A collaborative research program on the phylogenetics, taxonomy and evolution of euptychiine butterflies
One of the richest butterfly radiations in the Neotropical lowlands is the nymphalid satyrine subtribe Euptychiina, which is also one of the three most poorly studied groups of 'true butterflies' (Papilionoidea). In fact, few other animal groups that are so taxonomically challenging are as speciose, large, conspicuous and commonly encountered by researchers, students and naturalists. Hundreds of thousands of euptychiine specimens in museum collections hold untapped data for studies in evolution, biogeography and conservation. However, we estimate that almost half of these specimens cannot be confidently identified.|
More than 400 species in 42 genera are currently recognized, with at least 20% of species still undescribed, a remarkable fraction among butterflies. Molecular phylogenies suggest that Euptychia itself may not be related to remaining "euptychiines", and the generic classification is chaotic, with 65% of genera currently invalid. Modern monographs treat only five euptychiine genera (15%), with references for remaining genera now a century old. Because of their drab wing patterns, difficulties in identifying the often highly cryptic species and their sedentary life-style, most euptychiine collections are in disarray, and the biology of most species remains unstudied. In the field, however, euptychiines are a common and diverse component of communities from the rainforest to the cerrado. Reliable guides to the taxonomy and identification of euptychiines should therefore unlock the group's great potential for broader studies in evolution, ecology, biogeography and conservation.
The USA's National Science Foundation recently funded a three-year project to revise the systematics of Euptychiina, under the project title: "ARTS: Phylogeny and systematic revision of the diverse and cryptic Euptychiina (Lepidoptera, Nymphalidae, Satyrinae)" (DEB-1256742). The project has 5 main goals:
(1) to combine morphological and molecular data to produce the first 'total evidence' phylogenetic hypothesis for the subtribe, and to use this to reliably define genera;
(2) to complete monographs for the majority of the largest genera not recently revised, tentatively including: Euptychia, Taygetis, Magneuptychia, Cissia, Paryphthimoides, Pareuptychia, Erichthodes, Euptychoides, Hermeuptychia. A secondary goal is to describe, or provide support and/or data for collaborating researchers to describe, the 21+ known undescribed species in other genera;
(3) to provide training for butterfly systematists at the University of Florida and at the University of Campinas, Brazil;
(4) to use project results to test hypotheses about Euptychiina evolution;
(5) to develop collection and web-based electronic resources to facilitate communication and to build a collaborative research program on the group.
The project involves collaboration among many researchers in North and South America and Europe, and we would be pleased to hear from anyone with an interest in research on euptychiines (contact Keith Willmott at firstname.lastname@example.org). We hope to provide resources such as scannned original descriptions for euptychiine names, specimen databases, images of type and other specimens, images of morphology, DNA sequence data, and other data to facilitate research on the group.
