Morphological and molecular data from two prior publications are combined with new data resulting in a mostly well-supported cladistic hypothesis of relationships among subtribes and genera of Ithomiini, a diverse clade of Neotropical butterflies. A revised classification for relationships among the nine tribes and 46 genera comprising the group is proposed, and Athesitina new subtribe is described for Athesis Doubleday, 1847 and Patricia Fox, 1940. Oxapampa Brabant, 2004 is synonymized with Veladyris Fox, 1945. © 2014 The Trustees of the Natural History Museum, London.122133147DEB 1241056; NSF; National Science FoundationAckery, P.R., The danaid genus Tellervo (Lepidoptera:Nymphalidae)- A cladistic approach (1987) Zoological Journal Ofthe Linnean Society, 89, pp. 203-274Ackery, P.R., Hostplants and classification: A review ofnymphalid butterflies (1988) Biological Journal of the LinneanSociety, 33, pp. 95-203Ackery, P.R., De Jong, R., Vane-Wright, R.I., The butterflies:Hedyloidea, Hesperioidea and Papilionoidea (1999) Handbook OfZoology, 4 (35), pp. 263-300. , KRISTENSEN, N. P., Ed. Lepidoptera, Moths and Butterflies.1. Evolution, Systematics and Biogeography Lepidoptera. de Gruyter, Berlin, GermanyBaker, R.H., Desalle, R., Multiple sources of characterinformation and the phylogeny of Hawaiian Drosophil (1997) Systematic Biology, 46, pp. 654-673Bates, H.W., (1862) Contributions to An Insect Fauna of the AmazonValley, 23, pp. 495-566. , Lepidoptera: Heliconidae. Transactions of theLinnean SocietyBeccaloni, G.W., Ecology, natural history, and behaviourof ithomiine butterflies and their mimics in Ecuador (Lepidoptera:Nymphalidae: Ithomiinae) (1997) Tropical Lepidoptera, 8, pp. 103-124Beccaloni, G.W., Vertical stratification of ithomiine butterfly(Nymphalidae: Ithomiinae) mimicry complexes: The relationshipbetween adult flight height and larval host-plantheight (1997) Biological Journal of the Linnean Society, 62, pp. 313-341Beccaloni, G.W., Viloria, A.L., Hall, S.K., Robinson, G.S., Catalogue of the Hostplants of the Neotropical Butterflies/Cat-alogo de las Plantas Hu-esped de las MariposasNeotropicales (2008) Zaragoza, Spain: Sociedad Entomol-ogica Aragonesa(SEA)/Red Iberoamericana de Biogeograf-?a y Entomolog-?aSistem-atica (RIBES)/Ciencia y Tecnolog-?a Para El Desarrollo(CYTED)/Natural History Museum, London, UK(NHM)/Instituto Venezolano de Investigaciones Cient-?ficas,Venezuela (IVIC)., 8, pp. 1-536. , M3m-Monograf-?as Tercer Milenio 1 fig, 3 tabsBrower, A.V.Z., Phylogenetic relationships among theNymphalidae (Lepidoptera) inferred from partial sequencesof the wingless gene (2000) Proceedings of the Royal Society OfLondon B, 267, pp. 1201-1211Brower, A.V.Z., The how and why of branch support andpartitioned branch support, with a new index to assess partitionincongruence (2006) Cladistics, 22, pp. 378-386Brower, A.V.Z., Desalle, R., Vogler, A.P., Genetrees, species trees and systematics: A cladistic perspective (1996) Annual Review of Ecology and Systematics, 27, pp. 423-450Brower, A.V.Z., Desalle, R., Patterns of mitochondrialversus nuclear DNA sequence divergence among nymphalidbutterflies: The utility of wingless as a source of characters forphylogenetic inference (1998) Insect Molecular Biology, 7, pp. 1-10Brower, A.V.Z., Freitas, A.V.L., Lee, M.