Revisiting the Andean butterfly Eryphanis zolvizora group (Lepidoptera, Nymphalidae): one or several species?

  • Patrick Blandin Muséum National d’Histoire Naturelle, Entomologie, 57 rue Cuvier, 75005 Paris
  • Roger Bristow 2 Sid Bank, Sid Lane, Sidmouth, Devon EX10 9AW
  • Andrew Neild The Natural History Museum, Entomology Department, Cromwell Road, SW7 5BD, London
  • Juan Carlos de Sousa Museo del Instituto de Zoología Agrícola, Facultad de Agronomía, Universidad Central de Venezuela, Maracay, Apdo. 4579, Aragua
  • Yuvinka Gareca Museo de Historia Natural Alcide d’Orbigny, Entomologia, CP4324, Av. Potosi N°1458 (esq. Av. América), Cochabamba
  • Blanca Huertas The Natural History Museum, Entomology Department, Cromwell Road, SW7 5BD, London
Keywords: Morphinae, Brassolini, Eryphanis zolvizora group, subspecies, taxonomic inflation


Eryphanis zolvizora (Hewitson, 1877) is a rare Andean endemic butterfly, described from Bolivia, which has been historically classified either as a unique species, or as part of a group of three allopatric species from Bolivia, Ecuador and Colombia. In this paper, the group is revised using more than 200 specimens housed in 34 European, and North and South American public and private collections. For the first time, the presence of the group in Western Ecuador and Venezuela is confirmed, and important data on Peruvian populations are provided. In some populations, individual variations of genitalia are observed. Nevertheless, male genitalia allow the distinction of four geographical groups. Considering also habitus characters, eight taxa are distinguished and considered to be subspecies, of which five are new: Eryphanis zolvizora inca ssp. nov., Eryphanis zolvizora chachapoya ssp. nov., Eryphanis zolvizora casagrande ssp. nov.., Eryphanis zolvizora reyi ssp. nov., and Eryphanis zolvizora isabelae ssp. nov.  In the present state of knowledge, these taxa are allopatric, except for a possible geographic overlap in central Peru, where data are insufficient to prove sympatry. The “several subspecies vs. several species” dilemma is discussed, considering its impact for conservation action and policies.


Agapow P-M., Bininda-Edmonds O.R.P., Crandall K.A., Gittleman J.L., Mace G.M., Marshall J.C. & Purvis A. 2004. The impact of species concept on biodiversity studies. The Quaterly Review of Biology 79: 161-179.

Alström P., Rasmussen P.C., Olsson U. & Sundberg P. 2008. Species delimitation based on multiple criteria: the Spotted Bush Warbler Bradypterus thoracicus complex (Aves: Megaluridae). Zoological Journal of the Linnean Society 154: 291-307.

Barnes J.B. & Ehlers T.A. 2009. End member modes for Andean Plateau Uplift. Earth Science Reviews 97: 117-144.

Berlepsch H.G. von 1901. Mitteilungen über die von den Gebrüdern G. und O. Garlepp in Bolivia gesammelten Vogel und Beschreibungen neuer Arten. Journal für Ornithologie 49: 81-99.

Blandin P. 1977. La distribution géographique des Brassolinae (Lepidoptera-Satyridae). Faits et problèmes. In: Descimon H. (ed.) Biogéographie et évolution en Amérique tropicale: 161-218. Publications du Laboratoire de Zoologie de l’Ecole Normale Supérieure, Paris.

Blandin P. & Descimon H. 1975. Contribution à la connaissance des lépidoptères de l’Equateur. Les Brassolinae (Nymphalidae). Annales de la Société entomologique de France (N.S.) 13: 3-28.

Blandin P. & Descimon H. 1977. Contribution à la connaissance des lépidoptères de l’Equateur. Nouvelles données sur les Brassolinae (Nymphalidae) de l’Occidente. Annales de la Société entomologique de France (N.S.) 13: 75-88.

Blandin P. & Purser B. 2013. Evolution and diversification of Neotropical butterflies: Insights from the biogeography and phylogeny of the genus Morpho Fabricius, 1807 (Nymphalidae: Morphinae), with a review of the geodynamics of South America. Tropical Lepidoptera Research, 23(2): 62-85.

Brack Egg A. & Mendiola Vargas C. 2004. Ecología del Perú. Asociación Editorial Bruño, Lima.

Brown A.D. & Kappelle M. 2001. Introducción a los bosques nublados del neotrópico: una síntesis regional. In: Kappelle M. & Brown A.D. (eds) Bosques nublados del neotrópico: 27-40. Instituto Nacional de Biodiversidad, Costa Rica.

