Purpose of website
Background and aims
Why butterflies?
Why Ecuador?
State of knowledge
Citation and copyright



The main purpose of this website is to provide information about our past and current research on Ecuadorian butterflies. We also hope to promote a better understanding of this country's fauna by providing various information resources to encourage research work on Ecuadorian butterflies. We provide information on how to obtain research permits, on priority areas of the country to visit and inventory, study methods and suggested research projects. We also provide a list of publications relating to Ecuadorian butterfly taxonomy, biogeography, ecology and conservation, in both the bibliography section (containing PDF reprints of most of our papers) and student project section.

Although most of the information presented is our own unpublished data, we hope that eventually this site will become a general source of information on Ecuadorian butterfly research and conservation. We thus include several links to websites of other researchers, and Ecuadorian organisations, biological stations and lodges.

One of the main goal's of this website is to make accessible a database of Ecuadorian butterfly taxonomy and distribution.

Any comments or suggestions by e-mail would be welcome (





We first visited Ecuador in 1991. Three facts impressed us the most: the huge diversity of species and habitats, the difficulty in identification of most species, and the lack of published basic biological information. These impressions inspired us to begin research into the taxonomy, biogeography and natural history of Ecuadorian butterflies, with a view to eventually producing a field guide to the butterflies of the country. So far we have spent a combined total of 60 months in the field in Ecuador on this project.

To date our taxonomic work has been concentrated on the Riodinidae and Nymphalidae, resulting in 82 published papers of relevance to Ecuador, which include descriptions of 112 species and 20 genera (see state of knowledge). The remaining groups most in need of work are the Satyrinae and Lycaenidae.

Ecuador's biogeographical position (see below) and diversity of species has meant that intensive collecting in the country can also provide valuable information in understanding the taxonomy of entire neotropical genera (e.g., Hall, 1999; Willmott, 2003; Hall, 2005, see Bibliography). We also work therefore on butterfly taxonomy at the regional level.

Ecuador's diversity of habitats, species, and elevational and climatological gradients also make it a natural laboratory for testing broad ecological and evolutionary theories. What are the origins of species in montane habitats? Are historical or ecological processes more important for understanding species distribution and diversity? These sorts of questions can be addressed with abundant and accurate distributional data such as are beginning to become available for Ecuador.

Finally, such distributional data can be used not only to test how effectively Ecuador's butterfly species are conserved within current protected areas, but also to examine more theoretical conservation issues. Are there effective, rapid methods for sampling localities and estimating diversity and/or endemism, and is this information of conservation value? Does the geographic distribution of diversity and complementarity of species in one group provide an accurate indicator for other groups? We hope to look at such questions with butterfly data from Ecuador.

Current Research Work

Our research work has been funded in the past by many organisations, most significantly by The National Geographic Society (USA), the Leverhulme Trust (UK) and the National Science Foundation (USA), which has supported the project through two Biodiversity Surveys and Inventories grants (see Acknowledgements). Our work at present involves two components: field work in Ecuador to continue improving our knowledge of Ecuador butterfly distribution, diversity and ecology, and taxonomic study to provide a more stable long-term classification for Ecuador's species. Our current objectives are:

1) Conclude inventorying all important collections containing Ecuadorian butterfly material.

2) Make new collections from the most poorly known areas of the country, concentrating on areas that are particuarly rich in endemic taxa, and on rare and/or more inconspicuous taxa often overlooked in historical surveys.

3) Identify and classify all taxa known from Ecuador, and name all undescribed taxa, including those from other Andean areas if the taxonomic context is appropriate.

4) Build up extensive collections of dried and alcohol-preserved material (suitable for both traditional and molecular taxonomic study) in both Ecuador and the USA, promoting the development of a more comprehensive Ecuadorian national butterfly collection.

5) Promote understanding and appreciation of entomology and conservation in Ecuador through continuing to train research assistants and undergraduate students in techniques for butterfly collection and identification.