-M., Silva-Brandao, K.L., Whinnett, A., Willmott, K.R., Phylogeneticrelationships among the Ithomiini (Lepidoptera: Nymphalidae)inferred from one mitchondrial and two nuclear generegions (2006) Systematic Entomology, 31, pp. 288-301Brower, A.V.Z., Jeansonne, M.M., Geographical populationsand subspecies of New World monarch butterflies(Nymphalidae) share a recent origin and are not phylogeneticallydistinct (2004) Annals of the Entomological Society of America, 97, pp. 519-523Brower, A.V.Z., Wahlberg, N., Ogawa, J.R., Boppre, M., Vane-Wright, R.I., Phylogenetic relationships amonggenera of danaine butterflies (Lepidoptera: Nymphalidae) asimplied by morphology and DNA sequences (2010) Systematicsand Biodiversity, 8, pp. 75-89Brower, L.P., Chemical defence in butterflies (1984) The Biology of Butterflies, 1, pp. 109-134. , ACKERY,P. R. & VANE-WRIGHT, R. Academic Press, London, UKBrown Jr., K.S., Centros de evolu̧c̃ao, ref-ugios quatern-arios e conserva̧c̃ao de patrim^onios gen (1977) Acta Amazonica, 7, pp. 75-137Brown, K.S.J.R., Historical and ecological factors in thebiogeography of aposematic neotropical butterflies (1982) AmericanZoologist, 22, pp. 453-471Brown Jr., K.S., Chemical ecology of dehydropyrrolizidinealkaloids in adult Ithomiinae (Lepidoptera, Nymphalidae) (1985) Revista Brasileira de Biologia, 44, pp. 435-460Brown Jr., K.S., Chemistry at the solenaceae/Ithomiinaeinterf (1987) Annals of the Missouri Botanical Garden, 74, pp. 359-397Brown Jr., K.S., Freitas, A.V.L., Juvenile stages ofIthomiinae: Overview and systematics (1994) Tropical Lepidoptera, 5, pp. 9-20Brown Jr., K.S., Henriques, S.A., Chemistry, co-evolution,and colonisation of Solanaceae leaves by Ithomiinaebutterflies (1991) Solanaceae III: Taxonomy, Chemistry, Evolution, pp. 51-68. , HAWKES, J.G., LESTER, R.N., NEE M. &ESTRADA, N., Eds London, Linnean Society of London, UKBrown Jr., K.S., Trigo, J.R., Francini, R.B., De Morais, A.B.B., Motta, P.C., Aposematic insects on toxic hostplants:coevolution, colonisation and chemical emancipation (1991) Plant-Animal Interactions, pp. 375-402. , PRICE, P.W., LEWINSOHN, T.W., FERNANDES, G.W. &BENSON, W.W., Eds John Wiley& Sons, New York, USAChazot, N., Willmott, K.R., Paola, G.S.E., Hill, R., Jiggins, C., Elias, M., Mutualistic mimicry andfiltering by altitude shape the structure of Andean butterflycommunities (2013) American Naturalist, , doi: 10.1086/674100Cho, S., Mitchell, A., Regier, J.C., Mitter, C., Poole, R.W., Friedlander, T.P., Zhao, S., A highly conservednuclear gene for low-level phylogenetics: Elongation factor1a recovers morphology-based tree for heliothine moths (1995) Molecular Biology and Evolution, 12, pp. 650-656Clark, A.H., The interrelationships of the several groupswithin the butterfly superfamily Nymphaloide (1947) Proceedingsof the Entomological Society of Washington, 49, pp. 148-149. , 192 (erratum)D'Almeida, R.F., Catalogo dos Ithomiidae Americanos(Lepidoptera) (1978) Centro de Recursos Audiovisuales, , UniversidadeFederal do Paran-a, Curitiba, BrazilDasmahapatra, K.K., Lamas, G., Simpson, F., Mallet, J., The anatomy of r 'suture zone' in Amazonian butterflies: Acoalescent-based test for vicariant geographic divergenceand speciation (2010) Molecular Ecology, 19, pp. 4283-4301De-Silva, D.L., Day, J.J., Elias, M., Willmott, K., Whinnett, A., Mallet, J., Molecular phylogenetics of