Campos F. 1927. Catálogo preliminar de los lepidópteros del Ecuador. Primera parte. Ropalóceros. Revista del Colegio Nacional Vicente Rocafuerte (Guayaquil) 9: 3-106.

Casagrande M.M. 2004. Tribe Brassolini. In: Lamas G. (ed.) Checklist. Part 4A. Hesperioidea-Papilionoidea. Atlas of Neotropical Lepidoptera vol. 5A: 201-205. Association of Tropical Lepidoptera/Scientific Publishers, Gainesville.

Cracraft J. 2000. Species concepts in theoretical and applied Biology: A systematic debate with consequences. In: Wheeler Q.D. & Miller R. (eds) Species Concepts and Phylogenetic Theory: A Debate: 1-14. Columbia University Press, New York.

D’Abrera B. 1987. Butterflies of the Neotropical Region. Part III. Brassolidae, Acraeidae, & Nymphalidae (Partim). Hill House, Victoria.

Descimon H. & Mallet J. 2009. Bad species. In: Settele J., Shreeve T., Konvicka M. & Van Dyck H. (eds) Ecology of Butterflies in Europe: 219-249. Cambridge University Press, Cambridge.

Dognin P. 1894. Lépidoptères de Loja et environs (Equateur). Descriptions d’espèces nouvelles. Troisième livraison. Imprimerie F. Levé, Paris.

Fassl A.H. 1911. Tropische Reisen. III. Erforschung des Monte Tolima. Entomologische Zeitschrift 24: 250-252, 259-260, 263-264, 267-268.

Fassl A.H. 1914. Tropische Reisen. V. Das obere Caucatal und die Westcordillere. Entomologische Rundschau 31: (7) 35-38, (38) 42-46, (39) 50-52, (10) 57-58.

Fassl A.H. 1915a. Die vertikale Verbreitung der Lepidopteren in der Columbishcen West-Cordillere. Entomologische Rundschau 32: 9-12.

Fassl A.H. 1915b. Tropische Reisen. VII. Ost-Columbien und die Llanos. Entomologische Rundschau 32: 33-35.

Fassl A.H. 1920. Meine Bolivia-Reise. Entomologische Rundschau 37: 10-11, 15-16, 17-18, 22-23, 25-27, 29-30, 34-35, 41-43, 45-48.

Fruhstorfer H. 1912. 5. Familie Brassolidae. In: Seitz A. (ed.) Die Gross-Schmetterlinge der Erde 5: 285-332. Alfred Kernen Verlag, Stuttgart.

Gregory-Wodzicki K.M. 2000. Uplift history of the Central and Northern Andes: A review. GSA Bulletin 112: 1091-1105.<1091:UHOTCA>2.0.CO;2

Griffin F.J. 1932. An attempt to determine the true dates of publication of Hewitson (W.C.) Illustr. New spec. exot. Butterflies, 5 vols. [1852-] (1856) 1876. Novitates Zoologicae, a Journal of Zoology in connection with Tring Museum, Tring 37(2): 189-206.

Hemming A.F. 1967. The generic names of the butterflies and their type-species (Lepidoptera: Rhopalocera). Bulletin of the British Museum of Natural History (Entomology), Supplement 9: 1-509.

Huertas B.C. & Donegan T.M. (eds) 2006. Proyecto YARÉ: investigación y evaluación de las especies amenazadas de la Serranía de los Yariguíes, Santander, Colombia. Informe final. Colombian EBA Project Rep. Ser. 7. Available from [accessed 29 Oct. 2013]

Huertas B., Rios C. & Le Crom J.F. 2009. A new species of Splendeuptychia from the Magdalena Valley in Colombia (Lepidoptera: Nymphalidae: Satyrinae). Zootaxa 2014: 51–58.

Isaac N.J.B., Mallet J. & Mace G.M. 2004. Taxonomic inflation: its influence on macroecology and conservation. Trends in Ecology and Evolution 19: 464-469.

James F.C. 2010. Avian subspecies: Introduction. Ornithological Monographs 67: 1-5.

Kirby W.F. 1877. A Synonymic Catalogue of Diurnal Lepidoptera. Supplement. John Van Voorst, London.

Kirby W.F. 1879. Catalogue of the collection of diurnal Lepidoptera formed by the late William Chapman Hewitson of Oatlands, Walton-on-Thames; and bequeathed by him to the British Museum. John Van Voorst, London.

Lamas G. 1981. Introducción a la Historia de la Entomología en el Perú. II. Período de los viajeros, colectores y estudiosos especializados. Revista peruana de Entomología 23: 25-37.

