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The images displayed here depict examples of the major butterfly subfamilies occurring in Ecuador and principal habitats. For any use other than talks or educational material that is supplied free of charge, permission must be sought from K.R. Willmott or J.P.W. Hall prior to use. Copyright to all images is currently held by K.R. Willmott and J.P.W. Hall.

IMMATURE STAGES
  Ithomiinae

ADULT BUTTERFLIES
  Papilionidae
  Pieridae
  Nymphalidae
  Nymphalinae
  Charaxinae
  Biblidinae
  Limenitidinae
  Danainae
  Ithomiinae
  Heliconiinae
  Morphinae
  Brassolinae
  Satyrinae

HABITATS
  Dry forest
  Moist forest
  Northwest lowland pluvial forest
  West submontane forest (1000-2500m)
  West upper montane forest (>2500m)
  Páramo
  Eastern lowland wet forest
  Eastern foothills
  Eastern submontane (1000-2500m)
  Eastern upper montane (>2500m)
  Highland valleys

 

BIBLIOGRAPHY AND REPRINTS

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Listed here are all papers by us relevant to the taxonomy and ecology of Ecuadorian butterflies. Following this is a list of additional papers of particular relevance by other authors, including papers on butterfly taxonomy and biology, and conservation issues in Ecuador. Please note that the latter list is not comprehensive, and will be continually updated with both old and new references.

BY JASON HALL AND KEITH WILLMOTT:

2019. Nakahara, S., Barbosa, E. P., Willmott, K. R., Lamas, G., Freitas, A. V. L. Two new species of Taygetina Forster, 1964 from Ecuador, Peru and Brazil with a possible case of "juxta loss" in butterflies (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity 3(6): 9; 1-13. DOI: 10.1093/isd/ixz023 (online 20 Nov 2019).

2019. Nakahara, S., Zacca, T., Dias, F. M. S., Dolibaina, D. R., Xiao, L., Espeland, M., Casagrande, M. M., Mielke, O. H. H., Lamas, G., Huertas, B., Kleckner, K., Willmott, K. R. Revision of the poorly known Neotropical butterfly genus Zischkaia Forster, 1964 (Lepidoptera, Nymphalidae, Satyrinae), with descriptions of nine new species. European Journal of Taxonomy 551: 1-67. DOI: 10.5852/ejt.2019.551 (online 12 Sep 2019).

2019. Brévignon, C., Rosant, T., Lamas, G., Tyler, S., Willmott, K. R. Description d’une nouvelle espèce d'Euptychiina du genre Chloreuptychia (Lepidoptera, Nymphalidae, Satyrinae). Bulletin de la Société Entomologique de France 124(2): 127-138. DOI: 10.32475/bsef_2044 (June 2019).

2019. Chazot, N., Willmott, K. R., Lamas, G., Freitas, A., Piron-Prunier, F., Arias, C., Mallet, J., De Silva, D. L., Elias, M. Renewed diversification following Miocene landscape turnover in a Neotropical butterfly radiation. Global Ecology and Biogeography 2019; 1-15. DOI: 10.1111/geb.12919 (online 10 May 2019).

2019. Willmott, K. R., Marín, M. A., Nakahara, S., Pomerantz, T., Lamas, G., Espeland, M., Xiao, L., Hall, J. P. W., Robinson Willmott, J. I., Freitas, A. V. L. A revision of the new Andean butterfly genus Optimandes Marín, Nakahara & Willmott, n. gen., with the description of a new species (Nymphalidae: Satyrinae: Euptychiina). Tropical Lepidoptera Research 29(1): 29-44 (online 3 May 2019).

2019. Torres, K. P., Artieda, N., Salazar, P., Willmott, K. R., Hill, R. I. 2019. Life history descriptions of Adelpha attica attica, Adelpha epione agilla, and Adelpha jordani from an eastern Ecuador lowland forest. Tropical Lepidoptera Research 29(1): 19-28 (online 3 May 2019).

2019. Nakahara, S., Lamas, G., Tyler, S., Marín, M., Huertas, B., Willmott, K. R., Mielke, O. H. H., Espeland, M. 2019. A revision of the new genus Amiga Nakahara, Willmott & Espeland, gen. n., described for Papilio arnaca Fabricius, 1776 (Lepidoptera, Nymphalidae, Satyrinae). ZooKeys 821: 85-152. DOI: 10.3897/zookeys.821.31782 (31 Jan 2019).

2018. Espeland, M., Breinholt, J., Barbosa, E., Casagrande, M. M., Huertas, B., Lamas, L., Marín, M. A., Mielke, O. H. H., Miller, J. Y., Nakahara, S., Tan, D., Warren, A. D., Zacca, T., Kawahara, A., Freitas, A. V. L., Willmott, K. R. Four hundred shades of brown: Higher level phylogeny of the problematic Euptychiina (Lepidoptera, Nymphalidae, Satyrinae) based on hybrid enrichment data. Molecular Phylogenetics and Evolution 131: 116-124, DOI: 10.1016/j.ympev.2018.10.039 (online 10 Nov 2018).

2018. Checa, M. F., Donoso, D. A., Rodríguez, J., Levy, E., Warren, A., Willmott, K. R. Combining sampling techniques aids monitoring of tropical butterflies. Insect Conservation and Diversity DOI: 10.1111/icad.12328 (online 29 Oct 2018).

2018. Dufour, P. C., Willmott, K. R., Padrón, P. S., Xing, S., Bonebrake, T. C., Scheffers, B. R. Divergent melanism strategies in Andean butterfly communities structure diversity patterns and climate responses. Journal of Biogeography 45: 2471-2482. DOI: 10.1111/jbi.13433 (online 4 October 2018).