6) Make results widely available on the internet as well as through traditional publication of papers in scientific journals.




Insects make up a conservatively estimated 70% of all terrestrial organisms (Samways, 1994), yet our knowledge of their biology and true diversity remains remarkably fragmentary in comparison with more conspicuous groups, such as vertebrates (Wheeler, 1990; Hawksworth & Ritchie, 1993; Stork, 1997). Nevertheless, their outstanding evolutionary success in virtually all terrestrial habitats suggests they may be one of the more valuable study groups for understanding the distribution and origins of total biodiversity, and for developing efficient means to conserve this biodiversity (Wheeler, 1990; Brown, 1991, 1997). Unfortunately, much research into tropical insects is obstructed by the enormous diversity of species and the profound taxonomic difficulties in many groups. These difficulties have led to the suggestion that research might be most profitably concentrated on certain taxa of special significance, like butterflies (Brown, 1991, 1997; New, 1993; Sparrow et al., 1994; Samways, 1994).

1) The true butterflies (Papilionoidea) form a natural (monophyletic) insect group that is sufficiently speciose and ecologically diverse to be capable of revealing fine patterns in the distribution of diversity;

2) It has been an important study group in numerous ecological, biological and biogeographical studies (e.g. Vane-Wright & Ackery, 1984), and achieving a good understanding of the taxonomy, diversity and distribution of Ecuadorian butterflies will result in data of value to a wide range of potential research topics;

3) Members of the superfamily occupy most habitats, microhabitats and elevations;

4) Almost all species are herbivorous and therefore may closely mirror patterns of diversity and endemism in plants, widely regarded as the group exerting the greatest control on the distribution of total biodiversity;

5) Butterflies are familiar, large and conspicuous organisms, and therefore attractive to the public and suitable for communicating ideas and fostering support for research and conservation;

6) They require no costly equipment to sample or (in most cases) identify, and are therefore a suitable group for research by scientists or students with limited funds;

7) Although the Papilionoidea is relatively well studied taxonomically, providing a sound base for systematic study, the group's importance in diverse fields of research makes a critical taxonomic evaluation of most genera essential, particularly in the light of new material from the poorly collected Andean regions. As an example, the widespread and common genus Hypanartia has traditionally been thought to contain from 8-9 species, but since 1998 one highly distinctive new species has been discovered and a further four undescribed cryptic species recognized, all occurring in Andean habitats (Jasinski, 1998; Willmott et al., 2001). More poorly studied groups, such as the Lycaenidae, Riodinidae and Satyrinae, are likely to contain even higher proportions of new or unrecognized species (e.g., Johnson, 1992; Torres et al., 1995; Willmott & Hall, 1995a; Hall & Willmott, 1995a-c, 1996a,b, 1998; Pyrcz et al., 1999; Hall & Harvey, 2001; Harvey & Hall, 2002; Hall, 2002a,b).

8) Although there is a relatively large amount of butterfly material in collections, only a fraction of the associated distributional data are available in published form and therefore widely accessible;

9) Many taxa, especially Andean, are incorrectly identified in collections due to outdated taxonomy; paradoxically, since butterflies are perceived to be of little direct economic importance, there are relatively few professional taxonomists in the world, despite the group's enormous popular appeal.




Ecuador's geography is dominated by the Andes, which bisect the country on a roughly north-south axis. This mountain chain not only provides a barrier separating the western and eastern lowlands, helping to maintain their distinctive faunal characteristics, but also produces a huge range of different habitats for butterflies, as climate, and thus vegetation, change from sea-level to over 6000m elevation.

In global terms, biodiversity is not evenly distributed throughout the planet, and it has long been recognized that the tropics generally contain the most diverse biotas (Pianka, 1966; Rohde, 1992; Rosenzweig, 1995; Gaston & Williams, 1996). Within the tropics, it is the forests of South America that contain the richest diversity of plants, birds, mammals, reptiles, amphibians, butterflies, and probably most large groups of terrestrial organisms (McNeely et al., 1990; Robbins & Opler, 1997; Fjeldså & Rahbek, 1998).

