Research papers


Zacca, T., Casagrande, M. M., Mielke, O. H. H., Huertas, B., Barbosa, E. P., Freitas, A. V. L., Lamas, G., Espeland, M., Brévignon, C., Nakahara, S., Checa, M. F., Willmott, K. R. Systematics of the Neotropical butterfly genus Paryphthimoides Forster, 1964 (Lepidoptera: Nymphalida: Satyrinae), with descriptions of seven new taxa.

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.

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, in press.


2018: Freitas, A. V. L., Barbosa, E., Zacca, T., Marín, M., Beirão, M., Silva, A., Casagrande, M. M., Espeland, M., Willmott, K. R. 2018. Before it is too late: description of a new genus and species of butterfly from a highly threatened Brazilian biome. Revista Brasileira de Entomologia 62: 148-158, DOI: 10.1016/j.rbe.2018.02.003 (online 9 May).

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).

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. Freitas, A. V. L., Magaldi, L. M., Willmott, K. R. A new subspecies of Oleria gunilla (Nymphalidae: Danainae) from north Mato Grosso, Brazil. Journal of the Lepidopterists’ Society, 70(4): 295-301.

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. Willmott, K. R., Mielke, O. H. H., Freitas, A. V. L. A new subspecies and validation of the enigmatic species Adelpha herbita Weymer endemic to the Brazilian Atlantic forest (Nymphalidae: Limenitidinae). Tropical Lepidoptera Research, 26: 32-37 (June).

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.

2015. Garzón-Orduña, I. J., Silva-Brandão, K. L., Willmott, K. R., Freitas, A. V. L., Brower, A. V. Z. Incompatible ages for clearwing butterflies based on alternative secondary calibrations. Systematic Biology DOI:10.1093/sysbio/syv032.

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 (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): 1-4.

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.

2014. Warren, A. D., Tan, D., Willmott, K. R., Grishin, N. V. Refining the diagnostic characters and distribution of Hermeuptychia intricata (Nymphalidae: Satyrinae: Satyrini). Tropical Lepidoptera Research, 24: 44-51. Download reprint [PDF]

2014. Brower, A. V. Z., Willmott, K. R., Silva-Brandão, K. L., Garzón-Orduña, I. J., Freitas, A. V. L. Phylogenetic relationships of ithomiine butterflies (Lepidoptera: Nymphalidae: Danainae) as implied by combined morphological and molecular data. Systematics and Biodiversity, DOI: 10.1080/14772000.2014.899650, 1-15.

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.

2013. Chazot, N., Willmott, K. R., Paola G., 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.

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. Download reprint [PDF]

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. Download reprint [PDF]

2013. Pyrcz, T. W., Willmott, K. R., Garlacz, R., Boyer, P. Latitudinal gradient and spatial covariance in species richness of tropical Lepidoptera in the Andes.Insect Diversity and ConservationOnline version: DOI: 10.1111/icad.12058.

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

2013. Giraldo, C. E., Willmott, K. R., Vila, R. Uribe, S. I. Ithomiini butterflies (Lepidoptera: Nymphalidae) of Antioquia, Colombia. Neotropical Entomology.Online version: DOI: 10.1007/s13744-012-0102-4.

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.Online version: 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 s.l. (Lepidoptera: Nymphalidae; Ithomiini). Biological Journal of the Linnean Society, 106: 540-560. Online version: 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. Online version: 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. Abstract.

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

2011. Bolaños M., I. A., Zambrano G., 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. Download reprint [PDF]

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. Online version. Abstract

2010. Mullen, S. P., Savage, W., Willmott, K. R. 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 Online version

2010. Hall, J. P. W., Willmott, K. R. 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. DOI 10.18473/lepi.v64i3.a2

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

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

2009. Janzen, D. J., Hallwachs, W., + 39 authors, Willmott, K. R., + 3 authors. Integration of DNA barcoding into an ongoing inventory of complex tropical biodiversity. Molecular Ecology Resources, 9(Suppl. 1), 1-26. Online version

2009. Elias, M., Gompert, Z., Jiggins, C., Willmott, K. R. Phylogenetic community ecology needs to take positive interactions into account: Insights from colourful butterflies. Communicative & Integrative Biology 2(2): 113-116 [Article addendum]. Download reprint [PDF]

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., Jiggins, C. 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. DOI 10.4289/0013-8797-111.1.47

2008. Elias, M., Gompert, Z., Jiggins, C., Willmott, K. R. Mutualistic interations drive ecological niche convergence in a diverse butterfly community. PLoS Biology, 6(12):e300. Definitive online version or Download reprint [PDF]Supporting data S1 [PDF]Supporting data S2 [PDF]

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., Dasmahapatra, K., Willmott, K. R., Brower, A., Mallet, J., and C. Jiggins. 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] and supplementary documentation

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

2007. Willmott, K. R., and G. Lamas. 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. Download reprint [PDF]

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 (2000). 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. Online versionDownload PAGE PROOFS [PDF] or e-mail authors for reprint

2006. Jiggins, C. D., Mallarino, R., Willmott, K. R., and E. Bermingham. The phylogenetic pattern of speciation and wing pattern change in neotropicalIthomia butterflies (Lepidoptera: Nymphalidae). Evolution, 60: 1454-1466. Download reprint [PDF]

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. Whinnett, A., Willmott, K. R., Brower, A.V.Z., Simpson, F., Zimmermann, M., Lamas, G., and J. Mallet. Mitochondrial DNA provides an insight into the mechanisms driving diversification in the ithomiine butterfly Hyposcada anchiala (Lepidoptera: Nymphalidae, Ithomiinae). European Journal of Entomology, 102: 633-639. Download reprint [PDF]