2018. Rosser, N., Freitas, A. V. L., Huertas, B., Joron, M., Lamas, G., Mérot, C., Simpson, F., Willmott, K. R., Mallet, J., Dasmahapatra, K. A. Cryptic speciation associated with geographic and ecological divergence in two Amazonian Heliconius butterflies. Zoological Journal of the Linnean Society XX: 1-17 DOI: 10.1093/zoolinnean/zly046 (online 4 August 2018).

2018. Dias, F. M. S., Janzen, D., Hallwachs, W., Chacón, I., Willmott, K. R., Ortiz-Acevedo, E., Mielke, O. H. H., Casagrande, M. M. DNA Barcodes uncover hidden taxonomic diversity behind the variable wing patterns in the Neotropical butterfly genus Zaretis (Lepidoptera: Nymphalidae: Charaxinae). Zoological Journal of the Linnean Society XX: 1-61. DOI: 10.1093/zoolinnean/zly036 (online 1 August 2018).

2018. Nakahara, S., Willmott, K. R., Mielke, O. H. H., Schwartz, J., Zacca, T., Espeland, M., Lamas, G. Seven new taxa from the butterfly subtribe Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) with revisional notes on Harjesia Forster, 1964 and Pseudeuptychia Forster, 1964. Insecta Mundi 0639: 1-38 (online 27 July 2018).

2018. Willmott, K. R., Lamas, G., Radford, J., Marín, M. A., Nakahara, S., Espeland, M., Xiao, L., Hall, J. P. W. A distinctive new species of cloud forest Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) from Ecuador and Peru. Tropical Lepidoptera Research 28(1): 39-45 (online 13 July 2018).

2018. Zacca, T., Casagrande, M. M., Mielke O. H. H., Huertas, B., Barbosa E. P., Freitas, A. V. L., Magaldi, L. M., Espeland, M., Nakahara, S., Willmott, K. R. Systematics of the butterfly genus Cissia Doubleday, 1848 (Lepidoptera: Nymphalidae, Satyrinae) using an integrative approach. Arthropod Systematics and Phylogeny, 76(2): 349-376 (June).

2018. Benmesbah, M., Zacca, T., Casagrande, M. M., Mielke, O. H. H., Lamas, G., Willmott, K. R. Taxonomic notes on Papilio ocypete Fabricius, 1776 and Papilio helle Cramer, 1779 with description of two new similar species from South America (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa 4425(1): 115-145 (May).

2018. Finkbeiner, S. D., Salazar-Carrión, P., Nogales, S., Rush, C., Briscoe, A. D., Hill, R., Kronforst, M., Willmott, K. R., Mullen, S. P. Frequency-dependence shapes the adaptive landscape of imperfect Batesian mimicry. Proceedings of the Royal Society B 285: 20172786 (online 4 April 2018).

2018. Chazot, N., De Silva, D. L., Willmott, K. R., Freitas, A. V. L., Lamas, G., Mallet, J., Giraldo, C. E., Uribe, S., Elias, M. Contrasting patterns of Andean diversification among three diverse clades of Neotropical clearwing butterflies. Ecology and Evolution, 2017; 1-18, DOI: 10.1002/ece3.3622 (online 25 March 2018).

2018. Hall, J. P. W. A Monograph of the Nymphidiina (Lepidoptera: Riodinidae: Nymphidiini): Phylogeny, Taxonomy, Biology, and Biogeography. Washington, D.C., Entomological Society of Washington.990 pp., illustrations (some color), maps.

2017. Ortiz-Acevedo, E., Bonfantti, D., Casagrande, M., Mielke, O. H. H., Espeland, M., Willmott, K. R. Using molecules and morphology to unravel the systematics of Neotropical preponine butterflies (Lepidoptera: Charaxinae: Preponini). Insect Systematics and Diversity, 1: 48-56, DOI: 10.1093/isd/ixx002 (online 25 October).

2017. Nakahara, S., Zacca, T., Huertas, B., Neild, A. F. E., Hall, J. P. W., Lamas, G., Holian, L. A., Espeland, M., Willmott, K. R. Remarkable sexual dimorphism, rarity and cryptic species: a revision of the ‘aegrota species group’ of the Neotropical butterfly genus Caeruleuptychia Forster, 1964 with the description of three new species (Lepidoptera, Nymphalidae, Satyrinae). Insect Systematics and Evolution, 1-53, DOI: 10.1163/1876312X-00002167 (online 31 July).

2017. Willmott, K. R., Lamas, G., Hall, J. P. W. Notes on the taxonomy of Actinote intensa Jordan (Lepidoptera: Nymphalidae: Heliconiinae) and the description of a new sibling species from eastern Ecuador. Tropical Lepidoptera Research, 27(1): 6-15 (June).

2017. Willmott, K. R., Simon, M., Ortiz-Acevedo, E., Hall, J. P. W. First record of the enigmatic tribe Anaeomorphini (Lepidoptera, Nymphalidae, Charaxinae) outside of the Amazon basin: a new species of Anaeomorpha Rothschild, 1894, from the Chocó region of western Ecuador. Insecta Mundi, 0548: 1-10 (May).

2017. Willmott, K. R., Robinson Willmott, J. C., Elias, M., Jiggins, C. Maintaining mimicry diversity: optimal warning color patterns differ among microhabitats in Amazonian clearwing butterflies. Proceedings of the Royal Society B, 284: 20170744. DOI: 10.1098/rspb.2017.0744 (May).

