top of page


Peer-Reviewed Publications

Updated May 2023

311 citations, h-index: 12, i10-index: 13.

Described and named 70 new butterfly taxa (13 genera, 52 species, 5 subspecies)

PDFs of each papers are also available at


ヘッディング 2

List of published journals

- Arthropod Systematics and Phylogeny

- Bull. de la Société Ent. de France

- Cladistics

- Conservación Colombiana

- European Journal of Taxonomy

- Insect Systematics and Diversity

- Insect Systematics and Evolution

- Molecular Phylogenetics and Evolution

- Trans. Lepid. Society of Japan

- Neotropical Entomology

- Peer J

- Tropical Lepidoptera Research

- Journal of Research on the Lepidoptera

- Journal of the Lepidopterists Society

- ZooKeys

- Zootaxa

- Revista Colombiana de Entomología

- Insecta Mundi

- Tijdschrift voor Entomologie

- Invertebrate Systematics

- Journal of Natural History


Journal Front Cover



Nakahara, S., Kleckner, K., Barbosa, E. P., Lourenço, G. et al. Reassessment of the type locality of Euptychia stigmatica Godman, 1905, with the description of two new sibling species from Amazonia (Lepidoptera, Nymphalidae, Satyrinae, Satyrini). ZooKeys, in press


Espeland, M., Nakahara, S., Zacca, T., Barbosa, E.P., Huertas, B., Marín, M.A. et al. 2023. Combining target enrichment and Sanger sequencing data to clarify the systematics of the diverse Neotropical butterfly subtribe Euptychiina (Nymphalidae, Satyrinae). Systematic Entomology, 1– 73. 


Padrón, P. S., Brito-Vera, G. A., Palomeque-Briones, M., Dueñas-Galvis, E., Nakahara, S. 2023. An updated checklist of the butterflies of Guayaquil, Ecuador (Lepidoptera: Papilionoidea). Revista Chilena de Entomología, 49(1): 23-41.


Corahua-Espinoza, T., Nakahara, S., Baine, Q., et al. 2023. Immature stages and new host plant records for three species in the “Taygetis clade” of Euptychiina in southeastern Peru (Lepidoptera: Nymphalidae: Satyrinae). Neotropical Entomology, 52(1): 67-80. 


Nakahara, S., Piovesan, M., Baine, Q., MacKenzie, E. C., et al. 2022. A new species of Caeruleuptychia Forster, 1964 from the Amazon basin (Lepidoptera: Nymphalidae: Satyrinae: Satyrini). Tropical Lepidoptera Research, 33(1): 23-36.


Nakahara, S., Zilli, A., Calhoun, J. V., Espeland, M., Padrón, P. S., Grishin, N. V.  2022. Resolving two centuries of mistaken identity: Reinterpretation of Papilio marcus Fabricius, 1787 (Lepidoptera: Nymphalidae, Hesperiidae). Zootaxa, 5195(3): 241-255.


Nakahara, S., Janzen, D. H., Hallwachs, W., Huertas, B., Das, G. N., Willmott, K. R. 2022. Revision of the “celia clade” of Pseudodebis Forster, 1964, with two new species and notes on Papilio phorcys Fabricius, 1793 (Lepidoptera: Nymphalidae: Satyrinae). Neotropical Entomology, 51(4): 536-556. 


Corahua-Espinoza, T., Nakahara, S., Shellman, B., Tejeira, R., Ccahuana, R., Gallice, G. 2022. Early stage biology of three satyrine species in the Peruvian Amazon (Lepidoptera: Nymphalidae: Satyrinae: Satyrini). Tropical Lepidoptera Research, 32(1): 38-46.


Corahua-Espinoza, T., Nakahara, S., Kabir, J., Shellman, B., Tejeira, R., Ccahuana, R., Gallice, G. 2022. Immature stages and new host plant records for three satyrine species feeding on herbaceous bamboos in southeastern Peru (Lepidoptera: Nymphalidae: Satyrinae: Satyrini). Zootaxa, 5125(1): 37-62. 10.11646/zootaxa.5125.1.3.