Part of Ecuador's diversity is due to the high neotropical diversity, but Ecuador, along with Colombia, and to a lesser extent Venezuela, is also uniquely placed to draw upon three of the four main biogeographic regions of the neotropics: the Transandean region (Central America to western Ecuador), the Andes, and the Amazon basin. This is, in part, why these countries contain such a disporportionate amount of the Earth's biodiversity and have been labelled "megadiversity" countries (Mittermeier, 1988; Mittemeier et al., 1999). Such countries merit special international attention from biologists and conservationists alike (Mittermeier, 1988; McNeely et al., 1990; WCMC, 1992).

We estimate that Ecuador contains approximately 2700 Papilionoidea species, about 50-55% of all Neotropical butterfly species (25% of the World's species (Heppner, 1991; Robbins, 1993; Hall, Willmott & Robbins, unpubl. data)), making it one of the world's three most diverse countries, along with Colombia and Peru. An effective conservation program therefore has the potential to make a global impact. The allocation and management of resources for biodiversity conservation is still conducted at the country level (Mittermeier, 1988; McNeely et al., 1990; Bibby et al., 1992; Mittermeier et al., 1999); it therefore makes sense to concentrate survey efforts within a single country to provide comprehensive datasets of maximum use to local conservation bodies.

Montane forests are recognized as one of the most poorly known and severely threatened tropical bioregions (Churchill et al., 1995; Biodiversity Support Program et al., 1995; Dinerstein et al., 1995; Aldrich et al., 1997), and the Andes are no exception. Andean forests are small in extent, under increasing threat from explosive population growth, yet incredibly diverse with distinct butterfly faunas restricted to often very narrow altitudinal bands.

While not a biogeographic entity in itself, Ecuador contains an extraordinary range of Andean habitats, within a relatively small geographic area, that are readily accessible. It encompasses two of only a handful of global hotspot areas (sensu Myers, 1988, 1990), tropical habitats that contain exceptional concentrations of species with high levels of endemism, namely the western Andean Chocó region and western Amazonian uplands along the base of the eastern Andes. One of the principal Andean faunal boundaries, that between the central and northern Andean biogeographic regions, lies in southern Ecuador, while both slopes contain several smaller biogeographic regions containing variably differentiated populations. The western slope exhibits one of the most extreme environmental gradients on the continent, changing from pluvial forests with over 6m annual precipitation in the far north, to the dry deciduous woodlands and thorn scrub with less than 30cm annual rainfall in the south, over a distance of 400km.

As such, this broad diversity of environmental and biogeographic regions will allow our data to be used in investigating many biogeographic and ecological processes, including fine-scale population differentiation along latitudinal (eastern Ecuador - e.g., many ithomiines) and altitudinal (eastern Ecuador - e.g., many ithomiines) gradients, the effect of climatic gradients on diversity (north-west to south-west Ecuador), the relative diversity of the western and eastern Andean slopes through cross-Andean transects at a single latitude, and rates of species turnover both along altitudinal (throughout Ecuador) and latitudinal (southern to central Ecuador) gradients.




The first explorers to bring back butterfly specimens from mainland Ecuador were Alexander Von Humboldt and Aime Bonpland, in 1802, followed by Clarence Buckley (e.g. Hewitson, 1869, 1870a,b, 1877), whose collections laid the foundation of our knowledge of Ecuadorian butterflies. Buckley, and a succession of nineteenth century collectors, including Gaujón, de Mathan, Rosenberg and Haensch, managed to sample all the major regions of Ecuador except the north-east and south-west Andes and far eastern lowlands, but the central-eastern region was by far the most heavily collected and still remains the best known today. The most heavily sampled elevations are 400-600m and 1600-2000m on the east, and sea-level to 900m and 1600-2000m on the west, leaving middle and high elevations poorly sampled on both slopes.