2017. De Silva, D. L., Mota, L., Chazot, N., Mallarino, R., Silva-Brandão, K. L., Gómez P., L. M., Freitas, A. V. L., Lamas, G., Joron, M., Mallet, J., Giraldo S., C. E., Uribe S., S. I., Särkinen, T., Knapp, S., Jiggins, C. D., Willmott, K. R., Elias, M. 2017. North Andean origin and diversification of the largest ithomiine butterfly genus. Scientific Reports, 7: 45966, DOI: 10.1038/srep45966 (April).

2016. Nakahara, S., Tan, D., Lamas, G., Parus, A., Willmott, K. R. 2016. A distinctive new species of Hermeuptychia Forster, 1964 from the eastern tropical Andes (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera Research, 26: 77-84 (December).

2016. Nakahara, S., Vega, G., Willmott, K. R. Description of a new species of Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the western Andes. Zootaxa, 4184: 358-366. DOI: 10.11646/zootaxa.4184.2.7 (November).

2016. Huertas, B., Lamas, G., Fagua, G., Mallet, J., Nakahara, S., Willmott, K. R. A remarkable new butterfly species from western Amazonia (Lepidoptera: Nymphalidae: Satyrinae). Conservación Colombiana, 24: 1-5. (October).

2016. Chazot, N., Willmott, K. R., Condamine, F., de Silva, D. L., Freitas, A. V. L., Lamas, G., Morlon, H., Giraldo, C. E., Jiggins, C. D., Joron, M., Mallet, J., Uribe, S., Elias, M. Into the Andes: multiple independent colonizations drive montane diversity in the Neotropical clearwing butterflies Godyridina. Molecular Ecology DOI: 10.1111/mec.13773 (October).

2016. Chazot, N., Willmott, K. R., Freitas, A. V. L., De Silva, D. L., Pellens, R., Elias. Patterns of Species, Phylogenetic and Mimicry Diversity of Clearwing Butterflies in the Neotropics. In: Pellens, R., Grandcolas, P. (Eds.) Biodiversity Conservation and Phylogenetic Systematics. Topics in Biodiversity and Conservation 14, DOI: 10.1007/978-3-319-22461-9_17. Pp. 333-353.

2016. Nakahara, S., Barbosa, E. P., Marín, M. A., Freitas, A. V. L., Pomerantz, T., Willmott, K. R. Graphita gen. nov., a new genus for Neonympha griphe C. Felder & R. Felder, 1867 (Lepidoptera, Nymphalidae, Satyrinae). Neotropical Entomology, 17 pp, doi: 10.1007/s13744-016-0414-x (June).

2016. Freitas, A. V. L., Barbosa, E., Willmott, K. R., Wahlberg, N., Lamas, G. 'Species' from three different butterfly genera combined into one: description of a new genus of Euptychiina (Nymphalidae: Satyrinae) with unusually variable wing pattern. Revista Brasileira de Entomologia, doi:10.1016/j.rbe.2016.01.004 (March).

2015. Nakahara, S., Hall, J. P. W., Lamas, G., Willmott, K. R. Seven new species and one new subspecies of Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the tropical Andes. Tropical Lepidoptera Research 25(2): 63-79 (December).

2015. De Silva, D. L., Elias, M., Willmott, K. R., Mallet, J., Day, J. J. Diversification of clearwing butterflies with the rise of the Andes. Journal of Biogeography doi:10.1111/jbi.12611 (online September).

2015. Willmott, K. R. A new subspecies of Cithaerias pyropina (Salvin & Godman, 1868) from the Cordillera de Cutucú in southeastern Ecuador (Nymphalidae: Satyrinae). Tropical Lepidoptera Research, 25: 1-4 (July).

2015. Ebel, E. R., DaCosta, J. M., Sorenson, M. D., Hill, R. I., Briscoe, A. D., Willmott, K. R., Mullen, S. P. Rapid diversification associated with ecological specialization in Neotropical Adelpha butterflies. Molecular Ecology, 24: 2392-2405 (April).

2014. Checa, M. F., Rodríguez, J., Willmott, K. R., Liger, B. Microhabitat significantly affects the composition, abundance and phenology of butterfly assemblages in an Ecuadorian dry forest. Florida Entomologist, 97: 1-13 (March).

2014. Checa, M. F., Willmott, K. R. Hadas del Ecuador: una mirada a su diversidad. In: García, M., Parra, D. (Eds.) Ecuador: País de la Biodiversidad. Fundación Botánica de Los Andes/Ministerio del Ambiente/Fundación Ecofondo. Quito, Ecuador. Pp. 250-255.

2013. Chazot, N., Willmott, K. R., Paola, S. E., Toporov, A., Hill, R., Jiggins, C., Elias, M. Mutualistic mimicry and filtering by altitude shape the structure of Andean butterfly communities. American Naturalist, doi: 10.1086/674100, http://www.jstor.org/stable/10.1086/ 674100 (November).

2013. Willmott, K. R., Hall, J. P. W. A new species and two new subspecies of Adelpha Hübner, [1819] from the tropical Andes (Lepidoptera: Limenitidinae). Journal of the Lepidopterists’ Society, 67: 241-252 (November).

2013. Radford, J., Willmott, K. R. A distinctive new subspecies of Catasticta poujadei from the Cordillera del Cóndor in eastern Ecuador (Lepidoptera: Pieridae: Pierinae). Tropical Lepidoptera Research, 23: 86-91 (November).

2013. Pyrcz, T. W., Willmott, K. R., Garlacz, R., Boyer, P., Gareca, Y. Latitudinal gradient and spatial covariance in species richness of tropical Lepidoptera in the Andes. Insect Diversity and Conservation, doi: 10.1111/icad.12058 (November).