Nakahara, S., Rodríguez, M. M., Kleckner, K., Espeland, M., Casagrande, M. M., Barbosa, E., See, J., Tejeira, R., Gallice, G., Corahua-Espinoza, T., Lamas, G., Willmott, K. R. 2022. Systematic revision of the new butterfly genus, XXX Nakahara & Espeland, n. gen., with descriptions of three new taxa (Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity, 6(1): 2; 1-30. 10.1093/isd/ixab028.


Hurtado, T., Nakahara, S., Rodríguez, M. M., Tejeira, R., See, J., Ccahuana, R., Gallice, G. 2021. Complete life cycle of the euptychiine butterfly Taygetis cleopatra (C. Felder & R. Felder, 1867) (Lepidoptera: Nymphalidae: Satyrinae) in southeastern Peru. Tropical Lepidoptera Research, 31(3): 179-185.


Nakahara, S., Willmott, K.R., MacDonald, J.R., Thurman, A. 2021. Description of an enigmatic new species of Chloreuptychia Forster, 1964 from Panama (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera Research, 31(3): 145-150.


Nakahara, S., Barbosa, E.P., Nakamura, I., Lamas, G., Freitas, A.V.L. 2021. Description of a new species of Yphthimoides Forster, 1964 from Peru (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera Research, 31(3): 138-144.


Ccahuana, R., Corahua-Espinoza, T., Nakahara, S., Tejeira, R., Gallice, G. 2021. Immature stages and new hostplant record for Leucochimona hyphea (Cramer, 1776) in the Peruvian Amazon (Lepidoptera: Riodinidae: Riodininae). Tropical Lepidoptera Research, 31(3): 158-165.


Ccahuana, R., Tejeira, R., Hurtado, T., Nakahara, S.,  Rodríguez, M. M., Gott, R. J., See, J., Gallice, G. 2021. Immature stages of Ebusus ebusus ebusus (Cramer, 1780) in the Peruvian Amazon (Lepidoptera: Hesperiidae: Hesperiinae). Tropical Lepidoptera Research, 31(2): 90-95.


Tejeira, R., Ccahuana, R., Hurtado, T., Nakahara, S., See, J., Rodríguez, M. M., Corahua-Espinoza, T., Gallice, G. 2021. Immature stages of Chloreuptychia marica (Weymer, 1911) (Lepidoptera: Nymphalidae: Satyrinae: Satyrini). Tropical Lepidoptera Research31(2): 96-100.


Nakahara, S., Matos-Maraví, P., Willmott, K. R., Nakamura, I., MacDonald, J. 2021. Description of a new species of Pseudodebis Forster, 1964 from Central America. (Lepidoptera: Nymphalidae: Satyrinae).

Tropical Lepidoptera Research, 31(1): 35-41.


Nakahara, S., Matos-Maraví, P., Schwartz, J., Willmott, K. R. 2021. Assessing a generic synapomorphy of Pseudodebis Forster, 1964 (Lepidoptera: Nymphalidae: Satyrinae) and a recent speciation with a shift in elevation between two new species in the western Andes. Invertebrate Systematics, 35(2) 158-180.


Nakahara, S., Lamas, G., Willmott, K. R., Espeland, M. 2020. Description of a new genus and species for a common and widespread Amazonian satyrine butterfly (Lepidoptera: Nymphalidae: Satyrinae: Satyrini).  

Peer J, 8: e10324


Nakahara, S., Anderson, R. A., Nakamura, I., MacDonald, J. R. 2020. An integrative taxonomic approach supports a new species of Vettius from Central America (Lepidoptera: Hesperiidae: Hesperiinae). Journal of Natural History, 54 (27-28): 1699-1711, DOI: 10.1080/00222933.2020.1825857


Nakahara, S., Kleckner, K., Lamas, G., Huertas, B., Willmott, K. R. 2020. Contribution towards the systematics of Magneuptychia Forster, 1964: Caeruleuptychia probata (Weymer, 1911), n. comb. and Caeruleuptychia francisca (Butler, 1870), n. comb. (Lepidoptera: Nymphalidae: Satyrinae). Tijdschrift voor Entomologie, 163: 51-61.