Although local commercial dealers have been operating in the central-eastern region since the early 1900's, the next important lepidopterist to work in Ecuador was Frederick M. Brown, who made small collections of high altitude lycaenids and pronophiline satyrines (Brown, 1941a, 1943, 1944), and produced a comprehensive gazetteer of Ecuadorian collecting localities known at that time (Brown, 1941b). Following a period of inactivity, a small number of lepidopterists have concentrated their field studies on Ecuador or contributed significantly to our knowledge of the fauna, including:

  David Ahrenholz (USA: St Paul) (throughout country; Riodinidae)
  Ismael, Raul and Euclides Aldaz Villafuerte (Ecuador: Baños, Nanegalito, Riobamba) (throughout country; all families)
  Stéphane Attal (France: Paris) (largely central and south-east below 2000m; certain nymphalid genera, e.g. Perisama)
  Maurizio Bollino (Italy: Lecce) (throughout country; Papilionidae, Pieridae, Pronophilini)
  Pierre Boyer (France: Marseille) (throughout country; all families)
  Robert and George Busby (USA: Boston) (largely eastern and central western Andes; Lycaenidae, Riodinidae),
  Charlie Covell (USA: Gainesville) (largely central western and central eastern Andes; mainly Riodinidae, Lycaenidae)
  Philip DeVries (USA: Oregon) (Jatun Sacha and La Selva Lodges, Río Napo; mainly Nymphalidae, Riodinidae)
  Tom Emmel (USA: Gainesville) (largely central east and central west; all families)
  Artur Jasinksi (Poland: Kraków) (largely Andes; all families)
  Stan Nicolay (USA: Washington) (largely central western and central eastern Andes and lowlands; mainly Lycaenidae, Riodinidae)
  Francisco Piñas (Ecuador: Quito) (throughout country; all families)
  Tomasz Pyrcz (Poland: Kraków) (largely above 1500m: Satyrinae [Pronophilini])
  Tommaso Racheli (Italy: Rome) (largely central east and central west; mainly Papilionidae, Pieridae, Nymphalidae)
  Fabio Vitale (Italy: Lecce) (throughout country; Ithomiinae)

Aside from the collections made as part of this project, other major depositories of Ecuadorian material include the Natural History Museum, London, UK (BMNH), the National Museum of Natural History, Washington DC, USA (USNM), American Museum of Natural History (AMNH) (historical) and the Zoologische Museum für Naturkunde, Berlin, Germany (ZMHU). There is also important material in a number of other private collections in the USA and Europe, including those of the individuals listed above, and undoubtedly others of whom we are unaware and would very much like to contact.

Our research work to date suggests that approximately 2700 species of true butterflies (Papilionoidea) occur in Ecuador. There are a number of species that are expected to occur on the basis of ranges spanning the country or collection very close to a border, and these are listed under species to look for. It is quite probable that Ecuadorian specimens of some of these species, at least, are already in private collections, and we would like to hear from anyone who is able to confirm the presence of any of these species in the country.

When we began working in 1993 in Ecuador, a remarkable 430 species were undescribed, about 16% of the total fauna. Since then, 212 of these have been described, including 112 by ourselves and co-authors. The table below shows a breakdown of these figures by family.


Summary of Ecuadorian species described since 1993

undescribed at start (1993)
total described by us (end 2010)
total described by others (end 2010)
total described to date (end 2010)
remaining to be described



The map below shows a preliminary analysis of species richness for Papilionidae, Pieridae and Nymphalidae throughout Ecuador, in quarter degree grid squares.