2013. Ortíz-Acevedo, E., Willmott, K. R. Molecular systematics of the butterfly tribe Preponini (Nymphalidae: Charaxinae). Systematic Entomology, DOI: 10.1111/syen.12008 (February).

2012. Matos-Maraví, P. F., Peña, C., Willmott, K. R., Freitas, A. V. L., Wahlberg, N. Systematics and evolutionary history of butterflies in the “Taygetis clade” (Nymphalidae: Satyrinae: Euptychiina): towards a better understanding of Neotropical biogeography. Molecular Phylogenetics and Evolution, 66: 54-68 (November). DOI: 10.1016/j.ympev.2012.09.005.

2012. Hill, R. I., Elias, M., Dasmahapatra, K. K., Jiggins, C. D., Koong, V., Willmott, K. R., Mallet, J. Ecologically relevant cryptic species in the highly polymorphic Amazonian butterfly Mechanitis mazaeus sensu lato (Lepidoptera: Nymphalidae; Ithomiini). Biological Journal of the Linnean Society, 106: 540-560 (June). DOI: 10.1111/j.1095-8312.2012.01874.x

2012. Rosser, N., Phillimore, A. B., Huertas, B., Willmott, K. R., Mallet, J. Testing historical explanations for gradients in species richness in heliconiine butterflies of tropical America. Biological Journal of the Linnean Society, 105: 479-497 (January) DOI: 10.1111/j.1095-8312.2011.01814.x.

2011. Willmott, K. R., Elias, M., Sourakov, A. Two possible caterpillar mimicry complexes in neotropical danaine butterflies (Nymphalidae, Danainae: Danaini, Ithomiini). Annals of the Entomological Society of America, 104: 1108-1118 (December).

2011. Bolaños, I. A., Zambrano, G., Willmott, K. R. Descripción de los estados inmaduros de Pteronymia zerlina zerlina, P. zerlina machay, P. medellina y P. veia n. ssp. (Lepidoptera: Nymphalidae: Ithomiini) de Colombia y del Ecuador. Tropical Lepidoptera Research, 21: 27-33 (June).

2011. Pyrcz, T., Greeney, H. F., Willmott, K. R., Wojtusiak, J. A taxonomic revision of the genus Daedalma Hewitson with the descriptions of twenty new taxa and the immature stages of two species (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa, 2898: 1-68 (May).

2011. Gompert, Z., Willmott, K. R., Elias, M. 2011. Heterogeneity in predator micro-habitat use and the maintenance of Müllerian mimicry diversity. Journal of Theoretical Biology, 281: 39-46 (April).

2010. Mullen, S. P., Savage, W., and K. R. Willmott Rapid diversification and not clade age explains high diversity in neotropical Adelpha butterflies. Proceedings of the Royal Society of London B doi:10.1098/rspb.2010.2140

2010. Hall, J. P. W., and K. R. Willmott. Description of a new Lucillella species (Riodinidae: Symmachiini) discovered in the eastern Andes of Ecuador using the single rope canopy access technique. Journal of the Lepidopterists' Society, 64: 139-146. Download high (12 Mb) or low (4.7 Mb) resolution reprint [PDF].

2010. Willmott, K. R., and J. P. W. Hall. A new species of Dynamine Hübner, [1819] from northwestern Ecuador (Lepidoptera: Nymphalidae: Biblidinae). Tropical Lepidoptera Research, 20: 23-27. Download reprint [PDF]

2010. Hall, J. P. W., and D. H. Ahrenholz. A new species of Alesa (Riodinidae: Eurybiini) from eastern Ecuador. Tropical Lepidoptera Research, 20: 19-22.

2010. de-Silva, D. L., Day, J. J., Elias, M., Willmott, K. R., Whinnett, A., and J. Mallet. Molecular phylogenetics of the neotropical butterfly subtribe Oleriina (Nymphalidae: Danainae: Ithomiini). Molecular Phylogenetics and Evolution, 55:1032-1041. Download reprint [PDF]

2009. Elias, M., Gompert, Z., Jiggins, C., and K. R. Willmott. Phylogenetic community ecology needs to take positive interactions into account: Insights from colourful butterflies. Communicative & Integrative Biology 2(2): in press [Article addendum]

2009. Hall, J. P. W., and K. R. Willmott. Two new species of Euselasia (Riodinidae: Euselasiinae) from western Ecuador. Tropical Lepidoptera Research, 19(1): 52-55. Download reprint [PDF]

2009. Elias, M., Joron, M., Willmott, K. R., Kaiser, V., Silva-Brandão, K. L., Freitas, A. V. L., Arias, C. F., Gomez Piñeres, L. M., Uribe, S., and C. Jiggins. Out of the Andes: patterns of diversification in clearwing butterflies. Molecular Ecology, 18: 1716-1729. Download page proofs [PDF]

2009. Willmott, K. R., Freitas, A. V. L., Hall, J. P. W., Silva-Brandão, K. L., Paluch, M. A new species of Actinote Hübner from the eastern Andes of Ecuador (Lepidoptera: Nymphalidae: Heliconiinae). Proceedings of the Entomological Society of Washington, 111(1): 47-56. Download reprint [PDF] - NOTE: figures in published version are monochrome

2008. Willmott, K. R., Vitale, F. Taxonomic notes on Napeogenes (Lepidoptera: Ithomiinae) from Ecuador and Colombia, with the description of ten new subspecies. Genus, 19(3): 585-609. Download reprint [PDF]