Zacca, T., Casagrande, M. M., Mielke, O. H. H., Huertas, B., Espeland, M., Freitas, A. V. L., Willmott, K. R., Nakahara, S., Lamas, G. 2020. Revalidation of Vareuptychia Forster, 1964, description of Vanima gen n., and notes on Euptychia cleophes Godman & Salvin, 1889 (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa, 4858 (1): 001–034.


Nakahara, S., Hoffman, F.L.A., Hoffman, F. L., Gallice, G. 2020. Immature stages of Magneuptychia harpyia (C. Felder & R. Felder, 1867) (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera Research, 30(1): 33-38.


Zacca, T., Casagrande, M. M., Mielke, O. H.H., Huertas, B., Barbosa, E., Freitas, A. V. L., Lamas, G., Espeland, M., Brévignon, C., Nakahara, S., Checa, F., Willmott, K. R. 2020. Systematics of the Neotropical butterfly genus Paryphthimoides Forster, 1964 (Lepidoptera: Nymphalidae: Satyrinae), with descriptions of seven new taxa. Insect Systematics and Evolution, doi: 10.1163/1876312X-00001027


Nakahara, S., Gallardo, R. 2019. Scientific Note: Reinstatement of Euptychia sericeella Bates, 1865: Amiga sericeella stat. rev., with corrigenda to Nakahara et al. (2019) (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera Research, 29(2) 111-114. (Open Access Article: PDF)


Nakahara, S., Matos-Maraví, P., Barbosa, E. P., Willmott, K. R., Lamas, G., Freitas, A. V. L. 2019. Two new species of Taygetina with a possible case of “juxta loss” in butterflies(Lepidoptera: Nymphalidae: Satyrinae). Insect Systematics and Diversity, 3(6): 1-13. doi: 10.1093/isd/ixz023


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


Baine, Q., Polo-Espenoza, G., Nakahara, S., Gallice, G. 2019. Immature stages and new host record of Taygetis rufomarginata Staudinger, 1888 (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera Research, 29(2) 79-86. (Open Access Article: PDF)


Willmott, K. R., Marín, M. A., Nakahara, S., Pomerantz, T., Lamas, G., Huertas, B., Espeland, M., Xiao, L., Hall, J.P.W., Willmott, J.I.R., Freitas, A.V.L. 2019. 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, 19(1): 29–44. (Open Access Article: PDF)


Nakahara, S., Lamas, G., Tyler, S., Marín, M. A., 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. (Open Access Article: PDF)


Nakahara, S., MacDonald, J., Delgado, F., Padrón, S. P. 2018. Discovery of a rare and striking new pierid butterfly from Panama (Lepidoptera: Pieridae). Zootaxa, 4527 (2): 281–291. (Open Access Article: PDF; In MediaForbesSmithsonian Magazine)


Espeland, M., Barbosa, E., Breinholt, J., Casagrande, M. M., Huertas, B., Lamas, G., Marín, M. A., Mielke, O.H.H., Miller, J.Y., Nakahara, S., Tan, D., Warren, A.D., Zacca, T., Kawahara, A.Y., Freitas, A.V.L., Willmott, K.R.  2018. 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


See, J., Nakahara, S., Gallice, G. 2018 Immature stages of Splendeuptychia quadrina (Butler, 1869) (Lepidoptera: Nymphalidae: Satyrinae). Tropical Lepidoptera Research, 28 (2): 49–53. (Open Access Article: PDF)


Nakahara, S., Willmott, K. R., Mielke, O. H. H., Schwartz, J., Zacca, T., Espeland, M., Lamas, G. 2018. 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. (Open Access Article: PDF)


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


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


Warren A. D., Nakahara, S., Llorente-Bousquets, J. E., Luis-Martínez, A., Miller, J. Y. 2018. A new species of Cyllopsis R. Felder, 1869 from the highlands of Chiapas, Mexico (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa, 4403 (3): 570–577. (Open Access Article: PDF)