Lamas G., Robbins R.G. & Field W.D. 1995. Bibliography of Butterflies. An Annotated Bibliography of the Neotropical Butterflies and Skippers (Lepidoptera: Papilionoidea and Hesperioidea). Atlas of Neotropical Lepidoptera, Vol. 124. Association for Tropical Lepidoptera, Scientific Publishers, Gainesville.

Mallet J. 2007. Subspecies, semispecies, superspecies. In: Levin S.A. (ed.) Encyclopaedia of Biodiversity 5. Online update 1: 1-5. Elsevier, Oxford.

Mittermeier R.A., Robles Gil P., Hoffmann M., Pilgrim J., Brooks T., Goettsch Mittermeier C., Lamaoreux J. & Da Fonseca G.A.B. 2004. Hotspots Revisited. Cemex, Mexico City.

Neild A.F.E. 1996. The Butterflies of Venezuela. Part 1: Nymphalidae I (Limenitidinae, Apaturinae, Charaxinae). A comprehensive guide to the identification of adult Nymphalidae, Papilionidae, and Pieridae. Meridian Publications, Greenwich.

Neild A.F.E. 2008. The Butterflies of Venezuela. Part 2: Nymphalidae II (Acraeinae, Libytheinae, Nymphalinae, Ithomiinae, Morphinae). A comprehensive guide to the identification of adult Nymphalidae, Papilionidae, and Pieridae. Meridian Publications, London.

Patten M.A. & Unitt P. 2002. Diagnosability versus mean differences of Sage Sparrows subspecies. Auk 119: 26–35.;2

Paynter Jr. R.A. 1992. Ornithological gazetteer of Bolivia, 2nd Ed. Museum of Comparative Zoology, Harvard University, Cambridge, MA.

Paynter R. A. Jr. 1997. Ornithological Gazetteer of Colombia. Museum of Comparative Zoology, Harvard University, Cambridge, MA.

Penz C. 2008. Phylogenetic Revision of Eryphanis Boisduval, with a Description of a New Species from Ecuador (Lepidoptera, Nymphalidae). Insecta Mundi 0035: 1-25.

Piñas F. 2004. Vol. 11a. Familia: Nymphalidae: Subfamilias: Libytheinae, Danainae, Charaxinae, Morphinae, Heliconiiae. Mariposas del Ecuador. Compañía de Jesús, Quito.

Remsen J.V. 2010. Subspecies as a meaningful taxonomic rank in avian classification. Ornithological Monographs 67:62-78.

Rey F. & Pyrcz T.W. 1996. A perfect mimicry of two cloud forest pierids: Leptophobia eleone and Catasticta revancha n. sp. (Lepidoptera: Pieridae). Genus (Wroclaw) 7(3): 497-502, 3 figs.

Sclater P.L. & Salvin O. 1879. 2. On the Birds collected in Bolivia by Mr. C. Buckley. Proceedings of the Zoological Society of London 47: 588-645.

Silva X. 2011. Ecología de Mariposas del Ecuador. Universidad San Francisco, Quito.

Staudinger O. 1887. VIII. Brassoliden. In: Staudinger O. & Schatz E. (eds) Exotische Schmetterlinge 1: 211-218. G. Löwensohn, Fürth.

Stichel H. 1904. Lepidoptera Rhopalocera. Fam. Nymphalidae. Subfam. Brassolinae. Genera Insectorum 20: 1-48.

Stichel H. 1909. Brassolidae. Das Tierreich 25. R. Friedländer und Sohn, Berlin.

Stichel H. 1932. Brassolidae. In: Strand E. (ed) Lepidopterorum Catalogus 51. W. Junk, Berlin.

Vane-Wright R.I. 1991. A portrait of Clarence Buckley, zoologist. The Linnean 7(3): 30-33.

Weymer G. & Maasen J.P. 1890. Lepidopteren gesammelt auf einer Reise durch Colombia, Ecuador, Perú, Brasilien, Argentinien und Bolivien in den Jahren 1868-1877 von Alphons Stübel. A. Asher & Co, Berlin.

Zink R.M. 2004. The role of subspecies in obscuring avian biological diversity and misleading conservation policy. Proceedings of the Royal Society of London B 271: 561-564.

How to Cite
Blandin, P., Bristow, R., Neild, A., Carlos de Sousa, J., Gareca, Y., & Huertas, B. (2014). Revisiting the Andean butterfly Eryphanis zolvizora group (Lepidoptera, Nymphalidae): one or several species?. European Journal of Taxonomy, (71).