2008. Willmott, K. R., Lamas, G. A revision of the genus Megoleria (Lepidoptera, Nymphalidae, Ithomiinae). Tropical Lepidoptera Research, 18(1): 46-59.Download reprint [PDF]

2007. Elias, M., Hill, R., Willmott, K. R., Dasmahapatra, K., Brower, A., Mallet, J., Jiggins, C. Limited performance of DNA barcoding in a diverse community of tropical butterflies. Proceedings of the Royal Society of London B, 274: 2881-2889.Download reprint [PDF]

2007. Brehm, G., Hartmann, T., and K. R. Willmott. Pyrrolizidine alkaloids and pharmacophagous Lepidoptera visitors of Prestonia amabilis (Apocynaceae) in a montane rain forest in Ecuador. Annals of the Missouri Botanical Garden, 94(2): 463-473.Download reprint [PDF]

2007. Willmott, K. R., Lamas, G. A revision of Pachacutia, a new genus of rare Andean ithomiine butterflies (Nymphalidae: Ithomiinae), with the description of two new species. Annals of the Entomological Society of America, 100: 449-469.

2007. Hall, J. P. W., Willmott, K.R. Four new species of Symmachiini from Ecuador (Lepidoptera: Riodinidae). Tropical Lepidoptera, 16(1-2): 1-5.Download reprint [PDF]

2007. Hall, J. P. W. A phylogenetic revision of the new riodinid butterfly genus Minstrellus (Lepidoptera: Riodinidae). Annals of the Entomological Society of America, 100: 773-

2007. Hall, J. P. W., Harvey, D. J. Three new species in the riodinid tribe Symmachiini from Panama and Colombia (Lepidoptera: Riodinidae). Tropical Lepidoptera, 16: 7-10.

2007. Hall, J. P. W. Two new genera in the Nymphidiina (Lepidoptera: Riodinidae: Nymphidiini). Zootaxa, 1415: 35-42.

2007. Hall, J. P. W., Lamas, G. Four new symmachiine taxa from the eastern Andes of Peru and Ecuador (Lepidoptera: Riodinidae). Zootaxa, 1533: 41-52.

2006. Pyrcz, T. W., Willmott, K. R., Hall, J. P. W., and A. L. Viloria. A review of the genus Manerebia Staudinger (Lepidoptera: Nymphalidae: Satyrinae) in the northern Andes. Journal of Research on the Lepidoptera, 39: 37-79.Download reprint [PDF]

2006. Jiggins, C. D., Mallarino, R., Willmott, K. R., and E. Bermingham. Phylogenetic evidence for speciation caused by ecological adaptation in neotropical Ithomia butterflies (Lepidoptera; Nymphalidae). Evolution, 60: 1454-1466.Download reprint [PDF]

2006. Willmott, K. R., and A. V. L. Freitas. Higher-level phylogeny of the Ithomiinae (Lepidoptera: Nymphalidae): classification, patterns of larval hostplant colonisation and diversification. Cladistics, 22: 297-368.

2006. Willmott, K. R., and G. Lamas. A phylogenetic reassessment of Hyalenna Forbes and Dircenna Doubleday, with a revision of Hyalenna (Lepidoptera: Nymphalidae: Ithomiinae). Systematic Entomology, 31(3): 419-468. Download reprint [PDF]

2006. Brower, A. V. Z., Freitas, A. V. L., Lee, M.-M., Silva Brandão, K. L., Whinnett, A., and K. R. Willmott. Phylogenetic relationships among the Ithomiini (Lepidoptera: Nymphalidae) inferred from one mitochondrial and two nuclear gene regions. Systematic Entomology, 31: 288-301. Download reprint [PDF]

2005. Hall, J. P. W. A Phylogenetic Revision of the Napaeina (Lepidoptera: Riodinidae: Mesosemiini). Entomological Society of Washington, Washington, DC. 232 pp.

2005. Hall, J. P. W., and K. R. Willmott. A new species of Paiwarria (Lepidoptera: Lycaenidae: Eumaeini) from western Ecuador. Proceedings of the Entomological Society of Washington, 107(4): 960-967.

2005. Hall, J. P. W. Montane speciation patterns in Ithomiola butterflies (Lepidoptera: Riodinidae): are they consistently moving up in the World? Proceedings of the Royal Society of London B, 272: 2457-2466. Download reprint [PDF]

2005. Mallarino, R., Bermingham, E., Willmott, K. R., Whinnett, A., and C. D. Jiggins. Molecular systematics of the butterfly genus Ithomia (Lepidoptera: Ithomiinae): a composite phylogenetic hypothesis based on seven genes. Molecular Phylogenetics and Evolution, 34(3): 625-644. Download reprint [PDF]

2005. Hall, J. P. W., K. R. Willmott and R. C. Busby. Five new Penaincisalia species (Lepidoptera: Lycaenidae: Eumaeini) from the Andes of southern Ecuador and northern Peru. Zootaxa, 797: 1-20. Download reprint [PDF]

2005. Hall, J. P. W. A review of the Metacharis syloes group (Lepidoptera: Riodinidae), with the description of two new west Andean species. Proceedings of the Entomological Society of Washington, 107(1): 200-208. Download reprint [PDF]

2004a. Willmott, K. R., and J. Mallet. Correlations between adult mimicry and larval hostplants in ithomiine butterflies. Proc. R. Soc. Lond. B (Suppl.) 271: S266-S269. Download reprint [PDF]

2004b. Hall, J. P. W. The Genus Adelpha: its Systematics, Biology and Biogeography (Lepidoptera: Nymphalidae: Limenitidini). by Keith R. Willmott. Proceedings of the Entomological Society of Washington, 106(3): 728-730. [book review] Download reprint [PDF]