Current project activities1. Generic revisions or systematic studies are underway in a number of genera. The table below lists researchers that we currently know of who have an interest in euptychiines, and their current main projects:
|Researchers with an interest in Euptychiina|
|Eduardo Barbosa||Brazil||revision of Yphthimoides|
|Christián Brévignon||French Guiana||Guianan euptychiines|
|Andy Brower||USA||Euptychiina molecular phylogenetics|
|Marianne Espeland||USA||Coordinating total-evidence higher-level phylogeny, Euptychiina diversification|
|Steve Fratello||USA||Guianan euptychiines|
|André Freitas||Brazil||generic revisions, phylogenetics|
|Nick Grishin||USA||revision of Hermeuptychia|
|Blanca Huertas||United Kingdom||Revision of Splendeuptychia, Colombian euptychiines|
|Akito Kawahara||USA||Euptychiina molecular phylogenetics|
|Gerardo Lamas||Peru||Peruvian euptychiines, generic revisions|
|Jean-François LeCrom||Colombia||Colombian euptychiines|
|Mario Marin||Colombia||Euptychiina morphological phylogenetics, revision of Pareuptychia, Moneuptychia|
|Olaf Mielke||Brazil||Brazilian euptychiines|
|Jackie Miller||USA||Revision of Taygetis|
|Shinichi Nakahara||Japan||Guianan euptychiines, revisions of Euptychia, Magneuptychia|
|Andrew Neild||United Kingdom||Venezuelan euptychiines|
|Carlos Peña||Finland||Euptychiina phylogenetics|
|Tomasz Pyrcz||Poland||Andean and southern euptychiines, Forsterinaria|
|Noemy Seraphim||Brazil||revision of Hermeuptychia|
|Denise Tan||Singapore||Euptychiina behavior and speciation, systematics of Hermeuptychia|
|Stephanie Tyler||USA||revision of Chloreuptychia|
|Angel Viloria||Venezuela||Venezuelan euptychiines|
|Niklas Wahlberg||Finland||Euptychiina molecular phylogenetics|
|Keith Willmott||USA||Ecuadorian euptychiines, revisions of Hermeuptychia, Chloreuptychia|
|Thamara Zacca||Brazil||Revision of Cissia, Paryphthimoides, Magneuptychia|
2. Curation and databasing is ongoing in several priority collections, including the NHM (London, UK) and the FLMNH (Gainesville, Florida, USA), with other priorities including the UFP (Parana, Brazil), MUSM (Lima, Peru), USNM (Washington DC, USA) and AMNH (New York, NY, USA). We are focusing mainly on genera under revision, except for the NHM where photography of the numerous type specimens is an important foundation for work in other collections.
3. Field work is planned for Ecuador, Peru, French Guiana and Brazil this summer, with a focus on collecting material for the higher-level phylogenetic study.
4. Work in the molecular lab has involved testing of various approaches to allow us to begin a phylogenomic study of c. 100 species this Fall. DNA barcoding of several genera with particularly diverse cryptic species complexes, such as Hermeuptychia, is also continuing.
Review of Euptychiina Systematics and BiologyThe Satyrinae is one of the most diverse nymphalid subfamilies, with 2600 species occurring on all continents except Antarctica (Ackery et al., 1999), and the Euptychiina is an almost entirely Neotropical subtribe of the most diverse tribe, the global Satyrini (Peña & Wahlberg, 2008). Most euptychiines are small with dark brown wings marked by a few simple ocelli, although several genera have white or brilliant blue wings. The majority of euptychiines are lowland species, except for the largely Andean Forsterinaria (Peña & Lamas, 2005). Community diversity peaks in the W. Amazon, where 100 species can coexist (Lamas et al., 1991; Brown, 1996), while the Atlantic region of Brazil also has a diverse, endemic fauna. A single Asian genus, Palaeonympha, is also apparently a euptychiine (Miller, 1968; Peña et al., 2006, 2010).
Moderately detailed descriptions of early stages have been published for only 20 species (5% of the subtribe) (Müller, 1886; Singer et al., 1983; Young, 1984; Murray, 2001, 2003; Freitas, 2003, 2004a,b, 2007; Freitas & Peña, 2006; Kaminski & Freitas, 2008), and hostplant records are similarly scarce. Like most satyrines, larvae of most species feed on Gramineae, grasses and bamboo (Beccaloni et al., 2008), with some records on Cyperaceae and Marantaceae, except for Euptychia which feed on mosses and lycopsids (Singer et al., 1971; Singer & Mallet, 1986; DeVries, 1986; Brévignon, 2008). Development times are relatively long among tropical butterflies, up to three months or more (Freitas, unpub. data from > 50 species and 27 genera). Eggs are generally isolated and lack chorionic sculpturing. Larvae have bifid "tails" and may bear head horns that vary in size. Pupae are generally short, squat and smooth. Adults are rarely encountered at nectar, but often feed on decaying fruits or tree sap, and on carrion and animal feces. Some species are crepuscular (e.g. Taygetis, Murray, 2003), and males of forest species patrol territories in the understory, often on ridgetops, in the late afternoon (Peixoto & Benson, 2009; Willmott, pers. obs.). Males often bear tufts of hair-like scales or other androconia on the wings that presumably function in courtship.