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1997. Tropical Montane Cloud Forests: An Urgent Priority for Conservation. WCMC Biodiversity Bulletin No. 2. World Conservation Monitoring Centre, Cambridge. [20] pp.
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Brown, F. M.
1941a. Notes on Ecuadorian butterflies - I. Steroma, Pseudosteroma and Steremnia (Satridae [sic], Rhop.). Ann. Ent. Soc. Amer., 34(2): 432-436.
1941b. A gazetteer of entomological stations in Ecuador. Ann. Ent. Soc. Amer., 34(4): 809-851.
1943. Notes on Ecuadorian butterflies. III. The genus Lymanopoda Westwood (Satyridae). Ann. Ent. Soc. Amer., 36(1): 87-102.
1944. Notes on Ecuadorian butterflies. IV. The genus Penrosada, new (Lepidoptera, Satyridae). Ann. Ent. Soc. Amer., 37(2): 255-260.
Brown, K. S.
1991. Conservation of neotropical environments: insects as indicators. In: Collins, N. M., & J. A. Thomas (Eds.), The Conservation of Insects and their Habitats, pp. 349-404. London, Academic Press.
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Churchill, S. P., H. Balslev, E. Forero, and J. L. Luteyn (Eds.)
1995. Biodiversity and Conservation of Neotropical Montane Forests. New York, New York Botanical Garden.
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1995. A Conservation Assessment of the Terrestrial Ecoregions of Latin America and the Caribbean. Washington, The World Bank. xvii + 129 pp.
Fjeldså, J., and C. Rahbek
1998. Priorities for conservation in Bolivia, illustrated by a continent-wide analysis of bird distributions. In: Barthlott, W., and M. Winiger (Eds.), Biodiversity: a Challenge for Development Research, pp. 311-327. Berlin, Springer-Verlag.
Gaston, K. J., and P. H. Williams
1996. Spatial patterns in taxonomic diversity. In: Gaston, K. J. (Ed.), Biodiversity: a Biology of Numbers and Differences, pp. 54-76. London, Blackwell Science.
Hall, J. P. W.
2002a. A phylogenetic revision of Calydna and relatives (Lepidoptera: Riodinidae). Insect Systematics and Evolution, 33(2): 185-237.
2002b. 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.
Hall, J. P. W., and D. J. Harvey
2001. A 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.
Hall, J. P. W., and K. R. Willmott
1995a. Two new species of Mesene from western Ecuador (Lepidoptera: Riodinidae). Trop. Lepid., 6(2): 110-112.
1995b. Five new species and a new genus of riodinid from the cloud forests of eastern Ecuador (Lepidoptera: Riodinidae). Trop. Lepid., 6(2): 131-135.
1995c. Notes on the genus Argyrogrammana, with descriptions of five new species (Lepidoptera: Riodinidae). Trop. Lepid., 6(2): 136-143.
1996a. The genus Theope: four new species and a new subspecies (Lepidoptera: Riodinidae). Trop. Lepid., 7(1): 63-67.
1996b. Notes on the genus Argyrogrammana Part 2, with the description of a new species. Trop. Lepid., 7(1): 71-80.
1998. Nine new species and one new subspecies of Euselasia from Ecuador (Lepidoptera: Riodinidae).Trop. Lepid., 9(2)(suppl. 1): 27-35.
Harvey, D. J., and J. P. W. Hall
2002. Phylogenetic revision of the Charis cleonus complex (Lepidoptera: Riodinidae). Systematic Entomology, 27(3): 265-300.
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1993. Biodiversity and Biosystematic Priorities: Microorganisms and Invertebrates. Oxon, CAB International. 120 pp.
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1991. Faunal regions and the diversity of Lepidoptera.Trop. Lepid., 2(suppl. 1): 1-85.
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1869. Remarks on and descriptions of new species of butterflies collected by Mr. Buckley in Ecuador. London, John Van Voorst. 1: ii + 16 pp, 2: 17-32, 3: 33-48.
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1870b. Descriptions of twenty-two new species of Equatorial Lepidoptera. Trans. Ent. Soc. Lond., 1870(2): 153-163.
1877. Equatorial Lepidoptera Collected by Mr. Buckley. London, John Van Voorst. 5: 81-96.
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1992. Genera and species of the Neotropical "elfin"-like hairstreak butterflies (Lepidoptera: Lycaenidae: Theclinae). Rep. Mus. Nat. Hist., Univ. Wisconsin, 22(1): 1-135; (2): 136-279.
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1990. Conserving the World's Biological Diversity. Gland, IUCN. viii + 265 pp.
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1966. Latitudinal gradients in species diversity: a review of the concepts. Amer. Nat., 100: 33-46.
Pyrcz, T., K. R. Willmott, and J. P. W. Hall
1999. 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.
Robbins, R. K.
1993. Comparison of butterfly diversity in the Neotropical and Oriental regions. J. Lep. Soc., 46(4): 298-300.
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1997. Butterfly diversity and a preliminary comparison with bird and mammal diversity. In: Reaka-Kudla, M. L., D. E. Wilson, & E. O. Wilson (Eds.), Biodiversity II. Understanding and Protecting our Biological Resources, pp. 69-82. Washington, Joseph Henry Press.
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1992. Latitudinal gradients in species diversity: the search for the primary cause. Oikos, 65: 514-527.
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1995. Species Diversity in Space and Time. New York, Cambridge University Press.
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1994. Insect Conservation Biology. London, Chapman & Hall. xiii + 358 pp.
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1994. Techniques and guidelines for monitoring neotropical butterflies. Conserv. Biol., 8(3): 800-809.
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1997. Measuring global biodiversity and its decline. In: Reaka-Kudla, M. L., D. E. Wilson, & E. O. Wilson (Eds.), Biodiversity II. Understanding and Protecting our Biological Resources, pp. 41-68. Washington, Joseph Henry Press.
Torres, R., J. P. W. Hall, K. R. Willmott, and K. Johnson
1996. A new genus of " elfin" butterflies from the northern high Andes (Lepidoptera: Lycaenidae).Trop. Lepid., 7(1): 81-86.
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1992. Global Biodiversity: Status of the Earth's Living Resources. London, Chapman & Hall. 585pp.
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1990. Insect diversity and cladistic constraints. Ann. Ent. Soc. Amer., 83(6): 1031-1047.
Willmott, K. R., and J. P. W. Hall
1995a. Two new species of satyrines from Ecuador (Nymphalidae: Satyrinae).Trop. Lepid., 6(2): 103-105.
Willmott, K. R., J. P. W. Hall, and G. Lamas
2001. Systematics of Hypanartia (Lepidoptera: Nymphalidae: Nymphalinae), with a test for speciation mechanisms in the Andes. Syst. Ent. 26(4): 369-399.