2004c. Hall, J. P. W., D. J. Harvey, and D. H. Janzen. Life history of Calydna sturnula with a review of larval and pupal balloon setae in the Riodinidae. Annals of the Entomological Society of America, 97(2): 310-321. Download reprint [PDF]

2004d. Hall, J. P. W., R. K. Robbins, and D. J. Harvey. Extinction and biogeography in the Caribbean: new evidence from a fossil riodinid butterfly in Dominican amber. Proceedings of the Royal Society of London B, 271(1541): 797-801. Download reprint [PDF]

2004e. Hall, J. P. W., and G. Lamas. A new cloud forest species of Calydna (Lepidoptera: Riodinidae) from Peru, with a revised phylogeny of the C. hiria group. Proceedings of the Entomological Society of Washington, 106(4): 733-738. Download reprint [PDF]

2004f. Willmott, K. R., and Lamas, G. A new species of Tithorea (Lepidoptera: Nymphalidae: Ithomiinae) from the Chocó region of northwestern South America and Panama. J. New York Ent. Soc., 112(1): 1-17. Download reprint [PDF]

2004g. Hall, J. P. W., and D. J. Harvey. Five new riodinid species from the Napo region of Ecuador and Peru (Lepidoptera: Riodinidae). Tropical Lepidoptera,12(1-2): 35-41. Download reprint [PDF]

2004h. Willmott, K. R., and J. P. W. Hall. Taxonomic notes on the genus Zaretis, with the description of a new species (Lepidoptera: Nymphalidae: Charaxinae). Tropical Lepidoptera, 12(1-2): 25-38. Download reprint [PDF]

2003a. Hall, J. P. W. Phylogenetic reassessment of the five forewing radial-veined tribes of the Riodininae (Lepidoptera: Riodinidae). Systematic Entomology, 28(1): 23-37. Download reprint [PDF]

2003b. Hall, J. P. W., and C. J. Callaghan. A revision of the new riodinid butterfly genus Pseudotinea (Lepidoptera: Riodinidae). Journal of Natural History, 37(7): 821-837. Download reprint [PDF]

2003c. Willmott, K. R. Cladistic analysis of the neotropical butterfly genus Adelpha (Nymphalidae: Limenitidini), with comments on the subtribal classification of the tribe Limenitidini. Systematic Entomology, 28: 1-43. Download reprint [PDF]

2003d. Willmott, K. R. The Genus Adelpha: its Systematics, Biology and Biogeography (Lepidoptera: Nymphalidae: Limenitidini). Gainesville, Scientific Publishers. viii + 322 pp. Download pre-print [PDF]

2002a. Hall, J. P. W. A review of the new riodinid butterfly genus Panaropsis (Lepidoptera: Riodinidae: Symmachiini). Proceedings of the Entomological Society of Washington, 104(1): 63-72. Download reprint [PDF]

2002b. Hall, J. P. W. A review of Chalodeta with a revision of the Chalodeta chaonitis group (Lepidoptera: Riodinidae). Proceedings of the Entomological Society of Washington, 104(2): 376-389. Download reprint [PDF]

2002c. Hall, J. P. W. Phylogeny of the riodinid butterfly subtribe Theopeina (Lepidoptera: Riodinidae: Nymphidiini). Systematic Entomology, 27(2): 139-168. Download reprint [PDF]

2002d. Hall, J. P. W. A phylogenetic revision of Calydna and relatives (Lepidoptera: Riodinidae). Insect Systematics and Evolution, 33(2): 185-237. Download reprint [PDF]

2002e. Hall, J. P. W., and D. J. Harvey. A survey of androconial organs in the Riodinidae (Lepidoptera). Zoological Journal of the Linnean Society, 136: 171-197. Download reprint [PDF]

2002f. Hall, J. P. W., and D. J. Harvey. The phylogeography of Amazonia revisited: new evidence from riodinid butterflies. Evolution, 56(7): 1489-1497. Download reprint [PDF]

2002g. Hall, J. P. W., and D. J. Harvey. A phylogenetic review of Charis and Calephelis (Lepidoptera: Riodinidae). Annals of the Entomological Society of America, 95(4): 407-421. Download reprint [PDF]

2002h. Hall, J. P. W., and D. J. Harvey. A revision of the Neotropical butterfly genus Seco Hall and Harvey (Lepidoptera: Riodinidae). Proceedings of the Entomological Society of Washington, 104(4): 941-947. Download reprint [PDF]

2002i. Hall, J. P. W., and D. J. Harvey. Basal subtribes of the Nymphidiini (Lepidoptera: Riodinidae): phylogeny and myrmecophily. Cladistics, 18(6): 539-569.