The current classification includes more than 400 species (Lamas, 2004; Freitas, Willmott, unpub.; Huertas et al., 2009; Peña et al., 2010; Huertas, 2011; Pulido et al., 2011; Brévignon & Benmesbah, 2011; Matos et al., 2012; Freitas et al., 2011, 2012; Zacca et al., 2013) in 42 genera, with from 1 (10 genera) to 47 species. Remarkably, 80 (20%) of the known species are undescribed, a statistic exceeded among the true butterflies only by the Lycaenidae (34%) and Pronophilina (27%) (Lamas, 2004). Most known undescribed species are distinctive but poorly represented in historical collections, and many cryptic species (e.g. Willmott & Hall, 1995) surely remain to be discovered. For example, Miller (1974) described 11 of the 28 species of Cyllopsis, and Peña and Lamas (2005) described 12 of the 23 species in Forsterinaria. Most inventories contain undetermined species (e.g. Brévignon, 2005, 2007, 2008; Brévignon & Benmesbah, 2011), with recent fieldwork in Brazil revealing 8 new species in Yphthimoides, Splendeuptychia and Moneuptychia (Freitas 2004b, 2007). Many euptychiines are externally very similar, and some are apparently highly variable; with only five euptychiine genera (15%) the subject of modern revisions (Miller, 1972, 1974, 1976, 1978; Peña & Lamas, 2005), references for generic and species identification are urgently needed.
Miller (1968) defined the subtribe by external morphology, and molecular (Murray & Prowell, 2005; Peña et al., 2006) and morphological (Marin, 2010) studies have largely confirmed the current classification (Lamas, 2004), except for excluding Oressinoma and including Amphidecta and the Asian Palaeonympha. The position of Euptychia itself, however, is still unresolved (Murray & Prowell, 2005); the most comprehensive molecular phylogeny of Euptychiina to date (Peña et al., 2010), based on 4447 bp from 5 genes, for 108 euptychiine species and 18 outgroup taxa, did not recover a monophyletic Euptychiina. However, Marin (2010) did recover the subtribe as monophyletic and found a number of clades in common with Peña et al. (2010). A major goal of our research, therefore, will be to integrate and expand these data matrices to generate a robust higher-level phylogeny for the group.
If the relationships of Euptychiina within the Satyrini are still unclear, the generic classification is arguably the most chaotic for any butterfly group of similar diversity. Weymer (1910-11) classified the majority of Neotropical species (270 spp.) as "Euptychia", presumably because of their wing pattern homogeneity, and D'Abrera (1988) continued the trend, ignoring a large number of generic names that were introduced by Forster (1964). Forster's diagnoses were based partly on wing pattern and male genitalia, but also on overall appearance, and his characters are often unreliable. Lamas (2004) moved almost all "Euptychia" (sensu D'Abrera) into Forster's genera, which constitute 26 of the currently accepted 39 Neotropical genera. Although the Lamas classification is a critical framework for future work, recent cladistic analyses reveal much remaining paraphyly and polyphyly among genera (Murray & Prowell, 2005; Peña et al., 2006). Peña et al. (2010) tested the monophyly of 23 Euptychiina genera and found that only 8 (35%) are monophyletic. Often, supposed congeners occurred within different species groups, such as Splendeuptychia, whose species appear in the Splendeuptychia, Pareuptychia and Hermeuptychia clades. Marin (2010) found similar results using morphological characters; only 10 of 22 Euptychiina genera tested (45%) proved monophyletic. Recent taxonomic revisions based on a molecular phylogeny of the nine Taygetis-clade genera resulted in the synonymy of three genera, rearrangement of three genera and revelation of two undescribed genera (Matos et al., 2012). Based on an expanded higher-level phylogeny, we hope to revise generic limits and describe useful morphological characters for generic identification.
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