org/10.1007/s00040-010-0102-5
Mertl, A.L. & Traniello, J.F.A. (2009) Behavioral evolution in the major worker subcaste of twig-nesting Pheidole (Hymenoptera: Formicidae): does morphological specialization influence task plasticity? Behavioral Ecology and Sociobiology, 63, 1411–1426.
Moreau, C.S. (2008) Unraveling the evolutionary history of the “hyperdiverse” ant genus Pheidole (Hymenoptera: Formicidae). Molecular Phylogenetics and Evolution, 48, 224–239.
https://doi.org/10.1016/j.ympev.2008.02.020
O'Hara, R.J. (1993) Systematic generalization, historical fate, and the species problem. Systematic Biology, 42, 231–246.
https://doi.org/10.1093/sysbio/42.3.231
Pie, M.R. & Traniello, J.F.A. (2007) Morphological evolution in a hyperdiverse clade: the ant genus Pheidole. Journal of Zoology, 271, 99–109.
https://doi.org/10.1111/j.1469-7998.2006.00239.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
Risch, S., McClure, M., Vandermeer, J. & Waltz, S. (1977) Mutualism between three species of tropical Piper (Piperaceae) and their ant inhabitants. American Midland Naturalist, 98, 433–444.
https://doi.org/10.2307/2424991
Roger, J. (1863) Die neu aufgeführten Gattungen und Arten meines Formiciden-Verzeichnisses nebst Ergänzung einiger früher gegebenen Beschreibungen. Berliner Entomologische Zeitschrift, 7, 131–214.
https://doi.org/10.1002/mmnd.18630070116
Santschi, F. (1923) Pheidole et quelques autres fourmis néotropiques. Annales de la Société Entomologique de Belgique, 63, 45–69.
Santschi, F. (1929) Nouvelles fourmis de la République Argentine et du Brésil. Anales de la Sociedad Cientifica Argentina, 107, 273–316.
Sarnat, E.М., Fischer, G., Guénard, B. & Economo, E.P. (2015) Introduced Pheidole of the world: taxonomy, biology and distribution. ZooKeys, 543, 1–109.
https://doi.org/10.3897/zookeys.543.6050
Shorthouse, D.P. (2010) SimpleMappr, an online tool to produce publication-quality point maps. Available from: http://www.simplemappr.net. (accessed 3 December 2018)
Tepe, E., Vincent, M. & Watson, L. (2004) Phylogenetic patterns, evolutionary trends, and the origin of ant-plant associations in Piper section Macrostachys: Burger’s hypothesis revisited. In: L. A. Dyer & A. D. N. Palmer (Eds), Piper: a model genus for studies of phytochemistry, ecology, and evolution. Kluwer Academic Publishing, New York, pp. 156–178.
https://doi.org/10.1007/978-0-387-30599-8_9
Tschá, M.K. & Pie, M.R. (2018) Correlates of ecological dominance within Pheidole ants (Hymenoptera: Formicidae). Ecological Entomology, 44, 163–171.
https://doi.org/10.1111/een.12685.
Waxman, H.K., Muscedere, M.L. & Traniello, J.F. (2017) Behavioral performance and neural systems are robust to sensory injury in workers of the ant Pheidole dentata. Brain, behavior and evolution, 89, 195–208.
https://doi.org/10.1159/000470899
Wheeler, W.M. (1922) The ants of Trinidad. American Museum Novitates, 45, 1–16.
Wheeler, W.M. (1925) Neotropical ants in the collections of the Royal Museum of Stockholm. Arkiv för Zoologi, 17A (8), 1–55.
Wheeler, W.M. (1934) Neotropical ants collected by Dr. Elisabeth Skwarra and others. Bulletin of the Museum of Comparative Zoology, 77, 157–240.
Wilson, E.O. (2003) Pheidole in the New World: A Dominant, Hyperdiverse Ant Genus. Cambridge, Mass, Harvard University Press, 794 pp.