2002j. Harvey, D. J., and J. P. W. Hall. Phylogenetic revision of the Charis cleonus complex (Lepidoptera: Riodinidae). Systematic Entomology, 27(3): 265-300. Download reprint [PDF]

2001a. Hall, J. P. W. A revision of the new riodinid butterfly genus Dachetola (Lepidoptera: Riodinidae: Riodinini). Journal of the New York Entomological Society, 109(2): 183-195. Download reprint [PDF]

2001b. Hall, J. P. W., and D. J. Harvey. A phylogenetic analysis of the Neotropical riodinid butterfly genera Juditha, Lemonias, Thisbe, and Uraneis, with a revision of Juditha (Lepidoptera: Riodinidae: Nymphidiini). Systematic Entomology, 26(4): 453-490. Download reprint [PDF]

2001c. Hall, J. P. W., and D. J. Harvey. Phylogenetic revision of the Charis gynaea group (Lepidoptera: Riodinidae) with comments on historical relationships among Neotropical areas of endemism. Annals of the Entomological Society of America, 94(5): 631-647. Download reprint [PDF]

2001d. Hall, J. P. W., and D. J. Harvey. A reassessment of Calociasma with the description of a new genus and a new species (Lepidoptera: Riodinidae: Nymphidiini). Journal of the New York Entomological Society, 109(2): 196-205. Download reprint [PDF]

2001e. Hall, J. P. W., and G. Lamas. Five new riodinid species from northwestern dry forest and northeastern Andean cloud forest habitats in Peru (Lepidoptera: Riodinidae). Revista Peruana Entomologia, 42: 9-19. Download reprint [PDF]

2001f. Willmott, K.R., J. P. W. Hall, and G. Lamas. Systematics of Hypanartia (Lepidoptera: Nymphalidae: Nymphalinae), with a test for geographical speciation mechanisms in the Andes. Systematic Entomology, 26: 369-399. Download reprint [PDF]

2001g. Willmott, K. R., L. M. Constantino, and J. P. W. Hall. A review of Colobura (Lepidoptera: Nymphalidae) with comments on larval and adult ecology and description of a sibling species. Annals of the Entomological Society of America, 94(2): 185-196. Download reprint [PDF]

2001h. Hall, J. P. W., and K. R. Willmott. Ecuador. Out on a limb. National Geographic Magazine (Update from the field), 199(2): 86. Download reprint [PDF]

2001i. Willmott, K. R., and J. P. W. Hall. Winged gems of Ecuador. Biodiversity paradise. National Geographic Magazine (Geographica) 199(5): xxviii. Download reprint [PDF]

2000a. Hall, J. P. W., and K. R. Willmott. Patterns of feeding behaviour in adult male riodinid butterflies and their relationship to morphology and ecology. Biol. J. Linn. Soc., 69: 1-23. Download reprint [PDF]

2000b. Hall, J. P. W. Two new genera in the Neotropical riodinid tribe Nymphidiini (Riodinidae). Journal of the Lepidopterists' Society, 54(2): 41-46. Download reprint [PDF]

1999a. Hall, J. P. W. A Revision of the Genus Theope: its Systematics and Biology (Lepidoptera: Riodinidae: Nymphidiini). Scientific Publishers, Gainesville. vii + 127pp.

1999b. Hall, J. P. W., and J. B. Heppner. Lemoniadini, a corrected tribal name in the Riodininae (Lepidoptera: Riodinidae). Tropical Lepidoptera, 10(1): 30.Download reprint [PDF]

1999c. Pyrcz, T., K. R. Willmott, and J. P. W. Hall. Contribution to the knowledge of Ecuadorian Pronophilini. Part III. Three new species and five new subspecies of Lymanopoda (Lepidoptera: Nymphalidae: Satyrinae). Genus, 10(3): 497-522. Download reprint [PDF - 1] [PDF - 2]

1999d. Willmott, K. R., and J. P. W. Hall. Taxonomic notes on Ecuadorian Adelpha with the description of two new species and seven new subspecies (Lepidoptera: Nymphalidae: Limenitidinae). Tropical Lepidoptera, 10(1): 1-17. Download reprint [PDF - 1] [PDF - 2]

1998a. Hall, J. P. W. A review of the genus Sarota (Lepidoptera: Riodinidae), pp. 1-21. In: J. P. W. Hall (ed.) A Contribution to Riodinid Systematics. Tropical Lepidoptera, 9(2), Supplement 1. Download reprint [PDF - 1] [PDF - 2]

1998b. Hall, J. P. W. Six new species in the "foliorum group" of Theope (Lepidoptera: Riodinidae). Lambillionea, 98(4): 562-573. Download reprint [PDF]

1998c. Hall, J. P. W., and D. J. Harvey. A new genus of riodinid, with a new species from Panama (Lepidoptera: Riodinidae: Nymphidiini), pp. 36-41. In: J. P. W. Hall (ed.) A Contribution to Riodinid Systematics. Tropical Lepidoptera, 9(2), Supplement 1. Download reprint [PDF]

1998d. Hall, J. P. W., and K. R. Willmott. Four new riodinid species from eastern Ecuador (Lepidoptera: Riodinidae). Lambillionea, 98: 325-334. ownload reprint [PDF]

1998e. Hall, J. P. W., and K. R. Willmott. Nine new species and one new subspecies of Euselasia from Ecuador (Lepidoptera: Riodinidae). Tropical Lepidoptera, Suppl. 1, 9(2): 27-35. Download reprint [PDF]

1998f. Hall, J. P. W., and K. R. Willmott. Three new species of Riodinini from the cloud forests of Ecuador (Lepidoptera: Riodinidae). Tropical Lepidoptera, Suppl. 1, 9(2): 22-26. Download reprint [PDF]

1996a. Hall, J. P. W., and K. R. Willmott. The genus Theope: four new species and a new subspecies (Lepidoptera: Riodinidae). Tropical Lepidoptera, 7(1): 63-67. Download reprint [PDF]

1996b. Hall, J. P. W., and K. R. Willmott. Notes on the genus Argyrogrammana Part II, with the description of a new species. Tropical Lepidoptera, 7(1): 71-80. Download reprint [PDF]

1996c. Hall, J. P. W., and K. R. Willmott. Systematics of the riodinid tribe Symmachiini, with the description of a new genus and five new species from Ecuador, Venezuela and Brazil (Lepidoptera: Riodinidae). Lambillionea, 96: 637-660. Download reprint [PDF]