2005. Whinnett, A., Zimmermann, M., Willmott, K. R., Herrera, N., Mallarino, R., Simpson, F., Joron, M., Lamas, G., and J. Mallet. Strikingly variable divergence times inferred across an Amazonian butterfly ‘suture zone’. Proceedings of the Royal Society of London B, 272: 2525-2533. Download reprint [PDF]

2005. Whinnett, A., Brower, A. V. Z., Lee, M.-M., Willmott, K. R., and J. Mallet. The phylogenetic utility of tektin, a novel region for inferring systematic relationships amongst Lepidoptera. Annals of the Entomological Society of America, 98: 873-886. 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]

2004. Balcázar-Lara, M., and K. R. Willmott. A new subspecies of Adelpha erymanthis from Mexico, with a key to identification of similar taxa (Lepidoptera: Nymphalidae: Biblidinae). Tropical Lepidoptera, 12(1-2)(“2001”): 25-38. Download reprint [PDF]

2004. Willmott, K. R., and J. Mallet. Correlations between adult mimicry and larval hostplants in ithomiine butterflies. Proceedings of the Royal Society of London B (Suppl.) 271: S266-S269. Download reprint [PDF]

2004. 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]

2004. 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]

2003. Mallet, J. L. B., and K. R. Willmott. Taxonomy: renaissance or Tower of Babel? Trends in Ecology and Evolution, 18(2): 57-59. Online version

2003. 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]

2002. Robinson-Dean, J. C., K. R. Willmott, M. J. Catterall, D. J. Kelly, A. Whittington, B. Phalan, N. M. Marples, and D. R. S. Boeadi. A new subspecies of Red-backed Thrush Zoothera erythronota kabaena subsp. nov. (Muscicapidae: Turdidae), from Kabaena island, Indonesia. Forktail, 18: 1-10.

2001. 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]

2001. 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]

2000. 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]

1999. 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]

1999. 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]

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

1998. 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]

1998. 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]

1996. 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]

1996. 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]

1996. 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]

1996. 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]

1995. 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]

1995. 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]

1995. 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]

1995. 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]

1995. 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]

1993. Hall, J. P. W., and K. R. Willmott. Temporal and elevational additions of distribution for Costa Rican butterflies. Tropical Lepidoptera, 4(2): 139-141. Online version


2003. Willmott, K. R. The Genus Adelpha: its Systematics, Biology and Biogeography (Lepidoptera: Nymphalidae: Limenitidini). Gainesville, Scientific Publishers. viii + 322 pp. Download below:

NOTE: File sizes are quite large (2-10 MB). If you can afford it, please buy a copy of the book from Scientific Publishers by contacting Dr John Heppner at

Complete book (c. 45 Mb – compiled text sections and figures are available individually below)

Individual text sections



Book chapters

2014. Checa, M. F., Willmott, K. R. Diversidad e importancia ecológica de mariposas del Ecuador. In: García, M., Parra, D. (Eds.) Ecuador: país de la Biodiversidad. Fundación Botánica de Los Andes/Ministerio del Ambiente/Fundación Ecofondo. Quito, Ecuador.

2006. Gareca, Y., Forno, E., Pyrcz, T., Willmott, K. R., Reichle, S. Lista preliminar de mariposas diurnas de Bolivia. In: Gareca, Y., and S. Reichle (eds.),Mariposas Diurnas de Bolivia, pp. 1-66. Tarija, Bolivia, PROMETA.

2004. Willmott, K.RBrush-footed butterflies (Lepidoptera: Nymphalidae). In: Capinera, J. L. (Ed.), Encyclopedia of Entomology, Volume I A-E, pp. 379-384. Dordrecht / Boston / London, Kluwer Academic Publishers. Download reprint [PDF]

Popular publications

2004. Willmott, K. R., and J. P. W. Hall. Common Butterflies of Yuturi and Yarina lodge. Yuturi Lodge, Ecuador. 2 pp.

2001. 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]

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

Letters and reviews

2015. Willmott, K. R. Butterflies. By Orenstein, R. I., Marent, T. (2015). Tropical Lepidoptera Research 25(2): 104-105. [book review]

2014. Warren, A. D., Willmott, K. R., Grishin, N. V. Subtle satyrs: differentiation and distribution of the newly described Hermeuptychia intricata in the southeastern United States (Lepidoptera: Nymphalidae: Satyrinae). News of the Lepidopterists’ Society 56(2): 83-85.

2012. Willmott, K. R. Lepidoptera. Borboletas e Mariposas do Brasil. [Lepidoptera. Butterflies and Moths of Brazil.] By de Almeida, A. C., Freitas, A. V. L. (2012). Tropical Lepidoptera Research 22(2): 120. [book review]

2009. Willmott, K. R. Mariposas de Canandé. Sus Amenazas, Potencial y Futuro. [The Butterflies of Canandé. Their Threats, Potential and Future.] By Checa V., M. F. (2009). Tropical Lepidoptera Research 19(1): 60. [book review]

2008. Willmott, K. RCatalogue of the Hostplants of the Neotropical Butterflies/ Catálogo de las Plantas Huésped de las Mariposas Neotropicales. By Beccaloni, G. W., Viloria, A. L., Hall, S. K. & Robinson, G. S. (2008). Tropical Lepidoptera Research 18(2): 124-125. [book review]

1998. Willmott, K. R. A fair review? News of the Lepidopterist’s Society, 40(4): 92-93.

1997. Hall, J. P. W., and K. R. WillmottThe Butterflies of Venezuela. Part I. Nymphalidae I (Limenitidinae, Apaturinae, Charaxinae). A Comprehensive Guide to the Identification of Adult Nymphalidae, Ppailionidae and Pieridae. By Andrew F. E. Neild. Tropical Lepidoptera, 8(1): 46. [book review]