Purpose of website
Background and aims
Why butterflies?
Why Ecuador?
State of knowledge
Acknowledgements
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Although most of the information presented is our own unpublished data, we hope that eventually this site will become a general source of information on Ecuadorian butterfly research and conservation.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.
Any comments or suggestions by e-mail would be welcome (kwillmott@flmnh.ufl.edu).
Background
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 about 73 person-months in the field in Ecuador on this project.
To date our taxonomic work has been concentrated on the Riodinidae and Nymphalidae, resulting in 133 published papers of relevance to Ecuador, which include descriptions of 143 species and 23 genera (see state of knowledge). The groups still in need of most 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; Pyrcz et al., 2011; Hall, 2019, 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 now 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) Continue to build 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, undergraduate students and other professionals 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 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, 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 was traditionally 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; Hall et al., 2005; Hall, 2005; Nakahara et al., 2015; Hall, 2019).
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 3000 species in the families Papilionidae, Pieridae, Nymphalidae, Riodinidae and Lycaenidae, about 50-55% of all Neotropical species in these groups (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 3000 species of butterflies in the families Papilionidae, Pieridae, Nymphalidae, Riodinidae and Lycaenidae 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.
When we began working in 1993 in Ecuador, a remarkable 616 species were undescribed, about 20% of the total fauna. Since then, 347 of these have been described, including 143 by ourselves and co-authors. The table below shows a breakdown of these figures by family.
Summary of Ecuadorian species described since 1993
| Papilionidae | |||||
| Pieridae | |||||
| Nymphalidae | |||||
| Riodinidae | |||||
| Lycaenidae | |||||
| TOTAL |
The map below shows a preliminary analysis of species richness for butterflies (excluding skippers) throughout Ecuador, in half-degree grid squares.
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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.
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1998. Nine new species and one new subspecies of Euselasia from Ecuador (Lepidoptera: Riodinidae).Trop. Lepid., 9(2)(suppl. 1): 27-35.
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2002. Phylogenetic revision of the Charis cleonus complex (Lepidoptera: Riodinidae). Systematic Entomology, 27(3): 265-300.
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1995a. Two new species of satyrines from Ecuador (Nymphalidae: Satyrinae).Trop. Lepid., 6(2): 103-105.
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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; we thank all of them. We acknowledge, in particular, the following:
People
Thomas Emmel, Gerardo Lamas, Thomas Pyrcz, Shinichi Nakahara, Andrew Neild, Robert Busby, Mark Simon, Dave Ahrenholz, Pierre Boyer, Jean-Claude Petit, Fabio Vitale, Maurizio Bollino, Jamie Radford, Marianne Elias, Sofía Nogales, Santiago Villamarin, Fernanda Checa, Patricio Salazar, Sebastián Mena, Sebastián Padrón, Geoff Gallice, Julia and Jamie Robinson Willmott, Ismael Aldas, Raul Aldaz, plus all the curators and collectors who allowed us to examine the specimens in their collections, and the numerous students and volunteers who helped with databasing specimens.
Institutions
Instituto Nacional de Biodiversidad (formerly Museo Ecuatoriano de Ciencias Naturales), Pontificia Universidad Católica del Ecuador, Ministerio del Ambiente, Parque Nacional Yasuní, Fundación Arcoiris, Fundación Jocotoco, Mashpi Lodge, Napo Wildlife Center, Yuturi and Yarina Lodge, McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida.
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, (MECN), then the Instituto Nacional de Biodiversidad (INABIO), with the particular help of Santiago Villamarín, Germania Estévez, María de los Angeles Simbaña, Patricia Galiano, Gabriela Montoya and Sofia Nogales. Ismael and Raul Serafín Aldas Villafuerte provided assistance in the field.
Funding
National Science Foundation (DEB-0103746, DEB-063986, DEB-1256742, DEB-1342705), Leverhulme Trust (Project Grant F/00696/C), National Geographic Society, Darwin Initiative, McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, the University of Florida (Florida Agricultural Experiment Station).
Any information published from this website should be cited as:
From the website only: 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.
Willmott, K.R., and J.P.W. Hall (XXX). Butterflies of Ecuador. www.butterfliesofecuador.com/XXX
Where XXX is the relevant date and page.