1996d. Torres, R., J. P. W. Hall, K. R. Willmott, and K. Johnson. A new genus of brilliantly colored elfin butterflies from the high Andes (Lepidoptera: Lycaenidae). Tropical Lepidoptera, 7(1): 81-86. Download reprint [PDF]

1995a. Hall, J. P. W., and K. R. Willmott. Two new species of Mesene from western Ecuador (Lepidoptera: Riodinidae). Tropical Lepidoptera, 6(2): 110-112. Download reprint [PDF]

1995b. Hall, J. P. W., and K. R. Willmott. Five new species and a new genus of riodinid from the cloud forests of eastern Ecuador (Lepidoptera: Riodinidae). Tropical Lepidoptera, 6(2): 131-135. Download reprint [PDF]

1995c. Hall, J. P. W., and K. R. Willmott. Notes on the genus Argyrogrammana, with descriptions of five new species (Lepidoptera: Riodinidae). Tropical Lepidoptera, 6(2): 136-143. Download reprint [PDF]

1995d. Willmott, K. R., and J. P. W. Hall. Two new species of satyrines from Ecuador (Nymphalidae: Satyrinae). Tropical Lepidoptera, 6(2): 103-105. Download reprint [PDF]

1995e. Willmott, K. R., and J. P. W. Hall. Two new species of Adelpha from eastern Ecuador (Lepidoptera: Nymphalidae). Tropical Lepidoptera, 6(2): 106-109. Download reprint [PDF]

1994. Willmott, K. R., and J. P. W. Hall. Four new species of Riodinidae (Lepidoptera) from Western Ecuador. Tropical Lepidoptera, 5(2): 87-91. Download reprint [PDF]

 

BY OTHER AUTHORS:

Butterfly taxonomy and biology:

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Brabant, R.
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2000. Contributions to the knowledge of Ecuadorian Pronophilini. Part IV. New taxa of Pronophila Doubleday (Lepidoptera: Nymphalidae: Satyrinae). Genus, 11(1): 69-86.

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Pyrcz, T., K. R. Willmott, and J. P. W. Hall
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1998. Lepidoptera diversity of an Ecuadorian lowland rainforest (Lepidoptera: Papilionidae, Pieridae, Nymphalidae, Saturniidae, Sphingidae). Neue Ent. Nachr., 41: 95-117.

2001. An annotated list of Ecuadorian butterflies (Lepidoptera: Papilionidae, Pieridae, Nymphalidae). Fragmenta Ent., 33(2): 213-380.

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Vitale, F., and M. Bollino
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1999b. Errata. Lambillionea, 99(2)(2): 257.

2000a. Ithomiidae dell'Ecuador, secondo contributo. Su alcune specie del genere Hyalyris Boisduval, 1870, con descrizione di nuovi taxa (Lepidoptera: Ithomiidae). Lambillionea, 100(1)(2): 164-172.

2000b. Ithomiidae dell'Ecuador, terzo contributo. Il genere Ithomia Hübner, 1816. Descrizione di quattro nuove sottospecie (Lepidoptera: Ithomiidae). Lambillionea, 100(3)(1): 343-358.

2003. Una nuova sottospecie del genere Velamysta Haensch dal Nord Ecuador (Lep. Ithomiidae). Lambillionea, 103(2)(2): 325-326.

Vitale, F., and G. Rodriguez
. 2004. Nuevos Ithomiinae de Colombia y Ecuador (Lepidoptera: Nymphalidae). Lambillionea, 104(1)(1): 86-91.


Conservation:

Dodson, C. H., and A. H. Gentry
. 1991. Biological extinction in western Ecuador. Ann Miss. Bot. Gard., 78: 273-295.

Ecociencia
. 1994. Parques Nacionales y otras areas protegidas del Ecuador: una esperanza para el futuro. Ecociencia, Quito. 31 pp.

Best, B. J. (Ed.)
. 1992. The threatened forests of south-west Ecuador. Leeds, Biosphere Publications.

Best, B. J., and M. Kessler
. 1995. Biodiversity and conservation of Tumbesian Ecuador and Peru. Cambridge, BirdLife.

Cabarle, B. J., M. Crespi, C. H. Dodson, C. C. Luzuriaga, D. Rose, and J. N. Shores
. 1989. An Assessment of Biological Diversity and Tropical Forests for Ecuador. Washington, Center for International Development and Environment. 172 pp.

INEFAN
. 1998. Ecuador. Informe Interino a la Secretaria del Convenio de Diversidad Biológica, sobre la Aplicación del Articulo 6. Quito, INEFAN.

Parker, T. A., and J. L. Carr (Eds.)
. 1992. Status of Forest Remnants in the Cordillera de la Costa and Adjacent Areas of Southwestern Ecuador. RAP Working Papers 2. Washington, Conservation International. 172 pp.

Schulenberg, T. S., and K. Awbrey (Eds.)
. 1997. The Cordillera del Condor Region of Ecuador and Peru: A Biological Assessment. RAP Working Papers 7. Washington, Conservation International. 231 pp.

Suaréz, L.
1993. La conservación de la biodiversidad biológica en el Ecuador: prioridades de investigación. In: Mena, P. A., and L. Suaréz (Eds.), La Investigación para la Conservación de la Diversidad Biológica en el Ecuador, pp. 333-340. Quito, Ecociencia.

 

DNA MATERIAL

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Material of a number of species, stored in alcohol in the field or as dried, papered specimens, is available that is suitable for molecular sequencing. Researchers wishing to obtain material of particular species or groups should contact us.