A large number of people have generously given us their time, advice, and valuable information in the course of our research, and many individuals and institutions have provided us with financial assistance.

Collections access

For giving us access to the collections in their care and for the loan of material for study and dissection, we thank Lee and Jackie Miller (AME, Sarasota), Jim Miller and Eric Quinter (AMNH, New York), Gerald Legg (BMB, Brighton), Philip Ackery and Jim Reynolds (BMNH, London), John Heppner (FSCA, Gainesville), Philip Perkins (MCZ, Cambridge), Jacques Pierre (MNHN, Paris), Miguel Monné (MNRJ, Rio de Janeiro), Gerardo Lamas (MUSM, Lima), Marcelo Duarte (MZSP, São Paulo), George McGavin (OUM, Oxford), Mike Feyers (RPM, Reading), Wolfgang Nässig (SMF, Frankfurt), Christoph Haüser (SMS, Stuttgart), Matthias Nuß (SMTD, Dresden), Olaf Mielke (UFP, Curitiba), Robert Robbins and Don Harvey (USNM, Washington), Wolfram Mey (ZMHU, Berlin) and Axel Hausman (ZSBS, Munich). We also thank Julie Harvey and Lorna Mitchell at the BMNH entomological library for their help in retrieving many obscure works and for allowing us to photocopy or photograph important pages of several rare books.