Nakahara, S., Zacca, T., Huertas, B., Neild, A. F. E., Hall, J. P. W., Lamas, G., Holian L. A., Espeland, M., Willmott, K.R. 2018. Remarkable sexual dimorphism, rarity and cryptic species: a revision of the ‘aegrota species group’ of the Neotropical genus Caeruleuptychia Forster, 1964 with the description of three new species (Lepidoptera, Nymphalidae, Satyrinae). Insect Systematics and Evolution, 49(2): 130-182. doi: 10.1163/1876312X-00002167 (In Media: ForbesZME Science)


Nakahara, S., Barbosa, E. P., Freitas, A. V. L. 2017. A potentially endangered new species of Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the Atlantic coastal forest of Brazil. Neotropical Entomology, 46 (3): 302-309. doi: 10.1007/s13744-016-0466-y


Nakahara, S., Tan, D., Lamas, G., Parus, A., Willmott, K. R. 2016. A distinctive new species of Hermeuptychia Forster, 1964 (Lepidoptera: Nymphalidae: Satyrinae) from the eastern tropical Andes. Tropical Lepidoptera Research, 26(2): 77–84. (Open Access Article: PDF)


Nakahara, S., Vega, G., Willmott, K. R. 2016. Description of a new Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the western Andes. Zootaxa, 4184(2): 358–366. doi: 10.11646/zootaxa.4184.2.7


Huertas, B., Lamas, G., Fagua, G., Mallet, J., Nakahara, S., Willmott, K. 2016. A remarkable new butterfly species from western Amazonia (Lepidoptera: Nymphalidae: Satyrinae) Conservación Colombiana, (24): 5–11. (Open Access Article: PDF)


Costa, M., Viloria, A. L., Attal, S., Neild, A. F. E., Fratello, S. A., Nakahara, S. 2016. Lepidoptera del Pantepui. Parte III. Nuevos Nymphalidae Cyrestinae y Satyrinae.  Bulletin de la Société Entomologique de France, 121(2), 179–206. (Open Access Article: PDF)


Nakahara, S., Marín, M. A., Neild, A. F. E. 2016. A new species of Pareuptychia Forster, 1964 (Lepidoptera, Nymphalidae, Satyrinae) from the northwestern Amazon basin with characterisation of two potential synapomorphies for the genus. Tropical Lepidoptera Research, 25(1), 6–11. (Open Access Article: PDF)


Nakahara, S., Barbosa, E. P., Marín, M. A., Freitas, A. V. L., Pomerantz, T., Willmott, K. R. 2016. Graphita gen. nov., a New Genus for Neonympha griphe C. Felder & R. Felder, 1867 (Lepidoptera, Nymphalidae, Satyrinae). Neotropical Entomology, 45: 675-691.  doi: 10.1007/s13744-016-0414-x


Warren, A. D., Nakahara, S., Lukhtanov, V. A., Dally, K. M., Ferris, C. D., Grishin, N. V., Martin, C., Pelham, J. P. 2016. A new species of Oeneis from Alaska, United States, with notes on the Oeneis chryxus complex (Lepidoptera: Nymphalidae: Satyrinae). Journal of Research on the Lepidoptera, 49, 1–20. (Open Access Article: PDF) (In Media: National Geographic SocietyThe Washington PostFuturity)


Nakahara, S., Hall, J. P. W., Lamas, G., Willmott, K. R. 2015. 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. (Open Access Article: PDF)


Neild, A. F. E.*, Nakahara, S.*, Zacca, T., Fratello, S., Lamas, G., Le Crom J-F., Dolibaina, D. R., Dias, F. M. S., Casagrande, M. M., Mielke, O. H. H., Espeland, M. 2015. Two new species of Euptychia Hübner, 1818 from the upper Amazon basin (Lepidoptera, Nymphalidae, Satyrinae).  ZooKeys, 541, 87–108. (Open Access Article: PDF*contributed equally; joint first authors. (In Media: BBCNewsweek, Daily Mail UK)