the neotropicalbutterfly subtribe Oleriina (Nymphalidae: Danaini:Ithomiini) (2010) Molecular Phylogenetics and Evolution, 55, pp. 1032-1041Devries, P.J., Lande, R., Murray, D., Associations ofco-mimetic ithomiine butterflies on small spatial and temporalscales in a neotropical rainforest (1999) Biological Journal Ofthe Linnean Society, 67, pp. 73-85Ehrlich, P.R., The comparative morphology, phylogenyand higher classification of the butterflies (Lepidoptera: Papilionoidea) (1958) University of Kansas Science Bulletin, 39, pp. 305-370Elias, M., Gompert, Z., Jiggins, C., Willmott, K.R., Mutualistic interactions drive ecological niche convergencein a diverse butterfly community (2008) Public Library of ScienceBiology, 6, pp. 2642-2649Elias, M., Hill, R.I., Willmott, K.R., Dasmahaptra, K.K., Brower, A.V.Z., Mallet, J., Jiggins, C.D., Limitedperformance of DNA barcoding in a diverse community oftropical butterflies (2007) Proceedings of the Royal Society of London,B, 274, pp. 2881-2889Elias Joron, M., Willmott, K.R., Silva Brandao, K.L., Outof the Andes: Patterns of diversification in clearwing butterflies (2009) Molecular Ecology, 18, pp. 1716-1729Felder, C., Felder, R., Specimen faunae lepidopterologicaeriparum fluminis Negro superioris in Brasilia septentrionali (1862) Wiener Entomologische Monatschrift, 6, pp. 65-80Fox, R.M., A generic review of the Ithomiinae (Lepidoptera,Nymphalidae) (1940) Transactions of the American EntomologicalSociety, 66, pp. 161-207Fox, R.M., A monograph of the Ithomiidae (Lepidoptera) Part 1 (1956) Bulletin of the American Museum of Natural History, 111, pp. 1-76. , 71-79Freitas, A.V.L., Brown Jr., K.S., Phylogeny of theNymphalidae (Lepidoptera) (2004) Systematic Biology, 53, pp. 363-383Freitas, A.V.L., Trigo, J.R., Brown Jr., K.S., Witte, L., Hartmann, T., Barata, L.E.S., Tropane and pyrrolizidinealkaloids in the ithomiines Placidula euryanassa andMiraleria cymothoe (Lepidoptera: Nymphalidae) (1996) Chemoecology, 7, pp. 61-67Goloboff, P.A., Extended implied weighting (2013) Cladistics(early View), , Article first published online: 19 JUL 2013 jdoi: 10.1111/cla.12047Goloboff, P.A., Farris, J.S., Kallersjo, M., Oxelman, B., Ramirez, M.J., Szumik, C.A., Improvements to resamplingmeasures of group support (2003) Cladistics, 19, pp. 324-332Goloboff, P.A., Farris, J.S., Nixon, K.C., TNT: A freeprogram for phylogenetic analysis (2008) Cladistics, 24, pp. 774-786Hill, R.I., Elias, M., Dasmahapatra, K.K., Koongv, D.J.C., Willmott, K.R., Mallet, J., Ecologically relevantcryptic species in the highly polymorphic Amazonianbutterfly Mechanitis mazaeus sensu lato (Lepidoptera: Nymphalidae;Ithomiini) (2012) Biological Journal of the Linnean Society106, pp. 540-560Jiggins, C.D., Mallarino, R., Willmott, K.R., Bermingham, E., The phylogenetic pattern of speciation and wing patternchange in neotropical Ithomia butterflies (Lepidoptera;Nymphalidae) (2006) Evolution, 60, pp. 1454-1466Lamas, G., Atlas of neotropical lepidopter (2004) Checklist:Part 4A Hesperioidea-Papiionoidea, , Scientific Publishers/Association of Tropical Lepidoptera, Gainesville,Florida, USALewis, P.O., A likelihood approach to estimating phylogenyfrom discrete morphological character dat (2001) SystematicBiology, 50, pp. 913-925Mallarino, R., Bermingham, E., Willmott, K.R., Whinnett, A., Jiggins, C.D., Molecular