We are grateful to the following for providing information, and in some cases inviting us to visit and record data, from their private collections: Chris Jiggins, Andrew Neild, Mike Perceval, David Trembath (UK), Stéphane Attal, Pierre Boyer, Olivier Duviols (France), Gerrit ten Broek (Holland), Fabio Vitale, Maurizio Bollino (Italy), Tomasz Pycrz, Artur Jasinski (Poland), David Ahrenholz, Robert and George Busby, Philip DeVries, Tom Emmel, Richard Hesterberg, Andrei Sourakov, Jon Turner, James Williams (USA), Luis Constantino, Jean LeCrom, Julián Salazar, and the late Ernesto Schmidt-Mumm (Colombia). We especially thank Gerardo Lamas for his constant willingness to provide unpublished taxonomic and bibliographic information, providing help and advice on numerous questions and photographs of type specimens.

Field Research

In Ecuador, collecting and export permits were obtained through INEFAN, and subsequently the Ministerio del Ambiente, the Pontificia Universidad Católica and the Museo Ecuatoriano de Ciencias Naturales, Quito, with the particular help of Santiago Villamarín, Germania Estévez, María de los Angeles Simbaña, Patricia Galiano and Gabriela Montoya. Ismael and Raul Serafín Aldas Villafuerte provided assistance in the field.

Financial Assistance

Institutional support

1993-2000: KRW & JPWH: Tom Emmel and the University of Florida (Florida Agricultural Experiment Station supported publication costs), provided facilities and research assistantships, funded by the U.S. Fish and Wildlife Service and private donations.
: KRW: The Natural History Museum, London, UK; Leverhulme Trust (Project Grant F/00696/C
: KRW: McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville, FL, USA
: JPWH: National Museum of Natural History, Smithsonian Institution, Washington DC, USA; two postdoctoral Fellowships; and National Science Foundation (Biodiversity Surveys and Inventories Grant # 0103746)

Fieldwork and museum research

1993-1994: Agfa, Balfour-Browne Fund, Banbridge Academy, C. Bruce, Butler Charitable Trust, Christ’s College Cambridge University (KRW only), Catherine Cookson Foundation, Harry Crook Foundation, Douglas Heath Eves Charitable Trust, Equafor, David Exell, R. and M. Foreman Charitable Trust, Hickley Valtone Ltd., Lindeth Charitable Trust, Morton Charitable Trust, Peter Nathan Charitable Trust, Northern Bank, Phoenix Mountaineering, Poulton Fund Oxford University (JPWH only), Albert Reckitt Charitable Trust, Round Table Trust, Royal Entomological Society, Vera Trinder Ltd., Ian Willmott, Muriel Willmott, Worts Fund.
: Sigma Xi, The Scientific Research Society
: American Museum of Natural History, New York, Collections Studies Grant
: National Geographic Society (Research and Exploration Grant No. 5751-96)
: Leverhulme Trust (KRW only)
: National Museum of Natural History, Smithsonian Institution, Short-term Visitor’s Grant (KRW only)
: This material is based in part upon work supported by the National Science Foundation under Grants No. 0103746 and 063986 (JPWH, KRW). Any opinions, findings, and conclusions or recommendations expressed in this material are those of the author(s) and do not necessarily reflect the views of the National Science Foundation.




Any information published from this website should be cited as:

From the website only:
Willmott, K.R., and J.P.W. Hall
(XXX). Butterflies of Ecuador.
Where XXX is the relevant date and page.

Any images displayed on this website, should be published only with the written prior consent of K.R. Willmott or J.P.W. Hall. Images may be used for talks or in educational material that are supplied free of charge without written consent.