Fratello, S. A., Nakahara, S., Brévignon, C. R., Harvey, D. J. 2015. Two new species of Euptychia Hübner 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the Guiana Shield and notes on E. marceli Brévignon, 2005 and E. rufocincta Weymer, 1911. Journal of the Lepidopterists' Society, 69(4), 293–306. doi: 10.18473/lepi.69i4.a5


Nakahara, S., Le Crom, J-F., Lamas, G. 2015. A remarkable new Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae) from the Amazon basin of Peru and Colombia. Journal of Research on the Lepidoptera, 48, 58–61.(Open Access Article: PDF)


Nakahara, S., Llorente-Bousquets, J. E., Luis-Martínez, A., Miller, J. Y., Warren, A. D. 2015. Two new species of Euptychia Hübner 1818 (Lepidoptera: Nymphalidae: Satyrinae) from Mexico and Guatemala. Journal of Research on the Lepidoptera, 48, 51–57. (Open Access Article: PDF)


Nakahara, S., Janzen, D. H., Hallwachs, W., Espeland, M. 2015. Description of a new genus for Euptychia hilara (C. Felder & R. Felder) (Lepidoptera, Nymphalidae, Satyrinae). Zootaxa, 4012(3), 525–541. doi: 10.11646/zootaxa.4012.3.7


Nakahara, S., Marín, M. A., Ríos-M, C. 2015. Taxonomic status and redescription of Magneuptychia nebulosa (Butler, 1867) (Lepidoptera: Nymphalidae: Satyrinae) with a lectotype designation.  ZooKeys, 503,135–147. (Open Access Article: PDF)


Lamas, G., Nakahara, S. 2015. Nomenclatural status of Euptychia mollina Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae).  Zootaxa, 3946(4), 587–590. doi: 10.11646/zootaxa.3946.4.8


Nakahara, S., Fratello, S. A., Harvey, D. J. 2014. A new species of Euptychia Hübner, 1818 (Lepidoptera: Nymphalidae: Satyrinae: Satyrini) from Mount Roraima, Guyana. Zootaxa, 3881(3), 291–300. (Open Access Article: PDF)


Lewis, D. S., Sperling, F. A. H., Nakahara, S., Cotton, A. M., Kawahara, A. Y., Condamine, F. L. 2014. Role of Caribbean Islands in the diversification and biogeography of Neotropical Heraclides swallowtails. Cladistics, 31, 291–314. doi: 10.1111/cla.12092


Neild, A. F. E., Nakahara, S., Fratello, S. A., Harvey, D. J. 2014. A new species of Euptychia Hübner, 1818 (Nymphalidae: Satyrinae: Satyrini) from the Amazon basin and the  Guianas. Tropical Lepidoptera Research, 24(1): 4–9. (Open Access Article: PDF)


Nakahara, S., Highton, A., Jost, B., Mitsuhashi, W. 2013. Annotated new records for Nymphalid species from the Sierra de Imataca, Venezuela (Lepidoptera: Nymphalidae). Association for Tropical Lepidoptera Notes, June, 2013. (Open Access Article: PDF)


Nakahara, S., Marín, M. A., Mitsuhashi, W. 2012. First records for Cissia themis (Lepidoptera: Nymphalidae) from Colombia and Venezuela. Revista Colombiana de Entomología, 38(2), 359–362.


Nakahara, S., Blandin, P. 2010. A southern range extension for Morpho amathonte Deyrolle, 1860 (Nymphalidae, Morphinae) in western Ecuador and the description of a new subspecies. Tropical Lepidoptera Research, 20(2), 85–87. (Open Access Article: PDF)


Yago, M., Nakahara, S., Abe, W., Isohata, Y., Tomokuni, M., Rei, U. 2008. A Discovery of Acytolepis puspa (Lepidoptera, Lycaenidae) in the Kanto district, Japan : a geographic range extension, dispersal pathway inferred from ecology, morphology and molecular analyses. Transactions of the Lepidopterological Society of Japan, 60(1), 9-24.

bottom of page