systematics of the butterflygenus Ithomia (Lepidoptera: Ithomiinae): A composite phylogenetichypothesis based on seven genes (2005) Molecular Phylogeneticsand Evolution, 34, pp. 625-644Mielke, O.H.H., Brown Jr., K.S., (1979) Suplemento Aocat-alogo Dos Ithomiidae Americanos (Lepidoptera) DeRomualdo Ferreira d'Almeida, , Centro de Recursos Audiovisuales,Universidade Federal do Paran-a, Curitiba, BrazilMiller, J.S., Generic revision of the Dioptinae (Lepidoptera:Noctuoidea: Notodontidae) (2009) Bulletin of the AmericanMuseum of Natural History, p. 321. , 1-971 \+ 48 platesMuller, F., Ituna and Thyridia: A remarkable case of mimicryin butterflies (1879) Proceedings of the Entomological Societyof London, pp. 20-29Nixon, K.C., The parsimony ratchet, a new method forrapid parsimony analysis (1999) Cladistics, 15, pp. 407-414Nixon, K.C., Carpenter, J.M., On simultaneous analysis (1996) Cladistics, 12, pp. 221-241Rindal, E., Brower, A.V.Z., Do model-based phylogeneticanalyses perform better than parsimony? A test withempirical dat (2011) Cladistics, 27, pp. 331-334Ronquist, F., Huelsenbeck, J.P., MrBayes 3: Bayesianphylogenetic inference under mixed models (2003) Bioinformatics, 19, pp. 1572-1574Ronquist, F., Teslenko, M., Mark, P., Ayres, D., Darling, A., Hohna, S., Larget, B., Huelsenbeck, J.P., MrBayes 3.2: Efficient Bayesian phylogeneticinference and model choice across a large model space (2012) Systematic Biology, 61, pp. 539-542Swofford, D.L., (2000) PAUPPhylogenetic Analysis Using Parsimony(-and Other Methods), , Sinauer Associates, Sunderland,Massachusetts, USAUehara-Prado, M., Freitas, A.V.L., The effect of rainforestfragmentation on species diversity and mimicry ringcomposition of ithomiine butterflies (2009) Insect Conservationand Diversity, 2, pp. 23-28Wahlberg, N., Weingartner, E., Nylin, S., Towards abetter understanding of the higher systematics of Nymphalidae(Lepidoptera: Papilionoidea) (2003) Molecular Phylogeneticsand Evolution, 28, pp. 473-484Wahlberg, N., Leneveu, J., Kodandaramaiah, U., Pena, C., Nylin, S., Freitas, A.V.L., Brower, A.V.Z., Nymphalidbutterflies diversify following near demise at the Cretaceous/Tertiary boundary (2009) Proceedings of the Royal Societyof London B, 276, pp. 4295-4302Willmott, K.R., Freitas, A.V.L., Higher-level phylogenyof the Ithomiinae (Lepidoptera: Nymphalidae): Classification,patterns of larval hostplant colonization anddiversification (2006) Cladistics, 22, pp. 297-368Willmott, K.R., Lamas, G., A phylogenetic reassessmentof Hyalenna Forbes and Dircenna Doubleday, with a revisionof Hyalenna (Lepidoptera: Nymphalidae: Ithomiinae) (2006) Systematic Entomology, 31, pp. 419-468Willmott, K.R., Lamas, G., A revision of Pachacutia, anew genus of rare Andean ithomiine butterflies (Nymphalidae:Ithomiinae), with the description of two new species (2007) Annals of the Entomological Society of America, 100, pp. 449-469Willmott, K.R., Mallet, J., Correlations between adultmimicry and larval hostplants in ithomiine butterflies (2004) Proceedingsof the Royal Society of London B (Biology LettersSuppl.), 271, pp. S266-S269Zwickl, D.J., (2006) Genetic Algorithm Approaches for the Phylogeneticanalysis of Large Biological Sequence Datasets Undera Maximum Likelihood Criterion, , Ph.D. dissertation, The Universityof Texas at Austin, Austin, Texas, USA